A Monograph of the Frogs of the Family Microhylidae

A MONOGRAPH OF THE

FROGS O F THE FAMILY MICROHYLIDAE by

H. W. PARKER, B.A. ASSISTANT KEEPER I N THE DEPARTMENT OF ZOOLOGY

LONDON : PRINTED BY ORDER O F THE TRUSTEES OF THE BRITISH MUSEUM SOLD A T T H E BRITISH MUSEUM (NATURAL HISTORY), CROMWELL ROAD, S.W.7 AND BY B. QUARITCH, LTD., DULAU & CO. LTD., AND OXFORD UNIVERSITY PRESS

X A D E A N D PRINTED I N GREAT BRITAIN B Y JARROLD A N D S O N S LTD. NOBWlCH

PREFACE INCE the appearance of the second edition of the Catalogue of the Batrachia Salientia i n the collection of the British Museum (1882), a number of emended d , each in turn schemes of classification of the Order have been put f o r ~ ~ a rand has had to be still further emended or entirely abandoned. As a rule these proposed schemes have been the outcome of the examination of a relatively limited number of skeletons, and, as further material has become available for examination, onc or more of the characters regarded as crucial have proved untrust~vorthy. The present volume is intended t o be the first of a series of monographs, similar in scope, though not necessarily in arrangement, in which an attempt will be made to base the classification on the osteological examination of as many species as possible. The collections of the British Museum, although very rich, are scarcely adequate for such a task, and the work would be quite impossible without the generous assistance of the many institutions which have lent or given specimens, notably : Zoological hluseum, Amsterdam ; Natural History Museum, Basel ; Zoological Museum, Berlin ; Royal Belgian xatural History Museum, Brussels ; Hungarian National Museum, Budapest ; Indian Museum, Calcutta ; Field Museum of Natural History, Chicago ; State Museurns for Zoology and Ethnology, Dresden ; Museum of the Senckenberg Natural History Society, Frankfurt ; Museum of Natural History, Genoa ; Museum of Comparative Zoology, Harvard ; McGregor Museum, Kimberley ; Sarawak Museum, Kuching ; University Museum of Zoology, Michigan ; Zoological Collection of the Bavarian States, Munich ; fifuseum of Natural History, Paris ; Natural History Museum, Vienna ; United States National Museum, Washington. In addition to the authorities of these institutions the author desires to express his gratitude to the following ladies and gentlemen who havc either made investigations on his behalf or have shown him personal kindness during the course of the preparation of this monograph : Dr. E. Ahl, M. I?. Angel, Dr. T. Barbour, Dr. A. M. Boring, Mr. L. D. Brongersma, Prof. C. E. Burt, Prof. E. R. Dunn, the late Prof. Baron FejCrvBry, Dr. G. Hecht, Mr. A. Loveridge, Dr. R. Mell, Dr. R. Mertens, Prof. L. Miiller, Dr. G. K. Noble, Mr. J. H. Power, Dr. J. lloux, Prof. A. G. Ruthven, Dr. Malcolm Smith, Prof. D. Vinciguerra, Dr. 0.Wettstein, and M. G. de Witte. With the exception of the compilation in Das I'ierreich, no complete account of this family has appeared since the publication of the British hluseum Catalogue, and during the interval there have been great additions to our knowledge of this somewhat obscure group of frogs. I n 1882 only 43 species were known, of which 34 were represented in the British Museum (79 per cent) ; now 184 species and subspecies are recognised, of which 125 are to be found in London (68 pcr cent), 27 have been examined elsewhere and only 32 are known to the author solely frorn published descriptions. The decline in the proportion of species represented in the British hluseum is due to the greatly increased activities of workers in other countries ; for example the average number of authors writing on the systematics of the Amphibia Salientia in a five-year period prior t o 1882 was 11, whereas during a similar period recently the average number was more than 30. W. T. CALMAN BRITISH MUSEUM(NATURAL HISTORY) Keeper of Zoology March 7th, 1934

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MONOGRAPH OF THE FROGS O F THE FAMILY MICROHYLIDAE INTRODUCTION HISTORY O F THE FAMILY The frogs of this family, being for the most part insignificant in size and seldom to be found in any great congregations of individuals, do not appear t o have attracted the attention of the earlier European voyagers, and the first record of any one of them in scientific writings is that of Seba (1735), who figures a " Rnna rubeta Africans." This figure was the basis of the first species to be named in accordance with modern practice (Rana gibbosa), and was the only one known to Linnaeus. In the first century after the publication of the tenth edition of the " Systema Naturae " remarkably few additional species were discovered, Giinther in 1858 recognising but fourteen. Even in 1882 Boulenger only knew thirty-nine forms, but the succeeding fifty years have increased the number of recognisable species and races t o 184. The early systematists, after it became a common practice t o subdivide the Amphibia Salientia into families, grouped the few genera known to them wit,h others which to-day are recognised as being of very different affinities. Not until 1843 were any of them separated off from the rest, and then Fitzinger proposed the families Hylaedactyli and Gastrophrynae which contained the majority of the forms known t o him. The great diversity of form in the species which were being discovered a t that period was reflected in a multiplicity of families ; Giinther (1858) referred the fourteen species known to him to six different families, and, in the interval before the publication of the second edition of the British Museum Catalogue, four others were proposed t o receive different members of the group. Boulenger (1882), however, recognised only two families, the toothed Dyscophidae and the toothless Engystomatidae, t o which he subsequently (1890) added the Genyophrynidae, characterised by the presence of " teeth " on the lower jaw. It soon became apparent, however, that the presence or absence of teeth mas of trivial importance in indicating relationships, so that Gadow in 1901 suggested the union of these three as a single family (the Engystomatidae), the original groups each being given the rank of a subfamily ; this arrangement was followed by many systematists until 1926. The only other proposal of major importance was that of Mkhely (1901), who suggested that the Engjrstomatidae (sensu Boulenger) should be subdivided into the Symphygnathinae and Eleutherognathinae, according t o whether the maxillary bones made contact anteriorly or not ; the suggestion was not generally adopted. Although the family was recognised for practical purposes, it had long been evident that, like several other Salientian families, i t could not be satisfactorily defined. Very few constructive suggestions were made, however, until Noble in 1922 put forward an emended scheme of classification of the Salientia, based largely on an earlier piece of work by Nicholls (1916). I n this scheme teeth, which had previously been considered of primary importance, were practically abandoned as a character of value in defining families, and stress was laid on the four different types of vertebral column which are found in the Salientia, viz., Opisthocoelous, Anomocoelous, Procoelous and Diplasiocoelous. It was claimed that the " Engystomatidae," now renamed " Brevicipitidae," were an entirely diplasiocoelous family, and that certain procoelous South American genera1 hitherto included in the family, or in the Ranidae, were really more closely allied to arciferal forms and should be placed, together with some Braehyeephalus, Rhinoderma, Atelopus and Oreophrynella. 1

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2

MOISOGRAPH O F THE MICROHTLIDAE

others, in a separate group, the Brachycephalidae. Additional evidence in support of this disposition was derived from the myology of the thigh, and, later (Noble, 1925, 1927), i t was shown that, with the exception of three South African genera, those " Brevicipitidae " which pass through a larval stage have a type of tadpole unknown elsewhere in the Amphibia. The South African forms were admitted t o be more closely allied t o Ranid genera than t o any of the " Brevicipitidae," but their inclusion in the latter family, thus rendering it obviously diphyletic, was justified on palaeontological precedent (Abel, 1909) ; i t was believed that the " Brevicipitidae " were themselves derived from a Ranid stem. Latsky (1930) emphasised that Noble's " Brevicipitidae " were not a natural family, but did not consider the removal of the discordant genera t o their true position with the Ranidae ; instead, the union of the two families was advocated. The most recent consideration of the family as a whole is again due t o Noble (1931). His conception is essentially the same as before, and no change is made in the limits of the family ; but i t is recognised that within it there are many clearly marked groups of closely related genera, and fourteen such groups are given subfamily rank. The group is admitted not to be a natural one, for two of the subfamilies are believed to " have arisen independently from African Ranids." The remainder are characterised as being firmisternal, without ribs a t any stage of their life-history, diplasiocoelous, with more or less dilated sacral diapophyses, and either passing the larval stage within the egg capsule or hatching out t o form a very distinctive tadpole ; they are considered to be derived from a primitive stock now represented by the Ranidae. The various subdivisions are :(1) Dyscophinae. Large entire prevomers almost surrounding the choanae ;omosternum, clavicles, procoracoids and sternum present. Genera :-Dyscophus, Calluella, Pseudohemisus, Scaphiophryne and ? Callulina. (2) Rhombophryninae. Prevomers divided, with a posterior portion overlying or replacing the palatine bone. Genera :-Mantipus, Platyhyla, Platypelis, Plethodontohyla, Rhombophryne, Cophyla, Anodonthyla and Stumpfia. (3) Sphenophryninae. Characters of the Dyscophinae but with the omosternum absent. Genera :Liophryne, Xphenophryne, Oxydactyh, Oreophryne. (4) Cacopinae. Prevomer as in the Dyscophinae ; omosternum, clavicles and procoracoids usually absent. Genera :--Colpog1ossus,~ ~ l ~ ~ h o ~ l o sCalliglutus, sus, Cacopus, Phrynomantis, Copiula, Cophixalzcs, Choerophryne, Aphantophryne and Genyophryne. (5) Symphygnathinae. Similar to the Cacopinae, but with the maxillaries meeting in front of the premaxillaries. Genera :-Callulops, iiantophryne, Xenobatrachus, Xenorhina and Asterophrys. (6) Kalophryninae. Prevomers small, confined t o the anterior and mesial margins of the choanae; pro~oracoids and clavicles present, but the omosternum reduced or absent. Genera :Kalophrynus, Hypopachus, Otophryne, Chiasmocleis, Wectodactylus and Stereocyclops. (7) Microhylinae. Prevomers as in the Kalophryninae ; clavicle and, usually, the procoracoid absent. Genera :-Microhyla, Phrynella, Gastrophryne, Ctenophrylze and Dasypops. (8) Phrynomerinae. Procoracoid and clavicle absent ; intercalary phalanges present. A single genus :-Phrynomerus. (9) Kaloulinae. Clavicles absent; procoracoids and a rudimentary omosternum persistent. Genera :-Kaloula, Ramanella. (10) Melanobatrachinae. Prevomers small ; a calcified omosternum, clavicle and procoracoid present ; no palatal ridges, A single genus :-Melanobatrachus. (11) Brevicipitinae. Roof of the mouth very glandular; skeleton retains many of the primitive features of the Dyscophinae. Genera :-Spelaeophryne, Breviceps, ?Didynamipue and ? Callulina. (12) Hoplophryninae. Prevomers greatly reduced ; inner finger greatly reduced. Genera :Parhoplophryne and Hoplophryne. (13) Hemisinae. Snout very pointed ; procoracoid and clavicle present ; scapula articulating with the skull ; tadpole of the Ranid type ; palate not glandular. A single genus :-Hemisus. (14) Cacosterninae. No clavicle; procoracoid present or rudimentary; omosternum bony or cartilaginous ; no palatal ridges ; tadpole of the Ranid type. Genera :-Cacosternum and Anhydrophryne. Caeosternum, Anhydrophryne and Hemisus.

INTRODUCTION

STRUCTURE AND CLASSIFICATION Unfortunately this scheme has not been found wholly satisfactory. In the first place, the Hemisinae and Cacosterninae can no longer be admitted as members of the family (de Villiers, 1933), for whilst it seems highly probable that they are of Ranid descent, it is even more probable (see below) that the other genera are not, as was originally supposed, descended from the same group, but represent a much older stock. Again, the Phrynomerinae, though closely allied t o the rest of the family, differ from it in exactly the same u7ay as do the Hylidae from the Ceratophryidae (Leptodactylidae, Cystignathidae or Bufonidae auctorum), and the Rhacophoridae from the Ranidae ; accordingly, as the author has already suggested (Parker, 1931), they are accorded family status. Other genera which are excluded from the family are Pseudohemisus and Scaphiophryne. The exact status of these two is uncertain, but they seem to be in many ways related to the Ranidae. Thus they both have an entire ethmoid, a character unknown in all the genera included in the present family, and Angel (1931) has shown that the tadpole is of the Ranid type, with labial teeth, horny mandibles and a sinistral spiraculum. On the other hand, they retain a vestige of the anterior palatal fold, so common in the present family, have dilated sacral diapophyses and are reported (Trewavas, in press) to have a hyo-laryngeal apparatus very similar to that of Calluella and Microhyla. They differ from all other Madagascar genera of the present family, except Dyscophus, in the possession of a diplasiocoelous vertebral column and an undivided prevomer, but are utterly unlike Dyscophus, or any of its allies, in the absence of maxillary and vomerine teeth and in the reduction of the post-choanal portion of the prevomer. Didynamipus has already been shown , (rafker, i*$3"l'j t o 'be -pdwc~'rlrya A&~vaCL:smYA $ i e ' ~ f r ? x y ? ~ , . It is believed that, after these eliminations, the residue is, with one or two doubtful exceptions, a truly monophyletic assemblage, but one which in different geographical regions has developed along different lines and consequently can be subdivided into other, smaller, natural groups. Some of these groups are equivalent to the subfamilies already proposed, but in other instances it seems probable that subdivision has been carried too far ; members of the same genus sometimes appear in different subfamilies. The inter-relationships of the various genera and the position of the family in the Amphibia Salientia can best be appreciated after a brief general survey of its adult - and- larval morphology.

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A. STRUCTURE O F THE ADULT The following organs or structures show modifications which have been shown, or are believed, t o be of importance in defining groups :(I) Premaxillae. Of the same general shape as in other families and not showing. any appreciable diversity of structure ; they may, or may not, bear teeth. (2) Maxillae. In general similar t o those of other families, but in two genera, Asterophrys and Xenobatrachus the p a ~ fan'alis s is greatly prolonged and meets its fellow in front of the premaxillae (fig. 26 ;p. 55). A condition intermediate between this and the normal is shown by Genyophryne and Metoposti~a,where although the maxillae overlap the premaxillae to some extent they do not make contact. A " syrnphygnathine " condition of the maxillae is usually accompanied by a similar condition of the dentaries (q.v.). Teeth may be present or absent, but when present are usually well developed, and of the normal Salientian pattern ; in Xphenophryne cornuta alone have they been discovered in a vestigial condition. (3) Prevomer. This shows a greater amount of variation than any other single bone. Its most primitive form, judged by the criteria of being that from which all others may be most easily derived, of occurring in the Caudata, and of being found in those genera which retain the greatest number of other primitive characters, is that of a large, almost horseshoe-shaped plate which nearly surrounds the choana and overlies the palatine region (fig. 1 ;p. 22). Such a prevomer has not been described in any Ranid and is only rarely found elsewhere. It occurs in the primitive Discoglossidae

MONOGRAPH OF THE MICROHYLIDAE (Discoglossus pictus) and the Hylidae (e.g. Hyla venulosa and H. maxinza). The post-choanal portion may, or may not, be fused with the palatine, and, though often edentulous, sometimes bears true teeth, or sometimes odontoids ; these structures are never found on the pre-choanal portion. The post-choanal portion, which, even if edentulous, is often raised into a strong, crenulate ridge or a single prominent knob, is frequently greatly reduced or may be entirely absent, and this reduction appears t o have taken place more than once within the family. In the Malagasy genera, for instance, a series can be traced (Noble and Parker, 1926 ; Parker, 1931) from an entire primitive type of prevomer, such as is found in Dyscophus (fig. 1; p. 22), through a series of intermediate stages to Anodonthyla, where the post-choanal portion is completely lost. Here the first stage is the severance of the pre- and post-choanal parts, the latter being by far the larger and still bearing a long row of teeth (e.g. Plwtyhyla, fig. 9 : p. 36). The dentigerous portion is then reduced (Platypelis, fig. 12; p. 38), fuses with its fellow (Cophyla, fig. 1 4 ; p. 41) and finally disappears. I n the IndoPapuan region, however, the post-choanal part undergoes reduction before it becomes detached from the pre-choanal portion (e.g. Uperodon, figs. 31 and 32 ; p. 74). The anterior portion, which borders the choanae antero-mesially, is never completely lost. (4) Palatine. This, like the post-choanal portion of the prevomer, from which it frequently cannot be distinguished, is often reduced or absent. Only rarely does i t persist when the post-choanal prevomer is absent (e.g. certain species of &ilicrohyla). (5) Ethmoid. The normal, adult, condition of this bone corresponds with its early post-larval condition in other families. I n the majority of frogs the ethmoid first appears as a vertical element in the anterior part of the inner wall of each orbit, and, later, these two elements meet and fuse above and below t o form a complete girdle around the anterior part of the cranium. I n the present family, however, the latter condition is never attained, the ethmoid retaining its primitive paired condition even in the adult ; rarely (e.g. Kalophrynus, Breviceps, Probreuiceps, Spelaeophryne and Callulina) it is completely absent. Representatives of fifty-eight genera (often many species), belonging to every other Salientian family, have been examined, and the normal condition of the ethmoid is single ; exceptions were found in the following :Rhacophorus (2 spp.), Cornufer (2 spp.), Hyperolius (2 spp.), Crinia (1sp.), Pseudophryne (1 sp.), nlicrobatrachella (1 sp.). Other exceptions will doubtless be found as more species are examined, but it is significant that all except one of those forms previously included in the present family, but rejected on other grounds (e.g. tadpole and vertebral column), have an undivided ethmoid (Brachyceplzalus, Rhinoderma, Atelopus, Oreophrynella, Cacosternunt, Anhydrophryne, Pseudohemisus, Scaphiophryne and Ilenzisus). A further peculiarity of the ethmoid is a marked tendency t o develop laterally along the posterior border of the nasal capsule, in those forms in which the post-choanal prevomers and the palatine are absent. This tendency has already been pointed out in the genus Microhyla (Parker, 1928), and is even more marked in Melnrrobatrachus and Hoplophryne. I n the latter two genera the ethmoid is also completely fused with the parasphenoid. (6) Ear. The majority of genera have the middle ear fully developed, though the skin over the true tympanum may not be thinned so as t o produce the condition usually spoken of as " tympanum visible." In fMelanobatrachus, Hoplophryne (and presumably Parhoplophryne also) the middle ear is incomplete ; there is no tympanum, cavum tympani, annulus ty?npanicus, columellu auris or Eustachian tube. The operculum, however, is normal and retains its muscular connection with the scapula. This condition of the ear appears to be of very common occurrence in the Salientia, for i t has also been noticed in Liopelma, Bombina, Pelobates, Atelopus, Oreophrynella, Dendropl~ryniscus, Didynamipus, the African species of Nectophryne, Sooglossus, iiesomantis, Hemisus, Pseudophryne, Brachycephalus and in early post-metamorphic specimens of Hyperolius horstocki (du Toit and de Villiers, 1932). That all these genera are closely allied is scarcely credible, but certain groups of them obviously are, and so the condition has considerable taxonomic importance ; Dendrophryniscus, Oreophrynella, Atelopus and Bruchycephalus have already been recognised as a closely allied series in a triphyletic assemblage spoken of as the " Brachycephalidae." That Sometimes a rudimentary Eustachian tube is present.

INTRODUCTION Didynamipus, and its probable ancestor Nectophryne (N. afra, N. batesi and N. parvipalmata), have the same incomplete ear as the South American genera, which they so closely resemble in other characters, seems to indicate that the resemblances are not due, as was previously suggested (Parker, 1931), to convergence. The fact that, as reported by du Toit and de Villiers (1932), the developing ear of young specimens of Hyperolius horstocki passes through an almost exactly similar stage suggests strongly that the condition may be a primitive, rather than a degenerate one. (7) Dentary. As in most families this bone is not very extensive and never bears true teeth ; in Genyophryne, however, its upper edge is serrated. Normally the dentaries are separated by the full extent of the mento-meckelian bones, but in the subfamily Asterophryinae they overlap the anterior surface of these bones to a greater or less extent, and in Xenobatrachus and Asterophrys form a median suture. (8) Mento-meckelian Bones. Very many genera have a cartilaginous process extending backwards from these bones and running parallel with the lower jaw ; a similar structure has been described in Phrynomerus (de Villiers, 1930). (9) Hyoid and Larynx. Trewavas has (in press) given a full account of the known variations of these structures in the Salientia, and taxonomic importance is believed to attach t o some of them. (10) Shoulder Girdle. An enormous amount of variation is shown by some of the elements of this structure, particularly the clavicle, procoracoid a n d omosternum, and it seems probable that these changes are correlated with loss of function. In no arciferal frog, where the clavicle and procoracoid form the only rigid strut keeping the glenoids apart, is any reduction of clavicle or procoracoid known. But in the firmisternia, on the other hand, where the function of strutting the glenoids apart can be better undertaken by the much larger, stronger coracoids, the clavicles and procoracoids have been reduced time and again ; in the present family the clavicle appears to have been lost on a t least five different occasions. On three of these the reduction has taken place from the distal end of the bone, with the result that, in the intermediate stages, a short clavicle is found only on the mesial half of the procoracoid. Platyhyla and Platypelis in Madagascar (figs. 10 and 13; pp. 37 and 38), Chiasmocleis and Elachistocleis in South America (figs. 49 and 50 ; pp. 117 and 120) and Oreophryne in the Papuan region (fig. 56 ; p. 160) are representatives of such stages. That they have attained the same condition independently seems certain, for in every other character they are more closely akin to genera with a fully developed clavicle than they are to one another. Thus Platyhyla and Platypelis have much in common with Mantipus, the neotropical genera are very similar to Hypopachus, and Oreophryne can only be distinguished from Sphenophryne by the condition of its shoulder-girdle. In other genera, however, the reduction appears to have occurred in the reverse direction. In Calluella the clavicles are weak mesially, and in its ally Colpoglossus are reduced to lateral vestiges (figs. 4 and 6 ; pp. 27 and 31), and a similar state of affairs is to be found in the entirely unrelated Parhoplophryne and Hoplophryne (fig. 67 ; p. 199). Reduction of the clavicle and procoracoid is almost always accompanied by reduction of the omosternum, though this is sometimes absent even in forms with an otherwise complete shoulder girdle ; the omosternum has a bony style in Dyscophus, is calcified in Melanobatrachus, but otherwise is cartilaginous. The sternum never has a definite bony style, such as is common in the Ranidae ; in Aphantophryne it is said t o be entirely absent. Throughout the family the sternum is completely fused with the epicoracoid cartilages. Jones (1933) has recently shown that whenever the clavicle or procoracoid is well developed the rM. coracoradialisarises from the whole, or the greater part, of its length ; in the Ranidae and Rhacophoridae, on the other hand, the same muscle is either not attached to the procoracoid-clavicular bar at all or only arises from its extreme mesial end. Reduction of the omosternum, clavicle and procoracoid is not accompanied by any corresponding reduction of the pectoral musculature. The Indo-Malayan species previously referred to Nectophryne have the ear fully developed and so should be regarded as generically distinct. The name Pedostibes, Giinther (Type species, P. tuberculosus, Gthr.) is availablc for them.

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MONOGRAPH O F THE MICROHYLIDAE

(11) Vertebral Column. Hitherto this family has been considered to be uniformly diplasiocoelous, and the condition of the vertebral column has in recent years been considered diagnostic of higher groups than families. Unfortunately it is now apparent that sixteen out of the forty-three genera here considered are procoelous (six have not been examined), and consequently it becomes necessary to re-assess the value of the vertebral condition as a character for systematic purposes. The work of Nicholls (1916) and Noble (1922) makes it clear that in most families there is considerable constancy in the vertebral condition, but any attempt to subdivide the present family on this character alone produces some anomalous results. Thus, for example, the Papuan Genyophryne, Sphenophryne, Oreophryne and Cophixalus, being procoelous, would have to be grouped with the majority of the Madagascar genera, and the Madagascar Dyscophus urould be placed with the Papuan Asterophrys and its allies. Yet the affinity of Genyophryne with Asterophrys can scarcely be gainsaid. Nevertheless groups of genera which on other structural and geographical grounds appear to be allied have, in general, the same type of vertebral column, and consequently the character appears to be a useful guide to the relationships of genera within the family. (12) Sacral Diapophyses. These are always more or less dilated in a horizontal plane. It seems to have been tacitly assumed, but nowhere explicitly stated, that the cylindrical diapophysis is the primitive, and the dilated the secondary, condition. But the converse appears to be rather the case. In the first place, dilated diapophyses are much commoner than cylindrical ones throughout the Salientia, and are universally present in all the more primitive families. Secondly, it has been suggested (Parker, 1931) that the dilated diapophysis is mechanically suitable for crawling or swimming, and the cylindrical for jumping, frogs ; if the pro-Salientian stock was caudate, as seems highly probable (Noble, 1922), the primitive frogs are unlikely to have been saltatory. (13) Urostyle. This normally articulates with the sacrum by two condyles, but in Breviceps and Probreviceps the articulation is lost and the two bones are completely ankylosed. (14) Terminal Phalanges. In species with digital dilatations the distal phalanges are usually T- or, more rarely, Y-shaped ; even where the disc is greatly reduced or absent a T-shaped phalanx may persist. Normally the terminal articulates with the penultimate phalanx by a single condyle, but in Platyhyla a unique modification is found : the condyle is completely divided (fig. 11; p. 37). Another unique modification of the terminal phalanges is exhibited as a secondary sex character (q.v.) in Kaloula verrucosa. (15) Tongue. Two types of tongue are found in the family, the one an oval, thin one, with its posterior half free, the other subcircular, thick and not free behind ; the latter usually has a longitudinal median groove culminating in a more or less developed " pocket " posteriorly. To this latter structure considerable importance has been attached, but it seems quite probable that its presence, especially when very deep and with a plicate edge, is a post-mortem condition produced by a strong contraction of the hyoglossus muscle. It has been claimed that the adherent, grooved tongue has appeared haphazard on several occasions within the family. But it seems doubtful whether this is true, for its appearance seems to be correlated with other characters which suggest affinity. Thus, except in Genyophryne, Rhombophryne and ? Ctenophryne, it is not found in any procoelous forms, and among diplasiocoelous genera it is almost entirely confined to those which retain a large, entire prevomer. (16) Pharyngeal Folds. Across the palate in front of the pharynx two (rarely three) transverse dermal folds are usually t o be found ; the posterior is often denticulate, the anterior more rarely so. The anterior is also often reduced, and all conditions between two well-developed folds and only a single one (the posterior) can be traced in many genera. Folds of this nature are not known in any other family, and their presence is almost diagnostic of the group, for in only one subfamily are they constantly absent. Other, glandular, folds are sometimes present close behind, or between, the internal nares. (17) Thigh Muscles. Considerable importance in the diagnosis of families has been attached to the arrangement of the distal tendons of the thigh muscles (Noble, 1922),

INTRODUCTION

7

but Latsky reports (1930) that "there exists a greater divergence between the two investigated species of Breviceps than between Rana and most of the South African Brevicipitid genera." Nevertheless the condition found in the Ranidae and Rhacophoridae (the distal tendon of the semitendinosus passing dorsal to the gracilis) seems to be constantly present in this family and serves t o distinguish its procoelous members from the procoelous " Brachycephalids."

B. EGGS AND LARVAE The larval stages of relatively few species are known, but, even so, it seems possible to make certain generalisations, and there is every probability that the characteristic larvae of different groups are reliable guides to natural relationships. Three different larval types can be distinguished within the family, and each one of these is peculiar to the family. They are :(a) Free-swimming Tadpoles. Those forms which have a long free-swimming larval stage lay numerous small eggs, usually characterised by the presence of a ridge on, or an apical flattening of, the outer capsule. The tadpole has adhesive organs and external gills a t first ; later internal gills develop in the normal four, open, branchial clefts which are covered by an operculum. The peculiar and characteristic features are1:-(1) The spiraculum is median. (2) There is never any trace of horny mandibles or teeth, the lips always being soft and the lower usually somewhat extensible ; correlated with the lack of mandibles is the absence of the supra-rostra1 cartilage which, in other tadpoles, supports the upper jaw. (3) The external nares do not appear until shortly before metamorphosis. Tadpoles with these three peculiarities appear to be characteristic of the Asiatic and American genera ; they have been reported in Calluella, Glyphoglossus, Uperodon, Kaloula, Ramanella, Kalophrynus, Chaperina and Microhyla. It is also interesting to note that the Madagascar Dyscophus, which on anatomical grounds is separated from the other genera of that island and grouped with Calluella, has small ovarian eggs, suggesting a free larval stage. The only important modification of this type of tadpole is found in Microhyla achatina and M. heymonsi, where the lips are produced to form a large funnel, used, apparently, for collecting solid particles off the surface film of the water ; the feeding habits of others have not been fully investigated, but many are microphagous. Possibly connected with microphagy is the possession by some species of a terminal flagellum on the tail, which is bent round to lie parallel with the body and is maintained in constant vibration. I t has been suggested (Parker, 1928) that this movement sets up a forward current of the same momentum as the backwardly directed current of water which is being filtered for feeding purposes by the gills and passed out from the spiraculum ; in the absence of any such compensating forward current the spiracular &ream would force the animal ahead continuously. ( b ) Metamorphosing Within the Egg. Species 11~11ichhave no prolonged free-swimming tadpole stage either have a larva which hatches a t an advanced stage and undergoes metamorphosis within a definite, more or less enclosed, nesting site, or does not hatch until metamorphosis is completed. In either case the egg naturally has a much greater yolk-content than where there is a free-living tadpole. Where metamorphosis occurs within the egg-capsules, the eggs are usually, perhaps always, provided with a mucilaginous cord by which they are strung together or attached to some inanimate o b j e ~ t . ~ The encapsuled larva has no adhesive organs, external gills, branchial clefts, internal gills, operculum, spiracle or mouth armature. Nourishment is entirely derived from the yolk, and respiration is accomplished through a leaf-like, vascular, non-muscular tail which is apposed to the inner surface of the egg-capsule. Similar larvae have been described in the family Ceratophryidae (Eleutherodactylus), but the latter are distinguished by the presence of an egg-tooth ; the eggs, too, lack the mucilaginous cord of the present family. 1 I t has been claimed that the developing feet are fully webbed until after metamorphosis, but this is not universally true. 2 The eggs of Chaperina fusca, which has a free-swimming tadpole, also have a connecting corp.

MONOGRAPH O F THE MICROHYLIDAE This type of encapsuled larva has only been recorded in genera whose centre of origin and distribution appears to be the Papuan region, and it is significant that no other type of development has been recorded in forms from that area. Genera in which these peculiar larvae or eggs have been recorded are Sphenophryne (?), Oreophryne, Cophiaalus and Aslerophrys. (c) ~Metarnorphosing Within the Nesting Site. Other genera, widely separated geographically from the preceding, have dispensed with a free-swimming tadpole, but have not attained such perfect adaptation to a strictly terrestrial life. As already mentioned, the larva bursts the egg-capsulesbefore metamorphosis and the " tadpole " completes its development within the " nest." These terrestrial larvae are of two types :(i) In Breviceps (and it may reasonably be expected in the other African genera allied to it) the eggs, which are laid in an underground chamber, have no mucilaginous cord or apical flattening. The larva lacks adhesive organs, external gills, branchial clefts and internal gills (?), and a spiraculunl ; but an operculum within which the fore-limbs develop is still retained and the tail is muscular and not abnormally vascular. (ii) Hoplophryne lays large eggs, which adhere singly, in the internodes of bamboo stems or in the axils of banana leaves ; there is no mucilaginous cord and the eggs have no apical flattening. The tadpole, which hatches a t a late stage, has adhesive organs, but no external gills ; it resembles the free-swimming tadpole of the Asiatic and American genera in having a toothless, beakless mouth, no supra-rostra1 cartilage, an operculum with median spiracle, a muscular tail and nostrils closed until shortly before metamorphosis. It differs, however, in the possession of but a single functional branchial cleft, with very short filaments, and a pair of peculiar locomotor (?) flaps, one on each side of the branchial region. The life-histories of the Malagasy genera are, unfortunately, quite unknown. Except in Dyscophus, which appears on morphological grounds to be more closely allied to Asiatic forms than t o the rest of the Madagascar genera, wherever ovarian eggs have been recorded they have been large. This strongly suggests that they will be found to have a shortened free-larval stage or to resemble the Papuan genera in having dispensed with it entirely. It has been reported (Ahl, 1928) that in Mantipus the eggs within the body of the female show signs of development. But the inference that these frogs are viviparous, with internal fertilisation or parthenogenesis, seems rather improbable, and it is possible that the author was mistaken ; strangely enough, he has reported exactly the same remarkable condition in another genus of quite a different family (Tornierella). C. CLASSIFICATION AND PHYLOGENY Summarised, the characteristics of the various firmisternal families are :(1) ATELOPODIDAE, Fitzinger, 1843 (=Brachycephalidae, Giinther, 1858). (a) Vertebral column procoelous ; sacral diapophyses dilated or cylindrical. ( b ) Prevomer small, confined to the antero-mesial border of the choana ; ethmoid entire ; ear fuIly developed or middle ear incomplete. (c) No palatal folds ; Eustachian tubes present or absent. ( d ) Coraco-rudialis attached to the procoracoid-clavicular bar, in its mesial half ; insertion of the semitendinosus passing ventral to the gracilis. (e) Larva with horny mandibles and labial teeth ; supra-rostra1 cartilage present ; spiraculum sinistral. (f) No intercalary phalanges. 2) RANIDAE.

(a) Vertebral column diplasiocoelous ; sacral diapophyses cylindrical. ( b ) Prevomer moderately small, never plate-like nor almost surrounding the choana ; ethmoid entire ; ear fully developed. (c) No palatal folds ; Eustachian tubes present. (d) Coraco-radialis not attached to the procoracoid-clavicular bar, or only a t its

INTRODUCTIOK mesial extremity; insertion of the semitendinosus passing dorsal to the gracilis. ( e ) Larva with horny mandibles and labial teeth ; supra-rostra1 cartilage present ; spiraculum sinistral. (f) No intercalary phalanges. (3) RHACOPHORIDAE (=Polypedatidae nuct.).

(a) Vertebral column diplasiocoelous (rarely procoelous) ; sacral diapophyses cylindrical. (b) Prevomer moderately small, never plate-like nor almost surrounding the choana ; ethmoid entire, or, rarely, divided ; ear fully developed. (c) No palatal folds ; Eustachian tubes present. (d) Coraco-mdialis not attached to the procoracoid-clavicular bar, or only a t its mesial extremity ; insertion of the senzitendinoszts passing dorsal to the grucilis. (e) Larva with horny mandibles and labial teeth ; supra-rostra1 cartilage present ; spiraculum sinistral. (f) Intercalary phalanges present. (4) PHRYNOMERIDAE. (a) Vertebral column diplasiocoelous ; sacral diapophyses dilated.

(b) Prevomer confined to the antero-mesial border of the choana ; ethmoid divided ; ear fully developed. (c) Palatal folds present ; Eustachian tubes present. (d) Procoracoid-clavicular bar absent ; insertion of the semitendinosus passing dorsal to the gracilis. (e) Larva without horny mandibles, labial teeth or supra-rostra1 cartilage ; spiraculum median. (f) Intercalary phalanges present. (5) MICROHYLIDAE. (a) Vertebral column pro- or diplasio-coelous ; sacral diapophyses more or less dilated. ( b ) Prevomer often a large plate-like bone almost surrounding the choana ; ethmoid divided or absent ; ear fully developed or middle ear incomplete. (c) Palatal folds usually present ; Eustachian tubes present or absent. (d) Coraco-radialis attached to the whole width of the procoracoid-clavicular bar when the latter is present ; insertion of the semitendilzosus passing dorsal to the gracilis. (e) Larva without horny mandibles, labial teeth or supra-rostra1 cartilage ; spiraculum, if present, median. (f) No intercalary phalanges.

Unfortunately no fossil frogs are known which throw any light on the relationships of these groups. But from the scanty morphological evidence available it seems that the present family is most closely allied to the Phrynomeridae, and the latter ought, prd~a'rjly,t 0 ' ~ eregarheh as an o"frSn06cfrom 'fne same it cod^. 'lne rernSirirng+dires, the Ranidae are more nearly related to the Microhylidae than any other, but instead of the latter being a derivative of the former, as has previously been maintained (Noble, 1922, 1931), there is now a considerable amount of evidence which suggests that the Microhylidae retain a number of primitive features (e.g. large prevomer, divided ethmoid and dilated sacral diapophyses) which have been lost by the Ranidae. It is not unreasonable to suppose that the Microhylidae are an early offshoot from the same firmisternal stock, which, later, gave rise to the Rhacophoridae in the Ethiopian region and to the Ranidae in the Holarctic realm. Within the family certain groups of genera form natural assemblages, and it is possible to recognise a number of subfamilies. Noble (1931) has stressed the difficulty of distinguishing between groups whose common features are due to convergence and not to phylogeny, but the author believes that the following arrangement, which

10

MONOGRAPH OF THE MICROHYLIDAE

takes little or no account of the condition of that very variable element the procoracoidclavicular bar, expresses the relationships of the constituent genera of the family ; it has the merit of according with the facts of geography as well as of morphology, (1) DYSCOPHINAE. Diplasiocoelous forms retaining maxillary and vomerine teeth, together with a large undivided prevomer. There is a free-swimming larval stage. The three genera of this group, found in the Indo-Malayan region and in Madagascar, appear to be primitive in many respects, and may be the scattered remnants of the original stem from which the whole family is derived. (2) COPHYLINAE Procoelous genera with the prevomer divided and showing all stages in the progressive reduction of the post-choanal portion and of maxillary and vomerine teeth. Larva unknown. The eight genera of this group are confined to Madagascar, and, with the exception of Dyscop12us of the preceding subfamily, constitute the entire Microhylid fauna of the island. They show great diversity of form and structure and are the product of an isolated evolutionary centre. (3) ASTEROPHRYINAE. Diplasiocoelous, or rarely procoelous, genera without teeth, but retaining a primitive, large, undivided, plate-like prevomer ; vomerine odontoids often present and maxillae and dentaries often symphygnathine. Tongue adherent. Metamorphosis occurring within the egg ; embryo without operculum or other aquatic modifications. This is a small, but very varied, group of Papuan forms. Like the Cophylinae, they seem to be the product of an isolated evolutionary centre, for their range nowhere overlaps that of the Asiatic subfamily to which they are morphologically closely allied. (4) SPHENOPHRITNINAE. Procoelous genera with an undivided, plate-like prevomer. Teeth almost always absent, but vomerine odontoids sometimes present. Tongue free. Metamorphosis occurring within the egg. The three best-known genera of this group form a linear series. Their centre of distribution is the Papuan region, but they have reached northern Queensland (possibly in very recent times) and spread northwards t o the Philippines and Borneo. Thus their geographical range overlaps that of the Microhylinae, but they do not appear to intergrade with them. Very possibly this group should be united with the Asterophryinae. (5) MICROHYLINAE. Diplasiocoelous, or rarely procoelous, forms with the prevomer considerably reduced and usually confined to the antero-mesial border of the choana ; only in a few forms is the post-choanal portion persistent. Teeth absent. Tongue free or adherent. A free-swimming larval stage. This, the largest subfamily, ranges from Ceylon, southern and eastern India south to Celebes, north t o Manchuria, and from the south-eastern United States to the Rio de la Plata. The greatest number of genera occurs in the Asiatic portion of its range, but there seems to be complete intergradation between the Asiatic and American forms ; the genus Microhyla is common to both regions. Southwards along the chain of islands in the Malay Archipelago the number of forms rapidly diminishes, and the most south-easterly points reached are Celebes and Flores, where Kaloulu buleata is found. (6) BREVICIPITINAE. Diplasiocoelous genera with a large, plate-like prevomer. Ethmoid absent. Teeth absent. Tongue adherent. Metamorphosis taking place on land, the larva retaining an operculum but no other aquatic adaptations. The four genera of this subfamily are confined to southern and eastern Africa, are exceedingly closely allied, and have many features in common with the Asterophryinae and Microhylinae. The prevomer is smaller than in any Papuan genus, but better developed mesially than in any of the Microhylinae. The absence of an ethmoid is characteristic of all the genera, but is not diagnostic of the group, since this bone is also absent in Kalophrynus. It is the onIy subfamily in which a complete shoulder girdle is retained by all the genera. (7) MELANOBATRACHINAE. Procoelous genera without prevomers or teeth. Ethmoid fused with the parasphenoid. Ear incomplete. Palatal folds not developed. Metamorphosis taking place on land, the larva retaining an operculum, spiracle, adhesive organs and one pair of gill clefts.

INTRODUCTION The three genera of this group are confined to the mountains of south-western India and Tanganyika Territory. They are very incompletely known, but seem to be a natural, probably archaic, assemblage. Possibly they should be regarded as a distinct family, but the beakless, toothless mouth of the larva and its median spiraculum suggest affinity with the Nicrohylinae rather than with any other firmisternal group.

DISTRIBUTION Fossils being entirely unknown, there is no direct evidence available as t o the previous distribution of the family either in space or in time ; but from their present geographical distribution a few conjectures can be made. Various methods have been suggested for use in determining the centres of origin and dispersal of groups of animals and plants, many of them the subject of controversy and none of them universally applicable or generally accepted. But in the present instance there is one geographical region which fulfils three important criteria. It seems probable that, in the absence of insurmountable geographical or climatic barriers, dispersal from a centre of evolution will be radial. Climatic or other change may have exterminated a group a t its original centre, but the point of intersection of all the possible migrational routes along which it is known to have wandered will probably indicate its point of origin. The geographic centre of the range of the present family is south-eastern Asia. Prom such a centre it could spread southwards to New Guinea, westwards and southwards to Africa and Madagascar, and eastwards t o Central and South America ; whether a t one time it extended its range into Europe, Central Asia and North America there is no means of telling. The range of the family along these three probable routes is no longer always continuous, but that fact is of little significance since known climatic and geographical changes will account for the discontinuity. The African species are widely separated from the Oriental, but the desiccation of the " Eremian " region in comparatively recent geological times is sufficient to account for the absence of the family in that area. There is nothing to indicate, or suggest, that the African and Madagascan stocks reached their present range viii any land connexion that may have a t one time existed across the southern Indian Ocean. Similarly, the complete absence of the family from New Zealand and the islands of the Pacific lessens the probability of the American stock having reached that continent across the South Pacific, and the occurrence of the same genus in Asia and North America indicates that the North Pacific connexion was the route followed by the family. In addition to being the centre from which the migrational routes diverge, southeastern Asia is also the area in which the greatest differentiation has taken place. There can, of course, be no certainty that the rate of change of a group of animals is constant everywhere within its range, and it is by no means improbable that, occasionally, very rapid differentiation in a restricted area does take place. But where large areas containing all types of country are considered, and there is no likelihood of one containing a greater variety of " environmental niches " than another, that in which the largest number of genera is t o be found is, probably, the territory which has been longest occupied by the group. There can scarcely be any likelihood of greater diversity of possible habitats in south-eastern Asia than in, say, Africa or the Neotropical region, yet the number of genera in each of the major territorial divisions in which the family occurs is :S.E. Asia 13 ; Madagascar 9 ; Papua 9 ; America 7 ; Africa 6. The third, and least trustworthy, criterion of the centre of origin of a group is the presence of primitive forms. The most primitive Microhylidae occur in the Oriental region and in Madagascar, and it is, perhaps, significant that these are the two regions in which the greatest generic differentiation has occurred. The distribution of the family in time can, in the absence of fossils, only be a matter of conjecture. Nevertheless, certain salient features of their present distribution can be correlated with past geological changes. Thus their presence in the Papuan region and Madagascar is explicable only on the assumption either of fortuitous dispersal

DISTRIBUTION OF DYSCOPHINAE, Solid Black.

THE

SPIIENOPIIRYNINAE, Hatched.

SUBFAMILIES MELANOBATRACHINAE, Stippled (in circles)

DISTRIBUTION OF THE SUBFAMILIES COPRYLINAE, Solid Black.

ASTEROPHRYINAE, Hatched.

MICROHYLINAE, Stippled. BREVICIPITINAE, Crosses.

MONOGRAPH O F THE MICROHYLIDAE over the sea or of previous land connexions between the Australian and Oriental regions and between Madagascar and Africa or India. That frogs might be transported by floating rafts, or by wading birds, or by other haphazard means is possible, but, in the present instance, improbable. If i t did happen, it can have occurred but once in each region, for the present faunae are quite homogeneous, and it is rather remarkable that other, easier, passages, such as that from New Guinea to the Solomon Islands, have not been accomplished. If the migrations were overland, then it follows that the family was already differentiated by the end of the Mesozoic, for the Australian region, of which Papua forms part, is believed to have been separated from the Oriental not later than the Cretaceous or Paleocene. The abundance of the family in New Guinea and its almost entire absence from Australia probably reflects the difference in climatic conditions between the two, but might conceivably indicate a late arrival in the region and failure t o penetrate sufficiently far south before the formation of the Torres Straits. The subsequent geological events affecting the family may, roughly, have been as follows. Madagascar was isolated from Africa and South America from the north in Eocene times. Madagascar has probably only had transient connexions with Africa since that epoch and its Microhylid fauna has developed undisturbed. But South America was re-connected with the Nearctic region during the Miocene, whilst the northern trans-Pacific connexion was still unbroken, and i t seems probable that a further influx of northern forms took place, as well as, possibly, some reverse movement of the Neotropical elements. Otophryne and Chiasmocleis are relatively primitive forms which retain a fairly large clavicle, and are found only in the Neotropical region, but Microhylu, which has lost its clavicle and procoracoid, is found in both the Oriental and Nearctic regions, with but one or two species of uncertain relationships in the true Neotropical region. Hypopachus retains a clavicle and has one species widely distributed in South America, with four others in Central America, but whether it is a member of the original Neotropical fauna which has spread northwards, or whether, like Microhyla, it may be a recent immigrant, is uncertain. The trans-Pacific connexion was broken either in Miocene or Pliocene times, and the present distribution of the family was completed by the desiccation of northern Africa and Arabia during the Pleistocene.

HABITS Surprisingly little has been written on the habits of the members of this family, but there is a widespread impression that i t is predominantly a cryptozoic, myrmecophagous group. These two habits are certainly widespread among the better-known members of the family, but there can be no doubt that many of the species with well-developed digital discs are scansorial, if not definitely arborea1,l and others, again, especially in New Guinea and Madagascar, which by their slender habitus may reasonably be expected t o be " Grass-frogs." No species are known t o be wholly aquatic.

SECONDARY SEX CHARACTERS No great amount of sex dimorphism or dichromatism is constant,ly present, but, as in most other families, there is usually some disparity in the sizes of the sexes. Females, as a rule, tend to be larger than males, and, possibly connected with this increase in size, there is a distinct tendency in the Asterophryinae and Sphenophryninae for the female to develop vomerine odontoids to a greater extent than the other sex. In some forms the male is characterised by a distinct increase in the amount of digital webbing, but the most constant secondary sex characters are vocal sacs and nuptial pads. Vocal sacs are always, when present, of the " internal " type, with a slit-like opening on each side of the tongue, and correlated with their presence is the development of pigment in the gular skin. Nuptial pads are less frequently developed than vocal sacs, and they are usually of the diffuse type so common in the genus Rana. Kaloula conjuncta has been reported to ascend trees.

INTRODUCTION Enlarged, conical spines are rare, but have been reported in Kalophrynus robinsoni and in Hoplophryne ; the latter genus also has pectoral, mammiform rosettes of spines similar to those of Aelurophryne. Two characters of a type unrecorded in any other family are found in the genus Kaloula. In K. verrucosa the terminal phalanges of the fingers are beset with small bony projections ; these are much larger in the male and are accompanied by the development of a series of blister-like pustules on the dorsal surface of each finger-tip (figs. 35 and 36 ; p. 80). ;Males of the two most northerly species of the same genus, K. borealis and K. manchuriensis, have the aponeuroses of the obliquus muscles greatly thickened and coloured pink, a colour which is visible through the translucent skin of the abdomen.

TRENDS It has already been pointed out how, in different groups, the same kind of skeletal change has occurred, particularly in the reduction of the anterior ventral elements of the shoulder girdle and of the prevomers. In both instances these changes can be claimed to be degenerative and correlated with a loss of function, but other instances of parallelism occur in which no such correlation is apparent. In almost all the large genera there is a marked diversity of general habitus, and it is usually possible to trace a regular series from species with a " Ranid " habitus, with large eyes, short, blunt snouts, slender bodies and long legs and digits, to forms with long, pointed snouts, small eyes, obese bodies and short limbs and digits. This type of orthogenetic variation is particularly noticeable in the Asterophryinae, where it is accompanied by changes in the degree of development of the anterior palatal fold. Thus in Xenobatrachus there is an almost uninterrupted series from slender species such as X. ocellatus with two well-developed dermal ridges in front of the pharynx to a stout, pointedsnouted species like X. rostratus, in which there is only a very short posterior palatal fold and scarcely any trace of the anterior one. An exactly similar series can be traced in the genus Asterophrys from A. rufescens to A. oxycephala, but in the allied genus Metopostira only the extremes are known so that M. ocellatus bears a greater resemblance to species of Asterophrys, such as A. rufescens, than it does to its congener M. atra ; the latter, in its turn, is superficially very like A. oxycephala, with which, indeed, it has actually been confused. Similar series are known in the genus Microhyla and the subfamily Brevicipitinae shows a similar tendency. This group shows, also, a correlated reduction of the outer toe. In the relatively slender Callulina the outer toe extends beyond the third ; in Spelaeophryne and Probreviceps it does not extend as far as the third, but still extends well beyond the second, and in the globose Breviceps it never extends much beyond the second and is frequently shorter or even quite vestigial.

NOMENCLATURE The International Rules of Zoological Nomenclature, as proposed a t the ninth International Zoological Congress in 1913, and modified a t subsequent congresses, have been followed, except where they concern typography rather than nomenclature. These rules, however, are concerned more with generic, specific and subspecific names than with those of families and subfamilies, so that there is still considerable confusion in the methods used for selecting names for these higher groups. Articles 4 and 5 are the only ones dealing with the question, and the starting-point for the application of both of these is the " type genus," an entity which is not defined and concerning which no information is given. Nevertheless, it seems reasonable t o consider the " type genus " as the analogue of the "type species," in so far as this is possible ; and this principle has been adopted. Article 4 states : " The name of a family is formed to the stem of the name of its type genus." This by adding the ending -idae exact procedure cannot be expected to be retrospective, but the principle is an old one, for most of the old group-names are etymologically derived from those of one

...

MOKOGRAPH O F THE MICROHYLIDAE of their included genera, which, in accordance with the spirit of Article 4, must be regarded as the type genus. Unfortunately in the present instance the matter is complicated by the fact that two genera have almost equal claims to be so regarded. It was not until 1843 that the name of any genus of the present family was made the basis of the name of a higher group, but then Fitzinger, in the same work and on the same page, erected two families, the Hylaedactyli, based on Hylnductylus (=Kaloula), and the Gastrophrynae, based on Gastrophryne (=Microhyla) ; priority cannot be given t o either genus on the basis of the original publication. But, m accordance with Article 5 of the Rules, the name of the family would have to be changed whichever genus was selected, and would now become either Kaloulidae or Microhylidae. Preference is given to the latter, for, whereas no family or subfamily name was derived from Kaloula until 1931 (Noble), the name derived from Microhyla was used as long ago as 1858 (Giinther), and another synonym of the same genus was the basis of the family name Engystomatidae, which has been in common use for nearly half a century.

DESCRIPTIVE METHODS, ETC. This is primarily a systematic monograph, so that descriptions of habits and habitats have been reduced to a minimum unless they have a direct bearing on classification. The synonymies are, no doubt, often incomplete, but an attempt has been made to refer to all the more important papers dealing with each species ; citations of the same name ending in -i or -ii are not recorded sepnrately. The generic references are to all the names which have been used for the group and to the more important recent works on the genus or on its status. Descriptions are based on the material examined, with allowance for the recorded variations, except when these must be considered as pathological or abnormal. Unfortunately changes in proportions and form with growth and age have never been seriously studied in this family, so that the descriptions include the recorded variation a t all ages. This naturally gives a greater range than actually occurs a t any one stage for certain characters, but in most instances the serjes of specimens available for examination are so small that considerable deviations from the present descriptions must be expected. In tadpoles the amount of variation within a species is especially largeWright (1932) records tadpoles of the same species in which the tail may be twice as long as in others-and allowance must be made accordingly. The descriptive methods and phraseology which are here used call for some explanation, although not differing appreciably from those in general use. The length of a frog is the maximum distance in a straight line between the tip of the snout and the hinder side of the thighs behind the vent ; the dimensions given a t the end of the descriptions are those of the largest specimen examined or recorded. Descriptions of the shape of the snout refer, unless otherwise stated, t o its profile ; its length is measured in a straight line between its tip and the anterior corner of the eye. The size of the eye is measured from its anterior to its posterior corner. The relative size of the hind-limb is given, as usual, in terms of the position attained by one of its joints when the leg is laid forwards along the side of the body ; this method is usually quicker and more convenient than working out proportions by actual measurement, but a broken femur has often in the past resulted in a considerable error owing t o the limb bending a t the fracture rather than a t the hip-joint. The relative lengths of digits are obtained by laying them together in their natural plane parallel t o the axis midway between their normal axes ; for example, when the third and fifth are compared they are laid parallel t o the fourth, and whichever extends farthest distally is considered the longer. But when the length of a digit is to be compared with some other structure it is measured along its mesial side. Interdigital membrane is roughly measured as " half," " one-third," and so on in the usually accepted sense of these terms (Nieden, 1926), but when greater accuracy is desired the digits are laid side by side and the position of the edge of the membrane midway between them is described relative to a fixed

INTRODUCTION

17

point, such as a subarticular tubercle, on one or other of the nearest digits ; this method, it is believed, is more accurate, and less likely to be affected by bad preservation, than any attempt to describe the distal limits of the membrane on the sides of the toes. The specimens listed are those actually examined by the author ; the number preceding each is the register number of the institution indicated, and the collector's name is given where possible ; names in parentheses indicate the source from which the specimen was obtained, when the collector's name is uncertain. The indications " type " (=monotype or holotype), " paratype," " cotype " or " topotype," if unqualified, refer t o the specific name under which the specimen is now listed. In the generic diagnosis it will be found that the external characters common t o all the species are divided into two groups. There is very little theoretical justification for such a procedure, but in practice it is found that in large genera certain characters vary from species t o species, whereas in smaller, allied genera one or more of these same characters may be common t o all known species. These are precisely the features in which any new species which remains t o be discovered may be expected to differ from known forms ; they are not "generic characters " as usually understood, and are included in the generic diagnosis merely in the interests of brevity. But it must not be thought; that a newly discovered species which differs from known forms in one of the other " generic " characters must necessarily be referred to a distinct genus ; a reconsideration of the existing classification would, of course, be indicated, but each case will have t o be judged on its own merits.

LIST OF WORKS AND PAPERS REFERRED TO " Was

FITZINGER, GADOW, GUNTIIER, JONES,I.,

verstehen wir unter monophyletischer und polyphyletischer Abstammung ? " Verh. zoo1.-bot. Ges. Wien, LIX, pp. 243-249. " Beschreibung neuer Frosche aus Madagascar." Mitt. 2002. Mus. Berlin, XIV, pp. 469484. " Contribution 9. lY6tudede la Faune de Madagascar, Rept. et Batr." Faune Col. Frang., Paris. IV, 5, pp. 495-597. " Cat. Batr. Sal. Brit. Mus.," ed. 2, London. " Second Report on Additions to the Batrachian Collection in the Natural-History Museum." Proc. 2001. Soc. London, p. 327. " Systema Reptilium." Vindobonae. " Amphibia and Reptiles." Cambridge Nut. Hist., VIII. " Cat. Batr. Sal. Brit. Mus.," London. "Observations on the Pectoral Musculature of Amphibia Salientia." Ann. Mag. Nat. Hist. (lo), XII, pp.. 403420. " A Synopsis of Researches Relating to the Validity of the South African Brevicipitidae as an Autonomous Family of the Anura." S. Afr. Journ. Sci., Cape Town, XXVIII, pp. 451-455. " Beitrage zur Kentniss der Engystomatiden von Neu-Guinea." Termes. Fiizetelc, XXIV, pp. 169-271. " The Structure of the Vertebral Column in the Anura Phaneroglossa and its Importance as a Basis of Classification." PTOC. Linn. Soc. London, pp. 80-92. " Das Tierreich. Anura 11," Berlin and Leipzig. " The Phylogeny of the Salientia. I. The Osteology and Thigh Musculature ; their Bearing on Classification and Phylogeny." Bull. Amer. Mus. Nut. Hist., N.Y., XLVI, Art. I. " An Outline of the Relation of Ontogeny t o Phylogeny within the Amphibia, I." Amer. Mus. Novitates, N. Y., CLXV.

18

MONOGRAPH O F THE MICROHYLIDAE

" The Value of Life History Data in the Study of the Evolution of the Amphibia."

NOBLEand PARKER, PARKER,

1931. 1926.

Ann. N.Y. Acad. Sci., XXX, p. 113.

" Biology of the Amphibia." New York.

" A Synopsis of the Brevicipitid

Toads of Madagascar." Amer. Novitates, N. Y., CCXXXII. 1928. " The Brevicipitid Toads of the Genus Microhyla." Ann. Mag. Nat. Hist., (lo), 11, pp. 473499. " Parallel Modifications in the Skeleton of the Amphibia Salientia." Arch. Zool., Torino, XVI, pp. 1240-1248. SEBA, 1735. " Thesaurus," 11. DU TOITand 1932. " Die Skedelmorphologie van Hyperolius horstockii as voorbeeld DE VILLIERS, van die Polypedatidae." S. Afr. Journ. Sci., Cape Town, XXIX, pp. 449465. " New Aspects of Anuran Cranial Osteology and Osteogeny." S. Afr. Journ. Sci., Cape Town, XXVII, p. 485. " Breviceps and Probreviceps ; comparison of the Cranial Osteology of two closely related Anuran Genera." Anat. Anz., Jena, LXXV, pp. 257-304. " Life History of the Frogs of Okefinokee S~vamp,Georgia. North American Salientia (Anura) No. 2," New York. 1%.

SYSTEMATIC SECTION MICROHYLIDAE Bombinatorina (part), Gray, 1825, Ann. Philosophy, (2), x, p. 214. Bombinatoroidea (part), Fitzinger, 1826, Class. Rept., p. 39. Hylae (part), Ceratophrydes (part) and Bombinatores (part), Tschudi, 1840, Mem. Soc. Sci. Neuchatel, 11, 1839, pp. 27, 44. Hylaeformes (part) and Bufoniformcs (part), Dumeril and Bibron, 1841, Hist. Rept., VIII, pp. 491 and 640. Bufonoidea (part), Holbrook, 1842, N. American Herp., V, p. 5. Ranae (part), Hylaedactyli and Gastrophrynae (part), Pitzinger, 1843, Syst. Rept., pp. 31 and 33. Bufones (part) and Systomata, Lichtenstein, 1856, Nom. Rept., pp. 40 and 43 Asterophrydidae, Rhinodermatidae (part), Engystomatidae (part), Bufonidae (part), Micrhylidae, and Hylaedactylidae, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., ed. 1, pp. 37, 47, 51, 54, 121 and 123. Engystomidae (part) and Brachymeridae (part), Cope, 1865, Nat. Hist. Review, V, pp. 100 and 101. Brevicipitidae (part) and Phryniscidae (part), Cope, 1867, Journ. Acad. Philadelphia, (2), VI, pp. 192 and 195. Xenorhinidae and Discoglossidae (part), Mivart, 1869, Proc. Zool. Soc. London, pp. 289 and 294. Engystomatidae (part) and Dyscophidae, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, pp, 146 and 179. Cophylidae, Cope, 1889, Bull. U.S. Nat. Mus., x x x ~ v p. , 390. Genyophrynidae, Boulenger, 1890, Proc. Zool. Soc. London, p. 326. Brevicipitidae, Noble, 1922, Bull. Am. Mus. Nat. Hist., XLVI, p. 20; van Kampen, 1923, Amph. Indo-Austr. Arch., Leiden, p. 95. Engystomatidae, Nieden, 1926, Das Tierreich, Snura 11.

Firmisternal Anura, with a tongue ; eustachian tubes, if present, paired ; no ribs ; no intercalary phalanges ; vertebral column pro- or diplasio-coelous ; sacral diapophyses more or less dilated ; ethmoid divided or absent. Palatal folds usually present. Coraco-radiulis attached to the whole width of the procoracoid-cla~ricularbar when this is present ; insertion of the semitendinosus passing dorsal to the gracilis. Larva without horny mandibles, labial teeth or suprarostral cartilage ; spiraculum, if present, median. ARTIFICIAL SYNOPSIS O F THE GENERA

I. Clavicle and procoracoid present, well developed, both extending from the mid-line of the girdle to the scapula.

A. Maxillary teeth present. (1) Vomerine teeth present. (a) Pupil vertical ; omosternum with a bony style (1)Dyscophus, p. 22 (b) Pupil round or horizontal ; omosternum cartilaginous. (i) Fingers with small triangular dilatations ; toes free or with but a rudiment of web ; terminal phalanges Y- or T-shaped . . . . . , (2) Mantipus, p. 33 (ii) Fingers not dilated ; toes a t least 4 webbed ; terminal phalanges simple . . . . . . . (3) Calluella, p. 27 (2) Vomerine teeth absent ;pupil round ; omosternum cartilaginous ; fingers with large triangular dilatations ; toes free ; terminal . . . (4) Anodonthyla, p. 48 phalanges Y- or T-shaped . 19

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MONOGRAPH OF THE MICROHYLIDAE B. Maxillary teeth absent. (1) No definite folds across the palate in front of the oesophagus. (a) Terminal phalanges simple ; toes webbed a t the base ; tongue simple ; no eustachian tubes (5) Melanobatrachzis, p. 196 (b) Terminal phalanges T-shaped ; toes free. (i) Tongue with a transverse median furrow ; eustachian . . . . (6) Stumpflu, p. 50 tubes present . . . (ii) Tongue simple ; no eustachian tubes (7)Parhoplophryne, p. 198 (2) One or more transverse folds across the palate in front of the oesophagus. (a) A cartilaginous ornosternum present, even though small. (i) Terminal phalanges T-shaped. . . (8) Kalophrynus, p. 95 (a) Digits not dilated (p) Digits with small triangular terminal dilatations (9) Callulina, p. 180 (ii) Terminal phalanges simple. (a) Toes webbed ; coccyx and sacrum articulated, with 2 condyles . . . (10) Otophryne, p. 108 (p) Toes free. *No glandular area on the palate in front of the pharyngeal folds. ?Coccyx and sacrum articulated, with two condyles . . . (11) Spelaeophryne, p. 182 ttCoccyx and sacrum fused (12) Probrmiceps, p. 183 **A large glandular area on the palate in front of the pharyngeal folds ; coccyx and sacrum . . ,(13) Breuiceps, p. 186 fused . . (b) Ornosternum absent. (i) Terminal phalanges T-shaped, rarely club-shaped. (a) Vomer large, almost surrounding the choana, in contact with its fellow; vertebral column . . . (14) Sphenophryne, p. 151 procoelous (p) Vomer divided; vertebral column diplasiocoelous . . . . . (15) Chape~ina,p. 101 (ii) Terminal phalanges simple. (a) Clavicles strongly bent, following the curvature . . . (16) Ribeirina, p. 115 of the coracoids (p) Clavicles straight or but slightly bent (17)Hypopachus, p. 110

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11. Clavicle and procoracoid present, the former much reduced, not reaching the scapula and confined to the mesial portion of the procoracoid. A. Maxillary and vomerine teeth present. (1) Terminal phalanges with two articular condyles (18) Platyhyla, p. 36 (2) Terminal phalanges with a single articular condyle (19) Platypelis, p. 38 B. Maxillary and vomerine teeth absent. (1) Procoracoid and clavicle usually reaching the distal half of the coracoid. (a) Digits with distinct discs ; terminal phalanges T-shaped (20) Oreophryne, p. 159 (b) Digits not dilated or with only very small discs ; terminal phalanges simple . . . . (21) Chiusmocleis, p. 116 (2) Procoracoids and clavicles very small, not reaching the distal . . . (22) Elacl-listocEeis,p. 120 half of the coracoid

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111. Procoracoid present, extending from the scapula to the mid-line of the girdle, or nearly so far ; clavicle absent or reduced to one or two nodules lying on the procoracoid.

MICROHYLIDAE

21

A. &faxillaryand vomerine teeth present. (1) Omosternum present ; no rudiments of clavicles ; pupil hori. . . . . . . . . . (23) Pbthodontohyla, p. 42 zontal (2) Omosternum absent ; rudiments of the clavicles persisting as nodules mesially and distally ; pupil vertical (24) Colpoglossus, p. 30 B. Vomerine, but no maxillary, teeth present . . (25) Rhombophryne, p. 47 C. Neither vomerine nor maxillary teeth present. (1) Two transverse folds across the palate in front of the oesophagus ; tongue large, entire and scarcely free behind ; vomerine odontoids present ; eustachian tubes present . . (26) Genyophryne, p. 52 (2) One or two transverse folds across the palate in front of the oesophagus ; tongue oval, about half free behind ; no vomerine odontoids ; eustachian tubes present 1 (27) Gastrophrynoides, p. 104 c (28) fhficrobatrachus, p. 179 (3) No palatal ridges ; no vomerine odontoids ; tongue about half free ; procoracoid sometimes not reaching the mid-line of the girdle ; no eustachian tubes. (29) Hoplophryne, p. 198

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IV. Clavicle absent ; procoracoid present, but confined t o the mesial end of the coracoid and not reaching the scapula. A. Maxillary and vomerine teeth present. (30) Cophyla, p. 41 B. Maxillary and vomerine teeth absent. (1) Subarticular tubercles of the fingers greatly enlarged to form (31) Phrynella, p. 105 accessory adhesive organs (2) Subarticular tubercles not abnormally enlarged. (a) A bony ridge along the posterior border of each choana (32) Kaloula, p. 76 (b) A dermal ridge a little way behind each choana (33) Ramanella, p. 90

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.

V. Clavicles and procoracoids absent. Maxillary teeth absent. A. Sternum large, cartilaginous. ( a ) Maxillae suturally united in front of the premaxillae (Sym-

phygnathine) (i) One or two very large, steeple-shaped, vomerine odontoids (34) Xenobatrachus, p. 54 (ii) Vomerine odontoids, if present, not larger than normal vomerine teeth . (35) Asterophqs, p. 58 (b) Maxillae not meeting in front of the premaxillae (Eleutherognathine). (i) Tongue almost entirely adherent, and frequently with a median logitudinal furrow culminating posteriorly in a pouch. (a) Pupil vertical . . . (36) Ctenophryne, p. 122 (p) Pupil round or rhomboidal. *Toes, a t least, with small discs ; terminal phalanges . (37) Metopostira, p. 68 T-shaped ; digits free **Toes webbed, not dilated ; terminal phalanges simple. ?Mental region greatly distended ; tongue small (38) Glyphoglossus, p. 72 ??Mental region normal ; tongue moderately large . . . . . . . (39) Uperodon, p. 73 (ii) Tongue half free behind, without medium furrow or posterior pouch. (or) Subarticular tubercles of the fingers greatly enlarged to form accessory adhesive organs (40) Metaphrynella, p. 107

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MONOGRAPH OF THE MICROHYLIDAE (p) Subarticular tubercles of the fingers normal. +A single horseshoe-shaped prevomer almost surrounding the choana and in contact with its fellow . (41) Cophizalus, p. 170 mesially **Prevomer confined to the antero-mesial border of the choana ; not in contact with its fellow !42) Microlyla, p. 123 B. Sternum vestigial . . . . . . . . (43) Aphantopliryne, p. 178

DYSCOPHINAE, Boulenger Dyscophidae (part), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 179. Dyscophinae (part), Gadow, 1901, Cambridge Nat. Hist., p. 235. Dyscophinae (part) and Cacopinae (part), Noble, 1931, Biol. Amph., pp. 528 and 532.

Maxillary and vomerine teeth present ; prevomer large, entire, almost surrounding the choana (fig. 1) ; ethmoid divided ; ear fully developed ; vertebral column diplasiocoelous.

Fig. 1. Prevomers of Dyscophus antongili

The three genera of this group are sharply marked off from all the other members of the family. Maxillary and vomerine teeth are only known elsewhere in the Cophylinae and Sphenophryninae, but these subfamilies are characterised by a procoelous vertebral column. Both the presence of teeth and the characteristic prevomer must be accounted primitive characters, and the present discontinuous distribution of the group may also be taken as evidence of its antiquity.

SYNOPSIS O F THE GENERA I. Clavicles and procoracoids well developed ; omosternum bony ; pupil vertical . . . . . . . . . . . . . . . Dyscophus, p. 22 11. Clavicles and procoracoids present ; omosternum cartilaginous ; pupil circular . . . . . . . . . . . . . CaElueZEa, p. 27 111. Clavicles and procoracoids vestigial ; omosternum absent ; pupil vertical Colpoglossus, p. 30 -

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D YSCOPHUS, Grandidier Dyscophus, Grandidier, 1872, Ann. Sci. Nat., Zool., (5), xv, 20, p. 10 (Type species Dyscophus insularis, Grandidier) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 180; Noble and Parker, 1926, Amer. Mus. Novitates, ccrrxxIr, p. 5 ; Nieden, 1926, Das Tierreich, Anura 11, p. 86; Noble, 1932, Biol. Amph., p. 528. Kaloula (non Gray), Grandidier, 1875, Ann. Sci. Nat., Zool., 11, Art. 6. Phrynocara (part), Boettger, 1913, Voeltzkow, Reise in 0.Afrika, III, p. 283.

Maxillary teeth present; prevomer large, almost surrounding the choana, in contact with its fellow mesially, ankylosed t o the palatine and bearing a long row of teeth on its post-choanal portion (fig. 1). Clavicles and procoracoids present, the

DYSCOPHUS

23

former strong, usually bent ; procorawids usually in contact with the coracoids mesially ; sternum large, cartilaginous ; omosternum with a bony style (fig. 2). Vertebral column diplasiocoelous. Terminal phalanges simple.

Fig. 2.

Ventral elements of the shoulder-girdle of Dyscophw anlongili (ventral view)

Pupil vertical. Tongue large, oval, entire and half free behind. A denticulated dennai ridge across the palate in front of the pharynx, sometimes preceded by an indistinct curved one. Tips of digits not dilated ; toes webbed. Third toe extending well beyond the fifth ; outer metatarsal tubercle wanting. SYNOPSIS OF THE SPECIES

I. Nostril equidistant from the eye and the end of the snout. A. Metatarsal tubercle distinctly shorter than its distance from the tip of the inner toe, without distinct cutting edge. Large species ( 5 6 100 rnrn.). (1) First finger shorter than the second ; no lozenge-shaped dark outline on the back . . . . . . . . . D. antongili, p. 24 (2) First finger as long as, or slightly longer than, the second; a lozenge-shaped dark outline on the back . . . . D. guineti, p. 24 B. Metatarsal tubercle as long as, or longer than, its distance from the tip of the inner toe, with a distinct cutting edge. Small species ( 2 M 0 mm.). (1) Snout a little longer than the eye ; first finger shorter than the second . . . . . . . . . . . D. grandidieri, p. 25 (2) Snout not longer than the eye ; first and second fingers equal D. quinquelineatus, p. 28

MONOGRAPH O F T H E MICROHYLIDAE 11. Nostril nearer the tip of the snout than the eye. Small species. A. No lateral fold. Tarso-metatarsal articulation reaching the tympanic region. . . . D. beloensis, p. 26 B. A lateral fold. Tarso-metatarsal articulation reaching beyond the tympanic region . . . . . . . . . . . D. insularis, p. 26

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I.

Dyscophus antongili, Grandidier

Dyscophua insularis var. antongilii, Grandidier, 1877, Bull. Soc. Philom. Paris (7), I, p. 41 (Type locality Antongil Bay) ; D. antongilii, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 180 ; Boettger, 1892, Kat. Batr. Samm. Senckenberg, p. 25 ; Mocquard, 1895, Bull. Soc. Philom. Paris, (a), VII, p. 109 ; Mocquard, 1909, Nouv. Arch. Mus., Paris,(b), I, p. 71 ; Andersson, 1910, Ark. Zool. Stockholm, w, 7, p. 14 ;Methuen and Hewitt, 1913, Ann. Transvaal Mus., rv, p. 59 ; Noble and Parker, 1926, Amer. Mus. Novitates, CCXXXII, p. 8, fig. 2 ;Meden, 1926, Das Tierreich, Anura 11, p. 87. Dyscophus insuloris var. paUidus, Grandidier, 1877, Bull. Soc. Philom. Paris, (7), I, p. 42 (Type locality Andovoranto) ; D. antongilii var. pallidus, Mocquard, 1895, Bull. Soc. Philom. Paris. (8), w, p. 109. Dyscophus sanguineus, Boettger, 1880, Zool. Anz., p. 667 (Type locality Foizana) ; Boettger, 1881, Abh. Senck. Natf. Ges., xrr, p. 489, P1. m.

Head depressed, once and two-thirds as broad as long ; snout rounded, as long as, or a little shorter than, the eye ; canthus rostralis scarcely distinct ; loreal region very oblique ; nostril equidistant from the eye and the end of the snout ; interorbital space as broad as (dd), or broader than (9?), the upper eyelid ; tympanum hidden, or indistinct, 4 to % the diameter of the eye. First finger slightly shorter than the second, which is slightly shorter than the fourth ; toes nearly half webbed, the membrane more developed and with its free edge irregularly dentate in adult males ; subarticular tubercles well developed ; metatarsal tubercle rather prominent and com pressed. Tarso-metatarsal articulation reaching the anterior comer of the eye. Skin porous and feebly tubercular above ; upper eyelid and anal region strongly tubercular ; lower surfaces smooth, except the belly, which is granular, and the chin and throat of the adult male, which are pustular. A distinct fold of skin connecting the posterior edges of the upper eyelids ; a strong, glandular ridge from the posterior comer of the eye to the groin, with a short, curved branch behind the tympanum to the fore-limb. Magenta-red or olive-brown above, with a purplish black streak below the lateral fold ; flanks lighter ; yellowish white beneath, faintly mottled with brown. Length : 9 90 mm. d 63 rnm. t Male with a subgular vocal sac opening by a long slit on each side of the tongue and a diffuse nuptial pad covering the inner sides of the three inner fingers and the whole of the inner side of the fore-arm. DISTRIBUTION : N.E. Madagascar. B.M.

77.2.8. 21-23 1926.10. 27.8-11

2.

9

3 6 8, $?

Antongil Bay . Antongil Forest, Maroantsetra

(Geale) (Rosenberg)

Dyscophus guineti (Grandidier)

Kalula guineti, Grandidier, 1875, Ann. Sci. Net., Zool., (6), 11, Art. 6 (Type locality Sambava, coast N.E. Madagascar); Boettger, 1877, Abh. Senck. Natf. Ges., XI, p. 41. Dyscophus guineti, Steindachner, 1882, Sitzber. Ak. Wiss. Wieu, LXXXV,p. 191, P1. 111 ; Mocquard, 1895, Bull. Soc. Philom. Paris, (8), m, p. 109 ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), 1, p. 71 ; Noble and Parker, 1926, Amer. Mus. Novitates, CCXXXII, p. 8 ; Nieden, 1926, Das Tierreich, Anura 11, p. 87.

DYSCOPHUS Dyscophus imularis (non Grand., 1872). Grandidier, 1877, Bull. Soc. Philom. Paris, (7). I. p. 41 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 180 (in part) ; F. Miiller, 1885, Verh. Nat. Ges. Basel. vn, p. 136.

Head depressed, about once and two-thirds as broad as long; snout rounded, as long as, or a little shorter than, the diameter of the eye ; canthus rostralis obtuse ; loreal region very oblique ; nostril equidistant from the eye and the end of the snout ; interorbital space a little broader than the upper eyelid ; tympanum more or less distinct, about half the diameter of the eye. First finger as long as, or slightly shorter than, the second, which is a little shorter than the fourtli ; toes nearly half webbed ; subarticular tubercles well developed ; metatarsal tubercle slightly compressed, prominent. Tarso-metatarsal articulation reaching the anterior corner of the eye. Skin smooth or shagreened above ; smooth beneath ; upper eyelids granular ; a more or less distinct fold connecting the posterior corners of the upper eyelids, and a strong glandular ridge from the eye to the groin, with a short, curved branch running behind the tympanum to the fore-limb. Magenta to greyish above, with numerous small, regular, brown or reddish vermiculations ; a purplish black streak below the lateral fold, a vertical dark bar beneath the eye and an ill-defined lozenge- or diamond-shaped dark outline on the middle of the back. Yellowish white beneath, the chin and throat with dusty mottlings. Length : $' 95 mm. DISTRIB~IO : E. N Madagascar. Skin B.M. 72.7.2. 1 82.7.13. 8-9 99 1925.7.2. 93 9 Vienna Mus.

SPECIMENS EXAMINED Madagascar . . . . . Ancaye Forest . . . . Antsihanaka . . . . . Madagascar

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(Cutter) (Robihrd) (Rosenberg)

3 . Dyscophus grandidieri, Boulenger Dyscophus grandidieri, Boulenger, 1896, Ann. Mag. Nat. Hi&., (6), xvn, p. 404 (Type locality " S.W. Madagascar "); Mocquard, 1909, Nouv. Amh. Mus., Paris, (5), I, p. 71 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxu,p. 6 ;Nieden, 1926, Das Tierreich, Anura n, p. 88.

Head rather depressed, once and two-thirds as broad as long ; snout blunt, a little longer than the diameter of the eye ; canthus rostralis obtusely angular ; loreal region oblique ; nostril equidistant from the eye and the end of the snout ; interorbital space a little broader than the upper eyelid ; tympanum indistinct. First finger shorter than the second, which is about as long as the fourth ; toes 4 webbed ; subarticular tubercles of the fingers moderate, of the toes indistinct ; metatarsal tubercle large and compressed. Tarso-metatarsal articulation reaching the middle of the eye. Skin of the back porous anteriorly and rather warty posteriorly ; smooth beneath, except the h i d e r part of the abdomen and the thighs, which are granular. A welldefined fold from the eye to the groin, with a short, oblique branch above the tympanic region to the fore-limb. Grey' above, with a dark marking, truncate anteriorly and narrowed behind, connecting the upper eyelids and becoming indistinct on the occiput; a faint, dorsolateral, dark streak from the level of the fore-limbs to above the groin, and a wellmarked, dark, white-edged line beneath the lateral fold, continued forwards along the canthus rostralis. Lower surfaces yellowish white, closely mottled and stippled with brown. Length : 33 mm. DISTRIBUTION : S.W. Madagascar. SPECIMEN EXAMINED B.M. 92.11.26. 29 $ S.W. Madagascar . . . . . . . Last, TYPE The dorsum of the unique specimen is badly rubbed and appears chocolate-brown.

26

MONOGRAPH O F T H E MICROHYLIDAE

4 . Dyscophus quinquelineatus (Boettger) Phrynocara quingaeIineatum, Boettger, 1913, in Voeltzkow, Reise in 0. Afrika, 111,p. 283, P1. XXIII, figs. 11-17 (Type localities, W. Madagascar; Tsirnanampetso, S.W. Madagascar and Soalala, N.W. Mabgascar) ; Nieden, 1926, Das Tierreich, Anura n, p. 96. Dyswphus quinguelineatus, Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxrr,p. 6, fig. 1.

Head much broader than long ; snout rounded, as long as the orbit ; canthus rostralis short, but distinct ; loreal region steeply sloping ; nostril equidistant from the eye and the end of the snout ; interorbital space much broader than the upper eyelids ; tympanum indistinct or hidden, less than half the diameter of the eye. First, second and fourth fingers subequal ; toes webbed a t the base ; subarticular tubercles prominent ; metatarsal tubercle compressed, shovel-shaped. Tibio-tarsal articulation reaching to between the elbow and the shoulder. Skin smooth above, but beset with flat warts posteriorly and on the flanks ; a glandular fold from the eye to the insertion of the fore-limb. Belly and lower surfaces of the thighs granular. Silver-grey above ; a broad olive-black longitudinal band on the back, enclosing a silvery median streak posteriorly and broadening in front to form a triangular, anteriorly truncate, spot between the eyes ; a black dorso-lateral band from the level of the fore-limb towards the hind-limb and a lateral stripe from the tip of the snout, above the eye and tympanic region to the groin ; two parallel, oblique, silvery lines beneath the eye. Lower surfaces uniformly blackish brown, the chin often quite black ; some indistinct whitish dots on the breast and anterior portion of the belly. Limbs indistinctly cross-banded. Length : 25 mm. DISTRIBUTION : N.W. to S.W. Madagascar. This species may prove to be conspecific with the preceding.

5 . Dyscophus beloensis, Mocquard Dyscophus insularis (non Grandidier), Mocquard, 1900, Bull. Soc. Philom. Paris, (9), 11,p. 111. Dyswphus beloensis, Mocquard, 1902, Bull. Soc. Phibm. Paris, (9), N, p. 23, PI. 11, fig. 4 (Type locality Beb); Mocquard, 1909, Nouv. Arch. Mm., Paris, (5), I, p. 71 ; Noble and Parker, 1926, Amer. Mm. Novitates, ccxxxn, p. 6 ; Kieden, 1926, Das Tierreich, Anura rr, p. 89.

Head about once and two-thirds as broad as long ; snout rounded, a little shorter than the diameter of the eye ; canthus rostralis obtuse ; nostril nearer the tip of the snout than the eye ; interorbital space once and a half as broad as the upper eyelid ; tympanum indistinct. First finger shorter than the second, which is 8ubequal to the fourth ; toes # webbed ; subarticular tubercles well developed ; metatarsal tubercle compressed, shovel-shaped. Tarso-metatarsal articulation reaching the tympanic region. Upper surfaces fmely shagreened ; an indistinct fold from the eye to the fore-limb. Lower surfaces smooth, except the thighs, which are granular. Uniform dark brown above, with an indistinct, narrow, lighter lateral band from the eye to the anus ; grey, washed with brown beneath. Length : 38 rnm. DISTRIBUTION : Belo, W. Madagascar.

6. Dyscophus insularis, Grandidier Dyscophw insularis, Grandidier, 1872, Ann. Sci. Nat., Zool., ( 5 ) , xv, Art. 20, p. 10 (Type locality Antsouhy, near Trabounzy) ;Boettger, 1877. Abh. Senck. Natf. Ges., XI,p. 42 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 180 ;Mocquard, 1909, Nouv. Arch. Mus., Paris, ( 5 ) , I, p. 71 ; Noble and Parker, 1926, Amer. Mus. Novitates, CCXXXII,p. 8 ; Nicden, 1926, Dm Tierreich, Anura 11,p. 87.

CALLUELLA

27

Head broader than long ; snout rounded ; nostril much nearer the tip of the snout than the eye ; tympanum hidden. Toes webbed at the base ; metatarsal tubercle shovel-shaped. Tarso-metatarsal articulation reaching the eye. Skin smooth above, but granular on the sides ; a lateral fold and another from the eye to the shoulder ; belly granular. Reddish brown above, with darker brown, finely yellow-edged, spots and vermiculations ; flanks reddish ; lower surfaces reddish brown. Length 35 rnrn. DISTRIBUTION : Antsouhy, near Trabounzy. CALL UELLA,

Stoliczka

Calluella, Stoliczka, 1872, Proc. Asiatic Soc. Bengal, p. 146 (Type species Megalophrys guttulata, Blyth) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 181 ; Theobald, 1882, in Mason, Burma, I, p. 293 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 498 ; Nieden, 1926, Das Tierreich, Anura U, p. 89 ; Smith, 1930, Bull. Raffles Mus., m, p. 119 ; Noble, 1932, Biol. Amph., p. 528. Kdluella, Gee and Boring, 1930, Bull. D e p Biol. Yenching Univ., I, p. 25 ; Pope, 1931, Bull. Amer. Mus. Nat. Hist., LXI, p. 591. Dyscophina, van Kampen, 1905, Zool. Jahrb., Syst., XXII,p. 708 (Type species Dyswphina volzi van Katnpen); idem, 1923, Amph. Indo-Austr. Arch., p. 95 ; Nieden, 1926, Das Tierreich, Anura 11, p. 92.

Fig. 3. Anterior cranial elements of Calluella guttulata (seen from beneath)

Fig. 4.

Ventral elements of the shoulder-girdle of Calluella volzi (ventral view)

28

MONOGRAPH OF THE MICROHYLIDAE

Maxillary teeth present ; prevomer large, almost surrounding the choana, in contact with its fellow mesially, ankylosed to the palatine and bearing a long row of teeth on its post-choanal portion (fig. 3). Clavicles and procoracoids present, the former weak mesially ; sternum large, cartilaginous ; omosternum small, cartilaginous (fig. 4). Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil circular. Tongue large, oval, entire and slightly free behind. Two dermal ridges across the palate in front of the oesophagus. Tips of digits obtuse or slightly spatulate ; toes webbed. Canthus rostralis obtuse ; loreal region oblique, slightly concave ; nostril equidistant from the eye and the end of the snout. First finger shorter than the second ; third toe longer than the fifth. SYNOPSIS OF THE SPECIES I. Toes 4 webbed C. guttulata, p. 28 11. Toes webbed. A. Tibio-tarsal articulation reaching the tympanic region ; toes ending obtusely . . . . . . . . . . . C. yunnanensis, p. 29 B. Tibio-tarsal articulation reaching the eye ; tips of toes spatulate C. volzi, p. 30

. . . . . . . . . . . . .

I.

Calluella guttulata (Blyth)

Megdophrys gzlttulata, Blyth, 1855, Journ. Asiatic Soc. Bengal, xxrv, p. 717 (Type locality Pegu, Burma) ; Callula guttulata, Giinther, 1868, Proo. Zool. Soc. London, p. 490, P1. XL, fig. 1. Calluella gzlttulato,Stoliczka, 1872, Proc. Asiatic Soc. Bengal, p. 146 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 181 ; Theobald, 1882, in Mason, Burma, I, p. 293 ; Boulenger, 1890,Fauna Brit. India,Rept. and Batr., p. 498 ; Nieden, 1926,Des Tierreich, Anura n, p. 8 ; Smith, 1930, Bull. Raffles Mus., m, p. 119, fig. 9.

Snout rounded, a little longer than the diameter of the eye ; interorbital space about once and two-thirds as broad as the upper eyelid. Second finger not extending quite so far as the fourth. Toes ending obtusely, about 4 webbed, the membrane between the third and fourth toes reaching midway between the subarticular tubercles of the third toe ; subarticular tubercles well developed ; inner metatarsal tubercle compressed, shovel-shaped, outer indistinct. Tibio-tarsal articulation reaching the tympanic region. Skin smooth, or very feebly warty, above and beneath ; anal region and throat of male slightly granular ; a slight fold from the eye to the shoulder and sometimes one across the occiput. Pale brown or pinkish above, with darker brown markings. In juvenas the brown marking is usually in the form of a large spot which commences between the eyes connecting the upper eyelids, narrowing on the occiput, broadening on the nape, narrowing between the shoulders, broadening again and finally sending a diagonal branch across each groin. In adults this spot is broken up into coarse marblings which are narrowly edged with lighter. Hinder side of thighs dark brown or black, sharply outlined above, but shading into the ground colour beneath. Limbs with diagonal dark cross-bars. Lower surfaces white, or pale brown ; throat of the male infuscate. Male with a subgular vocal sac opening by a long slit on each side of the tongue. Eggs small ; a free-living tadpole. Length : 49 mm. A tadpole with well-developed hind limbs has the following characters : Head and body ovoid, rounded anteriorly, slightly depressed, once and two-thirds as long as broad. Nostrils about midway between a line connecting the anterior borders of the eyes, and the tip of the snout. Eyes lateral, 5 or 6 times as widely separated as the nostrils. Mouth terminal, half as wide as the head a t the level of the eyes, with a thin, downwardly curved, upper lip and a contractile lower one. Spiraculum median, opening about the centre of the gut ; spiracular flap broad, strongly emarginate in the centre so as to produce two lappets. Anus median, the anal tube

CALLUELLA

29

long, S-shaped. Tail twice, or more than twice, as long as the head and body, and about 6 times as long as deep ; acutely pointed posteriorly ; caudal membranes shallow, the upper much shallower than the lower anteriorly and not extending on to the body. Pale green, grey or brown, more or less translucent ; dark pigment is usually present around the nostrils, in the middle between the eyes and a t the base of the tail ; posterior half of the tail sooty black. Toes fully webbed, the membrane not being reduced to the adult condition until the tail has been completely resorbed. Total length of a well-grown example : 50 mm. ; head and body 16 ; tail depth 5-5. DISTRIBWION : Southern Burma to the northern part of the Malay Peninsula. SPECIMENS EXAMINED dd,?? Pegu . . . . . . . ? Burma . . . . . . . juv. ? . . . . . . . ? Chumpon, Siam . . . . d Ta Rua (approx. 15' N. x lOOo30'E.) . . . . . 1921.4.1. 286 juv. Chengmai, N. Siam . . . 1921.4kl. 287 JUV. Lat Bua Kao (approx. 15' N. x 100°30'E.) . . . . 1921.4.1. 288 juv. Koh Lak (approx. 13' N. x loooE.) . . . . . 1916.4.17. 2 ? Paknampo, Siam . . . . 1928.12.13. 38431arvae and Ta Rua, C. Siam . . . . young. (Series originally described)

Theobald Theobald Beavan M. A. Smith

B.M. 68.4.3. 93-96 65.11.5. 3 67.5.12. 45 1921.4.1. 284 1921.4.1. 285

2.

M. A. Smith M. A. Smith M. A. Smith M. A. Smith M. A. Smith M. A. Smith

Calluella yunnanensis, Boulenger

Callwlla yunnunensis, Boulenger, 1919, Ann. Mag. Nat. Hist., (9), m, p. 549 (Type locality Yun nanfn). KallueUa yunnunewis, Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., IV, 2 p. 25; Pope, 1931, Bull. Amer Mus. Nat. Hist., L ~ I ,p. 591.

Snout rounded, depressed, once and a third as long as the eye ; interorbital space about once and a half as broad as the upper eyelid. Second k g e r as long as, or a little longer than, the fourth. Toes slightly swollen distally, about webbed, the membrane between the third and fourth reaching beyond the level of the distal tubercle of the third toe in adult males ; in recently metamorphosed specimens the webbing is similar to that of C. guttulata ; subarticular tubercles moderate ; inner metatarsal tubercle compressed, the outer round and flat. Tibio-tarsal articulation reaching the tympanic region. Skin with a few feeble ridges corresponding with the edges of the dorsal markings ; smooth beneath ; anal region granular ; a distinct fold from the eye to the shoulder, and another across the occiput. Pale pinkish grey above, with traces of a dark lateral band commencing on the canthus rostralis, passing through the eye above the shoulder to the middle of the flanks ; a dark dorsal area commences between the eyes, where it connects the upper eyelids, narrows behind the head, broadens between the shoulders, narrows again and finally broadens on the middle of the back ; lateral branches from the dark dorsal area form dark spots in the groin. Limbs with dark, diagonal cross-bars. Lower surfaces dirty white, faintly mottled with brown especially on the throat. Male with a vocal sac. Length: 33.5 mm. ; d 29 mm. DISTRIBWION : Yunnanfu. SPECIMENS EXAMINED d and Hgr. Yunnanfu . . Graham, COTYPE~ B.M. 1905.5.30.47 and 1907.5.4. 30.

+

.

MONOGRAPH O F T H E MICROHYLIDAE

3. Calluella volxi (van Kampen) Dyswphina oolzi, van Kampen, 1905, Zool. Jahrb., Syst., xxn, p. 708, P1. XXVI (Type locality Tandjong Laut, Sumatra) ; idem., 1923, Amph. Indo-Austr. Arch., p. 96, fig. ; Nieden, 1926, Das Tierreich, Anura 11, p. 92.

Snout rounded, depressed, as long as the diameter of the eye ; interorbital space about once and a half as broad as the upper eyelid. Fingers ending obtusely, the second and fourth subequal. Toes spatulate distally, about 8 webbed, the membrane between the third and fourth toes reaching the level of the distal tubercle of the third toe ;the web extends as a slight fringe to the tips of all the toes ;subarticular tubercles moderate ; inner metatarsal tubercle compressed, the outer indistinct. Tibio-tarsal articulation reaching the eye. Skin with a few small scattered warts which are confluent on the sides to form a narrow fold from the corner of the eye to the groin ; an indistinct fold from the eye to the shoulder ; smooth beneath, the anal region granular. Reddish brown above, with short dark lines and small blotches ; lower surfaces lighter, marbled with brown beneath the limbs, which have indefinite diagonal crossbars above ; hinder side of thighs dark brown, this area being sharply defined above but merging into the ground colour beneath. Male with a vocal sac, and a plicate and infuscate throat. Length : 34 mm. : Tandjong Laut, Sumatra. DISTRIBUTION

Mus. Amsterdam

SPECIMENEXAMINED Tandjong Laut, Sumatra

. . . . .

COLPOGLOSSUS,

Volz, TYPE

Boulenger

Colpoglossus, Boulenger, 1904, Ann. Mag. Nat. Hist., (7), xrn, p. 42 (Type species C. brooksii, Boulenger) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 97 ; Nieden, 1926, Das Tierreich Anura 11, p. 90 ;Noble, 1932, Biol. Amph., p. 532. Calliglutus, Barbour and Noble, 1916, Proc. New Eng. Zool. Club, VI, p. 20 (Type species Calliglzltus smith;, Barbour and Noble) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 98 ; Noble, 1932, Biol. Amph., p. 532.

Fig. 5. Anterior cranial elements of Colpoglossus brooksi (seen from beneath)

Maxillary teeth present ; prevomer large, almost surrounding the choana, in contact with its fellow mesially, ankylosed to the palatine and bearing a long row of teeth on its post-choanal portion (fig. 5). Clavicles and procoracoids reduced to minute vestiges

COLPOGLOSSUS

31

(fig. 6)' ; sternum large, cartilaginous ; no omosternum. Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil vertical. Tongue large, oval, entire and slightly free behind, sometimes with a plicate pouch posteriorly (see p. 6). Two dermal ridges across the palate in front of the pharynx. Digits not, or but slightly, dilated ; toes with a rudiment of web.

Pig. 6. Ventral elements of the shoulder-girdle of Colpoglossus brooksi (ventral view)

SYNOPSIS OF THE SPECIES I. Tibio-tarsal articulation reaching the tympanic region ; toes not dilated terminally ; outer toe short, not reaching the proximal tubercle of the fourth . . . . . . . . . . . . . . C. brooksii, p. 31 11. Tibio-tarsal articulation reaching the anterior border of the eye ; toes slightly dilated terminally ; outer toe moderate, reaching the proxiC. smithi, p. 32 mal tubercle of the fourth, or a little beyond. . . . , I.

Colpoglossus brooksi, Boulenger

Colpoglossus brooksii, Boulenger, 1904, Ann. Mag. Nat. Hist., (7), xm, p. 42, P1. n (Type locality Bidi, Sarawak) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 97 ; Nieden, 1926, Das Tierreich, Anura n, p. 90. fig. .50.

Snout much depressed, not prominent, longer than the diameter of the eye ; canthus rostralis indistinct ; loreal region very oblique ; nostril nearer the tip of the snout than the eye ; interorbital space three times as broad as the upper eyelid ; tympanum hidden. Fingers short, obtusely pointed, the first shorter than the second, which is shorter than the fourth. Toes not dilated, with the merest trace of webbing a t the base and indications of lateral fringes ; fifth much shorter than the third ; subarticular tubercles indistinct ;a large prominent inner, but no outer, metatarsal tubercle. Tibiotarsal articulation reaching the tympanic region. Skin of back smooth ; head finely, flanks more coarsely, granulate ; smooth beneath, the chin and anal region alone granulate ; a fold from the eye to the shoulder. Pale, yellowish white above, with a bold reticulum of dark brown; a X-shaped dark brown, light-edged marking on the head and nape, the longitudinal branches commencing on the upper eyelids and bifurcating in front and behind ; two chains of These vestiges were not mentioned in the original descriptions of either Colpoglossus or Calliglutus, but have been found in both.

MONOGRAPH OF THE MICROHYLIDAE black spots, some light-centred, commencing a t the posterior end of the X and running almost to the vent. Limbs with brown reticulations. Lower surfaces immaculate white, except the throat, which is etched with brown. Male with a subgular vocal sac. Length : 50 mm. DISTRIBUTION :

Bidi, Sarawak. SPECIMEN EXAMINED

B.M.

1903.11.24. 3

2.

8

Bidi, Sarawak

. .

. . . .

. . Brooks, TYPE

Colpoglossus smithi (Barbour and Noble)

Calliglutus mithi, Barbour and Noble, 1916, Proc. New Eng. Zool. Club, VI, p. 20, P1. n (Type locality Limbang River District, hrawak) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 98.

Snout rounded, depressed, about as long as the diameter of the eye; canthus rostralis indistinct ; loreal region oblique ; nostril nearer the tip of the snout than the eye ; interorbital space about 3 times as broad as the upper eyelid ; tympanum hidden. Fingers short, not dilated, the first shorter than the second. Toes with a trace of webbing a t the base, the tips feebly dilated ; the fifth shorter than the third ; subarticular tubercles slightly distinct ; a large oval inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the anterior border of the eye. Skin entirely smooth. Brownish grey above, with a median, pink-edged, dark mark on the back, commencing between the eyes, narrowing on the nape, broadening between the shoulders, narrowing again and then broadening over the rest of the dorsum ; flanks with black, light-edged blotches, of which the most prominent are a circular spot on the tympanic region, connected to the eye by a short, narrow bar, and an irregular marking in the groin and on the anterior aspect of the thigh; hinder side of thighs black; limbs with an irregular dark reticulum above ; a bright red X -shaped mark above the vent. White beneath, the throat and sides washed with light brown and stippled with white. Length : 31 mm. (imm. 9). DISTRIBUTION : Limbang River District, Sarawak. SPECIMEN EXAMINED M.C.Z. 3798

(immature)

Limbang River, N. Sarawak

. . * SmitA, PARATYPE

COPHYLINAE, Cope Cophylidae, Cope, 1889, Bull. U.S. Nat. Mus., xxxm, p. 390. Rhombophryninae, Noble, 1932, Biol. Amph., p. 529.

Maxillary or vomerine teeth, or both, usually present (except Slumpfza) ; prevomer divided into two, a choanal and a post-choanal portion, the latter rarely reduced or absent (Anodonlhyla) ; ethmoid divided ; ear fully developed ; vertebral column procoelous. The combination of maxillary or vomerine teeth and a procoelous vertebral column is usually s d c i e n t to define this exclusively Malagasy subfamily. But a similar condition is also rarely found in the Sphenophryninae, a group to which the toothless Slumpfza approaches closely. All the Sphenophryninae, however, retain a primitive undivided prevomer, and it seems probable that the two groups are descendants in different geographical regions of some Dyscophus-like, but procoelous, stock.

MANTIPUS SYNOPSIS OF THE GENERA

I. Vomerine teeth present.

.

A.'Maxillary teeth present ; post-choanal prevomer a long plate overlying the palatine ; vomerine teeth in two groups or series. (1) Clavicle well developed, extending from the mid-line of the girdle to the scapula. Tips of digits with small dilatations Mantipus, p. (2) Clavicle confined to the mesial half of the procoracoid. Tips of digits with large discs. (a) Terminal phalanges of the fingers articulating by a double condyle . . . . . . . . . . . . Phtyhyla, p. (b) Terminal phalanges of the fingers articulating by a single Phtypelis, p. condyle (3) Clavicle absent . . . . . . . . . . . Pbthodontohyla, p. B. Maxillary teeth present ; post-choanal prevomer fused with its fellow and not overlying the palatine ; vomerine teeth in a single median . . . . . . . . . . . . Cophyla, p. group C. Maxillary teeth absent ; post-choanal prevomer a long plate overlying the palatine ; vomerine teeth in two series . Rhombophryne, p.

. . . . . . . . . . . .

11. Vomerine teeth absent. A. Maxillary teeth present ; digits with terminal dilatations B. Maxillary teeth absent ; digits not dilated .

33

36

38 42 41 47

. Anodonthyla, p. 48 Stumpfla, p. 50

MANTIPUS, Peters Pldhodoniohyla (part), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 473; Werner, 1903, Zool. Anz., xxvr, p. 351. Mantipm, Peters, 1883, Sitzber. Ak. Wiss. Berlin, p. 165 (Type species M . hildebmndti, Peters) ; Noble and Parker, 1926, Amer. Mus. Novitates, CCXXXII, p. 8 ; Nieden, 1926, Das Tierreich, Anura 11, p. 94 ; Noble, 1932, Biol. Amph., p. 530. Mantiphrp, Mocquard, 1895, Bull. Soc. Philom. Paris, (a), vrr, p. 134 (Type species Manliphrys luevipes, Mocquard =M . hildebrandti, Peters). Mantophrya (emend.), Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 72.

Fig. 7. Prevomers of M a n t i p a inguinalis

Maxillary teeth present ;prevomer divided, the post-choanal portion long, in contact with its fellow mesially, overlying the palatine and bearing a long row of teeth (fig. 7). Clavicles and procoracoids present, well developed, reaching the mid-line of the girdle ; sternum cartilaginous ; omosternum small, cartilaginous (fig. 8). Vertebral column procoelous. Terminal phalanges Y- or T-shaped. 3

MONOGRAPH O F THE MICROHYLIDAE

Fig. 8.

Ventral elements of the shoulder-girdle of Mantipus inguinalis (ventral view)

Pupil horizontal. Tongue large, oval, entire and half free behind. A lobulated ridge across the palate in front of the pharynx. Digits slightly dilated terminally. First finger shorter than the second, which is a little shorter than the fourth ; third toe longer than the fifth ; outer metatarsal tubercle wanting. SYNOPSIS OF THE SPECIES I. Tarso-metatarsal articulation reaching well beyond the tip of the snout. No fold along the sides of the body. A. Toes free. (1) Nostril eauidistant from the eve and the end of the snout M. inguinalis, p. 34 (2) Nostril much nearer the tip of the snout than the eye M. hildebrandti, p. 35 B. Toes with a rudiment of web . . . . . . . M. angecliferus, p. 36 11. Tarso-metatarsal articulation not reaching beyond the tip of the snout. A fold along the sides of the body to the groin . . . M. peclcher, p. 37 \

1

t

I.

Mantipus inguinalis (Boulenger)

Plethodontohylrr inguinalis, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 473 (Type locality, East Betsileo); Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 7 2 ; Nieden, 1926, Das Tierreich, Anura n, p. 91. M a n t i p inguinali.9, Noble and Parker, 1926, Amer. Ifus. Novitates, ccxxxrr, p. 10, fig. 4. Head broad and flat, once and a half as broad as long ; snout vertically truncate, not prominent, once and a quarter as long as the eye ; canthus rostralis angular ;loreal region slightly oblique; nostril equidistant from the eye and the end of the snout ;

interorbital space nearly twice as broad as the upper eyelid which, a t the anterior corner of the eye is produced into a small but distinct, semicircular, vertical flap ; tympanum indistinct, about half the diameter of the eye. Fingers free, with small, triangular dilatations. Toes free, slightly dilated distally ; subarticular tubercles of the fingers well developed, of the feet indistinct ; metatarsal tubercle oval, ill-defined. Tibio-tarsal articulation reaching between the eye and the end of the snout. Skin smooth above and beneath; sometimes a few small pustules are present on the snout and the flanks ; a well-marked fold from the upper eyelid to the fore-limb. Brown, grey or purplish above ; loreal region and sides of head beneath the supra-

MANTIPUS tympanic fold black, sometimes faintly white-bordered. Anterior half of dorsum usually with a large, dark blotch which is truncate anteriorly, where i t connects the upper eyelids,rand breaks up into irregular short lines and smaller blotches posteriorly; sometimes a small, dark, inguinal spot. Limbs with narrow, dark cross-bars. Lower surfaces spotted and stippled with brown or black, more markedly so on the throat and chest ; hinder side of thighs brown with small, round, white dots. Length : 64 mm. Male without a vocal sac ; a flat, semicircular, projecting pollex. DISTRIB~TION : E. and N.E. Madagascar.

.

SPECIMENS EXAMINED B.M. 82.3.16. 30 9 East Betsileo . . . . 1925.7.2. 94 Antsihanaka . 1925.11.10. 12 Hgr.3 Antsihanaka . . . 1928.5.9. 20-21 99 Brickaville . .

. . . . .

2.

. . .

. . . . . . . . . .

Cowan, TYPE (Rosenbeg) (Rosenberg) (Rosenberg)

Mantipus hildebrandti, Peters

Mantipus hildebrandti, Peters, 1883, Sitzber. Ak. Wiss. Berlin, p. 166 (Type locality " Interior of ~ada~ascar*') ; Mocquard, 1896, Bull. Soc. Philom. Paris, (8), v n , p. 132 ; Mocquard, 1902, Bull. Soc. Philom. Paris, (9),IV, p. 23 ; Mocquard, 1909, Nouv. Amh. Mus., Paris, (5),I, p. 72 ; Methuen and Hewitt, 1913, Ann. Transvaal Mus., IV, p. 60 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 10, fig. 3 ;Nieden, 1926, Das Tierreich, Anura n , p. 95. Blantiphrys laevipes, Mocquard, 1895, Bull. Soc. Philom. Park, (a), m, p: 134 locality Mt. Ambre). Mantophrys laevipes, Mocquard, 1909. Nouv. Arch. Mus., Paris, (5), I, p. 72. Head flat, broader than long ; snout obtusely pointed, somewhat longer than the eye ; canthus rostralis distinct ; nostril much nearer the tip of the snout than the eye ;

(me

interorbital space twice as broad as the upper eyelid; tympanum indistinct, about half the diameter of the eye. Fingers free, with triangular dilatations. Toes free, with smaller triangular discs ; subarticular tubercles of the feet indistinct ; metatarsal tubercle indistinct. Tibio-tarsal articulation reaching the middle of the eye. Skin smooth above and below, with only a few small granules on the hinder side of the thighs ; a distinct supra-tympanic fold from the eye to the shoulder. Brown above, sometimes with an indistinct angular marking, the apex of which lies on the occiput ; loreal and temporal regions blackish ; a grey cross-bar between the eyes; lips flecked with lighter. Brown beneath, flecked with lighter in the groin, behind the thigh and beneath the tibia. Limbs with dark cross-bars. Length : 62 mrn. DI~TRIB~TION : Madagascar. SPECIMEN EXAMINED 1 ad. Madagascar Hildebrandt, TYPE Mus. Berlin 10440

. . . . .

3. Mantipus (?) anguliferus (Werner) Plethodontohyla angdifera, Werner, 1903, Zool. Am., xxn, p. 251 (Type locality unknown) ;Nieden, 1926, Das Tierreich, Anura II, p. 92. S Mantipwr anguliferw, Noble and Parker, 1926, Amer. Mus. Novitates, coxxxn, pp. 8 and 10.

Head broad ; canthus rostralis distinct; interorbital space twice as broad as the upper eyelid ; tympanum half the diameter of the eye. Fingers free, with small, truncate dilatations. Toes with a rudiment of web and very small discs ; subarticular tubercles quite indistinct. Tarso-metatarsal articulation reaching well beyond the tip of the snout. Skin smooth above and beneath, but pustular on the snout ; a postocular dermal fold. Reddish brown above, with four darker angular markings on the back ; the points of these bars are directed forwards and the hindermost and smallest unites a pair of

36

MONOGRAPH OF THE MICROHYLIDAE

inguinal spots ;loreal region and temples darker. Limbs with dark cross-bars. Throat and lower surfaces of the limbs reddish brown ; belly white. Length : 25 rnrn. DISTRIBUTION : Unknown.

4. Mantipus pulcher, Ah1 m a n t i p pdcher, Ahl, 1928, Mitt. Zool. Mus. Berlin, XIV, p. 484 (Type locality Akkoraka). Head flat, depressed, broader than long ; snout acuminate, scarcely prominent, longer than the eye ; canthus rostralis distinct, angular ; loreal region nearly vertical, not concave ; nostril nearer the tip of the snout than the eye ; interorbital space about twice as broad as the upper eyelid ; tympanum indistinct, about 3 to 3 the diameter

of the eye. Fingers free, with moderately large, triangular dilatations. Toes free, with smaller discs than the fingers ; subarticular tubercles of the hands well developed, of the feet indistinct ;metatarsal tubercle indistinct. Tibio-tarsal articulation reaching the tympanum or the posterior corner of the eye. Skin smooth above and beneath, only the hinder side of the thighs being granular ; a narrow, very distinct, fold from the eye, along the flanks to the groin. Pale brownish grey above, with a series of dark, chevron-shaped cross-bars, of which the acute angle is directed forwards ; canthus rostralis, edge of upper eyelid and lateral fold white, the sides below being dark brown. A pair of minute, white, black-bordered ocelli on the sacrum and sometimes another pair anterior to these. Limbs with narrow, brown cross-bars. Dirty white beneath, males regularly powdered with brownish. Anal region dark brown, edged with white above ; sole dark brown. Length : 34 mm. Females with a few (about twenty) very large (3.5 mm. diameter) eggs, which are reported to show signs of embryonic development. : Akkoraka, Central Madagascar. DISTRIBUTION Mus. Berlin 31623

SPECIMENS EXAMINED 5 88, ?? Akkoraka

PLAT YH YLA,

. .

. .

Braun, COTYPES

Boulenger

Platyhyla, Boulenger, 1889, Ann. Mag. Nat. Hist., (6), IV, p. 247 (Type species P. grandis, Boulenger); Nieden, 1926, Das Tierreich, Anura 11, p. 93 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 10 (in part) ; Noble, 1932, Biol. Amph., p. 530. Plethodontohyh, Methuen and Hewitt, 1912, Ann. Transvaal Mus., (2), IV, p. 60 ; Methuen, 1919, Proc. Zool. Soc. London, p. 354.

Fig. 9. Prevomers of Platyhyla grandis

Maxillary teeth present ; prevomer divided, the post-choanal portion long, in contact with its fellow mesially, overlying the palatine and bearing a long row of teeth (fig. 9). Clavicle present, but much reduced, not reaching the mid-line of the girdle or the scapula ; procoracoid curved, broad, resting on the middle of the anterior border of the coracoid (fig. 10) ; sternum large, cartilaginous ; omosternum small. Vertebral column procoelous. Terminal phalanges broadly Y-shaped, articulating by two condyles (fig. 11). Pupil horizontal. Tongue large, oval. entire and half free behind. Palate without dermal folds. Tips of digits broadly dilated.

PLATYHYLA

Fig. 10. Ventral elements of the shoulder-girdleof Platyhykz grandis (ventral view)

Fig. 11. Terminal phalanx of third finger of Platyhykz grand& to show the double articular condyle

I.

Platyhyla grandis, Boulenger

Platyhykz grandis, Boulenger, 1889, Ann. Mag. Nat. Hist., (6), N, p. 247 (Type locality " Madagascar ") ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 73 ; Methuen and Hewitt, 1913, Ann. Transvaal Mus., (2), N,p. 60 ; Noble and Parker, 1926, Amer. Mu. Novitates, ccxxxn, p. 10, fig. 5 ; meden, 1926, Das Tierreich, Anura n, p. 93; Angel, 1931, Faune Col. Frang., Iv, 5. p. 541. Platyhykz vermcosa, Mocquard, 1901, Bull. Mus., Paris, vn, p. 254 (Type locality Fort Dauphin) ; Mocquard, 1902, Bull. Soc. Philom. Paris, (Q),N, p. 24, P1. n, fig. 5 ; Mocquard, 1909, Nouv. Arch. MuB.,Paris, (5), I, p. 73 ; Nieden, 1926, Das Tierreich, Anura n, p. 94; Noble, 1931, =ol. Amph., fig. 1 7 3 ~ . Platyhyla voeltzkowi, Boettger, 1913, in Voeltzkow, Reise in 0.Afrika, In,p. 282, P1. xxm, fig. 7 (Type localities Sakana and Anevoka, E. Madagascar) ; Nieden, 1926, Das Tierreich, Anura n, p. 94. Head depressed, once and a half as broad as long ; snout rounded, not prominent, as long as, or slightly longer than, the eye ; canthus rostralis obtuse ; loreal region very oblique, concave ; nostril a little nearer the tip of the snout than the eye ; interorbital space once and a half as broad as the upper eyelid ; tympanum hidden. Fingers

with very large triangular dilatations, that of the third finger broader than the diameter of the eye; first finger shorter than the second, which is a little shorter than the fourth. Toes about 4 webbed, with large triangular dilatations, the fifth a little shorter than the third ; subarticular tubercles moderate ; an ill-defined oval inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum or the eye. Skin more or less tubercular above, especially on the snout ; young specimens are, as a rule, rougher than older examples ; a fold from the eye t o the shoulder ; lower surfaces and hinder side of thighs granular. Olive-brown to cream above, with some irregular darker or lighter'markings of which the best defined are an irregular W between the shoulders and a cross-bar

38

MONOGRAPH O F T H E MICROHYLIDAE

between the eyes ; snout darker. White beneath, uniform or with a few scattered brown dots. Limbs with broad, dark, faintly light-edged cross-bars. Male with a vocal sac. Length : 102 mm. DISTRIBUTION : E. Madagascar. SPECIMENS EXAMINED B.M. 89.8.1.3132 Madagascar. . . . . . Majastre, COTYPES 92.3.7.53 Sahambendrana . . . . Majastre d, 2 yg. Antsihanaka . . . . . (Rosenberg) 1925.7.2.97-99 1928.5.9.22 ? Brickaville . . . . . . (Rosenberg)

dP

PLAT YPELIS, Boulenger Platypelis, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 474 (Type species Platypelis wwanii, Boulenger) ; Nieden, 1926, Das Tierreich, Anura n, p. 97 ; Noble and Parker, 1926, Amer. Mus. Novitetes, ccxxxn, p. 12 ;Noble, 1931, Biol. Amph., p. 630. Plethodontohyla ( m a Boulenger), Methuen end Hewitt, 1913, Ann. Transvaal Mus., (2), m, p. 66; Methuen, 1919, Proc. Zool. Soc. London, p. 354. Platyhyla (part), Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn,p. 10. Caphyh ( m n Boettger), Ahl, 1929, Mitt. Zool. Mus. Berlin, xrv,p. 483. Maxillary teeth present ; prevomer divided, the post-choanal portion long, in

contact with its fellow mesially, overlying the palatine and bearing teeth (fig. 12). Clavicle present, much reduced, not reaching the mid-line of the girdle or the scapula ; procoracoid broad, curved, resting on the middle of the anterior border of the coracoid ; sternum large, cartilaginous ; omosternum small and cartilaginous (fig. 18) or absent. Vertebral column procoelous. Terminal phalanges Y-shaped, articulating by their whole proximal surface. Pupil horizontal. Tongue large, oval, entire and half free behind. Palate without dermal folds. Tips of digits broadly dilated. Toes webbed. The first finger is very short, much shorter than the second, which is slightly shorter than the fourth ; outer metatarsal tubercle wanting.

Fig. 12. Prevomers of Platypelis cowuni

n

Fig. 13. Ventral elements of the shoulder-girdleof Platypelie m n i (ventral view)

PLATYPELIS SYNOPSIS OF THE SPECIES I. Outer toe shorter than the third and with a smaller disc. No enlarged prepollex. A. Vomerine teeth in long transverse series behind the choanae P. tuberifera, p. 39 B. Vomerine teeth in short series, not extending laterally beyond the choanae. P. cmani, p. 39 (1) Skin smooth . . . . . . . . . . . . (2) Skin tubercular . . . . . . . . . . . P. tuberculata, p. 40 11. Outer toe longer than the third and with a large disc. A very prominent prepollex . . . . . . . . . . . . . . P. pollicaris, p. 40

Platypelis tuberifera (Methuen)

I.

Plethodontohyla notostictu (non Giinther), Methuen and Hewitt, 1913, Ann. Tramvaal Mus., (2),IV, p. 60.

~letbdontoh~la tuberifera, Methuen, 1919, Roc. Zool. Soc. London, p. 354 (Type locality Ambatoharanana). Platyhykz tuberijera, Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxu, p. 11. Head much depressed, once and a quarter as broad as long ; snout rounded, slightly prominent, longer than the diameter of the eye ; canthus rostralis scarcely marked ; loreal region very oblique, slightly concave ; nostril slightly nearer the tip of the snout than the eye ; interorbital space once and a half to twice as broad as the upper eyelid ; tympanum about half the diameter of the eye. Fingers with large subtriangular dilatations; the first very short, much shorter than the second, which is a little shorter than the fourth. Toes free, with large discs ; fifth slightly shorter than the third. Subarticular tubercles very feebly developed ; an indefinite inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum or the eye. Skin quite smooth ; a feeble fold from the eye to the fore-limb, and a pair of glandular spots, which may form distinct tubercles, on the sacrum; posterior part of belly and hinder side of thighs sometimes faintly granular. Pale brown to cream above, irregularly washed and spotted with darker; supratympanic fold edged with black ; sacral glandular spots white, dark-edged ; a fine white line from snout to vent. Beneath uniform white. Length : 40 mm. : Central and N.E. Madagascar. DISTRIBUTION

B.M. 1919.8.15.1 1925.11.10.8-11 2.

? 8, 3 99

Ambatoharanana Antsihanaka

.

.

. . . . .

Methuen, COTYPE (Rosenberg)

Platypelis cowani, Boulenger

Platypelis manii, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 474 (Type locality East Betsileo) ;Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 74 ;Nieden, 1926, Das Tierreich, Anura n , p. 97 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 12, fig. 6. Head depressed, once and a quarter as broad as long; snout rounded, scarcely prominent, as long as the diameter of the eye ; canthus rostralis scarcely distinct ; loreal region oblique ; nostril equidistant from the eye and the end of the snout ; interorbital space broader than the upper eyelid ; tympanum indistinct, f the diameter of the eye.l Fingers with triangular-dilatations, very large except on the first finger. Toes with a rudiment of web and triangular dilatations smaller than those of the fingers ; fifth toe a little shorter than the third ; subarticular and metatarsal tubercles feebly developed. Tibio-tarsal articulation not quite reaching the tympanum. Skin smooth above and beneath ; a supratympanic fold from the eye to the fore-limb. 1 The

dimensions of the tympanum given in the original description are erroneous.

MONOGRAPH OF T m MICROHYLIDAE Olive brown above, irregularly marbled with darker; limbs with regular dark cross-bars. Lower surfaces uniformly white. Length : 29 mm. DIS~IBU-~ION : East Betsileo, Madagascar. SPECIMEN EXAMINED East Betsileo

. . . . . . .

B.M. 82.3.16.40

Cowan, TYPE

3, Platypelis tuberculata (Ahl) Cophyla tuberculata, Ahl, 1929, Mitt. Zool. Mus. Berlin, XIV, p. 483 (Type locality " N.W. Madagas-

car Head depressed, broader than long ; snout rounded, scarcely prominent, as long as the eye ; canthus rostralis rounded ; loreal region oblique ; nostril a little nearer the tip of the snout than the eye ; interorbital space as broad as the upper eyelid ; tympanum hidden. Fingers with large discs. Toes with a rudiment of web and smaller dilatations than the fingers ; third toe longer than the fifth ; metatarsal tubercle elongate, indistinct. Tibio-tarsal articulation reaching the shoulder or the ear. Skin of the dorsal surfaces with moderately large distinct warts, which are largest and most numerous on the head and anterior part of the back. Throat and chest smooth ; belly and thighs granular. A supratympanic fold from the eye to the shoulder. Grey above, with five broad, dark-brown, irregularly notched cross-bars, one on the snout, another between the eyes, the third on the scapular region, fourth on the sacrum, and the last just in front of the hind-limbs ; the warts whitish; limbs distinctly cross-banded ; sides of the head grey, with three broad brown stripes radiating from the eye. White beneath ; the throat and breast infuscate. Length : 26 mm. DISTRIB~ION: N.W. Madagascar. 'I).

Mus. Berlin

30645

SPECIMEN EXAMINED N.W. Madagascar

2 Ads.

. . . Hildebrandt, COTYPES

4 . Platypelis pollicaris, Boulenger Platypelis pollk7i8, Boulenger, 1888, Ann. Mag. Nat. Hist., (6), I, p. 106, PI. VI, fig. 3 (Type locality "Madagascar ") ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 74 ; Nieden, 1926, Das Tierreich, Anura II, p. 97 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxw, p. 12.

Head depressed, a little broader than long ; snout rounded, not prominent, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region oblique ; nostril nearer the end of the snout than the eye ; interorbital space a little broader than the upper eyelid ; tympanum scarcely distinct, about 3 the diameter of the eye. Fingers with large, subtriangular dilatations, except the first, which has a small rounded disc ; a very large, prominent, oval prepollex. Toes short, almost ) webbed, with large distal dilatations ; fifth toe longer than the third and with the largest disc ; subarticular tubercles feebly developed ; metatarsal tubercle oval, indistinct. Tibiotarsal articulation reaching to between the tympanum and the eye. Skin smooth above and beneath, except the hinder part of the belly, which is slightly granular ; an ill-defined supratympanic fold. Brownish above, with lighter and darker marblings which may form alight X-shaped figure on the anterior two-thirds of the body; supratympanic fold darker; limbs cross-barrkd ; thighs and lower surfaces uniform pale brown or whitish. Length : 28 mm. Male with a vocal sac. DISTRIBUTION : Madagascar.

COPHYLA B.M. 87.12.22.51 96.12.2.32-33

SPECIMENS EXAMINED Madagascar . . . . Ambohimitombo Forest

8 5 and Hgr.

COPHYLA,

41

. . . .

Baron, TYPE Major

Boettger

Cophyla, Boettger, 1880, Zool. Anz., p. 281 (Type species Cophylrc phylhlactyla, Boettger) ; Boettger, 1881, Abh. Senck. Natf. Ges., xn,p. 516; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 475 ; Nieden, 1926, Das Tierreich, Anura n, p, 98 ; Noble and Parker, 1926, Amer. Mus. Novitates, C C X X X I I , p. 15 ; Noble, 1931, Biol Amph., p. 631. Maxillary teeth present ; prevomer divided, the post-choanal portion small, fused with its fellow and bearing a small patch of teeth (fig. 14). Clavicle absent ; procoracoid

reduced to a small plate lying on the anterior edge of the coracoid mesially, not reaching the scapula (fig. 15); sternum large, cartilaginous ; omosternum present. Vertebral column procoelous. Terminal phalanges Y-shaped. Pupil horizontal. Tongue large, oval, entire and half free behind. Palate without dermal folds. Tips of digits broadly dilated. Toes webbed.

Pig. 14. Prevomer and palatines of Cophyla phyllodactyla

Fig. 15. Ventral elements of the shoulder-girdleof Cophyla phyllodactyla (ventral view)

I.

Cophyla phyllodactyla, Boettger

Cophyla phyllodactylrc, Boettger, 1880, Zool. Anz., p. 281 (Type locality Nossi BB) ; Boettger, 1881, Abh. Senck. Natf. Ges., xn, p. 516, P1. v, fig. 19 ; Boulenger, 1882, Cat. Batr. Sal. Brit. MuR., ed. 2, pp. 183 and 475 ; Boettger, 1892, Kat. Batr. Samm. Senckenberg., p. 26 ;Mocquard, 1896, Bull. Soc. Philom. Paris, (8), w, p. 132 ;Mocquard, 1909, Nouv. Arch. Mus., Paris, ( 5 ) , I, p. 74 ; Nieden, 1926, Das Tierreich, Anura 11, p. 98 ; Noble and Parker, 1926, Amer. Mus. Novitates, ooxxxn, p. 15, fig. 9.

MONOGRAPH O F T H E MICROHYLIDAE Head a little broader than long; snout truncate, not prominent, very slightly longer than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; nostril much nearer the tip of the snout than the eye ; interorbital space once and a half as broad as the upper eyelid ; tympanum rather indistinct, about 8 the diameter of the eye. Fingers with large truncate dilatations with a distal notch and a slight longitudinal median furrow ; first finger very short, with a smaller disc ; second and fourth fingers subequal ; a very prominent, oval, inner metacarpal tubercle. Toes nearly & webbed, with discs similar to those of the fingers ; outer toe a little longer than the third and with a larger disc ; subarticular and inner metatarsal tubercles indistinct ; no outer metatarsal tubercle. Tibio-tarsal articulation reaching the eye. Skin smooth above ; belly and hinder side of thighs granular ; a supratympanic fold from the eye to the shoulder. Greyish brown above, uniform or with broad, indefinite A- or M-shaped cross-bars on the head and back. White or flesh-coloured beneath, with some sharply defined sulphur-coloured dots in the groin, on the femur, the edge of the lower jaw and some of the belly granules. Limbs more or less distinctly barred with darker. Length : 30 mm. Male with a subgular vocal sac. DISTRIBUTION : Nossi BC and Madagascar (Mt. Ambr6). B.M. 86.2.25.29

8

Nossi BC

. . . . .

.

. . (Mus. Senckenberg.)

PLETHODONTOH YLA, Boulenger Callula (non Gray), Giinther, 1877, Ann. Mag. Nat. Hist., (4), X I X p. , 316. Calohyla (non Peters), Boettger, 1881, Abh. Senck. Natf. Ges., xn, p. 539. Cophyla (non Boettger), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 182. Plethodontohyla, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 473 (Type species Callula notoaticta, Giinther); Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 72 ; Nieden, 1926, Das Tierreich, Anura n, p. 90 (in part) ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 12 ; Noble, 1932, Biol. Amph., p. 530. Phrynocara, Peters, 1883, Sitzber. Ak. Wiss. Berlin, VI, p. 166 (Type species Phrynocara tuberdum, Peters) ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 72 ; Boettger, 1913, in Voeltzkow, Reise in 0.Afrika, rn,p. 282 ; meden, 1926, Das Tierreich, Anura n, p. 95. Dyacaphus (non Grandidier), Moquard, 1901, Bull. Mus., Paris, VII, p. 254.

Maxillary teeth present ; prevomer divided, the post-choanal portion long, overlying the palatine, in contact with its fellow mesially and bearing a long row of teeth (fig. 16). Clavicles absent ; procoracoid present, narrow, not resting on the coracoid, reaching the scapula and the mid-line of the girdle ; sternum and omosternum resent, cartilaginous (fig. 11). Vertebral column procoelous. Terminal phalanges $shaped or simple.

Fig. 16. Prevomers of Plethodontohyla notosticta

Pupil horizontal. Tongue large, oval, entire and half free behind. Palate without dermal folds. Digits with or without terminal dilatations. The first finger is much shorter than the second, the fifth toe shorter than the third and the outer metatarsal tubercle wanting.

PLETHODONTOHYLA

Fig. 17. Ventral elements of the shoulder-girdle of Pldhodanlohyb mlosticta (ventral view)

SYNOPSIS OF THE SPECIES I. Second finger distinctly shorter than the fourth ; digits with small, but moderately distinct, dilatations. A. A lateral fold extending almost to the moin ; no occipital fold P. notoslicta, p. 43 B. No lateral fold behind the shoulder ; a transverse occipital fold. (1) or the eve : a . , Tibio-tarsal articulation reaching the tmvanum large black inguinal ocellus .- . . . . . . . P . o'cellnta, p. 44 (2) Tibio-tarsal articulation not reaching beyond the tympanum ; no inguinal ocellus . . . . . . . . . . P. alluaudi, p. 45 11. Second finger not distinctly shorter than the fourth ; no digital dilatations. A. Nostril equidistant from the eye and the end of the snout . P . hevipes, p. 45 B. Nostril nearer the tip of the snout than the eye. (1) Snout once and a half as long as the eye ; skin warty above P. luberata, p. 46 ( 2 ) Snout as long as the eye ; skin smooth above . . P . laevis, p. 46

-

-

.

I.

Plethodontohyla notosticta (Giinther)

Callula mtosticta, Giinther, 1877, Ann. Mag. Nat. Hist., (4). XIX, p. 316, P1. XVI, fig. c (Type localities Mahanoro and Anzahamara). Calohyh mtosticta, Boettger, 1881, Abh. Senck. Natf. Ges., XII, p. 639. Cophyh mtostidu, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 182. Pletlwdontohyh mtosticta, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 473 ;Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 72 ; Nieden, 1926, Das Tierreich, Anura n, p. 91 ;Noblo and Parker, 1926, Amer. Mus. Novitates, ccxxxrr,p. 14, fig. 7.

Head a little broader than long ; snout acuminate, scarcely prominent, once and a quarter as long as the eye ; canthus rostralis angular ; loreal region slightly oblique ; nostril a little nearer the tip of the snout than the eye ; interorbital space considerably

MONOGRAPH O F T H E MICROHYLIDAE broader than the upper eyelid ; tympanum moderately distinct, to g the diameter of the eye. Fingers with distinct subtriangular dilatations ; second finger shorter than the fourth; a prominent semi-circular prepollex. Toes free, or with a faint rudiment of web, with small dilatations ; subarticular tubercles rather indistinct; metatarsal tubercle oval, indefinite. Tibio-tarsal articulation reaching to between the tympanum and the anterior comer of the eye. Skin smooth above and beneath; a narrow fold from the posterior corner of the eye, above the tympanum, almost to the groin. Pinkish brown above, with darker lines which may form a fairly regular reticulum or irregular chevron-shaped cross-bars; frequently a r-shaped marking on the anterior part of the back, the cross-bar connecting the upper eyelids ; canthus rostralis, supraciliary edge and lateral fold pink, or white, edged beneath with darker ; a pair of small, white, black-bordered sacral spots and, sometimes, a few similar but smaller spots scattered on the dorsum. Limbs cross-barred ; tarsus and foot blackish beneath. Lower surfaces white, the chin and chest brown-speckled in females, infuscate in males. Length : 4 0 mrn. Male with a vocal sac. DISTRIBUTION : Madagascar. B.M. 76.10.3.13 77.2.8.13.15 89.8.1.2630 1925.7.2.95

2.

SPECIMENS EXAMINED Mahanoro Anzahamara . .

6 9 and Hgr.

{igf'$) 9

. . . . . . Madagascar . . . . Antsihanaka. . . .

.

. . .

(Geale) (Geale)

)

cOTypEs

Majastre (Rosenberg)

Plethodmtohyla ocellata, Noble and Parker

Pldhodoniohyb ocelhta, Noble and Parker, 1926, Amer. Mus. Novitates, CCXXXII, p. 14 (Type locality Antsihanaka) ;Angel, 1931, Faune Col. Franp, N, 5, p. 512, fig. 4, PI. rx, fig. 5. Mantipa ocellutus, Noble, 1931, Biol. Amph., fig. 59.

Head much broader than long; snout vertically truncate, once to once and a quarter as long as the eye ; canthus rostralis obtusely angular ; loreal region moderately oblique ; nostril equidistant from the eye and the end of the snout ; interorbital space considerably broader than the upper eyelid ; tympanum hidden or indistinct, & to 3 the diameter of the eye. Fingers very slightly dilated distally, the second shorter than the fourth ; a prominent, oval, inner metacarpal tubercle. Toes free, very slightly dilated distally ; subarticular tubercles indistinct ; metatarsal tubercle oval, indefinite. Tibio-tarsal articulation reaching the tympanum or the eye. Skin smooth or shagreened above; a supratympanic fold from the eye to the shoulder and an occipital fold connecting the posterior comers of the upper eyelids. Lower surfaces smooth. Grey, magenta or brownish above, with darker markings in the form of irregular, wavy bands which may be chevron-shaped, with the apices directly forwards, or be confluent so as to leave only uniformly scattered light spots of the ground colour ; upper lip, loreal region and temples black ; a large, black, white-bordered inguinal ocellus ; sometimes a curved, light, black-bordered streak from above the tympanic region to the sacrum. Limbs light spotted, the spots transversely elongate on the femur ; an indefinite white stripe along the hinder side of each thigh ; under surfaces of tarsus and metatarsus black. Lower surfaces dirty white, more or less profusely mottled with brown (particularly on the chin and chest) and dotted with tiny white specks. Length : 55 mm. Male with a vocal sac and a diffuse nuptial pad on the outer side of the first finger and metacarpal tubercle ; a small patch a t the base of the second finger. : Central and N.E. Madagascar. DISTRIBUTION

PLETHODONTOHYLA B.M. 1925.11.10.1 92.1.21.25-27 92.3.7.51-52 1925.7.2.81-82 1925.11.10.7 1928.5.9.18-19

9 9 and Yg. 9 and Hgr. Hgr. and Yg. 9 6

$?

Antsihanaka . Madagascar . Sahambendrana Antsihanaka . Antsihanaka . Brickaville .

. (Rosenberg) TYPE . Roucard . Majastre I

. (Rosenberg) I . (Rosenberg)

. (Rosenberg) !

PARATypEs

3. Plethodontohyla alluaudi (Mocquard) Dyscophus alluaudi, Moequard, 1901, Bull. Mus., Paris, vrr, p. 254 (Type locality Fort Dauphin) ; Mocquard, 1902, Bull. Soc. Philom. Paris, (9), y, p. 22, PI. n , fig. 3 ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 71 ; Nieden, 1926, Das Tierreich, Anura u, p. 88. Plelhodontohylo alluaudi, Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 15.

Head depressed, much broader than long ; snout short, truncate, about as long as the eye ; canthus rostralis angular, curved ; loreal region oblique ; nostril a little nearer the tip of the snout than the eye ; interorbital space once and three-fourths the breadth of the upper eyelid ; tympanum indistinct. Fingers with small terminal dilatationsfthe second a little shorter than the fourth. Toes short, free, with small dilatations ; subarticular tubercles feebly developed ; a flat, oval metatarsal tubercle. Tibio-tarsal articulation reaching to between the shoulder and the tympanum. Skin smooth or finely granular above ; a supratympanic fold from the eye to the shoulder and an occipital fold connecting the posterior corners of the upper eyelids. Lower surfaces smooth. Greyish or brownish above, with darker markings ; loreal region and temples dark brown ; head in front of the occipital fold slightly darker than the dorsum. An indefinite dusky mid-dorsal marking commencing between the eyes, sending a pair of oblique branches from the scapular region to the flanks and another pair from the sacral region to the groin (the hinder part of this marking sometimes indistinct) ; two black bars, or a round dark spot in each groin ; limbs with indistinct oblique cross-bars ; hinder side of the thighs brown, with lighter spots ; belly, chin and throat profusely mottled and stippled with brown, or pale brown with light dots. Length : 46 mm. DISTRIBWION : Fort Dauphin, S. Madagascar.

B.M. 1932.10.2.89 Mus. Vienna

9 Hgr.

Fort Dauphin . FortDauphin.

. . . . . . . .

. . . . . . . .

Sykora Sykora

4 . Plethodontohyla brevipes, Boulenger Plethodontohyla b~evipea,Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 474 (Type locality East Betsileo) ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), 1 p. 72 ; Nieden, 1926, Das Tierreich, Anura 11, p. 91 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 14.

Head broader than long; snout rounded, not prominent, a little longer than the diameter of the eye ; canthus rostralis obtusely angular ; loreal region moderately oblique ; nostril equidistant from the eye and the end of the snout ; interorbital space considerably broader than the upper eyelid ; tympanum scarcely distinct, about 3 the diameter of the eye. Fingers short, not dilated, the second and fourth subequal. Toes free, not dilated ; subarticular tubercles indistinct ; metatarsal tubercle elongate, indistinct. Tibio-tarsal articulation reaching the posterior border of the tympanum. Skin shagreened above ; a supratympanic fold from the eye to the shoulder and an occipital fold connecting the posterior corners of the upper eyelids. Lower surfaces smooth.

MONOGRAPH O F T H E MICROHYLIDAE Uniform dark brown above ; white beneath, with some brown stippling ; throat washed with brown. Length : 3 4 mrn. DISTRIBUTION : East Betsileo, Madagascar. SPECIMEN EXAMINED 9 East Betsileo . , Cowan, TYPE B.M. 82.3.16.31

. .

. . . .

5. Plethodontohyla tuberata (Peters) Phrynocasa tubesaturn, Peters, 1883, Sitzber. Ak. Wiss. Berlin, VI, p. 167, fig. (Type locality "Interior of Madagascar ") ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 73 ; Nieden, 1926, Das Tierreich, Anura n, p. 95, fig. 53. Plethodontohyla tuber&, Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 15, fig. 8. Head broader than long ; snout rounded, not prominent, once and a half as long as the eye ; canthus rostralis obtusely angular ; loreal region oblique ; nostril nearer the tip of the snout than the eye ; interorbital space broader than the upper eyelid ;

tympanum scarcely distinct, about $ the diameter of the eye. Fingers short, not dilated, the second as long as the fourth ; inner metacarpal tubercle rounded, prominent. Toes short, free, not dilated ; subarticular tubercles indistinct ; metatarsal tubercle moderately prominent. Tibio-tarsal articulation reaching the posterior border of the ear. Skin uniformly warty above ; smooth or feebly granular beneath. A transverse occipital fold sometimes present. Brownish above, with darker flecks and spots ; inguinal region and sides with yellow flecks. Brown, with yellow spots beneath, the throat of the male uniformly brown. Limbs yellow-spotted beneath ; tibia and tarsus each with a broad, but not welldefined, dark cross-bar above. Length : 4 2 mm. Male with a vocal sac. : Madagascar. DISTRIBUTION SPECIMENS EXAMINED R.W. 91.9.14.4 9 Madagascar . . . . . . . . Peracca 1932.10.2.87-88 $? Hgr. Madagascar . . . . . . . . Sykora Mus. Vienna 2 99, Hgr. Madagascar . . . . . . . . Sykora

6. Plethodontohyla laevis (Boettger) Phrynoeasa h u e , Boettger, 1913, in Voeltzkow, Reise in 0.Afrika, I=, p. 282, PI. xxm, Bg. 10 (Type locality Sakana, E. Madagascar) ;Nieden, 1926, Daa Tierreich, Anma rr, p. 96. Plethdonioi~yla laeue, Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 15. Head much broader than long ; snout rounded, as long as the eye ; canthus rostralis distinct ; loreal region oblique ; nostril nearer the end of the snout than the eye ; interorbital space once and a half as broad as the upper eyelid ; tympanum hardly

visible, about half the diameter of the eye. Fingers not dilated, the second and fourth subequal ; inner metacarpal tubercle well developed and prominent. Toes free, not dilated ; subarticular tubercles feebly developed ; metatarsal tubercle indistinct. Tibio-tarsal articulation reaching the tympanic region. Skin smooth above, except for a few small warts on the upper surface of the snout and the upper eyelid ; a supratympanic fold from the eye to the shoulder. Smooth beneath ; the hinder side of the thighs slightly granular. Reddish grey above, wit11 dark grey flecks and spots which are more numerous on the flanks ; a large brown, white-bordered X-shaped marking between and behind the eyes, and some similarly coloured, irregular blotches on the anterior part of the back ; supratympanic fold black, narrowly edged above with reddish white. Whitish beneath, spotted and dotted with dark grey, except on the belly and thighs. Limbs

RHOMBOPHRYNE

47

cross-banded ; anal region black, white-dotted ; toes with a white ring round the dark tip. Length : 41 mm. DISTRIBUTION : Sakana, E. Madagascar.

RHOMBOPHR YNE, Boettger Rhombophryne, Boettger, 1880, Zool. Anz., p. 567 (Type species Rhombophryne testudo, Boettger) ; Boettger, 1881, Abh. Senck. Natf. Ges., xn, p. 494 ;Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 177 ; Boulenger, 1884, Ann. Mag. Nat. Hist., (5), XN, p. 388, fig.; Vaaant, 1885, Bull. Soc. Philom. Paris, (7), IX, p. 116, fig. ; Nieden, 1926, Das Tierreich, Anura 11,p. 8 ;Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxIr, p. 17 ;Noble, 1931, Biol. Amph., p. 530. Maxillary teeth absent ; prevomer divided, the post-choanal portion long, in

contact with its fellow mesially, overlying the palatine and bearing a long row of teeth (fig. 18). Clavicle absent ; procoracoid a very thin slip, not resting on the coracoid, extending from the mid-line of the girdle to the scapula ; sternum large, cartilaginous ; ornosternum a cartilaginous rudiment (fig. 19). Vertebral column procoelous. Terminal phalanges T-shaped.

Fig. 18. Prevomers of Rhombophryne testudo

Fig. 19. Ventral elements of the shoulder-girdle of Rhombophryne testudo (ventral view)

MONOGRAPH O F T H E MICROHYLIDAE Pupil horizontal. Tongue large, oval, entire and scarcely free behind, with a median longitudinal groove terminating in a plicate depression posteriorly. A smooth dermal fold across the palate in front of the pharynx, preceded by a rounded dermal papilla. Digits not dilated distally. Toes free. I.

Rhombophryne testudo, Boettger

Rhombophryne testudo, Boettger, 1850, Zool. Anz., p. 567 (Type locality Nossi BQ); Boettger, 1881, Abh. Senck. Natf. Ges., XII, p. 496, PI. IV, fig. 15 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 177; Vaillant, 1885, Bull. Soc. Philom. Paris, (7), IX, p. 116, fig. : Boulenger, 1886, Zool. Rec., 1885, Rept. and Batr., p. 24; Boettger, 1892, Kat. Batr. Samm. Senckenberg., p. 25 ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 69 ; Nieden, 1926, Das Tierreich, Anura n, p. 8 ; Noble and Parker, 1926, Amer. Mus. Novitates, CCXXXII,p. 17 ; Noble. 1931, Biol. Amph., fig. 348. Head twice as broad as long, depressed ; snout obtusely rounded, not prominent, nearly twice as long as the very small eye ; canthus rostralis obtuse ; loreal region very oblique ; nostril a little nearer the eye than the tip of the snout ; interorbital space twice and a half as broad as the upper eyelid; tympanum rather indistinct, vertically oval, its maximum diameter greater than that of the eye. Fingers short, tapering, depressed, with more or less distinct fleshy lateral fringes ; first finger shorter than the second which is subequal to, or a little longer than, the fourth ; inner metacarpal tubercle prominent. Toes short, depressed, spatulate distally ; fifth toe much shorter than the third ; subarticular tubercles indistinct ; two metatarsal tubercles, the inner moderately prominent. Tibio-tarsal articulation reaching t o between the shoulder and the tympanum. Skin of back with numerous fine, short folds arranged in more or less regular longitudinal series ; one long pair constantly present, commencing above the otic region and converging slightly posteriorly ; a supratympanic fold from the eye to the shoulder, continued anteriorly along the canthus rostralis ; tip of snout and edge of lower jaw beset with numerous pustules and short barbels ; limbs pustular. Belly granular or wrinkled. Reddish or brown above, with some dark dots and vermiculations ; the warts and folds sometimes lighter ; loreal and temporal regions darker ; limbs indefinitely cross-banded ; a dark streak along t h e hinder side of the thighs. Belly brown or yellowish, with or without white marblings. Length : 49 rnm. Male with a vocal sac. : Nossi BB and E. Madagascar. DISTRIBUTION

EXAMINED SPECIMENS B.M. 84.11.3. 5-6 86.2.25.30-32 86.2.25. 33 87.12.5. 19-20 95.10.29. 90

d?

99

Skel. hgr.

6

9

NossiBC Nossi BC Nossi BC NossiBC Imerina

. . . . . . . . . . . . . . . . . . . .

. . (Linnaea) . . (Mus. Senckenberg.) . . (Mus. Senckenberg.) . . Deyrolle . .Baron

ANODONTH YLA, Miiller Amdonthyla, Miiller, 1892, Verh. Nat. Ges., Basel, X, p. 198 (Type species A . boulengerii, Miiller) ; Nieden, 1926, Das Tierreich, Anura 11, p. 98. Amdontohyla (emendation) Mocquard, 1909, Nouv. Arch. Mus., Paris, (5),I, p. 74; Noble and Parker, 1926, Amer. Mus. Novitates, coxxxn, p. 18; Noble, 1931, Biol. Amph., p. 531. &fam'ella( m n Boulenger), Boettger, 1913, in Voeltzkow, Reise in 0. Afrika, 111, p. 280. Maxillary teeth present ; prevomer small, confined t o the anterior border of the choana (fig. 20). Clavicle and procoracoid present, well developed, reaching the scapula and the mid-line of the girdle; omosternum and sternum well-developed,

ANODONTHYLA

49

cartilaginous (fig. 21). Vertebral column ?. Terminal phalanges broadly Y- or T-shaped. Pupil horizontal. Tongue pear-shaped, slightly notched behind. Tips of digits dilated. The first finger is very short, much shorter than the second, which is a little shorter than the fourth ; toes free ; outer metatarsal tubercle wanting.

I

I

[AmNoble and Parker

Fig. 20. Prevomers and palatines of Ancdonthyla boulengeri

Fig. 21.

Ventral elements of the shoulder-girdleof Amdonthyla boulengeri (ventral view)

SYNOPSIS OF THE SPECIES I. Nostril equidistant from the eye and the end of the snout ; latter prominent with vertical lores ; tympanum indistinct, half the diameter of the eye A. boulengeri, p. 49 11. Nostril nearer the tip of the snout than the eye ; snout not prominent, with oblique lores ; tympanum distinct, two-thirds the diameter of , . . . . . A. montuna, p. 50 the eye . . , . . .

. .

I.

.

Amdonthyla boulengeri, Miiller

Amdonthyla boulengerii, Miiller, 1892, Verh. Nat. Ges. Besel, x, p. 198, PI. m, fig. 28 (Type lomlity Medagaacar) ; Mocquerd, 1909. Nouv. Arch. Mus., Paris, ( b ) , I, p. 74 ; Methuen and Hewitt, 4

MONOGRAPH OF THE MICROHYLIDAE 1913, Bnn. Tmnsvaal Mus., N, pp. 60 and 61 ; Nieden, 1926, Das Tierreich, Anura 11,p. 98; Noble and Parker, 1926, Amer. Mus. Novitates, C C X X X ~ p. , 18, fig. 11. Mantea0 pUiazn's, Boettger, 1913, in Voeltzkow, Reise in 0.Africa, m , p. 280, PI. x x m , fig. 8 (Type locality Anevoka, E. Madagascar).

Head a little broader than long. Snout acuminate, slightly prominent, as long as the diameter of the eye ; canthus rostralis short, rounded ; loreal region vertical and slightly concave ; nostril equidistant from the eye and the end of the snout ; interorbital space as broad as, or a little broader than, the upper eyelid; tympanum indistinct, about half the diameter of the eye. Fingers slender, the three outer with very large, subtriangular dilatations ; a large, flat, inner metacarpal tubercle. Toes scarcely dilated, the first very short ; subarticular tubercles flat ; metatarsal tubercles flat, sometimes indistinct. Tibio-tarsal articulation reaching the tympanum or the eye. Skin finely granular above and beneath, sometimes with several longitudinal rows of warty ridges between the eyes. Dark brown above, uniform or with a broad, dark, chevron-shaped marking on the back ; the apex of this mark may be white-edged, and lies on the occiput, whilst its posterior ends lie in the dirty-white groins ; thighs with lighter flecks a t their bases and on their hinder sides ; tibia and tarsus each with a broad, dark, oblique cross-bar. Greyish brown or whitish beneath, the chin and chest darker. Length : 22 mm. Ovarian eggs very large, those of a female 21 mm. in length measuring 2 mm. in diameter. DISTRIBUTION : E. Madagascar. 2.

Ansdsnthyla msntana, Angel

Amdonthyla montana, Angel, 1925, Bull. Mus., Paris, p. 62, figs. 1 4 . (Type locality Andringitra Mts., 2,600 metres) ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 18 ; Angel, 1931, Faune Col. Franp., N, 5, p. 544. Head as long as broad. Snout rounded, not prominent ; canthus rostralis indistinct ; loreal region oblique, slightly concave ; nostril nearer the tip of the snout than the eye ; interorbital space once and a half (juveniles) to twice (adults) as broad as the upper eyelid ; tympanum visible, 3 the diameter of the eye. Fingers with large subtriangular dilatations. Toes scarcely dilated, the third and fifth subequal ; subarticular tubercles feebly developed ; metatarsal tubercle large and flat. Tarso-

metatarsal articulation reaching the posterior corner of the eye or, in juveniles, a little farther. Skin smooth above; a slight supratympanic fold from the eye to tJle shoulder sometimes present ; smooth or faintly granular beneath, and behind the thighs. Yellowish brown above, with indistinct darker spots, or blackish with grey-green spots and vermiculations ; limbs with dark cross-bars. Yellowish or greenish white beneath, immaculate or dotted and spotted with brown. Length : 38 mm. DISTRIBUTION : Andringitra Mts., Madagascar.

STUMPFFIA, Boettger Stumpjia, Boettger, 1881, Zool. Anz., rv, p. 360 (Type species 8. psologlossa, Boettger) ; Boettger, 1881, Abh. Senck. Natf. Ges. xn,p. 521 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 146 ; Noble and Parker, 1926, Amer. Mus. Novitates, ccxxxn, p. 4, footnote.

Maxillary teeth absent ; prevomer divided, the post-choanal portion in contact with its fellow mesially, ankylosed to the palatine, edentulous (fig. 22). Clavicle and procoracoid present, slender, reaching the scapula and the mid-line of the girdle (fig. 23) ; no omosternurn ; sternum cartilaginous. Vertebral column procoelous. Terminal phalanges T-shaped.

-

STUMPFFIA

51

Pupil horizontal. Tongue oval, rounded or pointed behind, with a median transverse groove which divides it into two parts. No pharyngeal folds. Digits not, or but slightly, dilated.

Pig. 22. Anterior cranial elements of Stumpfia paobgloaaa (seen from beneath)

Fig. 23. Ventral elements of the shoulder-girdle of Stumpfia paobglossa (ventral view) I.

Stump$a psologlossa, Boettger

8tumpfla psologlossa, Boettger, 1881, 2001. Anz., N,p. 360 (Type locality Nossi BB) ; idem, 1881, Abh. Senck. Natf. Ges., xn,p. 522, P1. V, fig. 21 ; idem, 1892, Kat. Batr. Samm. Senckenberg., p. 21 ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 67 ; Boettger, 1913, in Voeltzkow, Reise in 0.m a , 111, p. 327. Head a little broader than long ; snout blunt, moderately prominent, once and a quarter as long as the eye ; canthus rostralis rounded ; loreal region almost vertical ; nostril a little nearer the tip of the snout than the eye ; interorbital space nearly twice as broad as the upper eyelid ; tympanum indistinct, about the diameter of the eye. Fingers free, the third relatively long and slender, with a slight terminal swelling; the first very short ; second subequal to, or shorter than, the fourth. Toes free, the tips feebly dilated ; first very short ; fifth much shorter than the thud ; subarticular tubercles indistinct ; a rudimentary inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the eye. Skin smooth above ; belly and hinder side of the thighs granular.

+

52

MONOGRAPH O F T H E MICROHYLIDAE

Pale brown to grey above, uniform or with dark dots and blotches which may form four longitudinal series, two on the back and one on each flank ; a dark interorbital bar ; limbs faintly cross-barred ; lips with dark vertical bars. Grey beneath, clouded or mottled with darker and lighter. Length : 16 mm. : Islands of Nossi BC and Sakatia. DISTRIBUTION B.M. 1930.11.18. 1-2 2.

SPECIMENS EXAMINED Nossi BC . . .

. . . . .

Voeltzkow

Stumpfia madagascariensis, Mocquard

Stumpfla rnadqascariensis, Mocquard, 1895, Bull. Soc. Philom. Paris, (a),vn, p. 132 (Type locality Mt. Ambre) ; Mocquard, 1909, Nouv. Arch. Mus., Paris, (5), I, p. 67 ; Angel, 1931, Faune Col. Franp., IV,p. 540.

Resembling S. psologlossa, but with the inner fmgers and toes very short, resembling tubercles rather than digits. Reddish brown above, darker on the back, with an ash-grey lateral stripe from the eye to the groin. Belly dotted with white. Length : 9.5 mm. : Mt. Ambre and Arnbongo, Madagascar. DISTRIBUTION

A S T E R O P H R Y I N A E , Giinther 1

Asterophrydidae, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 51. Xenorhinidae, Mivart, 1869, Proc. Zool. Soc. London, p. 289. Genyophrpidae, Boulenger, 1890, Proc. Zool. Soc. London, p. 326. Symphygnathina~and Eleutherognathinae (part),MBhely, 1901, TermBs. Fiizetek, XXIV,pp. 172-189. Genyophryninae, Gadow, 1901, Cambridge Nat. Hist., VIII, p. 236. Cacopinae (part), Noble, 1931, Biol. Amph., p. 531. Maxillary and vomerine teeth absent ; maxillae often overlapping the premaxillae

and usually in contact anteriorly; prevomer Large, entire, almost surrounding the choana, with a mesial, wing-like expansion which is in contact with its fellow ; postchoanal portion of the vomer frequently with odontoids ; ethmoid divided ; ear fully developed ; vertebral column diplasiocoelous (rarely procoelous). No free-swimming tadpole, metamorphosis taking place within the egg ; the larva lacks external gills, has no open gill clefts, internal gills, operculum or spiracle ; respiration is performed through a non-muscular, vascular tail. The entire vomer, with post-choanal portion and mesial expansion, absence of teeth, maxillary condition and diplasiocoelous vertebral column are diagnostic of this group of Papuan genera. SYNOPSIS OF TKE GENERA I. Maxillae not meeting in front of the pre-maxillae. A. Procoracoid present . . . . . . . . . . . Genyophryne, p. 52 B. Procoracoid absent . . . . . . . . . . . Metopostira, p. 6 8 11. Maxillae suturally united in front of the premaxillae. A. One or two very large, steeple-shaped vomerine odontoids on each side Xenobatmchus, p. 5 4 B. Vomerine odontoids, if present, small, rounded . . . Astemphrys, p. 5 8

GENYOPHR YNE, Boulenger Qenyophryne, Boulenger, 1890, Proc. Zool. Soc. London, p. 326 (Type species Qenyophryne thomoni, Boulenger) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 159 ; Nieden, 1926, Das Tierreich, Anura n, p. 99 ; Noble, 1931, Biol. Amph., p. 534.

GENYOPHRYNE

53

Maxillae overlapping the premaxillae anteriorly, but not in contact; prevomer undivided in contact with its fellow mesially, the post-choanal portion bearing a group of small, conical odontoids (fig. 24) ; edge of the dentary slightly crenulate ; roof of the skull involved in secondary ossification with the derm. Clavicles absent ; procoracoids present, reaching the mid-line of the girdle and the scapulae (fig. 25) ; no ornosternum ; sternum large, cartilaginous. Vertebral column procoelous. Terminal phalanges T-shaped.

Fig. 24. Anterior cranial elements of Benyophryne thomeoni (seen from beneath)

Fig. 25. Ventral elements of the shoulder-girdle of Oenyophryne thomoni (ventral view)

Pupil horizontal. Tongue large, oval, entire, scarcely free behind and with a small median pouch posteriorly. A denticulated dermal ridge in front of the pharynx, preceded by a shorter, curved, crenulate one. Digits free; toes with very small, scarcely distinct, terminal dilatations. This is the only procoelous genus included in this subfamily ; but its other characters all point to a closer relationship with Aste~ophrys,Metoposti~a,etc., than with any member of the Sphenophryninae.

MONOGRAPH O F THE MICROHYLIDAE I.

Genyophrym thomsoni, Boulenger

Genyophryne thomsoni, Boulenger, 1890, Proc. Zool. Soc. London, p. 327, P1. xxv, fig. 1 (Type locality Sud Eat Island) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 159 ; Nieden, 1926, Das Tierreich, Anura 11, p. 100; Parker, 1931, Archiv. Zool. Ital., XVI,p. 1243, fig. 1 ; Burt, 1932, Bull. Am. Mus. Nat. Hist. LXIU, p. 483.

Head broad and much depressed ; snout scarcely prominent, once and two-thirds as long as the eye ; canthus rostralis indistinct ; loreal region very oblique, concave ; interorbital space more than three times as broad as the upper eyelid ; tympanum indistinct, half the diameter of the eye, or hidden. Fingers very short, the first shorter than the second, which is a little shorter than the fourth ; third toe longer than the fifth ; no subarticular tubercles ; inner metatarsal tubercle indistinct, the outer absent. Tibio-tarsal articulation reaching the tympanic region. Skin shagreened above, smooth beneath. A pair of dermal folds along the back, commencing at the upper eyelids, converging about the middle of the back, diverging in the direction of the groins and then becoming indistinct ; a supratympanic fold from the posterior corner of the eye ; a small tubercle on the heel. Pale yellowish brown above, irregularly spotted and stippled with dark brown, the canthus rostralis and dennal folds usually being picked out in this colour ; hinder side of the thighs below the vent, and lower surfaces of the tarsi, dark brown ; sometimes dark, grey-brown patches are present a t the insertion of the fore-limb, in the groin and behind the knee. Lower surfaces white, the thighs and posterior part of the belly sometimes with fine, dark brown vermiculations. Length : 35 mm. Male with a vocal sac. DI~TRIBUTION : Sud Est Island, Ferguson Island and British New Guinea (Albert Edward Mts., 6,000 ft.). SPECIMENS EXAMINED B.M. 89.7.1. 1 2 1904.11.1. 44-48 1904.11.1. 5 0 1904.11.1. 7 2

d 33 d Skel. 3

Sud Est Island, New Guinea Albert Edward Mts., 6,000 ft. Albert Edward Mts., 6,000 ft. Ferguson Island, New Guinea

. . . .

Thornson, TYPE Rohu Rohu Meek

XENOBATRACHUS, Peters and Doria Xenobatrachus, Peters and Doria, 1878, Ann. Mus. Civ. Genova, x m , p. 432 ( ~ ~ ~ : s ~ e c iXenoes batrachua ophiodon, Peters and Doria); van Kampen, 1918, Bijdr. Dierk., xm, p. 54; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 126 ; Nieden, 1926, Das Tierreich, Anura u, p. 52 ; Noble, 1931, Biol. Amph., p. 535. Chamantha, Mbhely, 1898, Termbs. Piizetek, XXI, p. 175 (Type species Chanamntha rostrata, Mbhely). Xenorhina (non Peters), Mbhely, 1901, Termbs. Piizetek, xxrv, pp. 183 and 231 (in part).

Maxillae forming a median suture in front of the premaxillae, and dentaries in front of the mento-meckelian bones ; prevomer large, undivided, in contact with its fellow mesially, the post-choanal portion overlying the palatine region and bearing one or two very large steeple-shaped odontoids (fig. 26). Clavicles, procoracoids and omosternum absent ; sternum cartilaginous. Vertebral column diplasiocoelous. Terminal phalanges T-shaped or simple. Pupil horizontal ; tongue large, oval, entire and scarcely free behind, with a median furrow and a small depression posteriorly. A denticulated dermal ridge across the palate in front of the pharynx, sometimes preceded by a shorter smooth one. Digits free, not, or but slightly, dilated terminally. In all the known species the first finger is shorter than the second, the third toe

XENOBATRACHUS

55

longer than the fifth, the subarticular tubercles indistinct and the outer metatarsal tubercle absent.

b

Fig. 26. Anterior cranial elements of Xenobatrachus rostratw (seen from beneath)

SYNOPSIS OF THE SPECIES I. A single steeple-like odontoid behind each choana. A. Snout about twice as long as the diameter of the eye . X. rostratus, p. 55 B. Snout not more than once and a half as long as the eye. (1) Tarso-metatarsal articulation reaching the eye ; white (red in life) inguinal markings . . . . . . X. ocellatus, p. 56 (2) Tarso-metatarsal articulation reaching beyond the tip of the snout ;no light inguinal markings. X . macrops, p. 56 11. A pair of odontoids behind each choana. A. Tarso-metatarsal articulation not reaching beyond the eye ; size . . . . . . . . X . giganteus, p. 57 large (86 mm.) . B. Tarso-metatarsal articulation reaching the tip of the snout o r beyond ; size small (30 rnm.) (1) Eye moderate, the snout but little longer than its diameter; fingers moderately slender ; tibio-tarsal articulation reaching to between the eye and the tip of the snout X. ophiodon, p. 58 (2) Eye small, the snout distinctly longer (13) than its diameter ; fingers short ; tibio-tarsal articulation reaching to between the shoulder and the posterior border of the eye. X. bidens, p. 57

. . . . . .

.

.

.

. .

.

I.

Xenobatrachus rostratus (Mdhely)

Choamcsnthar rostrata, Mbhely, 1898, Term&. Fiizetek, XXI, p. 175, P1. XU (Type locality Erima, Astrolabe Bay, New Guinea). Xenorhina r o s t r h , Mbhely, 1901, Term&. Fiizetek, xxrv, pp. 185 and 233, P1. IX, fig. 10, P1. XI, figs. 1 and 2 ; Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 430 ;van Kampen, 1914, 2001. Jahrb., Syst., xxxw, p. 373; Xenobatrachus rostratus, van Kampon, 1918, Bijdr. Dierk., XXI, p. 54 ;van Kampen, 1923, Amph. Indo-Auatr. Arch., p. 127 ; Nieden, 1926, Das Tierreich, Anura II, p. 53, figs. 37 and 38 ;Burt, 1932, Bull. Amer. Mus. Nat. Hiat., Lxm, p. 485. Choamantha mdhhelyi, Boulenger, 1898, Ann. Mus. Civ. Genova, (2), x m , p. 709, P1. w, fig. 6. (Type locality Vikaiku).

A single tooth-like prominence behind each choana ; sometimes a tubercle-like anterior palatal ridge. Snout pointed, distinctly prominent, nearly twice as long as

MONOGRAPH O F T H E MICROHYLIDAE the diameter of the eye and pustulous on its lower, anterior surface ; canthus rostralis rounded ; loreal region oblique, concave ; interorbital space nearly three times as wide as the upper eyelid ; tympanum indistinct or hidden, about as large as the eye. Fingers short, not dilated, the second a little shorter than the fourth. Toes with very small discs, the third much longer than the fifth ; inner metatarsal tubercle indistinct. Tibio-tarsal articulation reaching the shoulder or the tympanum in adults, the loreal region in juveniles. Skin with a few scattered warts above ; smooth beneath ; a feeble supratympanic fold. Brown to dark grey above, sometimes with dark, light-centred spots in longitudinal rows ; lower surfaces whitish (red in life), more or less profusely marbled and spotted with brown ; hinder side of thighs dark brown ; a fine white line from snout to vent, intersected above the vent by another line which traverses the posterior side of the hind-limb to the foot ; a similar line occurs on the lower side of the forearm. Female with large eggs (ovarian 3.5 mm. in diameter). Length : 47 mm. : New Guinea. DISTRIBUTION Mus. Mus. Mus. B.M.

2 Ads. Ad.

Hung. 241416 Amsterdam Amsterdam 1921.11.11. 5

.

2.

,

d

9

.

.

Erima . . . Sermowai River . Hellwig Mts., 2,200m. Weyland Range

. . .

. . . . . . . . . . . .

Biro, COTYPES Pratt

Xenobatrachus ocellatus (van Kampen)

Xenorhim ocellata, van Kampen, 1913, Nova Guinea, IX, 3, p. 461, PI. XI, figs. P 5 (Type locality Hellwig Mts., 2,500 m.). Xenobatrachus ocellatus, van Kampen, 1918, Bijdr. Dierk., XXI,p. 54 ; idem, 1923, Amph. Indo-Austr. Arch., p. 128 ;Nieden, 1926, Das Tierreich, Anura n, p. 54.

A single tooth-like prominence behind each choana; a short, curved, anterior palatal ridge. Snout very slightly prominent, a little longer than the diameter of the eye, pustulous on its lower anterior surface ; canthus rostralis rounded ; loreal region oblique, concave ; interorbital space nearly twice the breadth of the upper eyelid ; tympanum distinct, nearly as large as the eye. Fingers not dilated, the second slightly shorter than the fourth. Toes not dilated ; inner metatarsal tubercle indefinite. Tibio-tarsal articulation reaching the shoulder. Skin smooth above and below ; a strong, curved, supratympanic fold. Brown above and beneath ; a large, white, brown-bordered ocellus in the groin ; hinder side of the thighs white-spotted. Female with large eggs (ovarian 5 mm. in diameter). Length : 42 mm. : Hellwig Mts., circa 2,500 m., Dutch New Guinea. DISTRIBUTION Mus. Amsterdam

2

99

SPECIMENS EXAMINED Hellwig Mts., c. 2,500 m.

. . . . .

COTYPES

3. Xenobatrachus macrops (van Kampen) Xenorhim rostrata (non Mhhely), van Kampen, 1909, Nova Guinea, IX, i, p. 39, P1. n, fig. 6. Xenorhina macrops, van Kampen, 1913, Nova Guinea, IX, 3, p. 460 (Type locality Hellwig Mts., c. 2,500 m.). Xenobatrachus nucerops, van Kampen, 1918, Bijdr. Dierk., XXI, p. 54 ; idem, 1923, Amph. Indo-Austr. Arch., p. 128 ;Nieden, 1926, Das Tierreich, Anura n, p. 53. A single, tooth-like prominence behind each 'choana ; a tubercle-like, anterior

palatal ridge. Snout pointed, distinctly prominent, slightly longer than the diameter of the eye, pustulous above and below ; canthus rostralis rounded ; loreal region oblique, slightly concave ; interorbital space twice the breadth of the upper eyelid ; tympanum not, or scarcely, distinct, ncarly as large as the eye. Fingers with scarcely

XENOBATRACHUS

57

perceptible discs, the second a little shorter than the fourth. Toes with small, but distinct discs ; inner metatarsal tubercle indefinite. Tibio-tarsal articulation reaching the tympanum or the posterior border of the eye. Skin smooth above and beneath ; a supratympanic, and sometimes an occipital, fold. Brown above ; lighter beneath and on the upper lip, with white rnarblings ; a h e white line along the hinder side of the tibia and tarsus. Female with large eggs (ovarian 3.5 mm. in diameter). Length : 53 mm. DISTRIBUTION : Dutch New Guinea. Mus. Amsterdam

2

99

SPECIMENS EXAMINED Hellwig Mts., c. 2,500m.

. . . . .

COTYPES

4 . Xenobatrachus giganteus (van Kampen) X e m h i n a gigantea, van Kampen, 1915, Nova Guinea, XIII, i, p. 40 (Type locality " Bijenkorfbiwak," near Lorentz River). Xenobatrachus giganteus, van Kampen, 1918, Bijdr. Dierk., XXI,p. 54 ; idem, 1923, Amph. Indo-Austr. Arch., p. 130, fig. 16 ;Nieden, 1926, Das Tierreich, Anura n, p. 54. Two tooth-like prominences behind each choana ; a tubercle-like anterior palatal

ridge. Snout slightly prominent, a little longer than the diameter of the eye, pustulous on its lower anterior surface ; canthus rostralis rounded ; loreal region oblique, slightly concave ; interorbital space more than twice the breadth of the upper eyelid ; tympanum moderately distinct, about as large as the eye. Fingers with scarcely perceptible discs, the second slightly shorter than the fourth. Toes with small discs, the fifth much shorter than the third; inner metatarsal tubercle feebly developed. Tarso-metatarsal articulation reaching the shoulder ( 8 ) or the temporal region ( 9). Skin thick, glandular and finely shagreened ; a curved supratympanic fold from the eye to the shoulder. Brownish purple above ; yellow, marbled and dusted with brown beneath. Ovarian eggs large (6 rnm.) ; male with a vocal sac. Length : 86 mm. DISTRIBUTION : Snow Mountains, 1,700 m., Dutch New Guinea. Mus. Amsterdam

8,$2

SPECIMENS EXAMINED Near Lorentz River, 1,700 m.

. . . .

COTYPES

5 . Xenobatrachus bidens (van Kampen) X e w h i n a bidena, van Kampen, 1909, Nova Guinea, Ix, i, p. 39 (Type locality Digul River) ;Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 430; van Kampen, 1913, Nova Guinea, IX, 3, p. 460. Xenobat~achusbidena, van Kampen, 1918, Bijdr. Dierk., XXI, p. 54 ; idem, 1923, Amph. IndoAustr. Arch., p. 129 ; Nieden, 1926, Das Tierreich, Anura n, p. 54. Two tooth-like prominences behind each choana ; no anterior palatal ridge. Snout pointed, moderately prominent, once and a half as long as the eye ; canthus rostralis rounded ; loreal region very oblique ; interorbital space twice as broad as the upper eyelid ; tympanum scarcely distinct, about as large as the eye. Fingers not dilated distally, the second and fourth subequal. Toes slightly dilated terminally ; inner

metatarsal tubercle elongate, indistinct. Tibio-tarsal articulation reaching the tympanum. Skin smooth or slightly warty above ; smooth beneath ; supratympanic fold absent or very indistinct. Pale brown above, with or without a few darker spots on the back and limbs; white beneath, uniform or faintly dusted with brown. Ovarian eggs large (3-5mm.). Length : 29 mm. DI~TRIBUTION : Dutch New Guinea.

MONOGRAPH OF THE MICROHYLIDAE

9

Mus. Amsterdam Mus. Amsterdam

Ad.

SPECIMENS EXAMINED Dim@ River . . . . Bivak Isld., Lorentz River

. . . . . . TYPE . . . . . .

6. Xenobatrachus ophiodon, Peters and Doria Xenobatrachus ophiodon, Peters and Doria, 1878, Ann. Mus. Civ. Genova, xm, p. 432, P1. VII, fig. 5 (Type locality Hatam, Arfak Mts.) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 172 ; Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 432 ;van Kampen, 1913, Amph. Indo-Austr. Arch., p. 131 ; Nieden, 1926, Das Tierreich, Anura u, p. 55. Two tooth-like prominences behind each choana ; no anterior palatal ridge. Snout

slightly prominent, very little longer than the diameter of the eye and slightly pustulous anteriorly ; canthus rostralis rounded ; loreal region very oblique, concave ; interorbital space broader than the upper eyelid ; tympanum rather indistinct, about the diameter of the eye. Fingers with very small discs. the second a little shorter than the fourth. Toes with slightly larger discs than the fmgers ; inner metatarsal tubercle indistinct. Tibio-tarsal articulation reaching the loreal region. Skin smooth above and below. Dark brown above, irregularly spotted and marbled with black ; white beneath, the throat and chest mottled with brown. Length : 30 rnrn. DISTRIBUTION : Arfak Mts., Dutch New Guinea.

+

Mus. Genova

Ad.

Hatarn, Arfak Mts.

. . . . .

.Beccari, COTYPE

ASTEROPHR Y S , Tschudi Asterophrys, Tschudi, 1838, Mem. Soc. Sci. Neuchbtel, u, p. 82 (Type species Ceratophrys turpicola, Miiller) ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 37 ;Cope, 1865, Nat. Hist. Review, p. 107 ; idem, 1866, Joum. Acad. Philadelphia, (2), VI, p. 80 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 443; van Kampen, 1918, Bijdr. Dierk. XXI., p. 5 4 ; idem, 1923, Amph. Indo-Austr. Arch., p. 133 ; Nieden, 1926, Das Tierreich, Anura n, p. 51 ; Noble, 1931, Biol. Amph., p. 535. Bombinator (non Merrem), Schlegel, 1858, Handl. Dierk., n, p. 58. X e w h i n a , Peters, 1863, Mon. Ak. Berlin, p. 82 (Type species Bombinator ozycephalua, Schlegel) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 179 ; van Kampen, 1923, Amph. IndoAustr. Arch., p. 131 ;Noble, 1931, Biol. Amph., p. 535. Phrynomantis, Peters. 1867, Mon. Ak. Berlin, p. 35 (Type species Phrynomantis fusca, Peters) ; Nieden, 1926, Das Tierreich, Anura n, p. 12 (in part); Noble, 1926, Amer. Mus. Novitates, CCHII, p. 19 ; Noble, 1931, Biol. Amph., p. 533. Eylophorbus, Macleay, 1878, Proc. Linn. Soc. N. S. Wales, n, p. 136 (Type species Hylophorbus rufescens, Macleay) ; F r y , 1923, Mem. Queensland Mus., n, p. 48 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 137 (in part) ;Nieden, 1926, Das Tierreich, Anura u, p. 57 (in part). CalZuZops, Boulenger, 1888, Ann. Mag. Nat. Hist., (6), I, p. 345 (Type species Callulops doriae, Boulenger) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 123 ; Nieden, 1926, Das Tierreich, Anura n, p. 52 ; Noble, 1931, %ol. Amph., p. 535. Mantophryne, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), XIX,p. 12 (Type species Mantophryne lateralis, Boulenger, = mfwcens, Macleay) ; MBhely, 1901, TermBs. Fiizetelr, xxrv, pp. 172 and 219 ; Noble, 1931, Biol. Amph., p. 535. Cfnathophryne, MBhely, 1901, TermBs. Fiizetek, xxrv, pp. 177 and 225 (Type species Mantophryne robusta, Bonlenger). Pomatops, Barbour, 1910, Proc. Biol. Soc. Washington, xxm, p. 89 (Type species Pomatops valvifera, Barbour) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 147 ; Nieden, 1926, Das Tierreich, Anura n, p. 63. Liophryne (non Boulenger 1897), Boulenger, 1914, Trans. Zool. Soc. London, xx,p. 252 ;van Kampen, 1923, Amph. Indo-Austr. Arch., p. 100 (in part) ;Nieden, 1926, Das Tierreich, Anura n, p. 50 (in part).

ASTEROPHRYS

59

Pseudengystom, de Witte, 1930, Ann. Soc. R. Zool. Belg., LX, p. 132 (Type species Pseudengystoaa bouwensi, de Witte) ;de Witte, 1930, Mem. Mus. R. Hiat. Nat. Belg., v, i, p. 6.

Maxillae forming a median suture in front of the premaxillae and dentaries in front of the mento-meckelian bones ; prevomer large, undivided, in contact with its fellow mesially (fig. 27) ; the post-choanal portion overlying the palatine region and sometimes, in females, bearing a few small odontoids which are not larger than normal vomerine teeth. Clavicles, procoracoids and omosternum absent ; sternum cartilaginous (fig. 28). Vertebral column diplasiocoelous. Terminal phalanges T-shaped or simple. Pupil horizontal. Tongue large, subcircular, entire and scarcely free behind, with a:more or less distinct median furrow. A denticulated dermal ridge across the palate

Fig. 27.

Anterior cranial elements of Aaterophrys rufeacens (seen from beneath)

Fig. 28. Ventral elements of the shoulder-girdle of Asterophrys doriae (ventral view)

60

MONOGRAPH O F T H E MICROHYLIDAE

in front of the pharynx, sometimes preceded by a shorter smooth one. Digits free, with or without terminal dilatations. I n all the known species the first finger is shorter than the second, the third toe longer than the fifth, and the enter metatarsal tubercle absent. SYNOPSIS O F THE SPECIES N.B.-Certain species which have not been examined are tentatively included here, but may ultimately prove t o be elcutheropathine and so referable to Metopostira. I. Discs of the fingers and toes large, triangular ; those of the fingers larger than those of the toes . . . . . . . . . . . A. boettge~.i,p. 61 11. Discs of the fingers and toes small, subequal . . . . A . amboinensis, p. 61 111. Discs of the fingers and toes small, or ahsent; if present, those of the toes are larger than those of the fingers. A. Snout not more than once and a half as long as the eye. (1) Tibio-tarsal articulation reaching the eye, or beyond. (a) Tympanum nearly as large as the eye ; a pair of tubercles beneath the chin ; snout slightly longer than the diameter of the eye . . . . . . . . . . A. rufescens, p. 62 ( b ) Tympanum about half the diameter of the eye ; no tubercles beneath the chin ; snout not longer than the eye. (i) Tibio-tarsal articulation reaching the tip of the snout A. louisiadensis, p. 62 (ii) Tibio-tarsal articulation reaching the centre of the eye A. duhin, p. 63 (2) Tibio-tarsal articulation not reaching beyond the posterior border of the eye. (a)Tips of the toes with small, but definite, dilatations ; tibiotarsal articulation reaching the tympanic region. (i) Upper eyelid without cutaneous spines. (a) Tympanum less than half the diameter of the eye. * Lower surfaces brown, white spotted . A. fusca, p. 63 ** Lower surfaces buff or white, immaculate ;adult females with a long row of palatine odontoids . . A. valznfera, p. 64 *** Lower surfaces greenish yellow, immaculate A mimotis, p. 64 (p) Tympanum half, or more than half, the diamekr of the eye. *Tympanum scarcely visible, about half the diameter of the eye ; snout not longer than the latter ; females with not more than one or two very small vomerinc odontoids A. ~obusta,p. 64 ** Tympanum distinct, a t least the diameter of the eye ; snout slightly longer than the latter ; females with a short row of moderately well-developed vomerine odontoids A. doriae, p. 65 *** Tympanum distinct, the diameter of the eye ; snout nearly once and a half as long as the latter ; females without vomerine odontoids A. bouwensi, p. 66 (ii) Upper eyelid with prominent cutaneous spines ;females with a row of well-developed vomerine odontoids A. turpicola, p. 66

+

ASTEROPHRYS

61

(b) Tips of toes not dilated ; tibio-tarsal articulation reaching the shoulder. . . A. minima, p. 67 B. Snout prominent, pointed, twice as long as the diameter of the eye A. oxycephala, p. 67

. . . . .

I.

. . . .

Asterophrys boettgeri (MChely)

Phrynixalus montanus (part), Boettger, 1901, Abh. Senck. Natf. Ges., xxv, p. 368. Gnathophryne boettgeri, MAhely, 1901, Term&. Fiizetek, xrv, pp. 181 and 229 (Type locality Halmaheira, 670 m.). Hylophorbus boettgeri, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 140; Nieden, 1926, Das Tierreich, Anura n, p. 61 ; Mertens, 1930, Zool. Mededeel., Leiden, x m , p. 147.

A curved, smooth, anterior palatal ridge. Snout rounded, a little shorter than the diameter of the eye ; canthus rostralis rounded ; interorbital space as broad as the upper eyelid ; tympanum moderately distinct about half the diameter of the eye. Fingers long, with large triangular discs, the second shorter than the fourth. Toes with distinct discs, which, however, are smaller than those of the fingers ; inner metatarsal t e r c l e indistinct. Tibio-tarsal articulation reaching the nostril. Skin with larger and smaller pustules on the upper eyelids, occiput, shoulders and sides of the body ; smooth beneath. Brown above, with a yellowish white vertebral line and some dark markings ; among these are aW-shaped mark behind the head, a marking between the eyelids and on the sides of the snout, and, here and there on the sides of the body, large reddish brown, irregularly rounded spots ; limbs with transversely elongate black spots. Belly light brown ; chin, throat, chest and lower surfaces of the limbs with large chestnut-brown spots ; a median white spot on the mandibular symphysis and a pair of lateral spots on the edge of the lower jaw. Length : 39 mm. DISTRIBUTION : Halmaheira, 670 metres. No specimens have been examined, but Mkhely's clear description of the symphypathine jaws makes it evident that the species should be referred to this genus ; " Phrynixalus " montanus, Boettger, suggested by van Kampen to be conspecific, has been examined by Dr. Mertens at the author's request and appears to be referable to Cophixnlus.

2.

Asterophrys l amboinensis (Mertens)

Hylophorbus amboinensis, Mertens, 1930, Zool. Mededeel., Leiden, x m , p. 145, fig. 1 (Type locality Amboina).

A smooth, curved, anterior palatal ridge. Snout short, rounded, somewhat prominent and rather longer than the diameter of the eye ; canthus rostralis rounded ; loreal region oblique, concave ; nostril much nearer the tip of the snout than the eye ; interorbital space less than the diameter of the eye ; tympanum tolerably d~stinct,about 8 the diameter of the eye. Fingers long, with small discs which measure 4 to the diameter of the tympanum ; second a little shorter than the fourth. Toes long, wlth discs as large as those of the fingers ; inner metatarsal tubercle feebly developed. Ski; smooth above ; flanks slightly tubercular ; a supratympanic fold ; smooth beneath. Dark brown above, with small, indistinct, irregular blackish flecks; flanks with some whitish dots ; belly dirty yellowish brown. Length : 37 mm. . .J~JSTRIBUTION : Amboina.

+

Not examined ; generic status uncertain.

MONOGRAPH OF THE MICROHYLIDAE

3 . Asterophrys rufescens (Macleay) Hybpbrbus mfescens, Macleay, 1878, Proc. Linn. Soc. N. S. Wales, u, p. 136 (Type locality Katow, N. Guinea) ; Fry, 1913, Mem. Queensland Mus., n, p. 48 ; van Kampen, 1923, Amph. IndoAustr. Arch., p. 140 ;Nieden, 1926, Das Tierreich, Anura 11, p. 58. Mantophryne lateralis, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), xrx, p. 12, PI. 11, fig. 3 (Type locality Mt. Victoria, Owen Stanley Range) ; Boulenger, 1898, Ann. Mus. Genova, (2), xvm, p. 16 ; MBhely, 1901, Termbs. Fiizetek, XXN, pp. 173 and 220, Pls. N, v, and x, fig. 4. Manthophrym lateralis, Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 429.

A curved, smooth, anterior palatal ridge. Snout rounded, slightly prominent, as long as, or a little longer than, the diameter of the eye ; canthus rostralis distinct, rounded ; loreal region nearly vertical, slightly concave ; interorbital space as broad as, or broader than, the upper eyelid ; tympanum moderately distinct, nearly as large as the eye. Fingers long, with very small discs, the second and fourth subequal. Toes with slightly larger discs than the fingers ; subarticular tubercles indistinct ; inner metatarsal tubercle elongate. Tibio-tarsal articulation reaching the eye. Skin usually smooth above, sometimes with small tubercles which are most numerous on the upper eyelids, the flanks and the posterior part of the back, where they may form fine longitudinal lines ; smooth beneath, except the belly, which is sometimes more or less granular ; a pair of small tubercles on the edge of the lower jaw close to the mandibular symphysis. Grey or pale brown above, with small darker dots and a broad, dark brown lateral streak from the eye to the groin, bordered beneath by a narrow silvery stripe; a pair of divergent, diagonal light streaks from the anterior and posterior corners of the eye; sometimes a black, narrowly light-edged, lumbar ocellus. Lower surfaces pale brown or whitish, dotted and splashed with dark brown ; mental warts sometimes white ; hinder side of the thighs with a dark brown streak. Ovarian eggs large (5 mrn.) ; male with a vocal sac. Length : 55 mm. : New Guinea. DISTRIBUTION The five cotypes of Mantophryne lateralis Boulenger are not all conspecific, but the name must be applied to the present species ; the description and figure are clearly drawn up from one of the specimens listed below :

B.M. 96.10.31.32-34

$!?

3 3,?

Mt. Victoria, Owen Stanley Range

Albert Edward Ranges, 6,000 ft. 1934.1.5.2-3 Kokoda (1,200 ft.) Brit. N. Guinea Mus. Hung.A.2414/5 3 specs. ; 2 dissected Sattelberg . . Anat. preparations 1901.11.27. 20

.

Anthony, COTYPES of Mantophnjne lateraus, Boul.

. . . .Rohu . . . .Cheesman . . . . Bir6

4 . Asterophrys louisiadensis s p . n . Type locality Rossel Island, Louisiadb Archip.

A curved, smooth, anterior palatal ridge. Snout rounded, scarcely prominent, as long as the diameter of the eye ; canthus rostralis distinct, rounded ; loreal region almost vertical, slightly concave ; interorbital space slightly less than the breadth of the upper eyelid ; tympanum moderately distinct, a little more than half the diameter of the eye. Fingers long and slender, with very small discs, the second and fourth subequal. Toes slender, with small, but distinct discs, which are a little larger tlltlu those of the fingers ; subarticular tubercles prominent ; inner metatarsal tukrcie distinct. Tibio-tarsal articulation reaching the level of the tip of the snout.

ASTEROPHRYS

63

Skin with a few small tubercles arranged in longitudinal rows on the back and flanks ; a very fine median longitudinal fold ; a supratympanic fold. Lower surfaces smooth. Pale brown above, the flanks and hind limbs faintly marbled with darker ; hinder side of the thighs lighter ; dirty white beneath, the throat spotted with pale brown. Length : 47 mm. ; fore limb 3 0 mm. ; hind limb 78 mm. DISTRIBUTION : Rossel Island, LouisiadC Archip. -

SPECIMEN EXAMINED B.M. 99.4.25. 22

Rossel Island

. . . . . . . . .

Meek, TYPE

5 . Asterophrys dubia (Boettger) Xenorhina dubia, Boettger, 1895, Zool. Anz., 471, p. 134 (Type locality North Halmaheira, up to 2,200 ft.) ; Boettger, 1901, Abh. Senck. Natf. Ges., xxv, p. 369, P1. XVI, fig. 10. Gnathphryne dubia, MBhely, 1901, TermBs. Fuzetek, xxrv, pp. 182 and 231. Hylolphorbus dubius, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 143 ; Nieden, 1926, Das Tierreich, Anura n, p. 61. A smooth 9nterior palatal ridge. Snout pointed, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region oblique ; interorbital space about once and a half the breadth of the upper eyelid ; tympanum feebly distinct, about half the

diameter of the eye. Fingers with very small discs, the second and fourth subequal. Discs of the toes small ; subarticular tubercles feebly developed ; inner metatarsal tubercle weak, compressed. Tibio-tarsal articulation reaching about the middle of the eye. Skin smooth above and beneath ; flanks somewhat warty ; a supratympanic fold. Greyish black above, uniform or indistinctly spotted and marbled with lighter ; limbs with a few indistinct, whitish grey bars, transversely elongate spots and rings ; lower surfaces brownish yellow, reticulated with brown, or brown, marbled and spotted with yellow. Length : 2 4 mm. DISTRIBUTION : Halmaheira, up to 2,200 ft.

6 . Asterophrys fusca (Peters) Phrynomantis fusca, Peters, 1867, Mon. Ak. Berlin, p. 35 (Type locality Amboina) ; Peters and Doria, 1878, Ann. Mus. Genova, xIn, p. 429; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 173 ;Nieden, 1926, Das Tierreich, Anura n, p. 15 ;Noble, 1926, Amer. Mus. Novitates, ccxn, p. 19, fig. 7. Oreophryne celebensis (part), van Kampen, 1923, Amph. Indo-Austr. Arch., p. 112.

A curved, smooth, anterior palatal fold. Snout truncate, not prominent, shorter than the diameter of the eye ; canthus rostralis rounded; loreal region slightly oblique, slightly concave ; interorbital space a little less than the breadth of the upper eyelid ; tympanum moderately distinct, rather less than half the diameter of the eye. Fingers with small discs, the second a little shorter than the fourth. Toes with slightly larger discs than the fingers ; subarticular tubercles feebly developed ; inner metatarsal tubercle elongate. Tibio-tarsal articulation reaching the hinder border of the tympanum. Skin smooth above and beneath ; a supratympanic fold. Dark brown above, flecked more or less profusely with white ; lower surfaces pale brown, flecked with white, darker beneath the throat and on the limbs. Length : 3 5 mm. : Arnboina and Batanta. DISTRIBUTION Mus. Genova

juv. $

Batanta

. .

. . . . . . . . Doria

MONOGRAPH O F T H E MICROHYLIDAE

7 . Asterophrys valvifera (Barbour) P d p a valuifera, Barbour, 1910, Proc. Biol. Soc. Washington, x x m , p. 89, P1. I (Type locality Fak-Fak, Dutch New Guinea) ;van Kampen, 1923, Amph. Indo-Austr. Arch., p. 147 ;Nieden, 1926, Das Tierreich, Anura 11, p. 63. Liophryne kampeni, Boulenger, 1914, Trans. Zool. Soc. London, XX, 5, p. 252, P1. x x v ~ ,figs. 448 (Type locality Mimika River, Dutch New Guinea) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 101 ;Nieden, 1926, Das Tierreich, Anura 11, p. 51.

A curved, rather fibrous, anterior palatal ridge. Snout rounded, not prominent, shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region oblique, slightly concave ; interorbital space slightly less than the breadth of the upper eyelid ; tympanum indistinct or hidden, slightly less than half the diameter of the eye. Fingers with small discs, the second a little shorter than the fourth. Toes with slightly larger discs than the lingers ; subarticular tubercles feebly developed ; inner metatarsal tubercle indistinct. Tibio-tarsal articulation reaching the tympanum or the posterior border of the eye. Skin smooth, or with a few very small pustules on the flanks ; a supratympanic fold. Slate grey to purplish brown above, uniform or with lighter mottling; white or buff beneath. Post-choanal portion of the prevomer forming a distinct ridge behind and mesial to each choana ; this ridge bears a row of small odontoids, a t least in adult females. Length : 55 mm. : Dutch New Guinea. DISTRIBUTION The " semilunar-shaped white spots " beneath each eye mentioned in the original description appear to be the translucent lower eyelids rendered opaque by immersion in spirit. The topotype examined formed part of the collection in which was the type. SPECIMENS EXAMINED B.M. 1909.4.30.16 juv. Fak-Fak . . . Pratt, TOPOTYPE 1913.10.31.349 $2 Mimika River. . TYPEof Liophryne kampeni, Boul.

8. Asterophrysl microtis (Werner) Manlophryne microtis, Werner, 1901, Zool. Anz., XXIV,p. 102 (Type locality " German New Guinea") ; Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 429 ;van Kampen, 1923, Amph. Indo-Auetr. Arch., p. 144. Snout truncate, a little shorter than the eye ; canthus rostralis indistinct ; loreal region oblique, slightly concave ; interorbital space twice as broad as the upper eyelid ; occiput somewhat swollen ; tympanum indistinct, ) the width of the eye. Fingers

with scarcely perceptible discs. Toes with distinct small discs ;subarticular tubercles indistinct ; inner metatarsal tubercle small, flat, elliptical. Tibio-tarsal articulation reaching the anterior border of the tympanum. Skin smooth above and beneath, with small pustules on the sides ; a supratympanic fold. Upper surfaces dark olive green ; snout and upper eyelid blackish ; sides and hindlimbs with scattered white dots. Lower surfaces greenish yellow. Length : 56 mm. DI~TRIBUTION : Known only from the type locality.

9. Asterophrys robusta (Boulenger) Mantophryne robwta, Boulenger, 1898, Proc. Zool. Soc. London, p. 480, P1. xxxv~n,fig. 4 (Type locality St. Aignan Island). Gnathphryne robusta, Mbhely, 1901, Termes. Fiizetek, xxrv, pp. 178 and 225, P1. IV, figs. 1-3, P1. IX, figs. 6-9 ; Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 429. Hybphorbus robustzta, van Kampen, 1918, Bijdr. Dierk., x x ~ p. , 54; van Kampen, 1 Not examined ; generic status uncertain.

ASTEROPHRYS

65

1923, Amph. Indo-Austr. Arch., p. 143 (in part) ; Nieden, 1926, Das Tierreich, Anura n, p. 60, k s . 4 2 4 3 ;Kinghorn, 1929, Rec. Austr. Mus., xvn, p. 76. Hylophorhs sp., Kinghorn, 1928, Rec. Austr. Mus., xm, p. 289.

A curved, smooth, anterior palatal ridge. Snout obtusely pointed, prominent, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space as broad as, or a little broader than, the upper eyelid ; tympanum indistinct, about half the diameter of the eye. Fingers short, with very small discs, the second a little shorter than the fourth. Toes short, with rather larger discs than the fingers ; subarticular tubercles moderately well developed, flat ; inner metatarsal tubercle not well defined. Tibio-tarsal articulation reaching the tympanum. Skin rather thick and glandular ; smooth, or with a few scattered tubercles above ; smooth beneath ; a supratympanic fold. Brown or purplish above, uniform or with white-centred black spots ; sometirnes a white spot in the groin. Lower surfaces whitish, Length : 68 mm. The eggs are large, connected by a mucilaginous cord and laid on land (e.g. in a decaying tree stump) ; there is no free lama1 stage, development being completed within the egg membranes. The developing embryo lacks external and internal gills, respiration taking place by means of a leaf-like, non-muscular, vascular tail ; there is no lama1 q u t h armature and no operculum. DISTRIBUTION: New Guinea (Sattelberg and Mt. Lamington District) and St. Aignan Island. SPECIMENS EXAMINED B.M.98.3.31. 9-11 $ 9 St. Aignan.Island . COTYPES Mus. Vienna New Guinea . . . . Rolle Mus. Hung. 239'711118 $ Sirnbang, New Guinea . . Bir6

. . . . . . . . . . . . .

10. Asterophrys

doriae (Boulenger)

Cdlulops doriae, Boulenger, 1898, Ann. Mus. Genova, (2), xvn, p. 709 (Type locality Milne Gulf, New Guinea) ; Vogt, 1911, Sitzber. Gee. Nat. Fr. Berlin, p. 431 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 123 ; Nieden, 1926, Dm Tierreich, Anura n, p. 52. ? Manthophryne neuhusai, Vogt, 1911, Sitzber. Gee. Nat. Fr. Berlin, pp. 425 and 429 (Type locality Sattelberg). ? Hylophorbus neuhaussi, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 144 ; Nieden, 1926, Das Tierreich, Anura n , p. 59. Hylophorbus robustus (part), van Kampen, 1923, Amph. Indo-Austr. Arch., p. 143.

A curved, smooth, anterior palatal ridge. Snout obtusely pointed, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region oblique ; interorbital space once and a quarter the breadth of the upper eyelid; tympanum distinct, to 1 the diameter of the eye.l Fingers short, with very small discs, the second and fourth subequal. Toes with slightly larger discs than the fingers ; subarticular tubercles flat ; an oval inner metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder or the tympanum. Skin very thick and glandular, smooth, or beset with small warts above ; a supratympanic, and sometimes an occipital, fold. Brown or purplish grey above, the head (from the level of the hinder comers of the eyes) sometimes lighter ; dorsal warts tipped with white. Lower surfaces pale brown or dirty white, faintly stippled with lighter ; sometimes a fine white line across the chest a t the level of the axillae, intersected by a similar median longitudinal line on the belly. Groin, anterior and posterior sides of the thighs, posterior sides of the tibiae and upper surfaces of the feet black, with numerous large, round, yellow spots which reduce the ground colour to a reticulum. Post-choanal portion of the prevomer forming a slight ridge behind and mesial to each choana. In adult females this ridge bears a few small odontoids, but is quite smooth in the other sex. The original description is incorrect with regard to the size of the tympanum. 6

MONOGRAPH O F THE MICROHYLIDAE Ovarian eggs large (6.5 mm.). Length : 100 mm. : S.E. New Guinea. DISTRIBUTION Milne Gulf . . . . . Haveri, British New Guinea 69 Albert Edward Mts., 6,000 ft. Skel. Albert Edward Mts., 6,000 ft. 6,Hgr. Astrolabe Mts., 4,000 ft. . .

9 6

B.M. 88.3.21. 10 97.12.10. 140 1901.11.27. 17-18 1901.11.27. 19

Mus. Vienna II.

. . Forbes, TYPE . . Loria . . Rohu

. . Rohu .

.

Asterophrys bouwensi (de Witte)

Pseudengystoma bouwensi, de Witte, 1930, Ann. Soc. R. 2001. Belg., LX, p. 132, PI. vx (Type locality Arfak Mts., 1,000 m.) ; de Witte, 1930, Mem. Mus. R. Hist. Nat. Belg., v, i, p. 6, figs. 3 and 4. Posterior palatal ridge short ; no anterior. Snout slightly prominent, bluntly pointed, nearly once and a half as long as the eye ; canthus rostralis rounded ; loreal region oblique, but not markedly concave ; interorbital space nearly twice as broad

as the upper eyelid; tympanum moderately distinct, 4 the diameter of the eye. Fingers short, not dilated, the second a little shorter than the fourth. Toes with small, but distinct, discs ; subarticular tubercles feebly developed ; inner metatarsal tubercle oval, ill-defined. Tibio-tarsal articulation reaching the tympanum. Skin for the most part smooth, but with a few small warts posteriorly and on the flanks, and with some minute pustules on the snout ; a shallow, longitudinal groove on the occiput ; traces of a fine cutaneous fold from snout to vent ; no supratympanic fold. Uniform blackish brown above ; lower surfaces white, heavily mottled with brown ; hinder side of the thighs with an indistinct, broad, lighter band from the vent to the back of the knee. Ovarian eggs relatively large (2 mm.). Length : 20 mm. : Arfak Mts., New Guinea, 1,000 m. DISTRIBUTION Mus. Brussels 9223 : I11 I 2.

9

Arfak Mts., 1,000 m.

. . . . . .

TYPE

Asterophrys turpicola (S. Miiller)

Ceratophrys turpicola, MuUer in Schlegel, 1837, Abbild. Amph., p. 30, PI. X, fig. 4 (Type locality " West Coast of New Guinea "). Asterophrys turpicola, Tschudi, 1838, Mem. Soc. Sci. Neuchitel, n, 1839, p. 82 ; Gunther, 1858, Cat. Batr. Sal. Brit. Mus., p. 37 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 444 ;WandoUeck, 1911, Abh. ZOO^. Anthr.-Ethn. Mus. Dresden, x m , p. 14 ;van Kampen, 1918, Bijdr. Dierk., XXI, p. 54 ; Nieden, 1926, Das Tierreich, Anura n, p. 51, fig. i. Cerdophryne turpicola, Schlegel, 1858, Handl. Dierk., n, p. 57. Asterophrys turpicula, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 133, fig. 1 8 ; Burt, 1932, Bull. Am. Mus. Nat. Hist., Lxm, p. 483. Asterophrys leopoldi, de Witte, 1930, Ann. Soc. R. 2001. Belg., LX, p. 131, PI. v (Type locality Arfak Mts., 1,000 m.) ; de Witte, 1930, Mem. Mus. R. Hist. Nat. Belg., v, 1, p. 4, figs. 1-2. ? Asterophr~ssteini, Ah!, 1932, Mitt. 2001. Mus. Berlin, X ~ I 6, , p. 894, figs. (Type locality Jobi Island).

Anterior palatal ridge reduced to an indistinct median tubercle. Snout truncate, scarcely prominent, as long as, or slightly longer than, the diameter of the eye ; canthus rostralis bluntly angular ; loreal region almost vertical, concave ; interorbital space nearly twice the breadth of the upper eyelid ; tympanum rather indistinct, 8 to 1 the diameter of the eye ; a prominent bony ridge (due to the squamosal) from behind the eye to above the tympanum. Fingers with small, scarcely perceptible discs, the second subequal to, or a little longer than, the fourth. Toes with small, but definite, discs ;

ASTEROPHRYS

67

subarticular tubercles distinct ; inner metatarsal tubercle oval, ill-defined. Tibio-tarsal articulation reaching the tympanum or the eye. Skin with scattered tubercles on the dorsum, these being most plentiful on the snout, the upper surfaces of the limbs, the posterior part of the back and the flanks ; upper eyelid with four or five prominent cutaneous spines, one near the centre of the free edge being the largest and most conspicuous. Lower surfaces smooth, or with some scattered minute warts. Brown above (green in life) with scattered irregular, darker spots ; sides of head darker ; groin darker with white vermiculations ; digits regularly barred with white and dark brown above. Lower surfaces pale brown, the belly and limbs spotted and marbled with white. The post-choanal portion of the prevomer forms a prominent curved ridge behind each choana, and, in adult females a t least, bears a row of small odontoids. Ovarian eggs 4 mm. in diameter. Length : 65 rnm. DISTRIBUTION : New Guinea ; Jobi Island (?). SPECIMEN EXAMINED Mus. Dresden 8 Torricelli Mts. . . . . . . . Schlaginhaufen. 6

I

3. Asterophrys minima sp . n.

Xenorhim ocellata (part), van Kampen, 1913, Nova Guinea, IX, 3, p. 461.

Anterior palatal ridge reduced to a median tubercle. Snout bluntly pointed, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region oblique, concave ; interorbital space about as broad as the upper eyelid ; tympanum very indistinct, a little smaller than the eye. Fingers without discs, the second shorter than the fourth. Toes also without discs, the third longer than the fifth; subarticular and inner metatarsal tubercles very indistinct. Tibio-tarsal articulation reaching the shoulder. Skin smooth above and below ; a few small, scattered warts on the sides ; a supratympanic fold. Pale brown above, stippled with darker ; a definite triangular marking from the posterior corner of the eye to the angle of the mouth, a smaller spot on the lip in front of the eye and a spot on the tip of the snout, clear white ; a fine white line from snout t o vent. Lower surfaces dusted with dark brown ; a white stripe along the edge of the lower jaw, continued to the shoulder; under surfaces of the limbs, hands and feet also white. Length : 27 mm. ; fore-limb 13 rnrn. ; hind-limb 30 mm. DISTRIB~~~ION : Dutch New Guinea. The so-called juvenile doubtfully referred by van Kampen to Xenorhinu ocellata is apparently fully adult and belongs to this new species. SPECIMENS EXAMINED Mus. Amsterdam. A.A.31 Went. Mts., 1,000-1,360 m., Oct. 12, 1909 TYPE A.A.30 Ad. Hellwig Mts., 2,500 m. . . . . . . PARATYPE

14. Asterophrys oxycephala (Schlegel) Bombinator oxycephalus, Schlegel, 1858, Handl. Dierk. u, p. 58, P1. m, fig. 74 (Type locality New Guinea). Xenorhinu oxycephala, Peters, 1863, Mon. Ak. Berlin, p. 82 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 179 ; Mbhely, 1901, Term&. Biizetek, xxrv, pp. 188 and 236 ; Vogt, 1911, Sitzb. Ges. Nat. Fr. Berlin, p. 430 ; van Kampen, 1914, Zool. Jahrb., Syst., xxxm, p. 374; idem, 1918, Bijdr. Dierk., sxr, p. 54 ; idem, 1923, Amph. Indo-Austr. Arch., p. 132, fig. 17 (part) ; Nieden, 1926, Das Tierreich, Anura n, p. 55 (part) ; Burt, 1932, Bull. Amer. Mus. Nat. Hist., L X ~ p. ,485. ? Xenorhinu stresemunni, Ahl, 1932, Mitt. Zool. Mus. Berlin, xm, 6, p. 897, fig. (Type locality Jobi Island).

MONOGRAPH OF THE MICROHYLIDAE Anterior palatal ridge reduced to a median swelling ; posterior short. Snout acutely pointed, prominent, twice as long as the diameter of the eye ; can thus rostralis indistinct ; loreal region very oblique, feebly concave ; interorbital space about 3 times as broad as the upper eyelid ; tympanum indistinct or hidden, about as large as the eye. Fingers short, with small discs, the second very slightly shorter than the fourth, Toes with slightly larger discs than the fingers ; subarticular and inner metatarsal tubercles feebly developed. Tibio-tarsal articulation reaching the tympanic region. Skin smooth or slightly roughened ; sometimes a few small pustules beneath the snout ; a supratympanic fold. Brown, grey, violet or reddish above with or without darker reticulations or a light vertebral stripe ; grey, brick-red, orange or yellow beneath, with dark brown reticulations ; hinder sides of thigh, tarsus and foot, black. Length : 47 mm. DISTRIBUTION : New Guinea ; Jobi Island (?). SPECIMENEXAMINED Mus. Amsterdam $! Mosso River

Xenorhina (non Peters), Giinther, 1896, Novit. Zool., rn, p. 184 ; MBhely, 1901, TermBs. Fiizetek, xxm, pp. 183 and 231 (part). Metopostira, MBhely, 1901, Termbs. Fiizetek, xxm, pp. 189, 238 (Type species Metopostira ocellata, MBhely). Hylophorbus (part), van Kampen, 1918, Bijdr. Dierk., XXI,p. 54 ; idem, 1923, Amph. Indo-Austr. Arch., p. 137 ; Nieden, 1926, Das Tierreich, Anura n, p. 57. Phrynomantis (part), Noble, 1931, Biol. Amph., p. 533.

Maxillae not meeting in front of the prernaxillae; prevomer large, undivided, in contact with its fellow mesially, the post-choanal portion overlying the palatine region but without odontoids (fig. 29). Clavicles procoracoids and ornosternum absent; sternum cartilaginous. Vertebral column diplasiocoelous. Terminal phalangesT-shaped. Pupil horizontal. Tongue large, subcircular, almost entirely adherent behind, with a more or less distinct median furrow. A denticulated dermal ridge across the palate in front of the pharynx, sometimes preceded by a curved smooth one. Digits free ; toes with small terminal dilatations. The second and fourth fingers are subequal, the third toe extends beyond the fifth and the outer metatarsal tubercle is absent.

Fig. 29. Anterior cranial elements of Metopostira ocellata (seen from beneath)

This genus is only distinguished from Asterophrys by the separation of the maxillae anteriorly and M. ocellatus and M . kopsteini appear on other characters to be more

METOPOSTIRA

69

closely allied t o A. mfescens than to M. atra, and this latter species has more in common with A. oxycephala than with either M. ocellatus or M. kopsleini. This may indicate that Metopostira is not tenable, but, as has already been pointed out (p. 15), both Asterophrys and Xenobatrachus show a complete transitional series between extremes analogous to the species here referred to Meloposlira ; consequently the apparent resemblances between pairs of species in Asterophrys and Metopostira may be due to parallelism rather than t o close relationship. SYNOPSIS OF THE SPECIES

I. Eye as long as the snout ; habit slender ; digits long and slender. A. Loreal region nearly vertical ; tibio-tarsal articulation reaching the eye or beyond the end of the snout . . . . . . M. ocellala, p. 69 B. Loreal region oblique ; tibio-tarsal articulation reaching the shoulder or the posterior corner of the eye . . . . . . . M. kopsteini, p. 70 11. Eye less than half the length of the snout ; habit stout ; digits very short M. alra, p. 70 I.

Metopostira ocellata, MChely

Metopostira ocellata, MBhely, 1901, TermBs. Fiizetek, XXN, pp. 190 and 239, PI. vu, P1. xu, fig. 1 (Type locality Sattelberg) ; van Kampen, 1909, Nova Guinea, IX, i, p. 40 ; Vogt, 1911, Sitzb. Ges. Nat. Fr. Berlin, p. 429 ; van Kampen, 1913, Nova Guinea, IX,3, p. 461 ; idem, 1914, Zool. Jahrb., Syst., xxxw, p. 375 ; idem, 1915, Nova Guinea, xm, i, p. 40. Hylophorbus ocellatus, van Kampen, 1918, Bijdr. Dierk., xxr, p. 40 ; idem, 1923, Amph. Indo-Austr. Arch., p. 141 ; Nieden, 1926, Das Tierreich, Anura n, p. 60. Metopostira m r a , van Kampen, 1906, Nova Guinea, v, i, p. 167, P1. VI, figs. 1 and 2 (Type locality " Am Moso und Tami ").

A curved, smooth, anterior palatal ridge. Snout obtusely pointed, prominent, shorter than the diameter of the eye ; canthus rostralis distinct, rounded ; loreal region nearly vertical, concave ; interorbital space as broad as, or a little broader than, the upper eyelid ; tympanum hidden in juveniles, distinct in adults, $ to 9 the diameter of the eye. Fingers with small discs. Toes with slightly larger discs than the fingers ; subarticular and inner metatarsal tubercles distinct. Tibio-tarsal articulation reaching the eye or somewhat beyond the tip of the snout. Skin smooth above, the hinder part of the back and the flanks sometimes with small pustules, which may be arranged in longitudinal series ; smooth beneath ; a supratympanic fold. Brown above (greenish brown in life), with scattered dark brown spots on the back and limbs ; a broad, dark, vertical bar beneath the eye ; a circular black spot in the groin, bordered by lighter (red anteriorly and yellow posteriorly in life), flanks lighter, spotted with dark brown. Lower surfaces yellowish with dark spots and marblings which are more pronounced on the throat and chest. Length : 43 mm. Ovarian eggs large (3 mm.) ; male with a vocal sac. : New Guinea. DISTRIBUTION B.M. 83.10.23.4 96.10.31. 35 1901.11.27. 21 1934.1.5.1

9 9 9

.

.

Katow . . . . . Mt. Victoria, Owen Stanley Range

Doria Anthony, COTYPE of Mantophryne laleralis, Boul. Albert Edward Ranges, 6,000 ft. Rohu Kokoda (1,200 ft.), British N. Cheesman Guinea

MONOGRAPH OF THE MICROHYLIDAE Mus. Amsterdam

Mus. Hung.

1 1 1 2 2 2 17 spec.

Hellwig Mts., 1,00&1,360 m. Lower Bewain River . . Kloof bivak, Lorentz River Went Mts., 800 m. . Digul River . Sabany . . . . . .

.

. . . .

.

. .

Versteeg Lorentz Koch Bir6, COTYPES

anat. preps. 2.

Metopostira kopsteini (Mertens)

Hyl~phorbuskopsteini, Mertens, 1931, Zool. Mededeel., Leiden, XIII,p. 147. fig. 2 (Type locality Sanana, Suh Islands). ? Hylophorbus ocellatua (non MBhely), Schijfsma, 1932, Treubia, X N , i, p. 70.

A c w e d , smooth, anterior palatal ridge. Snout rounded, somewhat promixient, as long as the eye ; canthus rostralis rounded ; loreal region oblique, slightly concave ; nostril much nearer the end of the snout than the eye ; interorbital space less than the diameter of the eye ; tympanum very distinct, Q the diameter of the eye. Fingers with very small discs (& to the size of the tympanum). Toes with larger discs than the fingers ; subarticular tubercles feebly developed ; inner metatarsal tubercle feeble. Tibio-tarsal articulation reaching the posterior corner of the eye in males, or the shoulder in females. Skin smooth on the back ; flanks and hinder part of the back with small, scarcely distinct pimples ; a supratympanic fold ; a very fine dermal fold from snout to anus and on the mid-line of the belly. Lower surfaces smooth. Dark brown above, with a more or less distinct, black, anteriorly white-bordered, ocellar spot above the groin ; flanks with indistinct lighter and darker flecks ; belly dirty grey-brown with indefinite light grey flecks. Length : d 41.5 mm. ; $2 45 mrn. Eggs large (ovarian 3.5 rnm.). DISTRIBUTION : Sanana, Sulu Islands ; ? Amboina.

+

3. Metopostira atra (Giinther) Xemrhina atra, Giinther, 1896, Novit. Zool., nI, p3 184, P1. vm, fig. 1 (Type locality British New Guinea) ;Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, p. 430. X e w h i n a ozycephala (part), van Kampen, 1923, Amph. Indo-Austr. Arch., p. 123 ; Nieden, 1926, Das Tierreich, Anura n, p. 55.

Anterior palatal ridge indistinct. Snout obtusely pointed, slightly proAinent, more than twice as long as the very small eye ; no canthus rostralis ; loreal region very oblique, convex ; interorbital space a t least 3 times as wide as the upper eyelid ; tympanum scarcely distinguishable, once and a half the diameter of the eye. Fingers very short, without discs. Toes with small discs and traces of lateral ridges; subarticular and inner metatarsal tubercles flat, indistinct. Tibio-tarsal articulation reaching the tympanic region. Skin smooth above and beneath ; an indistinct supratympanic fold. Brown above with darker flanks, the line of demarcation being sharply defined. Sometimes a curved row of 4 or more dark brown spots is present just in front of the hind-limbs, which have very narrow, indistinct, light, dark-bordered cross-bars. Lower surfaces pale brownish grey ; the hinder side of the thighs darker. Length : 42 mrn. Male with a vocal sac. DISTRIBUTION : British New Guinea. SPECIMENS EXAMINED B.M. 96.7.7.3 9 British New Guinea . . . . . . . Day, TYPE 1901.11.27.28 9 Albert Edward Ranges, 6,000ft. . . . Rohu 1904.11.1. 51 d Albert Edward Ranges . . Rohu

'

MICROHYLINAE

MICROHYLINAE, Giinther Gastrophrynae (part),Fitzinger, 1843, Syst. Rept., p. 33. Micrhylidae, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 121. Kalophrynina, Mivart, 1869, Proc. Zool. Soc. London, p. 289. Cacopinae, Kalophryninae, Microhylinae, Kaloulinae, Noble, 1931, Biol. Amph., pp. 532, 636, 537 and 538. Maxillary and vomerine teeth absent ; prevomer much reduced, usually divided and the post-choanal portion lost, never with a wing-like mesial expansion and never in contact with its fellow ; ethmoid divided, or absent (Kalophrynus) ; ear fully developed ; vertebral column diplasiocoelous (rarely procoelous). A free-swimming tadpole stage. SYNOPSIS OF THE GENERA I. Pre- and post-choanal portions of the prevomer fused and forming a single horseshoe-shaped bone surrounding the choana. Tongue circular, adherent, with a longitudinal median furrow. Kaloula, p. 76 A. Vestigial procoracoids and ornosternum persistent . B. No procoracoids or omosternum. (1) Mental region greatly distended . . . . . . Glypl~oglossus,p. 72 (2) Mental region normal . . . . . . . . . Uperodon, p. 73 11. Pre- and post-choanal portions of the prevomer separated and the latter frequently entirely lost. A. No clavicles, procoracoids or omosternum. (1) Subarticular tubercles of the fingers normal. (a) Tongue adherent, with a median furrow. Uperodon, p. 73 (i) Toes with a rudiment of web . (ii) Toes 2 webbed . . . . . . . . Ctenophryne, p. 122 (b) Tongue half free behind, without a median furrow, Microhyla, p: 123 (2) Subal-ticular tubercles of the fingers enlarged to form accessory adhesive pads . . . . . . . . . 1MetaphrynelEa, p. 107 B. No clavicles ; procoracoid reduced t o a rudiment lying on the mesial end of the coracoid. (1) Subarticular tubercles of the fingers enlarged to form accessory . . . . . . . . . . Phrynella, p. 105 adhesive organs (2) Subarticular tubercles of the fingers normal . . . . Ramanella, p. 90 C. No clavicles ; procoracoid a narrow slip extending from the mid-line of the girdle to the scapula . . . . . . . Gastrophrynoides, p. 104 D. Clavicles and procoracoids present. (1) Clavicles and procoracoids confined to the mesial extremities of . . . . . . . . . . Elachistocleis, p. 120 the coracoids (2) Clavicles and procoracoids not reaching the scapulae, but extending over a t least the mesial half of the coracoid Chiasmocleis, p. 116 (3) Clavicles and procoracoids extending from the mid-line of the girdle to the scapulae. (a) Omosternum absent. (i) Terminal phalanges simple. ( E ) Clavicles strongly bent, following the curvature of . . . . . . . Ribeirina, p. 115 the coracoids ( p) Clavicles straight . . . . . . Hypopachus, p. 110 (ii) Terminal phalanges T-shaped . . . . Chaperina, p. 101 (6) Omosternum present. (i) Terminalphalangessirnple . . . . . .Otophryne, p. 108 (ii) Terminal phalanges T-shaped . , . . Kalophrynus, p. 95

MONOGRAPH OF THE MICROHYLIDAE GL YPHOGLOSSUS, Giinther Cflyphgbsszls, Giinther, 1868, Pmc. Zool. Soc. London, p. 483 (Type species Clyphglosszls WWb8szl8) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 175 ; Theobald, 1882, in Mason, Burma, I, p. 292 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 497 ; Nieden, 1926, Das Tierreich, Anura n, p. 20 ; Noble, 1931, Biol. Amph., p. 533.

Prevomer undivided, the post-choanal portion overlying the palatine region and bearing, mesially, one or two knob-likeprominences (fig. 30). No clavicles, procoracoids or omosternum ; sternum cartilaginous. Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil circu1ar.l Tongue small, almost entirely adherent, with a median furrow and a central depression posteriorly. A fimbriated ridge across the palate in front of the pharynx, preceded by a median papilla. An indefinite, pigmented, dermal ridge behind each choana. Digits not dilated distally. Submental region developed to form a globular sac.

Fig. 30. Anterior cranial elements of Clyphglossus molossus (seen from beneath)

This genus ought, perhaps, to be united with Uperodon. It stands in the same osteological relation to Uperodon as Ramanella does to Kaloula, but, whereas no species definitely links these latter two genera, U . globulosum is almost exactly intermediate between the type species of Uperodon and Glyphoglossus.

.

r . Glyphoglossus molossus, Giinther Glyphglossus molosszls, Giinther, 1868, Proc. Zool. Soc. London, p. 483, PI. xxxvnr, fig. 1 (Type locality Pegu, Burma) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 175 ; Theobald, 1882, in Mason, Burma, I, p. 292 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 497 ; Smith, 1917, Journ. Nat. Hist. Soc. Siam, 11, 4, p. 269 ; Nieden, 1926, Das Tierreich, Anura n, p. 21, figs. 1 6 1 7 ;Noble, 1931, Biol. Amph., fig. 174. Snout short, rounded, not prominent, a little longer than the diameter of the eye ; nostril nearer the eye than the tip of the snout ; no canthus rostralis ; loreal region

very oblique; skin beneath the chin much thickened, distended by an underlying gland to form a globular sac, truncate anteriorly and as deep as the head above the mouth ; interorbital space once and two-thirds as broad as the upper eyelid ; tympanum hidden. Wingers short, pointed, the first nearly as long as the second, whlch is much longer than the fourth. Toes pointed, 2 webbed, the membrane reaching the The pupil of this, and several other, genera is sub-circular when fully dilated, but triangular, with the apex ventral, when contracted.

UPERODON

73

tips of all except the fourth ; fifth toe much shorter than the third and but little longer than the second ; subarticular tubercles weak ; a very large, oblique, sharp-edged, compressed, inner metatarsal tubercle, and a small, transverse outer one. Tibio-tarsal articulation reaching the shoulder. Skin feebly granular above and on the anal region ; smooth beneath ; anterior part of head, to the level of the hinder border of the eyes, and the chin, beset with numerous minute, regularly arranged pustules ; sometimes an occipital fold ; a strong groove behind the mental dilatation. Brown to purplish above, more or less profusely dotted and spotted with white, especially on the limbs and flanks ; sometimes a pale dorsolateral streak from the comer of the eye. Lower surfaces white, the chin and throat infuseate. Length : 68 mm. Male with a subgular vocal sac opening by a slit on each side of the tongue. Spawn laid in masses, which float on the water ; the fully formed tadpole (with well-developed hind-limbs) has the following characters : Head and body ovoid, rounded anteriorly, slightly depressed, once and two-thirds as long as broad. Nostrils about midway between a line connecting the anterior borders of the eyes and the tip of the snout, or slightly nearer the former. Eyes lateral, 4 or 5 times as widely separated as the nostrils. Mouth terminal, half as wide as the h e a d a t the level of the eyes, with a thin, downwardly curved upper lip and a contractile lower one. Spiraculum median, opening about the centre of the gut ; spiracular flap broad, strongly emarginate in the centre so as to form two lappets. Anus median, the anal tube long, S-shaped. Tail once and a half to once and threequarters as long as the head and body, about 5 times as long as deep, and acutely pointed posteriorly ; caudal membranes shallow, straight in profle, not extending on to the body. Pale green, grey or brown, more or less translucent ; dark pigment is usually present around the nostrils, in the middle between the eyes and a t the base of the tail ; posterior half of the tail, and its ventral half anteriorly, often black. A well-grown example measures 35 rnm., with a head and body of 13 mm. and a tail 4.5 mm. deep.1 DISTRIBUTION : Burma, Siam.

a

SPECIMENS EXAMINED B.M. 68.4.3. 102 6 Pegu . . . . . Theobald, TYPE 1914.8.7. 23-26 Paknampo, N.E. Siam M. A. Smith 3 66,? 1916.4.17.24-30 Larvae and Young Paknampo, N.E. Siam M. A. Smith 1927.9.1. 6 Cleared Ta Rua C. Siam . . M. A. Smith 1928.12.13.84-109 Larvae and Young Ta Rua, C. Siam . M. A. Smith (Original series described)

UPERODON, Durn. and Bibr. Rana (part), Schneider, 1799, Hist. Amph., I, p. 144. Bombinator (part), Merrem, 1820, Versuch. Syst. Amph., p. 178. Engystoma (part), Cuvier, 1829, Regne Animal, ed. 2, 11, p. 112. Bydoma (non Wagler), Tschudi, 1838, Mem. Soc. Sci NeuchLtel, n, 1839. p. 86 ; Cope, 1867, Journ. Acad. Philadelphia, (2), vr, p. 193. Uperodon, Dumbril and Bibron, 1841, Erpet. Gen., vm, p. 746 (Type species Engyatorna mrmo~atum,Cuvier = Rana systoma, Schneider); Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 48 Hyperodon (emend.), Jerdon, 1854, Journ. Asiatic Soc. Bengal, xxn, p. 533. C m p s , Giinther, 1864, Rept. Brit. India, p. 415 (substitute name for Uperodon, Dum. and Bibr.) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 173 ; idem, 1890, Fauna Brit. India, Rept. and Batr., p. 496 : Nieden, 1926, Das Tierreich, Anura n, p. 19 ; Noble, 1931, Biol. Amph., p. 532. The dimensions given by M. A. Smith probably refer to a tadpole of Calluella guttulato which is almost indistinguishable from that of the present species.

74

MONOGRAPH O F THE MICROHYLIDAE

~ach~bat&hz~e, Keferstein, 1868, Arch. f. Naturg., x x m , p. 273 (Type species Pachybat~achus pebersi, Keferstein = R a m a y s l m , Schneider). Prevomer entire or divided (figs. 31 and 32), the post-choanal portion overlying the

palatine region and sometimes bearing a raised knob a t its mesial end. Clavicles, procoracoids and omosternum absent ; sternum cartilaginous. Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil circular.1 Tongue subcircular, entire and scarcely free behind, with a median furrow. A denticulated dermal ridge, across the palate in front of the pharynx, preceded by a shorter curved one ; a short ridge culminating in a papilla a t its mesial end, or a pair of papillae, behind each choana. Digits not dilated distally.

Pig. 31. Anterior cranial elements of Uperodon globulosum (seen from beneath)

Pig. 32. Anterior cranial elements of U p e r o b sysloma (seen from beneath)

SYNOPSIS OF THE SPECIES I. Snout about equal to the diameter of the eye ; interorbital space about

twice as broad as the upper eyelid . . . . . . . . U . systoma, p. 75 11. Snout nearly twice as long as the diameter of the eye ; interorbital space nearly three times as broad as the upper eyelid . . . U. globulosum, p. 76 As in Glyphogh9~6.

UPERODON I.

Uperodon systoma (Schneider)

Rana systoma, Schneider, 1799, Hist. Amph., I, p. 144 (Type locality " Eastern India "); Shaw, 1802, Gen. Zoology, m, p. 171 ; Peters, 1863, Mon. Ak. Berlin, p. 82. Bombinator systoma, Merrem, 1820, Syst. Amph., p. 178. C m p w systoma, Giinther, 1864, Rapt. Brit. India, p. 415; Theobald, 1868, Journ. Asiatic Soc. Bengal (Cat. Rept. Mus. Asiatic Soc.), p. 82 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 174, fig. ; Thurston, 1888, Cat. Batr. Sal. S. India, p. 43, P1. IX ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 496. fig. ;Boettger, 1892,

Kat. Batr. Senck. Nat. Ges., p. 2 5 ; Sclater, 1892, Batr. Ind. Mus., p. 2 4 ; Ferguson, 1904, Journ. Bombay N. H. Soc., xv, p. 507, P1. B, figs. ; Devanesen, 1912, Notes on the Anatomy of Caeopw systoma, 55 pp. ; Rao, 1918, Rec. Ind. Mus., xv, p. 46 ; Nieden, 1926, Dss Tierreich, Anura n, p. 19, figs. 1 6 1 5 . Uperodon systoma, Parker, 1931, Archiv. Zool. Ital., XVI, p. 1243, fig. 3. Engystoma marnwratum, Cuvier, 1829, Regne Animal, ed. 2, n, p. 112 (Type locality "India ") ; Gubrin, 1829, Icon. Gen. Rept., PI. xxvn, fig. 3. Uperodon marmotaturn, Dum. and Bibr., 1841, Erpet. Gen., v ~ n p. , 749 ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 49. Hyperodon marmotaturn, Jerdon, 1854, Journ. As. Soc. Bengal, XXII, p. 533. Systoma marmaraturn, Steindachner, 1867, Reise Novara, Amph., p. 36 ; Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 194. Systoma breviceps (part), Tschudi, 1838, Mem. Soc. Sci. NeuchLtel, u, 1839, p. 86. Systoma lesckhdti (n.n.), Tschudi, 1838, Mem. Soc. Sci. NeuchLtel, n, 1839, p. 86. Caeops globulosus (part), Giinther, 1864, Rept. Brit. India, p. 416 ; ? Pearson, 1910, Spolia Zeylanica, w, p. 110. Pachybatrachus petersii, Keferstein, 1868, Arch. f. Naturg., p. 274, P1. VI,figs. 8-10 (Type locality " New South Wales ").

Prevomer divided. Snout rounded, not prominent, as long as, or a little longer than, the eye ; no canthus rostralis ; loreal region oblique, convex ; nostril equidistant from the eye and the end of the snout, or a little nearer the former ; interorbital space once and three-quarters to twice as broad as the upper eyelid ; tympanum hidden. Fingers free, the first shorter than the second, which is subequal to, or a little longer than, the fourth. Toes with a rudiment of web, the fifth very short, extending but slightly, or not a t all, beyond the second ; subarticular tubercles weak ; two shovelshaped metatarsal tubercles, the inner very large. Tibio-tarsal articulation not reaching the shoulder, the tip of the fourth toe reaching the level of the tip of the snout or a little beyond. Skin smooth or slightly tubercular above ; a fold from the eye to the shoulder, and sometimes one connecting the posterior corners of the upper eyelids ; smooth beneath, except the hinder part of the belly and the anal region, which are more or less granular ; chin sometimes pustular. Olive, to fawn, or pinkish above, marbled and spotted with dark brown ; lower surfaces white, immaculate, or the throat faintly mottled with brown. Length : 55 mm. Male with a vocal sac. Eggs small, laid in large, globular masses which a t first float on the surface of the water ; the fully formed tadpole, with well-developed hind-limbs has the following characters : Head and body ovoid, once and a half as long as broad, rounded anteriorly. Nostrils slightly nearer a line connecting the anterior borders of the orbits than the end of the snout. Eyes lateral, 5 times as widely separated as the nostrils. Mouth terminal, its width ) the width of the head a t the level of the eyes ; no mouth armature. Spiraculum median, opening through a transparent tube which terminates close in front of the anus. Tail acutely pointed, 39 times as long as deep, and 1 3 times as long as the head and body ; caudal membranes moderately deep, the lower the deeper of the two. Greenish brown, the sides lighter, mottled with white ; white beneath ; tail greenish brown, with darker rnarblings and an interrupted white edging on the crests. Toes webbed as in the adult. Total length : 27 rnrn. ; head and body 10 mm. ; tail depth 5 mm. DISTRIBUTION : Southern and Eastern India as far north as the valley of the Ganges ; Ceylon.

MONOGRAPH O F T H E MICROHYLIDAE SPECIMENS EXAMINED Madras . . . . Jerdon Madras . . . . Jerdon 9 India . . . . Parry 3 ? . . . . Sir A. Smith Russelconda, Madras Trail, COTYPEof Juv. Cacopus globuPresidency losus, Gunther Beddome Balarangam Hills 9 , 5 Young Cleared Balarangam Hills . Beddome 9 and Young Malabar . . . . Beddome Larvae and Young Trevandrum, Travancore . . . . Ferguson

B.M. 46.11.22. 56-58 46.11.22. 59 48.8.19. 30 58.11.25. 6 61.12.29.3

S , 2 Hgr. Skel.

.

Uperodon globulosum (Giinther)

2.

Cacopus globulosus, Giinther, 1864, Rept. Brit. India, p. 416 (Type locality Russelconda, Madras Presidency) ; Theobald, 1868, Cat. Rept. Mus. As. Soc. Bengal, p. 82 ; Anderson, 1871, Proc. Zool. Soc. London, p. 201 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 175 ; Thurston, 1888, Cat. Batr. Sal. S. India, p. 43, P1. x ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 497 ; Sclater, 1892, Batr. Ind. Mus., p. 24 ; Nieden, 1926, Das Tierreich, Anura n, p. 20 ; Mukerji, 1933, Journ. and Proc. Asiatic Soc. Bengal xxvr~,I, p. 97, P1. I. Systoma globulosum, Cope, 1867, Proc. Acad. Philadelphia, (2), vr, p. 194. Uperodon globulosum, Parker, 1931, Archir. Zool. Ital., XVI, p. 1243, fig. 2. Prevomer entire. Snout rounded, once and two-thirds as long as the eye ; canthus rostralis obtuse ; loreal region oblique, convex ; nostril equidistant from the tip of the snout and the eye, or a little nearer the former ; interorbital space 24 to 3 times as broad as the upper eyelid ; tympanum hidden. Fingers free, the first shorter than

the second, which is a little longer than the fourth. Toes with a rudiment of web, the fifth very short, scarcely extending beyond the level of the first ; subarticular tubercles weak ; two shovel-shaped metatarsal tubercles, the inner very large. Tibiotarsal articulation not reaching the shoulder, the tip of the fourth toe just extending beyond the tip of the snout. Skin smooth or slightly tubercular above,; smooth or wrinkled beneath ; anal region granular ; an occipital fold and an indistinct one from the eye to the shoulder. Uniform brown above ; dirty white beneath. Length : 76 mm. Ovarian eggs small and very numerous. DISTRIBUTION : Madras Presidency, Central Provinces and Bengal. 9 The juvenile cotype proves t o belong to U . systoma, but as the description was drawn up almost wholly from the adult female the name globulosum is applicable to the species which it represents. B.M. 62.12.29. 4 82.2.10. 25-26

$2

9, Hgr.

Russelconda Russelconda

. . . . . . . Trail, TYPE . . . . . . . Beddome

KALO ULA, Gray Kaloukz, Gray, 1831, Zool. Misc., I, p. 38 (Type species Kaloula pulchra, Gray) ;Giinther, 1858, Cat. , 16 ; van Kampen. Batr. Sal. Brit. Mus., p. 122 ; Stejneger, 1925, Proc. U.S. Nat. Mus., ~ x v rp. 1923, Amph. Indo-Austr. Arch., p. 147 ; Shaw, 1929, Bull. Fan. Mem. Inst., I, p. 91 ; Smith, 1930, Bull. Raffles Mus., m, p. 120 ; Noble, 1932, Biol. Amph., p. 538. Calohyla (emend.), Peters, 1863, Mon. Ak. Berlin, p. 454. Callula (emend.), Giinther, 1864, Rept. Brit. India, p. 436 (part) ; Cope, 1867, Journ. Acad. Philadelphia, (2), vr, p. 192 (part) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 167 (part) ; Theobald, 1882, in Mason, Burma, I, p. 292 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 493 (part) ; Boulenger, 1912, Fauna

KALOULA

77

Malay Penins., Rept. and Batr., p. 263 (part) ; Nieden, 1926, Das Tierreich, Anura II, p. 21 (part). Caloula, Stoliczka, 1870, Journ. Asiatic Soc. Bengal, xxx~x,p. 155. Bontbznator (non Merrem), Miiller, 1836, Verh. Batav. Genootsch., p. 96. Hyladactylus, Tschudi, 1838, Mem. Soc. Sci. Neuchatel, 11, 1839, p. 48 (Type species Bombinator baleatus, Miiller) ; Hyladactyla, Tschudi, 1838, tom. cit., p. 85. Hylaedoetylua, Peters, 1863, Mon. Ak. Berlin, p. 454. P l e c t r o p , Dumbril and Bibron, 1841, Erpet. Gen., VIU, p. 736 (Type species Pleetropus pictus, Dum. and Bibr.). Holonectes, Peters, 1863, Mon. Ak. Berlin, p. 454 (Type species Hylaedactylus (Holoneetes)conjunctus, Peters). Cacopoides, Barbour, 1908, Bull. Mus. Comp. Zool. Harvard, LI, p. 321 (Type species Cacopoides borealis, Barbour).

Prevomer undivided, the post-choanal portion overlying the palatine region and raised into a strong, sometimes crenulate, ridge (fig. 33). No clavicles ; procoracoids persistent as small vestiges lying on the mesial ends of the coracoids ; a small cartilaginous omosternum ; sternum large, cartilaginous (fig. 34). Vertebral column diplasiocoelous. Terminal phalages T- or Y-shaped. Pupil horizontal. Tongue oval, entire, and free behind ; two transverse ridges across the palate in front of the pharynx ; a strong ridge behind each choana. Tips of the digitswith, or without, discs ; toes more or less webbed. In all the known species the first finger is shorter than the second and the outer toe shorter than the third.

Fig 33. Anterior cranial elements of Kaloula pulchra (seen from beneath)

Fig. 34. Ventral elements of the shoulder-girdle of Kaloula pulchra (ventral view)

MONOGRAPH O F T H E MICROHYLIDAE SYNOPSIS OF THE SPECIES I. Distal joints of fingers not wider than the penultimate ; a small supernumerary tubercle on the palm of the hand close to the proximal subarticular tubercle of each finger. A. Fifth toe extending as far as the second, or beyond ; toes of females about. ) webbed. (1) Interdigital membrane not extending beyond the level of the proximal subarticular tubercle of the third toe in ?? ; reaching the level of the distal tubercle in 88 ; inner metatarsal tubercle much shorter than its distance from the tip of the inner toe ; terminal phalanges of fingers without bony prominences . . . . . . . . . . . . K. rigida, p. (2) Interdigital membrane l extending to midway between the subarticular tubercles of the third toe in both sexes ; inner metatarsal tubercle a t least as long as its distance from the tip of the inner toe ; terminal phalanges of fingers without bony prominences . . . . . . . . . K. picta, p. (3) Interdigital membrane l extending t o midway between the subarticular tubercles of the third toe in ; reaching beyond the distal tubercle in $8 ; inner metatarsal tubercle as long as its distance from the tip of the inner toe ; terminal phalanges of fingers with bony prominences (fig. 35) K . vflmosa, p. B. Fifth toe not extending as far as the second; toes half webbed K. mediolineata, p.

79

79

80 81

11. Distal joints of fingers not wider than the penultimate ; no supernumerary tubercles a t the base of the fingers. A. Distance of nostril from anterior corner of eye a t least 8 the internarial space ; belly white . . . . . . . . . . . K. borealis, p. 82 B. Distance of nostril from anterior corner of eye about half the internarial space ; belly brown, with numerous circular yellow spots K. manchuriensis, p. 83 111. Distal joint of fingers distinctly wider than the penultimate ; usually no supernumerary tubercles on the hands. A. Web between the third and fourth toes l not extending distqlly beyond the level of the proximal subarticular tubercle of the fourth toe. (1) Discs of fingers more than twice as broad as the penultimate . K. kalingensis, p. 84 joint ; loreal region vertical . (2) Discs of fingers 1+t o 19 times as broad as the penultimate joint ; loreal region oblique K. pulchra pulchra, p. 84

. . . . . . .

B. Web between thethird and fourthtoeslextending beyond the proximal subarticular tubercle of the fourth toe (if present). but not, in females, to the level of the distal tubercle of the third. (1) Second finger distinctly shorter than the fourth. Digital webbing the same in both sexes. (a) Dark area of the back fenestrated by light, narrowly blackbordered spots, or broken up into well-defined, dark-edged K. pulchra taprobanica, p. 86 markings . (b) Dark area of the back broken up into indefinite spots, or absent . . . . . . . . . . . K. baleata, p. 88 i.e. the edge of the web midway between the toes.

KALOULA (2) Second finger as long as the fourth ; toes of males fully webbed

K. rugifera, p. 8 7 C. Web between third and fourth toes extending to the level of the distal tubercle of the third toe in both sexes. . . . . .K. conjuncta, p. 8 9 I.

Kaloula rigida, Taylor

Kaloula rigida, Taylor, 1922, Philippine Journ. Sci., xn, p. 176, P1. m, 6gs. 1 and 2 (Type locality Balbalan, Kalinga, Mountain Province, Luzon).

Post-narial ridges strong, curving backwards mesial to the inner edges of the choanae. Snout not prominent, as long as, or a little shorter than, its distance from the eye ; canthus rostralis rounded ; loreal region scarcely oblique ; interorbital space once and a half as broad as the upper eyelid ; tympanum indistinct. Fingers not dilated distally, the second a little shorter than the fourth. Toes ending obtusely, less than f webbed in females, but nearly half webbed in males; the membrane between the third and fourth toes does not extend beyond the proximal tubercle of the third toe in the former sex, but reaches the level of the distal tubercle in the latter ; fifth toe extending well beyond the second ; subarticular tubercles well developed ; two metatarsal tubercles, the inner not compressed and much shorter than its distance from the tip of the inner toe. Tibio-tarsal articulation reaching the shoulder. Skin smodth or slightly corrugated above ; a fold from the eye to the fore-limb, the skin above it somewhat thickened ; sometimes an occipital fold is present ; belly smooth ; chin and hinder side of the thighs slightly granular. Brownish red to deep purple above, uniform or with darker markings in the form of a median spot commencing on the back of the head, narrowing on the nape, broadening abruptly between the shoulders, narrowing and bifurcating in the sacral region, with one branch running to each groin ; loreal region, temples, anterior aspect of femur and anal region sometimes black ; limbs black-spotted. Lower surfaces pale brown with indistinct lighter spots and reticulations. Male with a subgular vocal sac ; throat infuscate. Length : 4 7 mm. DISTRIBUTION : Luzon, Philippine Islands. SPECIMENS EXAMINED B.M. 1924.2.12. 3 $ Balbalan, Luzon . . . . Taylor 1928.3.15.2 $ Balbalan, Luzon . . . . Taylor . . . . Taylor Mus. Amsterdam Balbalan, Luzon 2.

Kaloula picta (Durn. and Bibr.)

Plectropus pietus, Dumbril and Bibron. 1841, Erpet. Gen., w, p. 237 (Type locality Manilla); Bibron, 1842, in Eyd. and Soul., Voy. Bonite, Rept., p. 152, P1. IX, Qs. 3 and 4 ;Peters, 1863, Mon. Ak. Berlin, p. 465. Kaloula pictu, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 123 ; Taylor, 1920, Phil. Journ. Sci., XVI, p. 322, P1. IX. fig. 4. C&L picta, Giinther, 1864, Rept. Brit. India, p. 436 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 168 ; Boettger, 1892, Kat. Batr. Senck. Nat. Ges., p. 23 ; Nieden, 1026, Das Tierreich, h a 11, p. 26.

Post-narial ridges moderately strong, curving backwards mesial to the inner edges of the choanae. Snout not prominent, subacuminate as seen from above, as long as, or a little shorter than, the diameter of the eye ; canthus rostralis obtusely angular ; loreal region slightly oblique ; interorbital space once and a quarter as broad as the upper eyelid; tympanum indistinct or hidden, about half the diameter of the eye. Fingers not dilated distally, the second subequal to, or a little shorter than, the fourth ; a small supernumerary tubercle just proximal to the basal subarticular tubercle of each finger. Toes about f webbed in both males and females, the web between the third and fourth toes reaching to midway between the tubercles of the third toe ; fifth toe extending well beyond the second ; subarticular tubercles well developed ; two compressed metatarsal tubercles, the inner very large. Tibio-tarsal articulation reaching the shoulder.

MONOGRAPH O F THE MICROHYLIDAE Skin smooth or slightly pustular above ; an indistinct fold from the posterior corner of the eye along the flanks, and another to the forearm ; the skin between these two folds thickened and glandular ; sometimes an indistinct transverse occipital fold. Smooth beneath except the throat, posterior part of the belly and hinder side of the thighs in the anal region, which are slightly granular. Purplish brown or dark grey above with the following dark pattern. A large dorsal spot commencing as a branch from each eyelid, which unite on the nape ; this spot broadens on the scapular region, narrows abruptly and breaks up into four branches on the sacral region. The two median branches run straight back to the vent, whilst the laterals run to the groin, whence they continue as oblique bars across the femur, tibia and metatarsus. A dark line along the canthus rostralis. Flanks below the dorsolateral fold darker ; an indefinite light streak below the supratympanic fold. Belly yellowish olive, the throat and chest marbled and spotted with darker. Male with a subgular vocal sac. : Philippine Islands (Luzon, Negros, Mindoro and ? Dumaran). DISTRIBUTION SPECIMENS EXAMINED B.M. 61.8.12.56-57 72.8.20.12-13 72.8.20. 7 8 92.8.4. 1 5 R.1929.5.20. 1

8 Hgr. Hgr.

? . . . . . Negros. . . . Laguna de Bay . Manilla . Philippine Islands

. . . . . . . . . . . .

Bowring Meyer . Meyer . Walker

. . . . . .

3 . Kaloula verrucosa, Boulenger Cdlula ~e:erm8a, Boulenger. 1904, Ann. Mag. Nat. Hist., (7). xm,p. 131 (Type locality Yunnan-fu) ; Men, 1922, h c h . f. Naturg., Lxx-, A, p. 130; Vogt, 1922, Arch. f. Naturg., L X X X ~ I ,A, p. 145 ; Werner, 1924, Denkschr. Ak. Wiss. Wien, xcIx. p. 51 ; Nieden, 1926, Das Tierreich, Anura n, p. 27. Ka.bula v e m o s a , Schmidt, 1927, Bull. Amer. Mus. Nat. Hist., m,p. 562 ; Gee and Boring, 1930, Bull. Dept. Biol. Yenching Univ., I, p. 26 ; Boring, 1930, Peking Soc. Nat. &t. Bull., v, p. 43, Map x ; Pope, 1931, Bull. Amer. Mus. Nat. Hist., LXI, p. 690.

Fig. 35. Terminal phalanx of finger of breeding male Kabula vermeosa

Fig. 36. Dorsal surface of tip of second finger of male KaIaula v e m w s a

Post-narial ridges moderately strong, curving slightly backwards mesial to the inner edge of the choanae. Snout not prominent, rounded when seen from above, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region oblique ; interorbital space once and a quarter the width of the upper eyelid ; tympanum hidden. Fingers slightly knobbed, the second subequal to the fourth ; a small supernumerary

KALOULA tubercle a t the base of each finger, just proximal to the basal subarticular tubercle. Toes about 4 webbed in females, the membrane between the third and fourth toes reaching to midway between the subarticular tubercles of the third toe ; in males the toes are almost fully webbed, the membrane between the third and fourth extending beyond the distal tubercle of the third toe ; outer toe a little longer than the second ; subarticular tubercles well-developed, except the proximal one of the fourth toe, which may be small or wanting. Two compressed metatarsal tubercles, the inner very large. Tibio-tarsal articulation reaching the shoulder. Skin with rows of large warts above ; an indistinct fold from the eye to the fore-limb, and an indefinite occipital fold connecting the posterior corners of the upper eyelids. Lower surfaces smooth ; anal region granular. Brownish grey above, uniform or with rows of indistinct darker spots ; uniformly white beneath, the throat of the breeding male infuscate. Male with a subgular vocal sac. Nuptial excrescences of a kind hitherto unknown in the Salientia are developed. These consist of one or two rows of small pustules on the upper surface of the tip of each finger (fig. 36) ; the number of pustules varies from three or four to eight or ten. Each has a bony core derived from the terminal phalanx (fig. 35), but there is no apparent thickening of the epidermis, and no development of horny spicules, glands or sensory apparatus. In females, also, the terminal phalanx bears more ox less developed processes of a similar nature which may produce slight irregularities of the skin on the upper surface of the finger-tip. These irregularities are, however, much smaller and scarcely distinguishable. Length : 9 61 mm. ; d 45 mm. DISTRIBUTION : Yunnan and Hunan, S.W. China. SPECIMENS EXAMINED B.M. 1904.1.26.18-20 69 Yunnan-fu . Graham, COTYPES 1904.11.29.40 1905.5.30.42-46 1907.5.4. 22-27 1907.5.4. 28 1905.5.30.41

1

.

dd*9q

Hgr. Yg. Skel.

?

. .

. . .

Yunnan-fu . . Yunnan-fu . Tongchuan-fu, Yunnan

. . .

Graham Graham Graham

4 . Kaloula mediolineata, Smith Callulu mediolineata, Smith, 1917, Journ. Nat. Hist. Soc. Siam, 11, p 224, PI. (Type locality Prachuap Kirikan, S.W. Siam).

Post-narial ridges strong, straight. Snout short, rounded, as long as, or a little shorter than, the eye ; canthus rostralis rounded ; loreal region oblique ; interorbital space once and a half as broad as the upper eyelid ; tympanum hidden. Fingers moderate, without discs ; the second as long as the fourth ; a small supernumerary tubercle a t the base of each finger just proximal to the basal subarticular tubercle. Toes about half webbed, the membrane midway between the third and fourth toes reaching nearly to the level of the distal tubercle of the third toe and continued almost to the tips of all except the fourth ; fifth toe short, not extending as far as the second ; l subarticular tubercles well developed ; two strong, compressed, sharp-edged metatarsal tubercles, the inner the larger. Tibio-tarsal articulation reaching the shoulder, or not quite so far. Skin smooth or with a few flat warts above ; an indistinct fold from the posterior corner of the eye to the shoulder. Smooth or slightly wrinkled beneath ; anal region and hinder side of thighs granular. A large, dark blotch covering almost the whole of the back, commencing between the eyes, narrowing slightly on the nape and then broadening steadily to the groin ; a lateral stripe of the same colour from behind the eye to the groin, the two separated by a clearly defined light streak ; a light bar connects the anterior halves of the upper eyelids, and there is-a short median white stripe from the anus along the coccygeal In juveniles the outer toe is relatively slightly longer and may extend as far as the second. 6

82

MONOGRAPH O F T H E MICROHYLIDAE

region. A light spot above the fore-limb and sometimes indistinct, irregular light spots on the flanks, hind-limbs and back. Lower surfaces white ; the throat of the male black. Male with a subgular vocal sac. Eggs small ; a free-living tadpole. Length : 6 3 mm. DISTRIBUTION : S.W. Siam (Prahcuap Kirikan, Palmampo, Korat, Ubon). B.M. 1916.4.17.3 1917.5.14.4144 1921.4.1.279-283

juv. juvs.

. . . ..

Nong Pling, Paknampo Prachuap Kirikan 3 3 3 , 2 $? $? Ban Nong, Korat

Smith

5 . Kaloula borealis, Barbour. Callula vermcosa (non Boulenger), Wolterstorff, 1906, Zool. Anz., xxx, p. 554 ; idem, 1906, Abh. Mus. Magdeburg, I, pp. 128, 131, 145, PI. I ; Barbour, 1909, Proc. Biol. Soc. Washington, XXII, p. 89 ; Namiye, 1913, Japan. Zool. Mag., XXV, pp. 416, 421423; Nieden, 1926, Das Tierreich, Anura 11, p. 27 (part). Cacopoides borealis, Barbour, 1908, Bull. Mus. Comp. Zool. Harvard, LI, p. 321 (Type locality Antung, , Manchuria) ; Barbour, 1909, Proc. Acad. Philadelphia, p. 401, P1. ; Slevin, 1925, Proc. California Acad. Sci., xrv, p. 91. Kaloula borealis, Noble, 1925, Amer. Mus. Novitates, o ~ x vp. , 6; Schmidt, 1927, Bull. Amer. Mus. Nat. Hist., LIV, p. 561 ; Shaw, 1929, Bull. Fan Mem. Inst., I, p. 91, fig. 12 ; Gee and Boring, 1929, Peking Sac. Nat. Hist. Bull., rv, 2, p. 25 ; Liu and Li, 1930, Peking Soc. Nat. Hist. Bull., rv, 4, pp. 67-91 ; Boring, 1931, Peking Soc. Nat. Hist. Bull., V, p. 39, Map 10 ; Liu, 1931, Peking Soc. Nat. Hist. Bull., v, pp. 52-58 ; Fang and Chsng, 1931, Contr. Biol. Lab. Sci. Soc. China, vm,Zool., p. 89, fig. 8 ;Chen, 1931, Misc. Notes Zool. Chekiang and Kiangsu (not seen) ; Pope, 1931, Bull. Amer. Mus. Nat. Hist., LXI, p. 587, fig. 36 ; Boring and Liu, 1932, Peking Soc. Nat. Hist. Bull, VI, 3, p. 19 ; Hsiao, 1932, China Journal, r v n , 4, p. 184, PI. ; Callula tornieri, Vogt, 1913, Sitzber. Ges. Natf. Fr. Berlin, p. 219 (Type locality Korea). Cacopokh tornieri, Mori, 1927, Prep. Dept. Keijo Imp. Univ., Seoul, Korea (not seen) ; Okada, 1928, Journ. Chosen Nat. Hist. Soc., VI, p. 25, P1. I fig. i ; Mori, 1928, Journ. Chosen Nat. Hist. Soc., VI, p. 48; Okada, 1931, Mon. Jap. Tailless Batr., pp. 6k68, PI. v, figs. 1-2, P1. xvm, fig. 5, P1. xxrv, fig. 1 ;idem, 1931, Tailless Batr. Jap. Emp., p. 75, P1. v, figs. 1-2, PI. xvm, fig. 5, P1. xxrv, fig. i. Kaloukz tornieri; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., N, (2), p. 26 ; Boring, 1931, Peking Soc. Nat. Hist. Bull., v, p. 47. Kaloda woltersto$, Stejneger, 1925, Journ. Washington Ac. Sci, xv, p. 151 (Type locality Nanking) ; Sohmidt, 1927, Bull. Amer. Mu. Nat. Hist., LIV, p. 561 ; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., N, 2, p. 26; idem, 1931, op. oit., v, p. 42, Map 10 (part); Pope, 1931, Bull. Amer. Mus. Nat. Hist., LXI, p. 587. K . wolterstorfi, Shaw, 1929, Bull. Fan. Mem. Inst., I, 5, p. 97.

Post-narial ridges strong, straight, dilated mesially. Snout short, rounded, about as long as the diameter of the eye, not prominent ; nostril nearer the tip of the snout than the eye, its distance from the latter more than 3 the internarial space ; canthus rostralis rounded ; loreal region oblique ; interorbital space about once and a half to twice the width of the upper eyelid ; tympanum hidden. Fingers not dilated, without supernumerary tubercles ; the second a little shorter than the fourth. Toes short, about half webbed, the membrane between the third and fourth reaching halfway between the subarticular tubercles of the third ; fifth toe extending a little beyond the second ; subarticular tubercles well developed, though the proximal one of the fourth toe may be absent ; two compressed metatarsal tubercles, sometimes almost contiguous, the inner very large, about twice as long as the inner toe. Tibio-tarsal articulation reaching the shoulder. Skin smooth, or with more or less small, scattered warts, particularly in males ; an indistinct fold from the eye to the shoulder, and sometimes one across the occiput connecting the posterior corners of the upper eyelids. Lower surfaces smooth ; anal region granular.

KALOULA Brown, greenish or grey above, uniform, or with some irregularly scattered dark spots ; occasionally a faint light W-shaped marking may be discerned in the scapular region. Flanks lighter with round yellow spots and marblings which may extend on t o the upper surfaces and on to the chin and lower surfaces of the limbs. Belly white, rarely with traces of the colour of the flanks spreading towards the middle. Hinder side of thighs, the tibiae, tarsi and feet light-spotted. Male with a vocal sac and infuscate gular region. In this sex the m. obliquus abdominis ezternus and m. obliquus abdominis internus are greatly thickened and form a very thick, double, bright pink aponeurosis with the rectus abdominis, and one or more of the tendinous inscriptions of the latter muscle may also have the same colour. This brightly coloured aponeurosis is visible through the skin of living specimens, at least in its middle and posterior parts ; anteriorly, where it runs beneath the m. pectoralis it is hidden. There does not appear to be any glandular structure, as has been alleged, associated with this remarkable secondary sex character. Length : 51 mrn. ; 54 mrn. DISTRIBUTION : Kiangsu, Shantung, Shansi, Hopei, Manchuria, Korea and Quelpart Island. Eggs small, 1.4 mm. ; laid a t night in shallow pools or transient puddles, where they form a thin floating sheet. The breeding season varies with the incidence of the rainy seasonb(July-August). The whole larval period may be less than two weeks. A well-grown tadpole may be 37 mm. in total length, and has the following characters : " Length of body about 1J times its width ; tail 1) times length of body, 24 times as long as deep. Nostrils nearer to line joining anterior corners of eyes than to tip of snout. Eyes lateral, the distance between them about 6 times that between the nostrils. Spiraculum median, large, opening under a transparent sheath below posterior end of body. Anus median, opening just below edge of subcaudal crest. Tail pointed ; crests convex, about equal in depth. Mouth terminal, lacking horny beak and teeth, its width not more than 3 the interocular space. Uniform dark above, white beneath, the belly skin translucent. Sides and tail crests mottled " (Pope). Newly transformed frogs measure 15-17 mm. in length. SPECIMENS EXAMINED ? Tsingtao, Shantung

B.M. 1905.11.27. 3 1925.11.4. 10-11 1927.2.11. 1-4 1928.2.27. 3-7

Boring Collection

3 3 2

d? bd 8, 2 $?$2 b 3 . 1 $?

99

2 d , 2 $?$?

. Kreyenberg (Descr. Wolterstorff) Kichowpeh, Hopei . Pope Tsinanfu, Shantung . Hindle Tsingtao, Shantung . Hindle Nanking, Kiangsu . Nr. Peking . . . Taian, Shantung . .

6 . Kaloula mamhurimis, Boring and Liu Kaloltla manchuriensia, Boring and Liu, 1932, Peking Soc. Nat. Hist. Bull., VI, 3, p. 21, P1. (Type locality Eastern Tomb, Mukden).

Post-narial ridges strong, straight, dilated mesially. Snout short, rounded, about as long as the diameter of the eye, not prominent ; nostril midway between the tip of the snout and the eye, its distance from the latter about half the internarial width ; canthus rostralis rounded ; loreal region oblique ; interorbital space once and a half the width of the upper eyelid; tympanum hidden. Fingers not dilated, without supernumerary tubercles ; the second and fourth subequal. Toes about half webbed, the membrane between the third and fourth reaching half-way between the subarticular tubercles of the third ; fifth toe extending a little beyond the second ; subarticular tubercles rather feeble ; two, contiguous, compressed metatarsal tubercles, the inner very large, about twice as long as its distance from the tip of the inner toe. Length of hind-limb (from vent to tip of fourth toe with the leg a t right angles to the body) equal to the distance from the vent to the anterior corner of the eye.

MONOGRAPH O F T H E MICROHYLIDAE Skin of upper surface much thickened and slightly warty ; a strong supratympanic fold, continuous with a transverse occipital one. Lower surfaces smooth. Brown above, with some small, scattered, dark flecks; flanks, venter and lower surfaces of the limbs brown, with numerous, regular, closely set yellow spots, the ground colour becoming lighter towards the middle of the belly. Male with a vocal sac and infuscate gular region. Aponeuroses of the m. obliqui abdominis and the tendinous inscriptions of the m. rectus abdominis thickened and coloured as in the preceding species. Length : 57.5 mm. DISTRIBUTION : Known only from the type locality. It seems probable that this species will be found t o be based on an abnormally large, unusually coloured example of the preceding species. The only morphological characters separating the two (position of nostril and interorbital width) are approached by the largest specimens of K. borealis examined (Kichowpeh, 48 mm. from snout t o vent), and the unique secondary sex character of the male is an additional reason for suspecting the two to be conspecific. Boring Coll.

8

Eastern Tomb, Mukden

. . . . . . . . Liu

TYPE

7 . Kaloula kalingmis, Taylor Kaloula kalingensis, Taylor, 1922, Philippine Journ. Sci., xxr, 2, p. 178, P1. rn, figs. 1 and 2 (Type locality Balbalan, Kalinga, Luzon). Snout shorter than the eye ; canthus rostralis rounded ; loreal region perpendicular ; interorbital space once and a half as broad as the upper eyelid ; tympanum distinct,

+

about the diameter of the eye. Fingers with large triangular dilatations which are twice as broad as the penultimate joint ; the second as long as the fourth. Toes slightly dilated distally, without, or with only a slight, rudiment of web ; fifth toe longer than the second ; subarticular tubercles well developed, but the proximal one of the fourth toe absent; two strong metatarsal tubercles, the inner the larger. Tibio-tarsal articulation reaching the shoulder. Skin with flattened pustular tubercles above and on the sides ; a fold of skin from the eye above the forearm, becoming thickened and glandular posteriorly ; another fold from the eye to the insertion of the fore-limb. Smooth beneath, except the belly and femur, which are strongly granular. Bluish black above with deep red markings on neck, sides and limbs ; minute cream markings on each side of the anus ; belly and under side of limbs mottled with creamy ' white ; subarticular tubercles white ; throat and chin blue-black. Male with a subgular vocal sac. Length : 36'5 mm. : Kalinga, Luzon, Philippine Islands. DISTRIBUTION

8. Kaloula pulchra pulchra, Gray Kdoula pukhrc, Gray, 1831, Zool. Misc., p. 38 (Type locality " China ") ;Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 123 (part) ; Bteindachner, 1867, Reise Novara, Amph., p. 68 ;Barbour, 1912, Mem. Mus. Comp. Zool. Harvard, xm, pp. 72 and 173, P1. vn, fig. 29 ; Annandale, 1917, Mem. As. Soc. Bengal, m, pp. 146, 152, P1. m, figs. 7-7b; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 150; Mertens, 1929, Zool. Anz., ~xxxvr,p. 63; idem, 1929, Bliitt. Aq. Terr., XL, 6, p. 100 ; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull, rv, 2, p. 25 ;Mertens, 1930, Abh. Senck. Nat. Ges., XLU, 3, p. 237 ; Smith, 1930, Bull. Raffles Mus., m, p. 121, fig. 10; Boring, 1931, Peking Soc. Nat. Hist. Bull., v, p. 43, Map 1 0 ; idem, 1932, op. cit., vr, 3, p. 19. Cdlula pulchra, Giinther, 1864, Rept. Brit. India, p. 437 ; Blanford, 1870, Journ. Asiatic Soc. Bengal, xxxrx, 4, p. 375 ; Theobald, 1882, in Mason, Burma, I, p. 293 ; Boulengcz, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 170 (part) ; Miiller, 1887, Verh. Nat. Ges. Bascl, vm, p. 251 (part) ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 494 (part) ; Sclater, 1892, Batr. Ind.

KALOULA

85

Mus., p. 23 ; Boettger, 1892, Kat. Batr. Senck. Nat. Ges., p. 2 4 ; Flower, 1896, Roc. 2001. Soc. London, p. 908 ; Mbhely, 1897, Termbs. Fiizetek, xx, p. 68 ; Flower, 1899, Proc. Zool. Soc. London, p. 906 ; Schenkel, 1901, Verh. Nat. Ges. Basel, xrrr, p. 152 ; P. and F. Sarasin, 1901, Mat. Naturg. Celebes, 111, pp. 72 and 7 6 ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 390 ; Wolterstorff, 1906, Abh. Mus. Magdeburg, I, p. 131 ; van Kempen, 1907, in Weber, Zool. Ergeb. Reise 0-Ind., rv, p. 412 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 264, fig. 73 ; Mell, 1922, Arch. f. Naturg., Lxxxvm, A, p. 131 ;Nieden, 1926, Das Tierreich, Anura 11, p. 22, fig. Caloula pulch~a,Stoliczka, 1870, Journ. As. Soc. Bengal, xxxix, 2, p. 155. Calohyla pulcka, Peters and Doria, 1878, Ann. Mus. Genova, xm, p. 429. Hylaedactylus bivittdus Cantor, 1847, Journ. Asiat.ic Soc. Bengal, XVI, p. 1064 (Type locality Malay Peninsula). Callula m ~ o d a c t y l a ,Boulenger, 1887, Ann. Mus. Genova, (2), v, p. 485, P1. vm, fig. 3 (Type locality Kawkariet, Tenasserim) ; idem, 1890, Fauna, Brit. India, Rept. and Batr., p. 494 ; Nieden, 1926, Das Tierreich, Anura 11, p. 25. Post-narial ridges strong, straight ; snout short, rounded, about as long as the eye ; canthus rostralis rounded ; loreal region oblique ; interorbital space one and a half to twice as broad as the upper eyelid ; tympanum hidden. Fingers with small, truncate dilatations distally which measure 1: to 1 4 times the diameter of the penultimate joint ; second a little shorter than the fourth. Toes short, scarcely dilated, with a

rudiment of web which, between the third and fourth toes, does not extend beyond the proximal subarticular tubercle of the fourth toe ; fifth toe extending a little beyond the second ; subarticular tubercles well developed ; two compressed metatarsal shovels, the inner as long as, or a little longer than, its distance from the tip of the first toe. Tibio-tarsal articulation reaching the shoulder. Skin smooth or with irregular flat warts above ; an indefinite fold from the eye to the forearm and sometimes one across the occiput. Lower surfaces smooth or faintly granular. A dark triangular spot occupying the whole of the back from the middle of the eyelids, and a lateral streak of the same colour from the posterior corner of the eye, the two being separated by a yellow dorso-lateral stripe ;juveniles with a light forehead and a light spot on each side of the vent, another in the groin, and others across the upper arm, wrist, knee, heel and instep. More or less distinct traces of some or all of these markings may persist in the adult. Lower surfaces yellowish, faintly mottled with light brown ; throat of the male infuscate. Male with.a vocal sac. Eggs small, laid in floating masses ; development very rapid. Description of Tadpole with well-developed hind-limbs.-Head and body once and a half as long as broad ; upper surface flattened ; snout broad and truncate. Nostrils (not visible in specimens without fully developed legs) close together on the upper surface of the snout, nearer a line connecting the anterior borders of the eyes than the tip of the snout. Eyes lateral, six to seven times as widely separated as the nostrils. Mouth very small, terminal, without beak or teeth. Spiraculum median, prolonged posteriorly as a transparent tube, attached to the belly dorsally and opening a t the level of the middle of the thighs. Anus median, the anal tube curving downwards and backwards and projecting below the lower caudal membrane. Tail nearly twice as long as the head and body, pointed, with subequal, convex crests whichat their deepest part are about as broad as the muscular portion of the tail a t its base. Toes with the same amount of webbing as the adult. Olive brown to black above, sometimes with golden speckles ; lighter, white spotted beneath; a dark line connecting the nostril with the anterior corner of the eye ; sometimes a light bar across the snout and light markings on the sides. Caudal membranes colourless or with small dark patches. Total length : 40 mm. ; head and body 14 mm. ; breadth of body 9 mm. ; depth of tail 7.5 mm. DISTRIBUTION : S. China, as far east as Hong Kong, Tonkin, Hainan, Burma, Bengal Siam, Annam, Malay Peninsula, Sumatra, Borneo (?), Celebes and Flores (?). The exact, natural distribution of the species is difficult to determine accurately; it is known to have been introduced into Singapore, and records of it in the Malay Peninsula are scanty. In the Malay Archipelago it is known definitely to occur in Sumatra, and in Celebes around Macassar, but the other records (Borneo and Flores)

MONOGRAPH O F T H E MICROHYLIDAE are unconfirmed ; it seems probable that the species may have been introduced into Macassar, and possibly into other localities, just as it was into Singapore. SPECIMENS EXAMINED B.M. R.1929.6.2.1-2 9,H g . China . Reeve China . . . . . . . R.1929.6.2. 3-4 juvs. Hgr. China . . . . . . . 51.12.27. 10 93.9.14. 6 9 Hong Kong . Walker 88,$2 Hoi-How, Hainan . . Neumann 93.1.3. 24-26 Toungyi, Shan States, 5,000 ft. Blakeway 91.11.26.46-49 ? , 3 Hgr. 66.7.10. 8 Hgr. Moulmein, Burma . . Beavan He. Burma . . Day 70.8.14. 11 Hgr. Toungoo District . . Oates 91.11.26. 50 8,0 Siam . Mouhot 59.7.1. 3 0 3 1 93.9.6. 10 juv. Bangkok, Siam . Greening 98.11.8.71-72 8 and juv. Bangkok . Flower Bangkok . M. A. Smith 191 6.4.17.4-13 larvae 8 Bangkok . M. A. Smith 1927.4.21-26 juvs. Chantaboon, Siam. . Flower 98.11.8. 73-76 ? Cambodia . . Mouhot 61.8.12.5 62.1.9. 10 Skel. Cambodia . . Mouhot Saigon . . HolbC 1920.1.20. 3669 8, 1920.1.20. 3822 2 9 , 2 juvs. Saigon . . . . . . . HolbC 8 Malacca. . Cantor 60.3.19. 1095 8 Jalor, F.M.S. . . Annandale and 1903.4.13. 124 Robinson 1903.4.13. 125 juv. Selangor, F.M.S. . . Annandale and Robinson 96.6.25. 88-93 8, Singapore . . Flower 96.12.9.105 $! Macassar, Celebes . . Sarasin 1926.8.20. 128 $? Macassar, Celebes . . M. A. Smith

9. Kaloula pulchra taprobanica subsp. n. Kaloula pulchra (part), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 123. Callula pulchra, Giinther, 1864, Rept. Brit. India, p. 437 (part) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 170 (part) ; Miiller, 1887, Verh. Nat. Ges. Basel, vm, p. 251 (part) ; Sarasin, 1910, Zool. Jahrb., Supp. XII, i, p. 29. Holotype a number 1905.3.25. 138 in the British Museum, from Punduloya, Ceylon (4,000 ft.). *

9,

Post-narial ridges strong, curving backwards slightly mesially. Snout short, rounded, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space once and a half as broad as the upper eyelid ; tympanum hidden. Fingers with weU developed truncate discs which are about twice as wide as the penultimate phalanges ; the second shorter than the fourth. Toes moderate, ending obtusely, about ) webbed, the membrane between the third and fourth reaching to midway between the proximal tubercle of the fourth and the distal tubercle of the third toes ; fifth toe extending well beyond the second ; subarticular tubercles well developed ; two strong compressed metatarsal tubercles. Tibio-tarsal articulation reaching to about the shoulder. Skin with a few, small, scattered pustules above ; granular beneath the throat, on the hinder part of the belly and in the anal region. Colour pattern derivable from that of K, pulchra, and consisting of a dark median area separated from a dark lateral stripe and a dark snout by pale dorso-lateral bands and an interorbital bar of the same colour ; the median area is, however, fenestrated and broken up by irregular spots and patches of the paler colour of the lateral stripes, these fenestrations being sharply defined and narrowly edged with black ; a diagonal light stripe from the eye t o beneath the fore-limb. Lower surfaces yellowish, closely

KALOULA

87

spotted and marbled with dull olive brown. Length from snout to vent 56 rnm. ; fore-limb 32 mm. ; hind-limb 63 mm. The 6 paratypes show little variation in general morphological characters, and only a little in their colouring. All have the same general type of colour pattern, but the degree of fenestration and breaking up of the dorsal dark patch varies. In the specimen in which it is least marked the dorsal spot fuses with the lateral stripe posteriorly, and there are a pair of large oval light spots on the sacral region ; in others the fenestration is more marked but the light interorbital bar and dorso-lateral stripes are always clearly defined anteriorly. Specimens from Calcutta appear t o be intermediate between this race and the typical form ; the webbing of the toes is similar to that of Siamese specimens, but the colour pattern is that of the Ceylon subspecies. Male with a subgular vocal sac. In its general characters,particularly the size of the digital discs, proportions of the toes and interdigital webblng K. p. taprobanica is very similar to K. baleata. The geographical range of taprobanica is, however, well separated from that of K. baleata, and though there is little by which to distinguish the two, the colour pattern appears , to afford a perfectly reliable diagnostic character ; in a series of 26 specimens of baleata there is not one which resembles taprobanica in the slightest degree. : India (south of the Ganges), and Ceylon. DISTRIBUTION SPECIMENS EXAMINED $2 Punduloya, Ceylon, 4,000 ft. Green. TYPE B.M. 1905.3.25. 138 56.1.17. 1 $ Ceylon . . . . . . Keelart 74.4.29.1235 5 Ceylon. . . . . . . Beddome 76.3.21. 20 $2 Ceylon. . . . . . . Ferguson PARATYPES 1908.7.2. 14-15 $2 $2 Kummgda, Ceylon . . . Green 74.4.29.212 $2 Cauvery River, Madras . . Beddome Mus. Ind. $ $2 . Calcutta . . . . . . Anderson

.

I

I o Kaloula rugifera, Stejneger Kaloula rugifera, Stejneger, 1924, Occ. Papers Boston Soc. Nat. Hist., v, p. 119 (Type locality Kiatung, Szechwan) ; idem, 1925, Proc. U.S. Nat. Mus., LXVI, 25, p. 16 ;Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., IV, 2, p. 25 ; Boring, 1931, Peking Soo. Nat. Hist. Bull., v, p. 42, Map 10.

Post-narial ridges strong, curving backwards mesially, sometimes in contact. Snout rounded, as long as, or a little longer than, the diameter of the eye ; canthus rostralis rounded ; loreal region oblique, slightly concave ; interorbital space once and a half as wide as the upper eyelid ; tympanum indistinct or hidden. Fingers slender, with small truncate discs ; the second subequal to the fourth. Toes slender, swollen distally, about webbed in females, fully webbed in males ; subarticular tubercles well developed ; two compressed metatarsal tubercles, the inner the longer, the outer transverse. Tibio-tarsal articulation reaching beyond the shoulder. Upper surfaces and sides rough, with numerous elongate, wrinkled tubercles; transversely wrinkled beneath (due to preservation ?) ; preanal region granular ; a slight dermal fold from eye to shoulder. Dark brownish grey above, uniform or with a broad pale band, interrupted in the middle, across the neck between the fore-limbs ; this band edged with a series of small black spots. Back with small black spots which form a narrow band across the supraorbital region ; a line on the upper lip and indications of cross-bars on the legs and feet ; underside light brownish grey, uniform or with numerous round white spots on the chin and throat ; subarticular tubercles whitish. Male with a vocal sac. Length : 42 mm. DISTRIBUTION : Sze-chwan, China. SPECIMENS EXAMINED Shin Kai Si, Mt. Omei, Sze-chwan Graham $ B.M. 1933.5.18. 1 U.S. Nat. Mus. 75368 $2 Kiatung, Sze-chwan . . . Graham (TOPOTYPE)

+

MONOGRAPH O F THE MICROHYLIDAE II.

Kaloula baleata (S. Miiller)

Bombinator baleatus, Miiller. 1836, Verh. Batav. Genootsch., p. 96 (Type locality Krawang, Java). Hylaductylus baleatus, Tschudi, 1838, Mem. Soc. Sci. Neuchitel, 11, 1839, p. 85; Hykzedactylus baleatus, Dum. and Bibr., 1841, Erpet. Gbn., vm, p. 734 ; Kaloula baleata, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 122 ; Steindachner, 1867, Reise Novara, Amph., p. 68 ;Barbour, 1912, Mem. Mus. Comp. Zool. Harvard, XLW, p. 72 ;Taylor, 1920, Philippine Journ. Sci., x n , p. 324 ; Smith, 1922, Journ. P.M.S. Mus., x, p. 282 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 148, fig. 21 ;Mertens, 1927, Senckenbergiana, IX, 6, p. 242 ;Dunn, 1928, Amer. Mua. Novitates, cxv, p. 4 ; Mertens, 1929, Senckenbergiana, XI, p. 2 6 ; idem, 1929, Bl%tt. Aqu. Terr., XL, 6, p. 100 ;idem, 1930, Abh. Senck. Natf. Ges., xrn, 3, p. 235 ; Smith, 1930, Bull. Raffles Mus., 111, p. 121 ;van Kampen and Brongersma, 1931, Treubia, xrn, i, p. 15. Callula baleata, Cope, 1867, Journ. Acad. Philadelphia, (2), n , p. 192 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 169 ; Boettger, 1892, Kat. Batr. Senck. Nat. Gea., p. 2 3 ; idem, 1900, Abh. Senck. Natf. Ges., xxv, p. 338, footnote ; Schenkel, 1901, Verh. Nat. Ges. Basel, xm,p. 152 : Boettger, 1903, Abh. Senck. Natf. Ges., xxv, p. 384 ; Roux, 1904, Verh. Nat. Gea. Basel, xv, p. 433; van Kampen, 1909, Nat. Tijdsch. Ned. Ind., LXIX,p. 47 ; idem, 1915, in Kleinweg de Zwaan, Die Insel Nias bei Sumatra, p. 280 ; Nieden, 1926, Das Tierreich, Anura 11, p. 26. Hykzdactylus lividus (nom. nud.), Bleeker, 1857, Nat. Tijdsch. Ned. Ind., xrv, p. 239. Hylaedactylus celebensis, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 122 (Type locality " India "). Calohyla celebensis, Peters, 1872, Mon. Ak. Berlin, p. 585. ? Bzlfo biporcatus (non Tschudi), Elbert, 1912, Sunda Exped. n, p. 101. Kaloula pulchra (non Gray):Dunn, 1927, Amer. Mus. Novitates, c c ~ x x x mp. , 5.

Post-narial ridges strong, curving sharply backwards from the inner edges of the choanae. Snout rounded, as long as, or a little shorter than, the diameter of the eye ; canthus rostralis rounded ; loreal region moderately oblique ; interorbital space 1$t o 1& times as wide as the upper eyelid ; tympanum indistinct or hidden. Fingers dilated to form large truncate discs which are once and a half to twice as wide as the penultimate phalanges ; the second a little shorter than the fourth. Toes feebly dilated distally, about 3 webbed ; the membrane between the third and fourth toes reaching beyond the level of the proximal subarticular tubercle of the fourth toe, but not beyond the distal tubercle of the third ; fifth distinctly longer than the second ; subarticular tubercles well developed, but the proximal tubercles of the fourth and fifth toes often indistinct or absent; two large metatarsal tubercles, the outer smaller and transverse. Tibio-tarsal articulation reaching the shoulder or the ear. Smooth or pustular above ; often a fold from the eye to the insertion of the fore-limb, and sometimes an indistinct one from the eye above the shoulder ; an occipital fold sometimes present ; lower surfaces smooth or granular. * Brown or purplish above, with or without traces of a colour pattern similar to that of K. pulchra, but with the dorsal and lateral spots always broken up into irregular, ill-defined smaller markings ; white spots often present in the groin, beside the anus, behind the knee, on the heel and foot, and on the upper arm. Lower surfaces pale brown, profusely marbled and spotted with darker, or sepia coloured, dotted with lighter. Male with a vocal sac. Eggs small. The tadpole agrees in the position of the eyes, vent and spiracle and in the characters of the mouth with that of K. pulchra. Length : 7 6 mm. DISTRIBUTION : Malay Peninsula and Archipelago (Nias, Sumatra, Java, Borneo, Celebes, Tanah Jampeah, Komodo, Sumbawa, Sumba, Lombok, Bali and Flores) and Philippine Islands (Luzon). Taylor (1920) has cast doubt on the records of K. baleata from Laguna de Bay, suggesting that the specimens were really collected in Celebes : since, however, all the other specimens in the particular consignment of Dr. Meyer's collection, which contained the specimens of Kaloula, were from the Philippines and all the material from Celebes was received a t an earlier date, this suggestion is not very probable.

KALOULA SPECIMENS EXAMINED . . . . . (Mus.Leyden) COTYPES? Java 44.2.22. 93 . . . (Mus. Leyden) COTYPE? ? Java 63.12.11. 138 Java . . Bleeker 85.12.31. 36 d Willis Mts., Java, 5,000 ft.. . . . . . von Huegel 1928.3.15. 1 Kuala Tahan, Pahang, Robinson and Kloss F.M.S. 84.5.7. 1 9 Palembang, Sumatra . Forbes 85.7.20. 9 s? Nias. . . . . . Schreiber 92.6.3. 18 Mt. Dulit, Borneo . . Hose 95.7.2. 3 1 4 363 , Akar River, Sarawak . Hose 72.8.20. 69-70 Hgr. Laguna de Bay, Philippines . . Meyer 97.6.21. 100 P Sumba . . Everett 72.4.6. 150 . Meyer N.Celebes . 72.4.6.145-149 2 3 3 , 3 $?9 N. Celebes . . Meyer 96.12.9. 103-104 ?,Hgr. Kema, Celebes. . Sarasin 96.4.29. 91 ? Indrulaman, S. Celebes, 6 2,000ft. . . . .Everett 97.6.21. 101 d Tanah Jampeah, Nr. Celebes . . Everett 48.8.19.23 ? (India) . . . . . Parry, COTYPE,Hylaedactylus celebensis 52.2.22.25 Hgr. (W. Africa). . (Fraser) Mus. Vienna d Java Steindachner (1880) Mus. Vienna juv. Pengalengan, Bondong, W. Java, 4,500-5,000 ft.

B.M. 44.2.22.93

d 9

?

6

6

.

I 2.

Kaloula conjuncta (Peters)

Hybdaetylus (Holonectes) conjunctus, Peters, 1863, Mon. Ak. Berlin, p. 455 (Type locality Luzon, Philippine Islands) ;Steindachner, 1864, Verh. Zoo1.-bot. Ges. Wien,m, p. 256, P1. xr, k s . 5a-d Callvla mjuncta, Cope, 1867, Journ. Acad. Philadelphia, (2), vr, p. 192 ; Boulenger, 1882, Cat. Bstr. Sal. Brit. Mus., ed. 2, p. 168 ;Boettger, 1892, Kat. Batr. Senck. Nat. Ges., p. 23 ; Nieden. 1926, Das Tierreich, Anura 11, p. 26. Kaloula wnjuncta, Taylor, 1920, Philippine Journ. Sci., XVI, p. 325. ? Kaloula negrosenis, Taylor, 1922, Philippine Journ. Sci., xxrr, p. 180, P1. In, figs. 3-4 (Type locality Hinigaran, Negros, Philippine Islands).

Post-narial ridges moderately strong, curving backwards mesially. Snout truncate, as long as, or a litt,le shorter than, the diameter of the eye ; canthus rostralis obtuse ; loreal region slightly oblique ; interorbital space 1+ t o 1%times as wide as the upper eyelid. Fingers with large truncate discs which are more than twice as wide as the penultimate joint; supernumerary tubercles present, just proximal to the basal subarticular tubercle of each finger; the second finger a little shorter than the fourth. Toes distinctly dilated, about half webbed in females, the membrane midway between the third and fourth toes extending beyond the distal tubercle of the third toe ; toes of the male fully webbed, the membrane reaching the discs of all of them ; outer toe extending beyond the second ; subarticular tubercles well developed, but the proximal one of the fourth toe usually absent ; two metatarsal tubercles, the inner the larger. Tibio-tarsal articulation reaching the ear or the posterior border of the eye. Skin smooth or slightly pustular above ; a slight fold from the eye to the insertion of the fore-limb and another above the shoulder ; lower surfaces smooth ; anal region granular. Greyish brown above, with a median dark spot which commences between the eyelids, narrows on the nape, broadens between the shoulders, narrows again and, about the middle of the back, bifurcates, the two branches running towards the groin

90

MONOGRAPH OF THE MICROHYLIDAE

and being produced as diagonal cross-bars. on the femur and tibia ; sometimes a circular light spot is enclosed in the darker area between the shoulders ; loreal region darker ; a dark streak from the eye, above the arm, to the middle of the flanks. Dusky beneath, with darker marblings and spots ; a dark line along the hinder side of the thighs, sharply defined above, but merging into the colour of the belly beneath. Male with a vocal sac. Length : 55 mm. DISTRIB~ION : Philippine Islands (Luzon, Negros, Mindanao and Bubuan). SPECIMENS EXAMINED 9 and Hgr. Philippine Islands . . . . . B.M. R.1929.6.2. 5-7 $? Philippine Islands . . . . . R.1929.6.2. 8 8 Dumaquete, Negros . . . . Taylor 1929.6.1. 33 Mus. Vienna $? ? . . . . . . . . . (Linnaea) Mus. Amsterdam Ad. Negros . . . . . . . . Taylor

RAMANELLA, Rao and Ramanna

v

Hylaedactylus (part), Jerdon, 1854, Journ. Asiatic Soc. Bengal, x m , p. 533. Cdlula (part), Giinther, 1864, Rept. Brit. India, p. 436 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 167 ;i&m, 1890, Fauna Brit. India, Rept. and Batr., p. 493 ; Nieden, 1926, Das Tierreich, Anura n, p. 21. K h l a (part),Rao, 1918, Rec. Ind. Mus., xv, p. 43. Ramanella, Rao and Ramanna, 1925, Proc. Zool. Soc. London, p. 1445 (Type species Ramanella symbioitica,Rao and Ramanna = Callula variegata, Stoliczka) ; Noble, 1931, Biol. Amph., p. 538.

Prevomer divided, the anterior portion small, bounding the narial opening anteromesially, the post-choanal portion reduced t o one or two slivers of bone lying free in the mucosa of the palatine region (fig. 37). No clavicles ; procoracoids small, apposed t o the mesial ends of the coracoids ;omosternum small, cartilaginous ; sternum cartilaginous. Vertebral column diplasiocoelous. Terminal phalanges T-shaped. Pupil horizontal. Tongue oval, entire or slightly emarginate, and slightly free behind ; two transverse ridges across the palate in front of the pharynx, the anterior short and curved ; a short transverse ridge a little way behind each choana. Tips of digits more or less dilated ; toes more or less webbed. In all the known species the tympanum is hidden, the first finger shorter than the second, which is a little shorter than the fourth, and the fifth toe shorter than the third, but extending well beyond the second.

Fig. 37. Anterior cranial elements of Ramanella triangularis (Giinther) (seen from beneath) E., ethmoids. Max., Maxilla. P., palatine? P.Mx., premaxilla. P.s., parasphenoid. P.V., posterior portion of Prevomer. V., Prevomer.

This genus differs from Kaloula, with which it was formerly confused, in the reduction and breaking up of the prevomer.

RAMANELLA

91

SYNOPSIS OF THE SPECIES I. Toes 4 webbed, or more. A. Snout longer than the eye ; toes of females about 4 webbed R. montana, p. B. Snout not longer than the eye ; toes of females about webbed R. ohscura, p. C. Snout not longer than the eye ; toes of females about half webbed R. palmata, p. 11. Toes less than 4 webbed. A. Toes with a rudiment of web ; belly immaculate R. variegata, p. B. Toes perfectly free ; belly brown, white spotted . . . R. triangularis, p.

+

. .

91 92 93 93 94

I . Ramanella montana (Jerdon) Eylaedactylus montanus, Jerdon, 1854, Journ. Asiatic Soc. Bengal, xxn, p. 533 (Type locality " Mnuntain streams in Wynaad "). Cdlula obscura (non Giinther), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 169, P1. x n ~ , fig. 3 (in part) ; Miiller, 1887, Verh. Nat. Gee. B a s 4 w, p. 261 (in part) ; Thurston, 1888, Cat. Batr. Sal. S. In&, p. 42, PI. m, fig. 3; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 493 (in part) ; Schter, 1892, Batr. h d . Mus., p. 23 ; Ferguson, 1904, Journ. Bombay N. H. Soc., xv, p. 506 ; Nieden, 1926,Das Tierreich, Anura II,p. 23, figs. 19-21 (in part).

Post-narial ridges strong, nearly in contact on the middle line. Snout truncate, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space about once and a half as broad as the upper eyelid. Fingers with large triangular discs, which are twice as broad as the penultimate joint. Toes with slightly dilated tips ; & webbed in females, the membrane between the third and fourth toes reaching to midway between the subarticular tubercles of the third toe (fig. 38a) ; half webbed in males, the membrane between the third and fourth reaching the level of the distal tubercle of the third (fig. 38h) ; subarticular tubercles moderately well developed ; two rounded metatarsal tubercles. Tibio-tarsal articulation reaching the shoulder or the posterior border of the eye.

Fig.38. Foot of Ramanella montaw, (a) and (b) 6

MONOGRAPH O F THE MICROHYLIDAE Skin smooth above or slightly pustular, especially on the head ; a supratympanic fold, with an indefinite parotoid-like thickening of the skin above it ; sometimes an occipital fold ; smooth beneath. Pale brown to chocolate above, the tip of the snout, lores and flanks darker; a median dark spot on the back, commencing on the upper eyelids (truncate in front), narrowing, or sometimes interrupted, behind the head, broadening between the shoulders, narrowing again and finally broadening and breaking up in the sacral region ; broad dark cross-bars on the forearm, femur, knee, tibia, tarsus and metatarsus, these markings sometimes being light-edged ; a triangular dark spot enclosing the vent. Some or all of these markings may be greatly reduced. Lower surfaces dark brown, with regular white spots and blotches. Length : 35 rrm. Male with a vocal sac. Tadpole free-swimming. A well-developed specimen of 38 mm., has the following characters : Head and body ovoid, depressed, once and a half as long as broad. Nostrils close together, about midway between the tip of the snout and a line connecting the anterior borders of the eyes. Eyes lateral, the interorbital space more than 5 times the internarial. Mouth terminal, twice as wide as the internarial space, the lower lip contractile, the upper nearly straight. Spiraculum median, opening under a transparent flap close to the hinder end of the body. Anus median, the anal tube curving downwards and opening a t the level of the hinder side of the thighs. Tail once and a half as long as the head and body, pointed but not flagellate, nearly 3 times as long as deep ; caudal crests subequal, about as deep as the muscular portion of the tail a t its base. Toes webbed as in the adult. Greenish brown above, spotted with darker ; sides and lower surfaces white, immaculate. Tail brown spotted. The largest tadpole measures 45 mm. over all, and a newly metamorphosed frog measures 14 mm. from snout to vent. DISTRIRUTION : S.W. India (Malabar, Cochin and Travancore). This species has hitherto been confused with R. obscura. SPECIMENS EXAMINED B.M. 74.4.29. 607 juv. Malabar . . . . Beddome 74.4.29.993-1008 8 8, 9 9 and Hgr. Malabar . . . . Beddome 74.4.29.1009 Cleared Malabar . . . . Beddome Anamallays . . . Beddome 74.4.29. 1795 Wynaad . . . . Jerdon, TYPE 72.4.17. 212 74.4.29. 257 ? Travancore . . . Beddome 97.10.1. 3 4 3 7 8 , 3 99 Devicolum, Travancore (4,000-7,000ft.) Fegguson 1903.9.26.34--38 Larvae and Yg. Trevandrum, Travan- Ferguson (Series core described by Ferguson)

d

2.

Ramanella obscura (Giinther)

Callula obscura, Giinther, 1864, Rept. Brit. India, p. 438 (Type locality Ceylon) ; Boulenger, 1882 Cat. Batr. Sal. Brit. Mus., ed. 2, p. 169 (in part) ; idem, 1890, Fauna Brit. India, Rept. and Batr., p. 493 (in part) ; Werner, 1893, Verh. Zoo1.-bat. Ges. Wien, p. 354; Nieden, 1926, Das Tierreich, Anura n, p. 23 (in part).

Post-narial ridges short, almost circular, widely separated. Snout short, truncate, not prominent, as long as, or a little shorter than, the eye ; canthus rostralis rounded, curved ; loreal region almost vertical ; interorbital space once and a half as broad as the upper eyelid. Fingers with large triangular dilatations, which are twice as wide as the penultimate phalanges. Toes ending obtusely, those of females about 4 webbed, the membrane between the third and fourth toes extending to about midway between the subarticular tubercles of the third toe (male not examined) ; subarticular tubercles well developed ; two small metatarsal tubercles. Tibio-tarsal articulation reaching the tympanic region or the posterior corner of the eye.

Skin slightly warty above ; smooth beneath ; a supratympanic fold with paratoidlike thickening above i t ; sometimes an occipital fold. Brown above, the snout, lores and flanks darker ; a median dark area commencing on the upper eyelids (truncate in front), narrowing on the nape, broadening between the shoulders, narrowing slightly and finally broadening and breaking up into two or three branches on the sacral region. Broad, dark cross-bars are present on forearm, femur, knee, tibia, tarsus and metatarsus, and a triangular dark spot encloses the anus. Lower surfaces dark brown, regularly spotted with lighter. Length : 33 mm. : Ceylon. DISTRIBUTION B.M. 58.11.28. 42 54.3.21. 16-17 71.12.16. 13

? ?

Hgr.

Ceylon. Ceylon. Ceylon.

. . . . . . . . . . . . . . . . . .

Sir A. Smith, TYPE Cuming Thwaites

3. Ramanella palmata sp. n. CaUulu obscuw (part), Miiller, 1887, Verh. Nat. Ges. Basel, VIU,p. 251.

Post-nariat ridges short, almost circular, well separated on the middle line. Snout short, truncate, not prominent, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical ; interorbital space once and a half as broad as the upper eyelid. Fingers with large, triangular dilatations nearly as wide as the penultimate joints, and a suspicion of web a t their bases. Toes of both sexes half webbed, the membrane between the third and fourth extending as far as the distal tubercle of the third toe ; subarticular tubercles well developed ; two metatarsal tubercles, the outer small. Tibio-tarsal articulation reaching to between the shoulder and the tympanum. Skin tubercular above ; smooth beneath ; a supratympanic fold. Greyish above, the sides of the head brown ; an indefinite dark bar between the upper eyelids and a median dark area on the back, commencing behind the head, broadening between the shoulders, narrowing slightly and finally broadening on the sacral region and becoming indistinct. Broad, indistinct cross-bars on the forearm, tibia, tarsus and metatarsus ; anal region dark brown. Lower surfaces dark brown, with small, regular, white spots. Length : 36 mm. ; fore-limb 22 mm. ; hind-limb 28 mm. DISTRIBUTION : Newera Ellya, Ceylon, 7,000 ft. SPECIMENS EXAMINED Newera Ellya, 7,000 ft. 90.11.8. 63 $ Newera Ellya, 7,000 ft. Mus. Base1 1414-1415 $, imm. Newera Ellya, 7,000 ft. B.M. 90.11.8. 62

.

. .

Green, TWE Green PARATYPES Sarasin

}

4 . Ramanella variegata (Stoliczka) x m , p. 533 (Type locality "the Carnatic "). Cdlula variegata, Stoliczka, 1872, Proc. Asiatic Soc. Bengal, p. 111 (Type locality Ellore) ; Blanford, 1879, Journ. Asiatic Soc. Bengal, x~vnx,p. 116 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 494 ; Sclater, 1892, Batr. Ind. Mus., p. 23 ; Sarasin, 1910, Zool. Jahrb., Supp. XII, 1, p. 29 ; Nieden, 1926, Das Tiemeich, Anura 11, p.24. Kaloulu vare'egata, Rao, 1918, Rec. Ind. Mus., xv, p. 43, figs. CaUula olivaceo, Giinther, 1875, Proc. Zool. Soc. London, p. 576, P1. LXN, fig. B (Type locality Yellaghemy Hills) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 171 ; Miiller, 1887, Verh. Nat. Ges. Basel, vnx, p. 251; Thurston, 1888, Cat. Batr. S. India, p. 42, PI. wr, fig. 4. Ramadla symbwiticu, Rao and Ramana, 1925, Proc. Zool. Soc. London, 1925, p. 1445, P1. (Type locality Bangalore). ? Hykzedactylus carnaticus, Jerdon, 1854, Journ. Asiatic Soc. Bengal,

MONOGRAPH O F T H E MICROHYLIDAE Post-narial ridges feeble, slightly oblique, sometimes pigmented. Snout short, rounded, slightly prominent, as long as, or a little shorter than, the eye ; canthus mstralis rounded ; loreal region slightly oblique ; interorbital space once and a half as broad as the upper eyelid. Fingers with triangular dilatations nearly twice as wide as the penultimate joint. Toes scarcely dilated, with a rudiment of web which does not extend beyond the proximal tubercle of the third toe ; subarticular tubercles well developed ; two metatarsal tubercles. Tibio-tarsal articulation reaching the shoulder. Skin smooth or slightly pustlllar above ; sometimes a supratympanic fold with the skin above it somewhat thickened. Lower surfaces smooth ; anal region feebly granular. Brown above, with lighter marblings and spots which are sometimes arranged to give an indication of the typical colour pattern of the other species of the genus. Lower surfaces white, immaculate except the chin and throat, which may be faintly brown stippled. Length : 36 mm. Male with a vocal sac. Tadpole free-swimming :l head and body ovoid, depressed, once and a half as long as broad ; nostrils close together, on the upper surface of the head, about midway between the tip of the snout and the eyes. Eyes lateral, the interorbital space 7 times as broad as the internarial. Mouth terminal ; both lips contractile. Spiraculum median, opening under a broad notched flap ; anus median. Tail t%ce as long as the head and body, pointed, but not flagellate, 3 times as long as deep ; caudal crests subequal. Body brownish or grey, with minute black spots ; occasionally a blue spot in the groin ; muscular portion of the tail blotched with black. Length : 45 mm. ; tail 30 mm. DISTRIBUTION : S. India (as far north as the Godaveri Valley) ; Ceylon.

77.8.6. 12 1908.1.29. 26 87.11.2. 22 74.4.29.1351-2

EXAMINED SPECIMENS Godaveri Valley . S.E. of Bhadrachalam S.E. of Bhadrachalam Hgr. Chanda . . . . $8 Yellagheny Hills. .

1925.9.18.1-2

$9

B.M. 74.11.12. 2

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Bangalore

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. . . . . . .

. Blanford, COTYPE? . Blanford . Blanford

. Blanford

. Beddome, COTYPESof C. olivacea, Giinther . Rao, COTYPESof R. symbioitica, Rao and Ramanna 9

Ramanella triangularis ( Giinther)

Callula triangularis, Giinther, 1875, Proc. 2001. Soc. London, p. 576 (Type locality, Malabar) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 171, PI. xm, fig. 4 ;Thurston, 1888, Cat. Batr. S. India, p. 42, P1. v m , fig. 5 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 495; Sclater, 1892, Batr. Ind. Mus., p. 24 ; Boettger, 1892, Kat. Batr. Senck. Nat. Gea., p. 24 ; Ferguson, 1904, Journ. Bombay N. H. Soc., XV,p. 507 ; Nieden, 1926, Das Tierreich, A m a n, p. 24. K a h l a triangularis, Rao, 1918, Rec. Ind. Mus., xv, p. 44.

Post-narial ridges moderately strong and narrowly separated. Snout short, rounded, not prominent. as long as the diameter of the eye ; ,canthus rostralis rounded ; loreal region oblique ; interorbital space once and a quarter the breadth of the upper eyelid. Fingers with small triangular dilatations, not more than once and a half as wide as the penultimate joints. Toes free, not dilated distally ; subarticular tubercles well developed ; two metatarsal tubercles. Tibio-tarsal articulation reaching the shoulder or the tympanic region.2 Description according to Rao, who remarks, incidentally, that the lower lip may have a rim." Not the tarso-metatarsal articulation as sometimes stated.

"

horny

KALOPHRYNUS

95

Skin smooth, or with a few flat warts on the back ; a fold from the eye to the forelimb : sometimes an occipital fold. Lower surfaces smooth. Pale brown or olive above, with a lateral dark streak commencing on the loreal region ; a median dark blotch on the back commencing at the level of the fore-limbs and breaking up, on the coccygeal region, into two branches, which run almost at right angles to the main axis into the groin ; a broad dark band across the forearm, femur, tibia, tarsus and foot; a triangular dark spot enclosing the anus. These markings are usually sharply defined, but the dorsal blotch may be fenestrated by small spots of the ground colour. Lower surfaces dark brown, with small, regular, whitish spots. Length : 4 0 mm. Male with a vocal sac. A free-swimming tadpole : Head and body ovoid, depressedi once and a third as long as broad ; nostrils equidistant from the eyes and the tip of the snout. Interorbital space 6 times as broad as the internarial. Mouth small, terminal. Spiraculum median, inconspicuous, without notched flap. Anus median. Tail once and a half as long as the head and body, pointed, but not flagellate, and nearly 3 times as long as deep ; caudal crests subequal, narrower than the muscular portion of the tail. Transparent, with an oval or horseshoe-shaped dark mark on the back. Length : 5 2 mm. ; tail 1 9 mm. DISTRIBUTION : S.W. India. SPECIMENS EXAMINED B.M. 68.4.3. 126 HgrNilgiri Hills . . . Theobald 68.4.3. 127 Q cleared Nilgiri Hills . Theobald 87.8.26. 2 6 Nilgiri Hills, 6,500-: 7,000 ft. . . . . Thurston 74.4.29.891-900 3, Q, Hgr. Malabar . . . . Beddome1 74.4.29. 1010 Hgr. Malabar . . . . Beddomej

KALOPHR YNUS, Tschudi Kalophrynus, Tschudi, 1838, Mem. Soc. Sci. Neuchitel, 11, p. 86 (Type species Kalophrynus plearostigma, Tschudi) ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 54 ; Stejneger, 1908, Proc. U.S. Nat. Mus., xxxm, p. 575 ;Taylor, 1920, Philippine Journ. Sci., xvr, 3, p. 328 ;Noble and Parker, 1926, h e r . Mus. Novitates, CCXXXII, p. 19 ;Smith, 1930, Bull. Raffles Mus., 111, p. 123 ;Noble, 1931, Biol. h p h . , p. 536. Ca2ophynus (emend.), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 157 ;idem, 1890, Fauna Brit. India, Rept. and Batr., p. 490 ; idem, 1912, Fauna Malay Penins., Rept. and Batr., p. 256; Prashad, 1918, Rec. Ind. Mus., xv, p. 101 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 102 ;Nieden, 1926, Das Tierreich, Anura n, p. 15 (part). Engyslomo ? (mFitzinger), Blyth, 1855, Journ. Asiatic, Soc. Bengal, xxm, p. 732. Berdmorea, Stoliczka, 1872, Proc. Asiatic Soc. Bengal, p. 146 (Type species Engystoma ? interlineaturn, Blyth). Diplopelma (non Giinther, 1858), Giinther, 1895, Novit. Zool., n, p. 501. Bufo (non Laurenti), Isenschmid, 1903, Mitt. Naturf. Ges. Bern, p. 27. Microhyla (part), van Kampen, 1923, h p h . Indo-Austr. Arch., p. 154; Nieden, 1926, Das Tierreich, Anura n, p. 34. Prevomer divided, the post-choanal portion absent ; palatine absent (fig. 3 9 ) ;

ethrnoids minute or absent. Clavicles and procoracoids present, well developed, reaching the mid-line of the girdle and the scapulae ; omosternum small, cartilaginous ; sternum large, cartilaginous (fig. 40). Vertebral column diplasiocoelous. Terminal phalanges T-shaped. Pupil horizontal. Tongue oval or subcircular, entire and half free behind. A strongly denticulated ridge across the palate in front of the pharynx, preceded by a shorter, curved, nearly smooth one ; a smooth ridge behind each choana. Digits not dilated distally. Skin thick and glandular.

MONOGRAPH OF THE MICROHYLIDAE

Fig. 39. Anterior cranial elements of Kalophrynus pleurosligma (seen from beneath)

Fig. 40. Ventral elements of the shoulder-girdle of Kdophrynw pleuroatigma (ventral view)

KALOPHRYNUS SYNOPSIS OF THE SPECIES I. Third toe longer than the fifth ; second finger longer than the fourth. A. Fourth finger moderately well developed. (1) Toes frequently more than ) webbed ; free portion of the fifth toe shorter than the distance of the nostrils from the tip of the snout . . . K . pleurostigma pleurostigma, p. 97 (2) Toes never more than $ webhed; free portion of the fifth toe longer than the distance between the nostrils and the tip of the snout . K. pleu~ostigmainterlineatus, p. 99 B. Fourth finger very short, almost reduced to a knob. (1) Third finger as long as the eye ; nuptial excrescences of the male in the form of a thickened pad beset with numerous . . . . . . . . K. bunguranus, p. 100 small spines (2) Third finger distinctly shorter than the eye ; nuptial excrescences of the male in the form of comb-like rows of very large, conical spines . . . . . . . . . . K. robinsoni, p. 100 11. Third toe not extending beyond the fifth ; fourth finger as long as the K. punctatus, p. 101 second . . . . . . . . . . . . . .

I' . Kalophrynus pleurostigma pleurostigma, Tschudi Kalophynus pleu~ostigma,Tschudi, 1838,Mem. Soc. Sci. Neuchitel, n, p. 86 (Type locality Sumatra); Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 54 ;Smith, 1930, Bull. Raffles Mus., p. 123. Cabphrynus pleurostigma, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 158 ; Fischer, 1885, Arch. f. Naturg., p. 46 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 490 (part) ; Mocquard, 1892, Mem. Soc. Zool. France, v, p. 195 ; Flower, 1896, Proc. Zool. Soc. London, p. 908; idem, 1809, Proc. Zool. Soc. London, p. 900 (part) ; Butler, 1904, Journ. Bombay N. H. Soc., x v , p. 387 ; Robinson, 1905, Journ. F.M.S. Mus., I, i, p. 23 (part) ; van Kampen, 1905, Zool. Jahrb., Syst., xxn, p. 707 ;Boulenger, 1912, Fauna Mahy Penins., Rept. and Batr., p. 258 (part) ; Smith, 1916, Journ. Nat. Hist. Soc., Siam, 11, p. 168 ;van Kampen, 1923, Amph. lndo-Austr. Arch., p. 102 (part) ;Nieden, 1926, Das Tierreich, Anura 11, p. 16 (part). Calophrynus acutimstris, Boettger, 1897, Zool. Anz., xx., p. 165 (Type locality " Culion or Samar," Philippine Islands) ;Mertens, 1922, Senckenbergiana, N, p. 165 ;Nieden, 1926, Das Tierreich, Anura n, p. 17. Kabphrynus aeutirost~is,Taylor, 1920, Philippine Journ. Sci., XVI, p. 331. Bufo duderi, Isenschmid, 1904, Mitt. Naturf. Ges. Bern, 1903, p. 77, P1. v, fig. 2 (Type locality Palembang, Sumatra). Kalophrynw stellatus, Stejneger, 1908, Proc. U.S. Nat. Mus., XXXIII, p. 575 (Type locality Basilan, Philippine Islands) ; Taylor. 1920, Philippine Journ. Sci., xvr, p. 329, Pl. IX, fig. 2 ; Taylor, 1922, Philippine Journ. Sci., XXI, p. 280. Calophrynus stellatus, Nieden, 1926, Das Tierreich, Anura 11, p. 18.

Snout prominent, pointed, as long as, or slightly longer than, the diameter of the eye ; canthus rostralis distinct ; loreal region almost vertical ; interorbital space broader than the upper eyelid ; tympanum distinct, f to 8 the diameter of the eye. Fingers free, the first shorter than the second, which is much longer than the fourth ; third as long as, or shorter than, the snout. Toes f to 2 webbed, the degree of webbing being very variable but roughly correlated with size ; in juvenile and half-grown specimens they are rarely more than f to 3 webbed, but in large breeding males the web may reach to the tips of all except the fourth. The free portion of the fifth toe is never longer than the distance of the nostrils from the tip of the snout. Subarticular tubercles moderately distinct ; an inner metatarsal tubercle, the outer either absent or very indefinite. Tibio-tarsal articulation reaching the tympanum or the posterior border of the eye. Skin granular above, often with numerous, small horny spines ; coarsely granular on the belly and hinder side of the thighs. A more or less distinct parotoid gland, bordered above by a feeble fold extending from the posterior corner of the eye almost to the groin. Sometimes a fold connecting the posterior corners of the upper eyelids. Grey, brown or pinkish above, the loreal region and flanks below the dorso-latera 7

MONOGRAPH OF THE MICROHYLIDAE fold, black. Dorsal pattern consisting primitively of a darker area commencingbetween the eyes, connecting the two eyelids and truncate anteriorly; this narrows on the nape and then broadens and bifurcates, the two branches running to the groin, where, by darkening of their colour, they give rise to two rounded ocellar spots ; continuations of these two stripes appear as oblique transverse bars across the femur and tibia. This marking may be entirely absent, broken up into spots or well defined with, as a secondary pattern, further lines parallel with its edges. Anal region and proximal part of the hinder side of the thighs black, light-bordered above. Belly dirty white ; the throat and chest infuscate, with, sometimes, white glandular spots ; sometimes two, broad, dark, longitudinal stripes may be indicated on the chin ; in breeding specimens the throat, chest and back may be washed with pink or claret. Length : 6. 49 rnm. ; 8 53 mm. Male with a vocal sac and diffuse nuptial pads on the dorsal surfaces of the first, second and third fingers. Ovarian eggs, very small. A short developmental series, ascribed to this species, was reared by Hewitt a t Kuching in November. The first stage, labelled " 2nd day " (? after hatching), measures 6 mm., and is an unpigmented, very immature larva with the head distinct from the elongate body and a short, rounded tail ; there are three pairs of very short external gills, incipient opercula, a pair of very prominent, circular, adherent organs behind the mouth, and shallow nasal pits. Ry the third day the head and body have become compacted together, the opercular folds have covered the branchiae, but not yet formed a spiracle, the tail has lengthened and rudiments of the hind-limbs are apparent ; total length 7.8 mm. ; body 3 mm. The tadpole on the fifth day, the last stage of the series, is 9 mm. long, with the body ovoid, rather square anteriorly and once and a half as long as broad, with a small ventro-terminal mouth ; openings from the opercular folds are still separate, the anal tube is long, opening well behind the body, and the tail is acutely pointed, 4 times as long as deep, nearly twice as long as the body and with subequal crests. DISTRIBUTION : Malay Peninsula as far north as Patiyu, Sumatra, Borneo, Natuna Islands, Philippines. Despite the enormous amount of variation in the degree of webbing of the toes, it does not appear possible to subdivide this form. The greatest amount of webbing was found in specimens from the Natuna Islands, but Bornean specimens approach them very closely, and a series of specimens from the latter island shows all intermediate conditions between % webbing and $. A specimen from the Philippine Islands (apparently referable to K. sfellatus, Stejneger) and a series of specimens from the Malay Peninsula fall within the limits of variation shown by the Rornean series, and Boettger's Ii. acutirostris, so far as one can judge from the description, does not differ in any characters which can be regarded as of specific or subspecific value. 9 SPECIMENS EXAMINED Patiyu. Peninsular Siam . . M. A. Smith R.M. 1916.6.22.34-37 6.6. 1928.1.11. 22 cleared Patiyu, Peninsular Siam . . . M. A. Smith 97.11.15. 12 juv. Foot of Gunong Pulai, Johore Flower 1906.2.28. 53 Gunong Tahan, 3,000 ft. . . . Robinson 96.6.25. 99 JUV. Singapore Flower 91.5.1. 95-98 6.6. Paulo Laut, Natuna Islands . . Hose 92.6.3. 17 . C. Hose w e Mt. Dulit, Borneo . 92.9.2. 9 ? Baram River, N. Borneo . . . Everett Mus. Vienna c?? Baram River . . . . . . C. Hose B.M. 94.6.30. 133 ? Paitan, N. Borneo. . . . . Everett 95.7.2. 26-28 c?? Akar River, Sarawak . . . . Hose 1902.12.12.20-21 88 Bidi Caves, Sarawak . . . . Brooks 1914.5.12.49-60 larvae Kuching, Sarawak . . . . . Hewitt 86.5.15. 5-6 Pagat and Meratus Mts., S.E. Borneo . . Fischer 44.6.15. 3 ? Borneo . . . . . . 1924.2.12.2 9 Port ~ o l l a n dBasilan , Island Taylor

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KALOPHRYNUS 2.

99

Kalophrynus pleurostigma interlineatus (Blyth)

Engystoma ( 9 ) i7lterlineatum, Blyth, 1855, Journ. Asiatic Soc. Bengal, xxm, p. 732 (Type locality Pegu, Burma) ;Blyth, 1856, Journ. Asiatic Soc. Bengal, xxrv, p. 720 ; Anderson, 1871, Proc. Zool. Soc. London, p. 202. Engystoma interlineutum, Mason, 1860, Burma, p. 325. Berdmorea interlineata, Stoliczka, 1872, Proc. Asiatic Soc. Bengal, p. 146. Cdophrynus interlineata, Theobald, 1882, in Mason, Burma, I, p. 291. Calophrynus pleuroatigma, var. sinensis, Peters, 1867. Mon. Ak. Berlin, p. 33 (Type locality Hong Kong.) Calophrynus pleuroatignza, Giinther, 1873, Zool. Rec., VIII, 1871, p. 86 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 158 (part) ; idem, 1890, Fauna Brit. India, p. 490 (part) ;Sclater, 1892, Batr. Ind. Mus., p. 22 ; Flower, 1899, Proc. Zool. Soc. London, p. 900 (part) ; Werner, 1904, Abh. Bayer. Akad. Miinchen, xxn, p. 369 ; Robinson, 1905, Journ. F.M.S. Mus., I, p. 23 (part) ; Boulenger, 1912, Fauna Malay Pcnins., Rept. and Batr., p. 258 (part) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 102 (part) ; Nieden, 1926, Das Tierreich, Anura 11, p. 16 (part) ; Kalophrynua plenrostignza, Gee and Boring, 1929, Peking Soc. Nat. Hiat. Bull., rv, 2, p. 25 ; Pope. 1931, Bull. Amer. Mus. Nat. Hist. LXI,p. 586.

Snout pointed, only slightly prominent, as long as the diameter of the eye ; canthus rostralis distinct ; loreal region almost vertical ; interorbital space broader than the upper eyelib, tympanum distinct, 2 to $ the diameter of the eye. Fingers free, the first shorter than the second which is longer than the fourth ; third as long as, or longer than, the snout. Toes with a rudiment of web, never more than webbed ; the free portion of the fifth toe always longer than the distance from the nostrils to the tip of the snout. Subarticular tubercles distinct ; an inner, and frequently an outer, metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder, or between this point and the posterior border of the tympanum. Skin tubercular above ; coarsely granular on the belly and hinder side of the thighs. A more or less distinct parotoid gland, bordered above by a feeble fold extending from the posterior corner of the eye almost to the groin. Grey, brown or pinkish above, the loreal region, and flanks below the dorso-lateral fold, black. Dorsal pattern consisting primitively of a narrow dark streak commencing on the tip of the snout and bifurcating behind the occiput ; the two branches run diagonally to the groin, where, by darkening of their colour, they give rise to two rounded ocellar spots ; continuations of these two stripes appear as oblique transverse bars across the femur and tibia. This marking may be entirely absent or represented only by the ocellar spots ; associated with it there may be a varying number of paler, narrower lines parallel with its edges, and these, like the primary marking, may be broken up into spots; in extreme instances of this nature the whole back may be uniformly dotted with small, round spots. Anal region and proximal part of the hinder side of the thighs black, light-bordered above. Belly dirty white, the throat and chest infuscate, with, sometimes, white glandular dots. Frequently two broad, dark, longitudinal stripes may be present on the chin. Length from snout to vent : 8 38 mm. ; 9 44 mm. Male with a vocal sac ; nuptial excrescences are of the diffuse ranid type on the dorsal surfaces of the first, second and third fingers. Ovarian eggs very small.

+

DISTUIBUTION : Hong Kong, Kuangtung, Yunnan, Hainan, Tonkin, Burma, Siam and Cambodia. This subspecies is the Indo-Chinese form of K. pleurostigma, and may usually be differentiated from the southern subspecies by the smaller amount of interdigital webbing; but the two intergrade in this and some other respects. The constant differences between them are the shorter 6gures and toes of pleurostigma, reflected in the empirical characters used in the key, and t.he colour pattern. In interlineatus the dorsal marking is a narrow line commencing on the tip of the snout, whereas in the Malayan subspecies it commences between the upper eyelids as a broad stripe, truncate in front, which narrows behind the occiput and then broadens again before bifurcating.

MONOGRAPH OF THE MICROHYLIDAE R.M. 1903.4.29.75-83 1903.4.29. 8 4 68.4.8. 128-9 91.11.26. 62 97.10.8. 1 0 1

SPECIMENS EXAMINED 33,$2$2 hgr. Man-Son Mts., Tonkin skel. $2 Man-Son Mts., Tonkin and hgr. Pegn . . . . . ? Toungoo . . . . hgr. Rortong, Siam . .

. . . . .

Fruhstorfer Fruhstorfer Theohald, TYPES(?) Oates Flower

3 . Kalophrynus bunguranus (Giinther) Diplopelma bunguranum, Gunther. 1895, Novit. Zool., 11, p. 501 (Type locality Gt. Natuna Island), Gunther, 1896, Novit. Zool., m, p. 185, P1. vm, figs. 2-3. Microhyh bungurana, van Kampen, 1924, Amph. Indo-Austr. Arch., p. 154 ; Nieden, 1926, Das Tierreich, Anura 11, p. 34. Kabphrynus bunguranus, Parker, 1926, Ann. Mag. Nat. Hist., (lo), 11. p. 473. Cabphrynus heberochirus, Boulenger, 1900, Roc. Zool. Soc. London, p. 186, P1. XVII,fig. 4 (Type locality Limbang, Borneo); van Kampen, 1924, Amph. Indo-Austr. Arch., p. 104 ; Nieden, 1926, Das Tie~eich,Anura n, p. 17. Snout slightly prominent, as long as the diameter of the eye ; canthus rostralis angular ; loreal region almost vertical ; interorbital space broader than the upper eyelid ; tympanum moderately distinct, $ the diameter of the eye. Fingers free. very short, the first shorter than the second, and the fourth almost rudimentary ; third

about as long as the diameter of the eye. Toes short, & webbed, the third extending beyond the fifth ; subarticular tubercles indistinct ; two flat, rather indefinite metatarsal tubercles. Tibio-tarsal articulation reaching tlie posterior border of the eye. Skin indistinctly granular above, sometimes beset with minute spines; smooth beneath, except the posterior portion of the belly and hinder side of the thighs, which are granular. An indefinite parotoid gland whose lower limit forms an S-shaped ridge from the posterior corner of the eye, above the tympanum, to the fore-limb. Uniform dark brown above with a light hand around the head, along the canthus rostralis and the edges of the upper eyelids, and extending backwards along the flanks in the direction of the groin ; limbs with dark cross-bars ; flanks, groin and hinder side of thighs white (pink in life), with yellow spots enclosed in a brown reticulum ; sometimes a brown, ocellar, inguinal spot. Lower surfaces dirty white, the throat and chest brown, white-spotted. Length : 3 24 mm. : 2 7 mm. Male with a vocal sac ; nuptial excrescences in the form of a thickened pad of skin on the dorsal surface of the hand and three inner fingers ; this pad beset with relatively small conical spines which are also present over the antero-ventral surface of the whole arm. Ovarian eggs numerous, small. DISTRIBUTION : Borneo (Limbang) and Gt. Natuna Island. 9 SPECIMENS EXAMINED (33,?? Gt. Natuna Island . Everett, COTYPES R.M. 95.5.1. 105-108 1909.8.18. 6-7 9,hgr. Limbang, Borneo . (Sarawak Mus.), COTYPES Calophrynus heterochirus, Boulenger

4 . Kalophrynus robinsmi, Smith Kabphrynus robinsoni, Smith, 1922, Journ. F.M.S. Mus., X, p. 280 (Type locality Gunong Tahan) ; Smith, 1930, Bull. Raffles Mus., 111, p. 123.

Snout slightly prominent, as long as the diameter of the eye; canthus rostralis obtuse ; loreal region almost vertical ; interorbital space broader than the upper eyelid ; tympanum moderately distinct, % the diameter of the eye. Fingers free, very short, the first shorter than the second. and the fourth almost rudimentary; third distinctly shorter than the diameter of the eye. Toes short, 3 webbed, slightly fringed, the third extending beyond the fifth ; subarticular tubercles indistinct ; metatarsal tubercles two, ill-defined. Tibio-tarsal articulation reaching the tympanum.

CHAPERINA Skin slightly tubercular above, the tubercles sometimes aggregated to form a definite dorso-lateral ridge from the eye to the groin or another along the hinder side of the thighs. Posterior part of belly and thighs coarsely granular; throat finely granular ; a curved fold from the eye above the tympanum to the shoulder. Light brown above ; the sides of the head and flanks dark brown, the two colours being sharply defined ; an X-shaped dark mark on the back, the anterior branches short and running to the upper eyelids, the posterior long and terminating in the groin ; continuatio~~s of this mark form transverse bars across the femur and tibia. In addition the dorsum may be irregularly dotted or vermiculated with darker. Anal region and proximal part of thighs darker. Lower surfaces yellowish, spotted with brown, particularly on the throat and chest. Male with a subgular vocal sac. Nuptial excrescences in the form of comb-like rows of very large, conical tubercles on the dorsal surfaces of the fingers ; three to five on the third finger, two or three on the second, and one on the first. Length : 6 17 mm. ; 0 18 rnrn. DISTRIBUTION : Gunong Tahan and Kuala Teku, Malay Peninsula. The type and two paratypes all prove to be breeding males, with well-developed nuptial excrescences, and not juveniles, as originally stated. SPECIMENS EXAMINED 3 Gunong Tahan, F.M.S., R.M. 1983.5.14. 29 3,300 ft. . . . . Robinson TYPE 1923.5.14.22-23 6 6 Gunong Tahan, F.M.S., 3,300ft. . . . . Robinson PARATYPES 1906.2.28.54 Gunong Tahan, F.M.S., 3,000 ft. . . . . Robinson

5. Kalophrynus punctatus, Peters Calophrynus puctatus, Peters, 1871, Mon. Ak. Berlin, p. 579 (Type locality Sarawak) ; Peters, 1872, Ann. Mus. Genova, rn, p.43 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.,ed. 2, p.158 ;Boulenger, 1894, Ann. Mus. Genova, (2), xrv, p. 617 ; van Kampen, 1905, Zool. Jahrb., Syst., xu, p. 714 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 104; Nieden.1926, DasTierreich, Anura 11, p. 17. Snout slightly prominent, pointed, as long as the diameter of the eye ; canthus rostralis distinct ; loreal region nearly vertical ; interorbital space broader than the upper eyelid ; tympanum distinct, the diameter of the eye. Fingers free, the first shorter than the second, which is subequal to the fourth ; third almost as long as the snout. Toes ) webbed, the third and fifth subequal ; free portion of the fifth toe not longer than the distance of thenostrils from the tip of the snout ; subarticular tubercles distinct on the first to fourth toes, absent on the fifth ; a large, rounded inner, and an

+

indistinct outer, metatarsal tubercle. Tibio-tarsal articulation reaching the posterior border of the eye. Skin granular above ; coarsely granular on the belly and lower surface of the thighs ; a transverse fold connecting the posterior comers of the upper eyelids ; parotoid gland not well defined. Dark brown. spotted with black above ; lower surfaces dusky, with round yellowish white spots. Male with a vocal sac. Length : 27 mm. : Sarawak, Mentawei Islands. DISTRIBUTION SPECIMENS EXAMINED 6 Sarawak . . . . . . . . . . Doria, TYPE Mus. Genova

CHAPERlNA, Mocquard Chaperinu, Moquard, 1892, Le Naturaliste, (2), VI, p. 35 (Type species Chaperina fusca, Mocquard) ; Mocquard, 1892, Mem. Soc. Zool. France, v, p. 194 ; Taylor, 1920, Philippine Journ. Sci., XVI, p. 332.

102

M O N O G W H OF T H E MICROHYLIDAE

Microhyla ( m n Tschudi), Boulenger, 1899, Ann. Mag. Nat. Hist., (7), m, p. 278 ;van Kampen, 1923, Amph. Indo-Austr. Arch., p. 162 (part) ;Nieden, 1926, Daa Tierreich, Anura n, p. 27 (part). Nedophryne ( n m Buchholz and Peters), Smith, 1921, Journ. F.M.S. Nus., x, p. 198 ; van Kampen, 1923, Amph. Indo-Awtr. Arch., p. 66 (part). Sphemphryna ( n m Petera and Doris), van Kampen, 1923, Amph. Indo-Austr. Arch., p. 104 (part) ; Nieden, 1926, Daa Tierreich, Anura n, p. 43 ; Smith, 1930, Bull. Raffles Mus., In, p. 124 ; Noble, 1931, Biol. Amph., p. 632.

Prevomer divided, the post-choanal portion slender, overlying the palatine region, not raised into ridges or knobs (fig. 41). Clavicles and procoracoids present, reaching the mid-line of the girdle and the scapulae, curved ; no omosternum ; sternum cartilaginous (fig. 42). Vertebral column diplasiocoelous. Terminal phalanges simple. Ethmoid and parasphenoid fused. Pupil horizontal. Tongue small, oval, entire and free posteriorly. A single smooth dermal ridge across the palate in front of the pharynx. Digits dilated distally.

Fig. 41. Anterior cranial elements of Chapen'nu f w c a (seen from beneath)

Fig. 42. Ventral elements of the shoulder-girdle of Chaperina fusea (ventral view)

CHAPERINA I.

Chaperina fusca, Mocquard

Chuperinu fusca, Mocquard, 1892, Le Naturaliste, (2), VI, p. 35 (Type locality Sintang, Borneo) ; Mocquard, 1892, Mem. Soc. Zool. France, v , p. 194, PI. VII, figs. 2-2b. Sphenophryne fusca, Nieden, 1926, Das Tierreich, Anura n, p. 46 (in part) ; Smith, 1930, Bull. Raffles Mus., 111, p. 124,fig. 12 ; Burt, 1932, Bull. Amer. Mus. Nat. Hist., ~ x mp., 484. Microhyla leucoatigma, Boulenger, 1899, Ann. Mag. Nat. Hist., (7), rn, p. 275, PI. XII, fig. 1 (Type locality Larut, Perak, 3,500 ft.) ; Flower, 1899, Proc. Zool. Soc. London, p. 905 ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 388 ; Boulenger, 1912,Fanna Malay Penins., Rept. and Batr., p. 260 ; Barbour, 1920, Occ. Pap. Mus. Zool. Michigan, LXXVI, p. 3 ; van Kampen, 1923,Amph. Indo-Austr.Arch., p. 156 ;Nieden, 1926,Das Tierreich, Anura n, p. 45 ; de Witte, 1933,Bull. Mus. Hist. Nat. Belg., nr, 24, p. 7. Sphenophryne leucostigma, Smith, 1925,Journ. Sarawak Mus., rn, i, p. 12. Chuperina beyeri, Taylor, 1920, Philippine Journ. Sci., XVI, 3, p. 333 (in part), PI. 111, fig. 3 (Type locality Upper Agusan, Mindanao, between Agusan and Simulao rivers (probably in Davao or near the Davao-Agusan line)). Sphenophryne beyera', van Kampen, 1923, Amph. Indo-Austr. Arch., p. 110. Nectophryne picturata, Smith, 1921,'~ourn.F.M.S. Mus., x, p. 198, P1. n, fig. 2 (Type locality Mt. Dulit, British N. Borneo) ; van Kampen, 1923,Amph. Indo-Austr. Arch., p. 72. Snout rouqded, as long as, or slightly longer than, the diameter of the eye, with indistinct canthus rostralis and almost vertical loreal region ; nostril equidistant from the eye and the tip of the snout, or slightly nearer the latter ; interorbital space much broader than the upper eyelid ; tympanum rather indistinct, its diameter slightly more than half that of the eye. Fingers free ; first very much shorter than the second, which is shorter than the fourth. Toes slightly webbed a t the base, and with more or less distinct fleshy lateral fringes ; subarticular tubercles moderate ; an elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the anterior border of the eye. Skin smooth above and below ; a slight fold from the eye, above the tympanum, to the shoulder ; a small, delicate, dermal spine usually present on both the heel and the elbow. Black or dark brown above, with or without small yellow dots ; hind-limbs lighter, more or less distinctly barred with darker. Yellow and dark brown beneath, the yellow as large oval spots in the meshes of a bold, brown reticulum. Male with a subgular vocal sac. Length : 27 mm. DISTRIBUTION : Malay Peninsula, Borneo, Palawan, Philippine Islands. As has already been pointed out by Smith,' the tadpoles described by Taylor for this species are probably \those of some Ranid. The developmental series, collected in Sarawak by Hewitt, from which Smith has described the tadpole, almost certainly belongs to Chaperina fusca; dissection of a specimen with four limbs reveals the presence of a pair of strongly curved clavicles, and T-shaped terminal phalanges ; these two characters occur together in no Bornean Microhylid other than Chaperina. The eggs, each enclosed in a transparent, membranous capsule, are strung together on a thin cord, and adhere strongly wherever they are in contact. Each measures about 1-6 mm. in diameter and one pole is slightly pigmented. Hatching apparently occurs when the larva measures about 3.5 mm. ; a t this stage the external gills are very small, there is still no sign of the opercular fold, and the adhesive organs have the form of two very large and prominent tubercles close behind the mouth. A tadpole of a total length of 24 mm., the head and body measuring 9, may be described as follows :length of head and body 14 to 1+timesits breadth ; body considerably depressed, the snout broadly rounded. Eyes supero-lateral, the interocular distance twice as great as the internarial. Nostnls midway between the level of the eyes and the tip of the snout. Spiraculum median ventral, situated 3 times as far from the anterior as from the posterior end of the head and body. Mouth terminal, small, without mandibles or labial teeth. Anus dextral. Tail obtusely pointed, once and a half as long as the head and body ; caudal crests moderate, subequal. Olive-brown above, paler beneath. Bull. Raffles Mus., 111, 1930, p. 125.

'

MONOGRAPH O F

8

THE

MICROHYLIDAE

SPECIMENS EXAMINED Larut, Perak, 3,500 ft.

. . Flower,

COTYPESof Microhyla leucostigma 1900.6.14.33 0 Larut Hills, Perak, 4,000 ft. Flower 99.12.9. 4 0 Baram, Borneo . . . . Hose 1921.4.1. 289 9 Mt. Dulit, N. Borneo . . M. A. Smith, TYPEof Nectophrynepicturata 1914.5.12. 19-38 larvae Kuching, Sarawak . . . Hewitt 1927.9.1. 1 cleared Kuching, Sarawak . . . M. A. Smith 94.6.30. 135 juv. Palawan . . . . . . Everett 1924.2.12.4 0 Jolo Island, Philippines . Taylor, as Chaperina bey& 1927.3.7.49 0 Jolo Island, Philippines . Taylor, as Chaperina beyeri

B.M. 98.9.22.112-113

GASTROPHR YNOIDES,Noble Engystoma (non Pitzinger), Boulenger, 1897, Ann. Mag. Nat. Hist., f6), XIX,p. 108 ; Nieden, 1926, Das Tierreich, Anura 11,p. 68 (part). Gadrophryne, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 125 (part). Qa$trophrynoides,Noble, 1926, Amer. Mw. Novitates, oxn, p. 22, figs. 8-9 (Type species Engystoma burneeme, Boulenger) ; Noble, 1931, Biol. Amph., p. 538.

Prevomer divided, the post-choanal portion absent ; palatine absent. Clavicles and omosternum absent ; procoracoids well developed, reaching the mid-line of the girdle and the scapulae ; sternum large, cartilaginous (fig. 43). Vertebral column procoelous. Terminal phalanges T-shaped. Pupil round. Tongue oval, entire, half free behind ; a wide dermal ridge across the palate in front of the pharynx, preceded by a shorter one, both being smooth. Digits with small discs.

Fig. 43. Ventral elements of the shoulder-girdle of Gastrophrynoides borneense (ventral view)

I.

Gastrophrynoides bornemse (Boulenger)

Engystoma bmeeme, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), xnr, p. 108 (Type locality Baram District, Sarawak). Enggstoma borneensis, Nieden, 1926, Das Tierreich, Anura 11, p. 68.

PHRYNELLA Gastrophryne borneensia, Stejneger, 1910, Proc. Biol. Soc. Washington, XUI, p. 166. Gastrophryne bmneense, Banks, 1924, Rept. Sarawak Mus., p. 9. Gastrophrymidea b m e m e , Noble, 1926, Amer. Mus. Novitates, CXU, pp. 20, 22. Snout pointed, very prominent, twice and a half as long as the diameter of the eye ; nostril midway between the eye and the tip of the snout ; canthus rostralis rounded ; loreal region oblique, slightly concave ; interorbital space 3 times as broad as the upper eyelid ; tympanum hidden. Fingers short, the third and fourth with small, distinct discs, the first and second ending obtusely ; the first finger shorter than the second, which is shorter than the fourth ; a fleshy fringe extending from the base

half-way along the inner side of the second and third fingers. Toes moderate, with distinct discs ; the third longer than the fifth. Subarticular tubercles moderately well developed; a small inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the eye. Skin smooth above and below. Dark brown above, with small, scattered, white dots ; uniformly pale brown beneath. Length : 39 mm. DISTRIBUTION : N. Borneo.

?

B.M. 97.3.4. 26 1914.5.12. 18

juv.

. . . . Hose, TYPE . . . . . Hewitt

Baram District, Sarawak . Kuching, Sarawak. .

PHR YNELLA,

Boulenger PhryneLla, Boulenger, 1887, Ann. Mag. Nat. Hist., (5), xrx, p. 346 (Type species Phr?lnella pulchra, Boulenger) ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 265 (part) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 124 (part) ; Nieden, 1926, Das Tierreich, Anura XI, p. 37 (part) ; Noble, 1931, Biol. Amph., p. 537 (part). K a h l a (non Gray), Smith, 1926, Ann. Mag. Nat. Hist., (9), x v m , p. 81.

Prevomer divided, the post-choanal portion absent and the pre-choanal part with a backwardly directed mesial process (fig. 44); palatine absent. No clavicles ; procoracoids small, confined to the mesial ends of the coracoids ; a small cartilaginous ornosternum ; sternum cartilaginous (fig. 45). Vertebral column procoelous. Terminal phalanges broadly Y-shaped. Pupil horizontal. Tongue oval, entire, scarcely free behind. Two transverse dermal ridges across the palate in front of the pharynx, the anterior sometimes divided into three lobes ; a strong dermal ridge behind each choana. Digits strongly dilated ; subarticular tubercles of the hands enlarged to form accessory adhesive organs.

Pig. 44. Anterior cranial elements of Ph,rynella plchra (seen from beneath)

MONOGRAPH OF T H E MICROHYLIDAE

Fig. 45. Ventral elements of the shoulder-girdle of Phrynella pulchra (ventral view)

I.

Phrynella pulchra, Boulenger

Phrynella pukhra, Boulenger, 1887, Ann. Mag. Nat. Hist., (5), xrx, p. 346 (Type locality Malacca) ; idem, 1890, Proc. Zool. Soc. London, p. 37; idem, 1894, Ann. Mus. Genova, (2), XN, p. 617 ; Flower, 1896, Proc. Zool. Soc. London, p. 910 ; idem, 1899, op. cit., p. 908; Schenliel, 1901, Verh. Nat. Ges. Basel, xm, p. 152 ; Isenschmid, 1904, Mitt. Naturf. Ges. Bern, 1903, p. 75, P1. v, figs. 1-la ;Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 392 ;van Kampen, 1905, Zool. Jahrb., Syst., xxn,p. 707 ;Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 265 ; Smith, 1922, Journ. F.M.S. Mus., x, p. 282 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 125 ;Nieden, 1926, Das Tierreich, Anura n,p. 37 ;Noble and Jaeckle, 1928, Journ. Morph. Phys., x ~ vp. , 280, fig. 16 ; Smith, 1930, Bull. Raffles Nus., m, p. 123, fig. 11. Kaloula boulengeri (substitute name), Smith, 1926, Ann. Mag. Nat. Hist., (9), xvm, p. 81.

Snout short, truncate, prominent, a little longer than the diameter of the eye; canthus rostralis rounded ; loreal region nearly vertical ; interorbital space once and two-thirds as broad as the upper eyelid ; tympanum hidden. Fingers with a rudiment of web, dilated distally into large, triangular discs which are nearly twice as broad as the penultimate joints ; subarticular tubercles enlarged to form accessory adhesive pads, one each on the two inner fingers and two each on the two outer ; first shorter than the second, which is considerably shorter than the fourth. Toes feebly dilated distally, 2 webbed, the membrane reaching almost to the discs of all except the fourth toe ; webbing slightly more emarginate in females than in males ; fifth toe shorter than the third ; subarticular tubercles large, the basal one of the fourth toe absent ; a small inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanic region or the eye. Skin smooth or slightly pustular above ; feebly granular or smooth beneath ; a fold from the eye to the fore-limb, the skin above this slightly thickened. Brown above, with regular, insuliform, darker spots, which are sometimes blackbordered and longitudinally elongate. Usually an indistinct cross-bar connecting the upper eyelids and often a distinct light dorso-lateral line from the posterior corner of the eye to the middle of the flanks or beyond ; a light streak from beneath the eye on to the upper arm ; limbs brown-spotted like the dorsum ; belly, groin and lower and hinder side of thighs pale yellow ; vent in a triangular dark spot, which is separated

METAPHRYNELLA from the dark of the upper surfaces of the hind-limbs by a zone of yellow ; throat and chest obscurely mottled with brown. Length : 42 mrn. Male with a subgular vocal sac. At the breeding season diffuse, pigmented nuptial pads are developed on the inner metacarpal tubercle and a t the bases of the first and second fingers, on their upper surfaces. DISTRIBUTION : Malay Peninsula, Sumatra and the Mentawei Islands. B.M. 86.12.28. 3 9 4 0 89.11.12. 31 94.12.18. 17 1927.9.1. 2

M.C.Z.

69 6 6 cleared 6

Malacca . . . . . . Deli, Sumatra . . . . Sereinu, Mentawei Islands Siberut, Mentawei Islands Sereinu, Mentawei Islands

.

. . . .

Hervey, COTYPES Moesch Modigliani M. A. Smith Modigliani

ME TAPHR YNELLA Gen. nov. Calohyla, Peters, 1867, Mon. Ak. Berlin, p. 35 (emend. of Kaloulo Gray). Callula (non Gray), Roux 1910, Zool. Anz., xxxv, p. 716 ;Nieden, 1926, Das Tierreich, Anura n, p. 25 (part). Kaloula (non ~ra~):van Kampen, 1923, Amph. Indo-Auatr. Arch., p. 151 (part). Neetophryne (part), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 281. PhryneZlcs (non Boulenger), Giinther, 1887, Ann. Mag. Nat. Hist., (5), XX, p. 313 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 265 (part) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 124 (part) ;Nieden, 1926, Das Tierreich, Anura n, p. 37 (part).

Prevomer divided, the post-choanal portion absent or represented by one or two bony fragments lying free in the mucosa ; palatine absent. No clavicles, procoracoids or omosternwn ; sternum, cartilaginous. Vertebral column procoelous. Terminal phalanges Y-shaped. Pupil horizontal. Tongue oval, slightly notched and scarcely free behind. Two transverse ridges across the palate in front of the pharynx ; a strong dermal ridge behind each choana. Digits strongly dilated ; subarticular tubercles of the hands enlarged to form accessory adhesive organs. Type species : Phnjnellu pollimris, Boulenger. I.

Metaphrynella pollicaris (Boulenger)

Phrynella pulchra (non Boulenger), Giinther, 1887, Ann. Mag. Nat. Hist., (6). XX, p. 313, P1. XVI, fig. B. Phrynella plliulris, Boulenger, 1890, Proc. Zool. Soc. London, p. 37 (Type locality Perak, 3,000 ft.) ; Flower, 1896, Proc. Zool. Soc. London, p. 910 ;idem, 1899, op. cit., p. 908 ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv p. 392 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 266 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 126 ; Nieden, 1926, Das Tierreich, Anura n, p. 38, fig. 27 ; Smith, 1930, Bull. Raffles Mus., m, p. 123. Snout short, scarcely prominent, a little longer than the diameter of the eye ;canthus rostralis obtuse, curved ; loreal region nearly vertical ; interorbital space once and a

half the breadth of the upper eyelid; tympanum hidden. Fingers short, with a rudimentary, fleshy web and triangular dilatations which are about once and a half as wide as the penultimate joint ; f i s t shorter than the second, which is a little shorter than the fourth ; male, a t least, with a strong, bony prepollex ; subarticular tubercles enormously enlarged to form accessory adhesive organs, one on each of the two inner fingers and two on each of the outer. Toes about half webbed, the membrane between the third and fourth toes reaching the level of the distal tubercle of the third toe ; small terminal discs and large subarticular tubercles present ; fifth toe a little shorter than the third ; an elongate inner, but no outer, metatarsal tubercle present. Tibiotarsal articulation reaching the tympanic region. Skin smooth or pustular above ; a fold from the posterior corner of the eye to the fore-limb, the skin above it somewhat thickened ; belly rather granular.

MONOGRAPH OF THE MICROHYLIDAE Olive-brown above ; an oblique yellow line from the eye to the angle of the mouth ; a pale, olive-yellow mark across the forehead, through the eyes and down the sides of the body to the thigh ; this band minutely brown-spotted. A yellow, dark-centred spot on the anal region ; limbs lighter, with a single diagonal cross-bar across the forearm, femur, tibia, tarsus and metatarsus. Lower surfaces brownish, irregularly and indistinctly spotted with white. Length : 38 mm. Male with a subgular vocal sac and infuscate throat. DISTRIBUTION : Malay Peninsula (mountains from 3,000 to 6,000 ft.) and Sumatra.

8

B.M. 87.7.30. 16 1930.2.27. 1

$

2.

SPECIMENS EXAMINED Perak, 3,000 ft. . . . . . . . . Wray, TYPE Cameron Highlands, Pahang, F.M.S., 5,000-6,000 ft. . . . . . . . Robinson

Me taphrynella sundana (Peters)

Calohyla sundam, Peters, 1867, Mon. Ak. Berlin, p. 35 (Type locality Pontianak, Borneo). CaElula sundana, Roux, 1910, Zool. Anz., xxxv, p. 716. KaZoula (?) sundana, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 151. Nectophryne sundana, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 281. Phrynella pollicaris (non Boulenger), Shelford, 1902, Rept. Sarawak Mus., 1901-1902, p. 14. Snout short, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded : loreal region nearly vertical ; interorbital space once and two-thirds to twice as broad as the upper eyelid ; tympanum hidden. Fingers short, with a distinct rudiment of web and large subtriangular terminal dilatations, which are about once and a half as wide as the penultimate joint ; first shorter than the second, which is shorter than the fourth ; no enlarged prepollex ; subarticular tubercles enormously enlarged to form accessory adhesive organs, one on each of the two inner, and two on each of the two outer, fingers. Toes $ webbed, the membrane between the third and fourth reaching well beyond the distal tubercle of the third toe ; distinct terminal discs, smaller than those of the fingers, present ; subarticular tubercles well developed ; third and fifth toes subequal ; metatarsal tubercles absent, or the inner faintly indicated only. Tibio-tarsal articulation reaching the tympanic region. Skin smooth above or with some small, indistinct warts ; a curved supratympanic fold ; belly smooth or feebly granular. Brown above, uniform or indistinctly flecked with darker ; belly yellowish, more or less heavily stippled with brown. Length : 23 mm. Male with a subgular vocal sac. DISTRIBUTION : W. Borneo. SPECIMENS EXARIINED Mus. Berlin 5635 2 ads. Pontianak, Borneo . Hofft, COTYPES Mus. Saram-ak Da. 2.6.1. d-f 8 Kuching . . . , Shelford

. .

.

0TOPHRYNE, Boulenger ?

Otophryne, Boulenger, 1900, Trans. Linn. Soc. London, V I ~ I , p. 55 (Type species Otophryne robusta, Boulenger) ; Nieden, 1926, Das Tierreich, Anura 11, p. 73 ; Noble, 1931, Biol. Amph., p. 536. Prevomer divided, the post-choanal portion absent ; palatine absent. Clavicles and procoracoids well developed, reaching the mid-line of the girdle and the scapulae ; a small cartilaginous ornosternum ; sternum cartilaginous (fig. 46). Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil round. Tongue oval, entire and free behind ; two strong, curved, crenulate ridges in front of the pharynx. Digits not dilated.

OTOPHRYNE

Fig. 46.

Ventral elements of the shoulder-girdle of Otophryne robusta (ventral view)

I.

Otophryne robusta, Boulenger

Otophryne robusta, Boulenger, 1900, Trans. Linn. Soc. London, VIII, 2, p. 55, PI. V, fig. 5 (Type locality Mt. Roraima, 3,500 ft., British Guiana) ; Nieden, 1926, Das Tierreich, Anura 11, p. 73 ; Crawford, 1931, Ann. Carnegie Mus., XXI, 1, p. 38. Snout pointed, prominent, a little longer than the diameter of the eye ; canthus rostralis angular ; loreal region vertical, slightly concave ; nostril beneath the canthus rostralis, equidistant from the eye and the tip of the snout ; interorbital space twice and a quarter as broad as the upper eyelid ; tympanum very distinct, circular, its

diameter equal to the length of the eye, with which it is in contact. Fingers short, obtuse ; the first shorter than the second, which is shorter than the fourth. Toes short, spatulate distally, about & webbed, the fleshy membrane extending as a fringe along each digit ; outer much shorter than the third. No distinct subarticular or metatarsal tubercles. Tarso-metatarsal articulation reaching the tympanum. Skin thick ; smooth above and beneath ; a supratympanic fold, from the posterior corner of the eye to the fore-limb, and a lateral fold from this to the inguinal region. Dark brown above, obscurely spotted with darker ; groin and hinder side of thighs orange, spotted with black ; some obscure black cross-bars on the tibia and foot. Chest and throat dark brown ; belly and lower surfaces of limbs whitish, with some scattered dark brown dots.

110

MONOGRAPH OF THE MICROHYLIDAE

Male with a vocal sac. Length : 53 mm. DISTRIBUTION : Mt. Roraima, British Guiana. The infuscate throat and digital webbing may prove to be secondary sex characters of the male, and further specimens will probably show a colour pattern of dark marbling and spotting on the back. SPECIMEN EXAMINED

B.M. 99.3.25. 18

$

Mt. Roraima, 3,500 ft., British Guiana

. McConnel

and Quelch, TYPE

H YPOPACH U S , Keferstein Engystoma (part), Cope, 1866, Proc. Acad. Philadelphia, p. 131 ; Brocchi, 1877, Bull. Soc. Philom. Paris, (7), I, p. 189 ; Boettger, 1885, Zeitschr. Nat. Halle, LVIII, p. 241. Systoma (part), Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 194. Hypopachus, Keferstein, 1867, Nachricht. Ges. Giittingen, p. 351 (Type species Hypopachus seebachii, Keferstein =Engystoma variolosum, Cope) ; idem, 1868, Archiv. Naturg., x x x ~ v ,1, p. 293 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Rlus., ed. 2, p. 159 ; Cope, 1889, Bull. U.S. Nat. Mus., XXXIV, p. 387 ; Giinther, 1902, Biol. Centr. Amer., Rept. and Batr., p. 211 ; Nieden, 1926, Das Tierreich, Anura 11, p. 70 ; Mir. Ribeiro, 1926, Arch. Mus. Nac. Rio de Janeiro, XXVII, p. 188 ; Parker, 1927, Occ. Papers Mus. Zool. Michigan, CLXXXVII,p. 2, fig. 3 ; Noble, 1931, Biol. Amph., p. 536 ; Kellogg, 1932, Bull. U.S.Nat. Mus., CLX, p. 183. Stereocyclops, Cope, 1871, Proc. Amer. Phil. Soc., XI, 1869, p. 165 (Type species Stereocyclops incrassatus, Cope) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 158 ; Nieden, 1926, Das , 191 ; Tierreich, Anura n, p. 73 ; Mir. Ribeiro, 1926, Arch. bfus. Nac. Rio de Janeiro, x x v r ~ p. Noble, 1931, Biol. Amph., p. 537. Dermatonotus, MBhely, 1904, Ann. Mus. Nat. Hung., n, p. 208 (Type species Engystoma mulleri, Boettger=Stereocyclop8 incrassutus, Cope) ; Nieden, 1926, Das Tierreich, Anura 11, p. 71. Prevomer divided, the post-choanal portion absent ;palatine absent. Clavicles

and procoracoids present, almost straight, reaching the m~d-lineof the girdle and the

Fig. 47.

Ventral elements of the shoulder-girdle of Hypopachus variolosus (ventral view)

HY POPACHUS scapulae ; omosternum absent ; sternum cartilaginous (fig. 47). Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil round or rhomboidal. Tongue large, oval, entire and half free behind. Two smooth dermal ridges across the palate in front of the pharynx, the anterior much shorter than the posterior. Digits not dilated. In all the known species the tympanum is hidden, the first finger shorter than the second and the third toe longer than the fifth. SYNOPSIS OF THE SPECIES

I. A single metatarsal tubercle 11. Two metatarsal tubercles.

. . . . .

. . .

. H.

incrassatus, p. 111

A. Outer metatarsal tubercle rounded, not compressed. Snout not . . . . . H. inguinalis, prominent B. Outer metatarsal tubercle compressed. Snout prominent. (1) Toes of male 4 webbed, those of female 4 ; belly brown with . . . . . H . variolosus, large, round, yellowish spots (2) Toes of male webbed, those of female 1: ; belly white with a regularpalebrownreticulum . H.oqhri,nus, (3) Toes of male not $ webbed, not more than in the female ; belly immaculate, or very indefinitely reticulated . . I$. cuneus,

. . .

. . .

+

I

. Hypopachus

.

. .

. .

p. 112

p. 112 p.113

p. 114

incrassatus (Cope)

Stereocyclops incrassatus, Cope, 1871, Proc. Amer. Phil. Soc., XI, 1869, p. 165 (Type locality SLo Matheos, Brazil) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 159 ; Nieden, 1926, Das Tierreich, Anura 11, p. 73 ; Mir. Ribeiro, 1926, Arch. Rlus. Kac. Rio de Janeiro, XXVII,pp. 193, 226, figs. 103, 104. Engystoma miilleri (or muelleri), Boettger, 1885, Zeitschrift f. Naturwiss., LVIII,p. 241 (Type locality Paraguay) ; idem, 1892, Kat. Batr. Senck. Nat. Ges., p. 22 ; Boulenger, 1894, Ann. Mag. Nat. Hist., (6), XIII, p. 347. Hypopachus miilleri, Peracca, 1895, Bol. Mue. Torino, x, 195, p. 23 ; idem, 1897. op. cit., XII, 274, p. 16 ; Andersson, 1906, Ark. Zool. Stockholm, 111, 12, p. 4 ;Mir. Ribeiro, 1926, Arch. Mus. Nac. Rio de Ja,neiro, XXVII,pp. 189, 226, fig 101. Dermatonotus mulleri, MBheIy, 1904, Ann. Mus. Nat. Hung., 11, p. 208, P1. xnI, figs. 1-3 ; Nieden, 1926, Das Tierreich, Anura 11, p. 72. Gastrophryne muelleri, Stejneger, 1910, Proc. Biol. Soc. Washington, SXIII, 1910, p. 166. Snout pointed, prominent, once and a third the length of the eye ; canthus rostralis rounded, curved ; loreal region slightly oblique, not concave ; nostril much nearer

the tip of the snout than the eye ; interorbital space once and a half to twice as broad as the upper eyelid. Fingers short, free, the second slightly longer than the fourth ; fourth sometimes rather thickened. Toes short, free in females, but with a slight rudiment of web in males ; subarticular tubercles rather feeble ; an oval, uncornpressed, inner, but no outer, metatarsal tubercle. Tarso-metatarsal articulation reaching the shoulder, or a little beyond. Skin thick, smooth above and beneath ; a transverse fold connecting the posterior corners of the upper eyelids and descending to the angle of the jaw. Olive, brown, or purplish above, with a few scattered dark vermiculations ; flanks, temples, groin and limbs dark brown, with numerous, small, rounded, light spots ; sometimes a fine white line from snout t o vent; pale brown, to blackish beneath, with small, regular, rounded white spots which may run together ; hinder side of thighs similar to the flanks. Throat of male infuscate. Male with a vocal sac and some minute, scattered spinules beneath t h e edge of the lower jaw, on the throat and on the inner surfaces of the forearm and fingers. Length: '72 mm. DISTRIRUTION : Paraguay, S.E. Bolivia, S.W. and N.E. Brazil.

PIIONOGRAPH OF THE IVIICROHYLIDAE

9,hgr. and juvs. 8

B.M. 94.3.14. 89-105 98.6.3.14

?

F.M.N.H. 49673 Mus. Vienna

hgr.

? 2.

Asuncion, Paraguay . Bohls Puerto 14 de Rlayo, Bahia Negra, N. Chaco . . . Boggiani Natal, X.E. Brazil Heath Rio das Velhas , . Steindachner ? . Fiebrig

.

. . .

. . .

Hypopachus inguinalis, Cope

Hypopachus inguinalis, Cope, 1869, Proc. Amer. Phil. Soc., XI, 1869, p. 166 (Type locality Coban, Vera Paz) : Brocchi, 1882, Miss. Sci. Mex., Batr., p. 91, P1. X, figs. 3-4 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Nus., ed. 2, p. 160 ;Giinther, 1902, Biol. Centr. Amer., Rept. and Batr., p. 211 ; Nieden, 1926, Das Tierreich, Anura 11, p. 71. Engystoma inguinalis., Brocchi, 1877, Bull. Soc. Philom. Paris, (7), I, p. 189. Hppopachus variolosus inguinalis, Cope, 1887, Bull. U.S. Nat. Mus., XXXII, p. IS.

Snout short, rounded, or vertically truncate, not prominent, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region oblique, not concave ; interorbital space once and a half as broad as the upper eyelid. Fingers free, the second a little shorter than the fourth. Toes with a rudiment of web in females and nearly a quarter webbed in males ; subarticular tubercles well developed ; two prominent metatarsal tubercles, the inner oval, the outer rounded, neither with a pronounced cutting edge. Tarso-metatarsal articulation reaching the eye. Skin smooth above and beneath ; a transverse fold connecting the posterior corners of the upper eyelids and descending to the angles of the jaw, and another from the posterior corner of the eye to beneath the fore-limb. Dark reddish brown above, sometimes pinkish posteriorly and on the thighs ; some obscure darker markings which may have the form of a broad mid-dorsal stripe, wider posteriorly than anteriorly ; temporal region and flanks darker ; a round, black, inguinal spot, partially covered by the folded hind-limbs, and a broad diagonal bar, sometimes broken up, across the femur, tibia and tarsus. Yellowish beneath, the throat and chest heavily stippled with brown ; the belly and lower surfaces of the limbs with large, bold, dark brown spots and reticulations. A fine, light line from snout to vent dorsally, meeting, above the vent, another which runs along the hinder side of the limbs to the heel ; frequently a fine mid-ventral line meeting, on the chest, another which runs to the wrist. A light streak from the eye t o beneath the fore-limb. Length : 37 mm. DISTRIBUTION : Guatemala. There has been no confirmation of the doubtfill record of this species from Ecuador.

R.M. 60.6.19. 1 5 60.12.1. 18-22 $, 64.1.26. 169 2Q 1920.1.20, 541

illus. Vienna

9 hgr. 9,hgr.

9,hgr., yg.

[Ecuador] . . . Coban, Guatemala . Coban, Guatemala . Vera Paz, Guatemala Coban, Guatemala .

.

.

.

. Fraser

. . . . . . , . . . . . .

Salvin Braconnier

. . . Schneider

3 . Hypopachus variolosus (Cope) Engyslstorna variolosum, Cope, 1866, Proc. Acad. Philadelphia, p. 131 (Type locality Arriba, Costa. Rica) ; Brocchi, 1877, Bull. Soc. Philom. Paris, (7), I, p. 192. Systoma variolosum, Cope, 1867, ,Tourn. Acad, Philadelphia, (2), VI, p. 194. Hypopachus variolosus, Cope, 1871, Proc. Amer. Phil. Soc. XI, 1869, p. 166 ; Brocchi, 1882, Miss. Sci. Mex., Batr., p. 92, P1. x, fig. 2 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 3, p. 159 ;Giinther, 1902, Biol. Centr. Bmer., Rept. and Batr., p. 211 (p-art); Kieden, 1926, Das Tierreich, Anura 11, p. 70 (part) ; Mir. Ribeiro, 1926, Arch. Mus. Nac. Rlo de Janeiro, XXVII,pp. 191,226, fig. 102; Kellog, 1932, Bull. U.S. Nat. ~ ~ U S . , C L Xp., 184(part).

HYPOPACHUS

113

Hypopachus saebachii, Keferstoin, 1867, Nachricht. Ges. Wiss. Gottingen, p. 352 (Type localit'y Costa Rica) ; idem, 1868, Arch. Naturg., p. 293, P1. IX,figs. 1-2. Snout pointed, prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region oblique, not concave ; interorbital space once and a half as broad as the upper eyelid ; fingers free, the second subequal to, or a little shorter than, the fourth. Toes not dilated ; the web between the third and fourth toes extends slightly beyond the proximal tubercle of the third toe in females : toes about half webbed in males, the fleshy membrane extending almost to the tip of the outer toe ; subarticular tubercles well developed ; two large, compressed, metatarsal tubercles, the outer transverse, and the inner measuring 8 to 2 its distance from the tip of the inner toe. Tarso-metatarsal articulation reaching the posterior corner of the eye. Skin smooth above and beneath, or uniformly beset above and on the chin with numerous minute spinules in males ; a transverse fold connecting the posterior corners of the upper eyelids and descending obliquely to beneath the arm. Brownish or pinkish above ; a dark lateral streak from the tip of the snout through the eye to the groin ; sometimes more or less distinct traces of a dark mid-dorsal area, with undulating borders, from between the eyes to the vent, wider posteriorly than anteriorly ; oblique transverse dark bars across femur, tibia and tarsus, and one or more dark blotches in the inguinal region. Pale brown beneath, with numerous, moderately large, rounded, yellowish spots. Hinder side of thighs dark brown with large yellow spots. Sometimes a fine light mid-dorsal and mid-ventral line and transverse light lines from heel to heel and wrist to wrist. Male with a vocal sac. Length : 48 mm. DISTRIBUTION : Costa Rica. The record of this species from Para is certainly incorrect ; the specimen on which it was based is really H. cuneus and, presumably, wrongly labelled for locality. Kellogg has recently regarded H. oxyrhinus, Boulenger, as a synonym of this species. But the differences in the webbing of the feet of the males, as well as the colour, make this improbable. It is, however, quite possible that H. variolosus, H. oxyrhinus and N. cuneus are all local races of the same species, for they form a nicely graded series. 11. variolosus, the most southerly form, has the most marked difference between the sexes, so far as webbing is concerned, and has a dark, light-spotted venter. In H. oayrhinzcs the disparity between the sexes is less marked, though still pronounced, and the dark pigment of the venter is reduced to a fine reticulum. H. czcneus, the most northerly form, has no appreciable difference in the webbing of the sexes, and the lower surfaces are almost completely immaculate white. Future collecting may show that each of these forms grades insensibly into the next, but in the absence of intermediates they are accorded full specific status. SPECIJIENS EXARIISED B.M. 68.8.17. 7 9 Costa Rica . , . . . . . . Salvin 74.8.13. 3-4 $28 Costa Rica . . , . . . . . 94.1 1.15. 24-25 Bebedero, Costa Rica . . . . . Underwood 1907.6.28.17-19 $, $2 9 San Jos6, Costa Rica . . . . . Underwood Mus. Vienna S! Costa Rica . . . . . . . . Gliickauf ad. San Jose, Costa Rica . . . . . Underwood

4. Hypopachus oxyrhinus, Boulenger Hypopachus oxyrhinus, Boulenger, 1883, Ann. Mag. Nat. Hist., (5) XI, p. 344 (Type locality Presidio de Mazatlan) ; Boettger, 1892, Kat. Batr. Senck. Kat. Ges., p. 22.

Hypopachus variolosus (part), Gunther, 1902, Bjol. Centr. Amer., Rept. and Batr., p. 21 1 ; Nieden, 1926, Das Tierreich, Anura 11, p. 70 ; Kellogg, 1932, Bull. U.S. Nat. Mus., CLX, p. 184. Snout pointed, prominent, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique, not concave ; interorbital space once and

MONOGRAPH OF THE MICROHYLIDAE three-quarters as broad as the upper eyelid. Fingers free, the second subequal to, or a little shorter than, the fourth. Toes moderate ; females with a rudiment of web, which, between the third and fourth toes extends to the level of the proximal tubercle of the third toe ; males with slightly more webbing, which extends as a distinct fringe a t least on the proximal half of each toe ; subarticular tubercles well developed ; two large, compressed, metatarsal tubercles, the outer almost transverse and the inner measuring 8 to 2 its distance from the tip of the inner toe. Tarso-metatarsal articulation reaching the middle of the eye or a little beyond. Skin smooth or slightly shagreened above ; smooth beneath ; sometimes a transverse fold connecting the posterior corners of the upper eyelids and descending to the angles of the mouth, and another from the posterior corner of the eye to beneath the arm. Male beset with microscopic spinules above, beneath the chin, and on the hands and feet. Purplish brown to dark grey above ; a more or less distinct dark, lateral stripe from the tip of the snout through the eye to the groin ; sometimes a darker mid-dorsal area, commencing between the eyes and broadening posteriorly, with darker, pronounced, undulating lateral borders which form a rounded spot in each inguinal region and are continued as oblique transverse bars on the femur, tibia and tarsus ; an oblique light streak from the posterior corner of the eye to beneath the fore-limb. Yellowish beneath with a regular, pale brown reticulum. Throat of male infuscate. Male with a vocal sac. Length : 38 mm. DISTRIBUTION : IV. Mexico (Sinaloa, Jalisco, RIichoacan), Guatemala.

B.M. 82.12.5. 8 83.4.5. 3 92.2.8. 65 1914.1.28. 150 1914.1.28. 151 1914.1.28. 152

Mus. Vienna

$2 $2

9 8 Q

$ hgr.

Presidio de Mazatlan, Sinaloa . . Forrer Presidio de Mazatlan, Sinaloa . . Forrer COTyP"S Agua Delgada, Guadalajara, Jalisco Buller San Salvador, Michoacan, 2,500 ft. . Gadow Buena Vista, Michoacan, 2,000 ft. . Gadow Cofradia, Michoacan, 700 ft. . . Gadow Coban, Vera Paz, Guatemala . Schneider

)

.

5. Hypopachz~cuneus, Cope f1ypo;uachus variolosus (part), Boulenger, 1882, Cat. Batr. Sal, Brit. Mus., ed. 2. p. 159 ; Giinther, 1902, Biol. Centr. Amer., Rept. and Batr., p. 211 ; Baumann, 1912, Zool. Jahrb., Syst., XXXIII, pp. 122, 123, 145, 161, 166 ; Nieden, 1926, Das Tierreich, Anura 11, p. 70. Hypopachus cuneus, Cope, 1889, Bull. U.S. Nat. Nus., x x x ~ v ,p. 388, fig. 98 (Type locality San Diego, Duval County, Texasj ; idem, 1889, Proc. Amer. Phil. Soc., XI, p. 392 ; Boulenger, 1890, Zool. Rec., 1889, Rept. and Batr., p. 21 ; Dickerson, 1,906,Frog Book, p. 169, Col. P1. 11, fig. 1, PI. LXI ; Strecker, 1908, Proc. Biol. Soc. Washington, x x ~ p. , 58 ; idem,, 1915, Baylor Bull., XVII;, 4, p. 47 ; Stejneger and Barbour, 1923, Check List N. Amer. Amph. and Rept., ed. 2, p. 38 ; Kellogg, 1932, Bull. U.S. Nat. Mus., c ~ xp., 153.

Snout pointed, prominent, as long as, or slightly shorter than, the diameter of the eye.; canthus rostralis rounded ; loreal region almost vertical, not concave ; interorb~t-a1space once and a half as broad as the upper eyelid. Fingers free, the first shorter than the second, which is subequal to, or slightly shorter than, the fourth. Toes moderate ; webbing rudimentary in both sexes, not extending beyond the proximal tubercle of the third toe ; subarticular tubercles well developed. Two compressed metatarsal tubercles, the outer almost transverse and the inner measuring about $ its distance from the tip of the inner toe. Tarso-metatarsal articulation reaching the hinder border of the eye. Skin smooth or slightly shagreened above ; smooth beneath ; sometimes a transverse fold connecting the posterior corners of the upper eyelids and descending to the angles of the jaw, and another from the posterior corner of the eye to beneath the arm. Greenish, yellowish or greyish brown above ; sometimes a dark lateral stripe from the tip of the snout through the eye to the groin, but this is usually only represented

RIBEIRINA

115

by a series of spots and blotches corresponding to its upper margin ; similarly, a dark mid-dorsal area commencing between the eyes, very narrow on the nape, broadening abruptly between the shoulders and then gradually widening caudad, is often represented by lines and dots corresponding with its lateral borders ; these spots are pronounced and larger in the groin, and continue as single or double cross-bars on the femur, tibia and foot ; hinder side of thighs brown dotted ; usually a fine white line from snout t o vent and an oblique light streak from the posterior corner of the eye to beneath the fore-limb. Lower surfaces yellowish, the belly immaculate, or with a very indefinite, pale brown reticulum ; throat brown stippled or, in males, infuscate. Male with a vocal sac. Length : 44 mm. : Texas. DISTRIBUTION

B.M. 73.4.30.8 87.1.4.42-43

...

. . . . . $3,$?9 Duval County, Texas . . . . (Para) .

(Berlin Mus.) Taylor

92.10. 24.43-44

RIBEIRINA, nom. nov. Ernydops (non Broom, 1912), Mir. Ribeiro, 1920, Rev. Mus. Paulista, 111, p. 286 (Type species Emydops hypomelas, Mir. Ribeiro) ; idem, 1926, Arch. Mus. Nac. Rio de Jsneiro, XXvIi, p. 187 ; Parker, 1927, Occ. Papers Mus. Zool. Michigan, c~xxxmr,p. 2, fig. 2.

[After Mir. Ribeiro

Fig. 48. Ventral elements of the shoulder-girdle of Ribeirina hypom,elas (ventral view)

MOKOGRAPH OF THE MICROHYLIDAE Prevomer 'r. Clavicles and procoracoids present, well developed, reaching the scapulae and the mid-line of the girdle, strongly bent and following the curvature of the coracoids ; no omosternum ; sternum cartilaginous (fig. 48). Pupil horizontal. Tongue oval, entire and free behind ; a bony, toothless ridge behind each choana ; a dermal fold across the palate in front of the pharynx. Digits not dilated, free. The bony post-choanal ridges, if correctly described, would appear to distinguish this genus from any of its S. American relatives.

I

. Ribeirina

hypomelas (Mir. Ribeiro)

Emydops hypomelas, Mir. Ribeiro, 1920, Rev. Mus. Paulista, XII,p. 287, PI. 11, figs. 4,5 and 6. (Type locality Porto Cachoeira, Espirito Santo, Brazil) ; idem, 1926, Arch. Mus. Nac. Rio de Janeiro, XXVII, pp. 188 and 226. Head moderately depressed ; snout rounded, slightly prominent, longer than the diameter of the eye ; nostril nearer the tip of the snout than the eye ; tympanum hidden. Fingers moderate, the first, second and fourth subequal. Toes slender, the third much longer than the fifth ; subarticular tubercles small ; an inner metatarsal tubercle. Skin smooth above and beneath ; a transverse occipital fold connecting the posterior corners of the upper eyelids and descending to the angles of the jaw. Cinereous above, with interrupted, longitudinal black streaks ; a sinuous streak from the nostril, above each eye to the nape ; sides of the head as far as the posttympanic region, and all the lower surfaces, black. Limbs marbled. Fine white mid-dorsal and mid-ventral lines from the tip of the snout to the anus ; the former meeting a similar line which extends from heel to heel along the hinder sides of the legs, and the latter a line connecting the first fingers of each hand. Length : 45 mm. DISTRIBUTION : Known only from the type locality, Porto Cachoeira, Espirito Santo, Brazil.

CHIASMOCLEIS, Mkhely Engystoma (non Fitzinger), Boettger, 1885, Zeitschr. f, Nat. Halle, LxvnI, p. 240 (part); Boulenger, 1888, Ann. Mag. Nat. Hist., (6), I, p. 416 ; Mir. Ribeiro, 1920, Rev. Mus. Paulista, XII,p. 281 (yart) ; idem, 1926, Arch. Mus. Nac. Rio de Janeiro, xxvn, p. 182 (part); Nieden, 1926, Das T~erreich,Anura a,p. 63 (part). Chiasmodeis, MBhely, 1904, Ann. Mus. Nat. Hung., 11,p. 210 (Typespecies Engystoma albopunclatum, Boettger) ; Mir. Ribeiro, 1920, Rev. Mus. Paulista, XII, p. 285 ; idem 1926, Arch. Mus. Nac. p. 187 ; Nieden, 1926, Das Tierreich, Anura rr, p. 72 ; Parker, 1927, Rio do Janeiro, XXVII, Occ. Papers Mus. Zool. Michigan, CLXXXVII, p. 3, fig. 4; Noble, 1931, Biol. Amph., p. 536. Gastrophryne (part), Stejneger, 1910, Proc. Biol. Soc. Washington, x x ~ np. , 166. Nectodactylus, Mir. Ribeiro, 1924, Bol. Mus. Nac. Rio de Janeiro, IV, p. 256 (Type species Bectodactylus spinulosus, Mir. Ribeiro) ; idem, 1926, Arch. Mus. Nac. Rio de Janeiro, xxvrr, p. 182 ; Noble, 1931, Biol. Amph., p. 536. Prevomer divided, the post-choanal portion wanting ; palatine absent. Clavicle and procoracoid present, short, reaching the mid-line of the girdle, but, meeting the coracoid in its lateral half, do not reach the scapula ; no omosternum ; sternum cartilaginous (fig. 49). Vertebral column diplasiocoelous. Terminal phalanges simple. Pupil round. Tongue oval, entire and free behind ; two smooth dermal ridges across the palate in front of the pharynx, the anterior much shorter than the posterior. Digits without, or with very small, terminal discs. In all the known species the tympanum is hidden, the first finger much shorter than the second, the outer toe shorter than the third and the outer metatarsal tubercle wanting.

CHIASMOCLEIS

Fig. 49. Ventral elements of the shoulder-girdle of Chiasmocleis albopunctata (ventral view)

SYNOPSIS OF THE SPECIES I. Epicoracoids long ; maxilla and jugal widely separated. Fingers with more or less distinct fleshy webbing or fringes. A. Toes 8 webbed . . . . . . . . . . . . C. spinulosa, R. Toes not more than ;5 webbed. (1) Canthus rostralis well marked, angular C. bicegoi, (2) Canthus rostralis rounded. (a) Toes with a distinct rudiment of web ; eye a little longer than its distance from the nostril . . . C. leucosticta, (b) Toes quite free ; eye a little shorter than its distance from the nostril . . . C. alhopunctata, 11. Epicoracoids short ; maxilla and jugal in contact. Fingers without any trace of webbing or fringes . . . . . . . . . C. boliviana,

. . . . .

p. 117 p. 118

p. 118 p. 118

p. 119

Kectodactylus spinulosus, Mir. Ribeiro, 1924, Bol. Mus. Nac. Rio de Janeiro, rv, p. 256 (Type locality near Humboldt, Sta. Catharina) ; idem, 1926, Arch. Mus. Nac. Rio de Janeiro, XXVII, pp. 186, 225, fig. 100. Snout rounded, scarcely prominent, once and a half as long as the eye ; canthus rostrdis rounded ; Ioreal region oblique, not concave ; nostril much nearer the tip of the snout than the eye ; interorbital space 29 times as broad as the upper eyelid.

Fingers short, knobbed, all bound together in a thick, fleshy web which extends to their tips ; the second subequal to the fourth. Toes slightly dilated distally, g webbed, the thick web extending as a fringe to their tips ; subarticular tubercles moderate ; a small inner metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder. Skin beset with minute spinules above, beneath the throat and on the hands and feet, particularly on the edges of the interdigital membranes ; smooth beneath ; a feeble fold from the eye to the anterior side of the fore-limb.

118

MONOGRAPH OF THE MICROHYLIDAE

Uniform brown above ; flanks yellow spotted : lower surfaces yellow, heavily blotched with brown. Throat of male uniform dark brown. Male with a vocal sac. Length : $ 22 mm. ; 9 25 mm. DISTRIBUTION : Sta. Catharina, Brazil. This species is said to live in the water in the axils of scitaminaceous plants. SPECIMENS EXAMINED B.M. 1925.11.6. 1-2 $$ Sta. Catharina . . . . . . . . Erhardt

.

2 . Chiasmocleis bicegoi, Mir Ribeiro Chiasmocleis bicegoi, Mir. Ribeiro, 1920, Rev. Mus. Paulista, xrr, p. 286 (Type locality 0s Perus, Siio Paulo, Brazil) ; idem, 1926, Arch. Mus. Nac. Rio de Janeiro, XXVII, pp. - - 187, 226. Snout prominent, pointed, once and a half as long as the eye ; canthus rostralis well marked, angular. Hand as in Alelopzcs ; fingers slightly knobbed distally, the first shorter than the second, which is shorter than the fourth. Subarticular tubercles indistinct ; metatarsal tubercles indistinct. Tibio-tarsal articulation not quite reaching the tympanic region. Skin smooth above and beneath ; no occipital fold ; a supratympanic fold to the shoulder. Ash-grey above ; flanks and lower surfaces vermiculated with yellow ; a white line from the tip of the snout meeting, on the coccyx, another which traverses the hinder side of the thighs. Length : 16 mm. DISTRIBUTIOX : 0 s Perus, Sao Pamulo,Brazil.

3. Chiasmocleis leucosticta (Boulenger) En,gystoma Zeucostictum, Boulenger, 1888, Ann. Mag. Nat. Hist., (6), I, p. 416 (Type locality Sierra do Catharina, Sta. Catharina, Brazil) ; Peracca, 1904, Bol. Mus. Torino, XIX, 460, p. 12 : Baumann, 1912, Zool. Jahrb., Syst., xxxr~r,pp. 143, 145, 148, 161 ; Mir. Ribeiro, 1920, Rev. Mus. Paulista, XTT, p. 282 ; idem, 1926, Arch. Mus. Nac. Rio de Janejro, XXVII,pp. 185, 225 ; Nieden, 1926, Das Tierreich, Anura 11, p. 68. Gastrophryne leucosticta, Stejneger, 1910, Proc. Biol. Soc. Washington, XXIII,p. 166. Snout rounded, scarcely prominent, once and a half as long as the diameter of the eye ; canthus rostralis rounded 3 loreal region oblique, not concave ; nostril much nearer the tip of the snout than the eye ; interorbital space about 24 times the width of the upper eyelid ; Fingers short, knobbed distally, with a trace of fleshy webbing a t the base and indications of lateral fringes ; the second subequal to the fourth. Toes knobbed, with a distinct rudiment of webbing and traces of fleshy lateral fringes ; subarticular tubercles moderate ; a small inner metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder. Skin smooth above and beneath ; a feeble fold from the eye to the anterior side of the arm. Uniform dark brown above ; yellowish beneath, more or less heavily blotched with brown. Length : 25 mm. DISTRIBUTION : Sta. Catharina, Brazil. SPECIMENS EXAMINED Q Sierra do Catharina, Sta. Catharina Michaelis, TYPE B.M. 88.4.23.3 9 Therezopolis, Sta. Catharina Friihstorfer 88.9.21. 20

. .

.. ..

4. Chiasmocleis albopunctata (Boettger) Engystoma albopuncfatum, Boettger, 1885, Zeitschr. f, Nat. Halle, LVITI, p. 240 (Type locality Paraguay) ; Peracca, 1904, Bol. Mus. Torino, xrx, 460, p. 12 ;Mir. Ribeiro, 1920, Rev. MUS. Paulista,

CHIASMOCLEIS XII,p. 282 ; idem, 1926, Arch. Mus. Nac. Rio de Janiero, XXVII, pp. 185, 225. Chiasniocleis albopunctaia, MBhely, 1904, AM. Mus. Nat. Hung., 11. p. 210, P1. XIII,figs. 4-5 ; Nieden, 1926, Das Tierreich, Anura 11, p. 72. Gastrophryne albopunclata, Stejneger, 1910, Proc. Biol. Soc. Washington, XXIIT, p. 166. Snout rounded, slightly prominent, once and a half as long as the eye ; canthus rostralis rounded ; loreal region oblique, not concave ; nostril much nearer the tip of the snout than the eye ; interorbital space 23 times as broad as the upper eyelid. Fingers not dilated, short, depressed, with more or less distinct fleshy lateral fringes ; the second subequal t o the fourth. Toes not dilated, depressed, free, but with traces of fleshy lateral fringes ; subarticular tubercles not well developed ; a small inner metatarsal tubercle. Tibio-tarsal articulation not quite reaching the shoulder. Skin smooth above and beneath ; an indistinct fold from the posterior corner of the eye to the anterior side of the arm. Dark brown, grey or blackish above, regularly marked with small whitish or silvery spots which may form an irregular streak along the canthus rostralis, the edge of the upper eyelid and for a short distance behind the eye. Dark yellow, brown or blackish beneath, regularly spotted with lighter. Throat of male black. Male with a vocal sac. Length : 29 mm. DISTRIBU~ION : Paraguay, Rfatto Grosso.

B.M.94.3.14.94-9s Jlus. Vienna

$, 4

6

$!

9

Asuncion, Paraguay Chaco, Paraguay .

. . . . . . . . . .

Bohls Fiebrig

5. Chiasmocleis boliviana, Parker Chiasmocleis bolizliana, Parker, 1927, Occ. Papers Mus. Zool. Michigan, CI,XXXVII, p. 3., fig. 5 (Type locality Buena Vista, Bolivia). Snout obtuse, scarcely prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region very oblique, not concave ; nostril nearer the tip of the snout than the eye ; interorbital space twice as broad as the upper eyelid. Fingers moderate, free, knobbed distally, the second a little shorter than the fourth. Toes free, knobbed distally ; subarticular tubercles large and flat ; an oval inner metatarsal tubercle. Tibio-tarsal articulation reaching the tympanic region. Skin smooth above and below ; a transverse fold connecting the posterior corners of the upper eyelids and descending to the angles of the jaw. Grey or pinkish above ; a broad, brown, dorsal stripe commencing between the upper eyelids, narrowing on the nape, broadening behind the scapulae, narrowing again, broadening on the sacral region and sending a diagonal branch towards each groin ; continuations of these branches form diagonal cross-bars on the femora, tibiae and tarsi, but only their outlines may persist ; loreal region black ; a dark lateral stripe, distinctly outlined above but merging into the ground colour of the belly beneath, runs from the upper eyelid, through the groin to the anterior surface of the knee ; a similar, but more distinct stripe extends from above the vent to the back of the knee ; a distinct, round, black inguinaI spot, not covered by the folded hind-limbs. A fine white line from snout to vent. White beneath, with brown spots and vermiculations which become denser towards the flanks ; lower surfaces of tarsus and foot black. Length : 39 mm. DISTRIBUTION : S.E. Bolivia. Possibly the differences in the skull and shoulder girdle which distinguish this species from the others of Chiasmocleis are suficient to justify its removal to a separate genus. Buena Vista, Bolivia $' B.M. 192'7.8.1.1 Mus. Zool. Michigan ad., hgr. Buena Vista, Bolivia

. . Steinbach, HOLOTYPE . . Steinbach, PARATYPES

120

MONOGRAPH OF THE MICROHYLIDAE

T h e following specimen closely resembles, a n d m a y ultimately prove t o belong to, t h e preceding species, b u t it is t o o y o u n g (length 14 mm.) for a c c u r a t e specific identification ; since it w a s f o u n d 1,700 miles a w a y f r o m t h e k n o w n r a n g e of C. boliviana i t does n o t seem advisable t o include it i n t h a t species. B.M. 1934.1.4. 1 juv. U p p e r Cuyuni River, Brit. Guiana Carter

. . . .

ELACHISTOCLEIS, Parker Rana (part), Schneider, 1799, Hist. Amph., I, p. I l l ; Merrem, 1820, Tent. Syst. Amph., p. 174. Bufo (part), Daudin, 1803, Hist. Rainettes, p. 71 ; idem, 1803, Hist. Rept., VIII, p. 137. Microps (non Megerle, 1823), Wagler, 1828, Isis, p. 744 (Type species JIicrops unicolor, Wagler = Rana ovalis, Schneid.) ; idem, 1830, Syst. Amph., p. 200. ~'tenocephalus(non Latreille, 1825), Tschudi, 1838, Mem. Soc. Sci. Neuchiite!, 11, 1839, pp. 49, 86 (Type species Stenocephalus microps, Wagler =Rana ovalis Schneid.). Oxyrhynchus (non Spix), Valenciennes, 1838, in GuBrin MBneville, Icon. GCn. RBgne Animal, I, Rept., P1. xxvrr, fig. 2 ; idem, 1844, op. cit., III, Rept. p. 17. Bastrophryne (part), Fitzinger, 1843, Syst. Rept., p. 33 ;Stejneger, 1910, Proc. Biol. Soc. Washington, xxrrr, p. 166. Engystoma (part), Durn. and Bibr., 1841, Erpet. GBn., VIII, p. 738 ; Giinther, 1858, Cat. Batr. Snl. Brit. Mus., p. 51 ;Cope, 1867, Journ. Acad. Philadelphia, VI, p. 194 ;Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 160 :Mir. Ribeiro, 1920, Rev. Mus. Paulista, XII, p. 281 ; idem, 1926, Arch. Mus. Nac. Rio de Janeiro, xxvn, p. 182 ; Nieden, 1926, Dns Tierreich, Anura 11, p. 63. Hypopachus (non Keferstein), Ruthven, 1914, Proc. Biol. Soc. Washington, XXVII, p. 77. Elachistocleis, Parker, 1927, Occ. Papers Mus. Zool. Michigan, c ~ x x x v r r ,p. 4, fig. 6 (Type species Rana ovalis, Schneid.). P r e v o m e r divided, t h e post-choanal portion lost ; palatine absent. Clavicle and procorncoid present, m u c h reduced, curved, resting o n t h e mesial half of t h e coracoid ; o m o s t e r n u m a b s e n t ; s t e r n u m cartilaginous (fig. 50). Vertebral column diplasiocoelous. Terminal phalanges simple. P u p i l round, t o n g u e oval entire a n d free behind. Two smooth, or slightly crenulate, d e r m a l ridges i n f r o n t of t h e p h a r y n x , t h e a n t e r i o r s h o r t e r a n d curved. Digits free, t h e t i p s n o t dilated.

Fig. 50. Ventral elements of the shoulder-girdle of Elachistocleis ovalia (ventral view)

ELACHISTOCLEIS I

.

Elachistocleis ovalis (Schneider)

Rana ovalis, Schneider, 1799, Hist. Amph., p. 131 (Type locality not cited) ; Shsw, 1802, Zool., 111, p. 111. Bufo ovalis, Daudin, 1803, Hist. Rainettes, p. 92 ; idem, 1803, Hist. Rept., VIII, p. 187. Engystoma ovale, Fitzinger, 1826, N. Class. Rept., p. 65 ; Dumhril and Bibron, 1841, Erpet. Ghn., VIII, p. 741 ; Giinther, 1858, Cat. Batr. Sal. Brit. hlus., p. 51 ; Steindachner, 1864, Verh. Zoo1.-bot. Ges. Wien, p. 285, PI. XVIII, fig. 4 ;Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 163 ; Boettger, 1885, Zeitsch. f. Nat. Halle, LVIII, p. 240 ; Boulenger, 1886, Ann. Mag. Nat. Hjst., ( 5 ) , xvrII, p. 439 ; Cope, 1887, Proc. Amer. Phil. Soc., x x ~ v ,p. 53 ; Boettger, 1892, Kat. Batr. Senck. Nat. Ges., p. 22 ; BouIenger, 1894, Ann. Mag. Eat. Hist., (6), xIrI, p. 347 ; Peracca, 1895, Bol. Mus. Torino, x, 195, p. 23 ; Berg, 1896, Ann. Mus. Buenos Aires, v, p. 159 ; Werner, 1899, Verh. Zoo1.-bot. Ges. Wien, p. 0 ; Peracca, 1904, Bol. Mus. Torino, xrr, 460, p. 11 ; Mhhely, 1904, Ann. Mus. Nat. hung.,^^, p. 211 : Lidth de Jeude, 1904, Notes Leyden Mus., XXV, p. 93 ; Baumann, 1912, Zool. Jahrb., Syst., XXXIII, pp. 129, 143, 148, 161; Marelli, 1924, Elenco Sist. Fauna Buenos Aires, p. 585 ; Mir Ribeiro, 1926, Arch. MUS.Kac. Rio de Janeiro, xxvn, pp. 184, 225 ; Lutz, 1927, Mem. Inst. Oswaldo Cruz, PI. VIII, fig. 2 ; idem, 1928, Est. Zool. Par. Venezuela, p. 92, P1. XVI, fig. 2. Engystoma ovalis, Nieden, 1926, Das Tierreich, Anura 11, p. 66. Gastrophryne ovaiis, Stejneger, 1910, Proc. Biol. Soc. Washington, XXITI, p. 166. G . ovale, Ruthven, 1922, Misc. Pub. Mus. 2001. Michigan, VIII, p. 51. Engystoma ovale ovab, Mir. Ribeiro, 2920, Rev. Mus. Paulista, XII, 1920, p. 282. Elachistocleis ovale ovale, Parker, 1927, Occ. Papers Mus. Zool. Michigan, cLxxxvn, p. 4 ; Elachistocleis ovale, Mertens, 1929, Blatt. Aq. Ten., XI,, 16, p. 286. Elachistocleis ovalis, Crawford, 1931, Ann. Carnegie Mus., XXI,1, p. 38. Bvfo surinamensis, Daudin, 1803, Hist. Rainettes, p. 91, PI. XXXIII, fig. 2 (Type locality Surinam) ; idem, 1803, Hist. Rept., VIII, p. 186. Rana bufonia, Merrem, 1820, Tent. Syst. Amph., p. 177 (substitute name for Bujo surinamensis, Daudin). ili'icrops unicolor, Wagler, 1830, Syst. Amph., p. 200 (substitute name for Rana ovalis, Schneid.). Qmtrophrpe unicolor, Fitzinger, 1843, Syst. Rept., p. 33. Stenocephalus microps, Tschudi, 1838, Rlem. Soc. Sci. Neuchltel, 11, 1839, p. 86 (substitute name for Rana oaalis, Schneid.). Oxyrhynchus bicolor, Valenciennes, 1838, in Gu8rin Mhneville, Icon. G8n. RBgne Animal, I, Rept., PI. XXVII, fig. 2 ; idem, 1844, op. cit., p. 17 (Type locality S. America). Engystorna ovale var. bicolor, Boulenger, 1885, Ann. Mag. Nat. Hist., (5), xv, p. 195 ; idem, 1885, op. cit., (5),xvr, p. 296 ; Boettger, 1885, Zeitsch. f. Nat. Halle, LvIrI, p. 240. Engystoma ovule bicolor, Mir. Ribeiro, 1920, Rev. Mus. Paulista, XII, p. 283 ;idem, 1926, Arch. Rlus. Nac. Rio de Janeiro, XXVII, p. 225 ; Elachistocleis ovale bicolor, Parker, 1927, Occ. Papers Mus. Zool. Michigan, CLXXXVII, p. 4, Hypopachus pearsei, Ruthven, 1914, Proc. Biol. Soc. Washington, xxvn, p. 77 (Type locality Fundacion, Sta Marta Mts., Colombia) ; idem, 1922, hfisc. Pub. Mus. Zool. Michigan, VIII, p. 50, P1. xr, fig. 3. Engystoma ovale lineata, Mir. Ribeiro, 1020, Rev. Mus. Paulista, XII, p. 284 (Type locality R. da Serra, Sfo Paulo). Engystorna ovale caesarii, Mir. Ribeiro, 1920, loc. cit. (Type localities Piquete, 0 s Perus, Alto da Serra (SBo Paulo), Cubatao (Santos) and Ypiranga). Engystoma ovale conwlor, Mir. Ribeiro, 1920, loc. cit. (Type locality Ypiranga). Engystoma caesarii mottae (nom. mud.), Mir Ribeiro, 1920, loc. cit., footnote (Type locality Siio Paulo). Snout pointed, very prominent, 1% to 1s times as long as the eye ; canthus rostralis rounded ; loreal region oblique, scarcely concave ; interorbital space about twice as broad as the upper eyelid ; tympanum hidden. Fingers very short, the first much shorter than the second, which is a little shorter than the fourth ; latter more or less

distinctly knobbed distally. Toes short, sometimes depressed and with rudimentary fleshy lateral fringes ; third much longer than the fifth ; subarticular tubercles moderately well developed ; an oval inner, but no outer, metatarsal tubercle. Tibiotarsal articulation reaching the shoulder or a little beyond. Skin smooth above and beneath ; a fold connecting the posterior corners of the upper eyelids and descending to the angles of the jaw; sometimes, in males, the dorsum, hands, feet, chin and throat are beset with microscopic spinules. Brown, grey or purplish black above, uniform, faintly stippled with darker and

RlOXOGRAPH O F THE ll1ICROHkTLIDAE

122

lighter, or, rarely, with a broad, dark mid-dorsal stripe from the interorbital region to the vent, very slightly broader posteriorly than anteriorly. Lower surfaces yellowish (red in life), uniform, or more or less heavily stippled and dotted with brown, sometimes with brown marhlings. A pronounced yellow (red in life) or white line along the hinder side of the thighs from the anus to the knee, sometimes continued, but Jess distinctly along the inner edge of the tibia and beneath the tarsus ; the width of the femoral streak is variable, and when it is broad there are usually one or more yellow (red in life) spots in the groin ; associated with this type of colouring the belly is usually more or less spotted and the feet and legs yellow spotted. Sometinles a fine white line from snout to vent. Throat of the male infuscate. Male with a vocal sac. Length : 47 mm. DISTRIBUTION : Panama southwards, east of the Andes, to Buenos Aires ; Trinidad. The numerous described colour varieties can, roughly, be grouped in two categories, the typical form with an immaculate belly, narrow white femoral stripe and no inguina; spots, and the bicolor type with a more or less profi~selyspotted lower surface, broad femoral stripe and some light spots in the groin, behind the knees and on the feet. These may ultimately prove to be two distinct species, but Mertens (1929) maintains that they are merely colour mutants of the same species. B.M. R.1931.2.13. 4 66.8.14. 268-269 69.5.8. 1 97.7.23. 110-111 92.11.22. 13-16 1902.2.10. 7-9 1927.8.1. 61 1927.8.1. 62 1927.8.1. 63 1927.8.1. 64 94.3.14. 89-93 82.10.4. 7-9 88.2.7. 3-4

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CTENOPHR YNE, Mocquard Ctenophryne, Mocquard, 1904, Bull. Mus., Paris, p. 308 (Type species Ctenophryne geayi, 3Iocquard); Nieden, 1926, Das Tierreich, Anura Ir, p. 69 ;Noble, 1931, Biol. Amph., p. 537. Prevomer ?. No Clavicles, procoracoids or omosternum ; sternum cartilaginous.

Vertebral column ?. Terminal phalanges dilated distally. Pupil vertical. Tongue large, oval, notched behind and entirely adherent, with a median furrow. Two dermal ridges across the palate in front of the pharynx, the anterior shorter than the posterior, which is denticulate. Digits slightly dilated distally.

I. ,

Ctenophryne geayi, Mocquard

Ctenophryne geayi, Mocquard, 1904, Bull. Mus., Paris, p. 308 (Type locality Sarare River, Colombia). Ctenophryne geagi (typ. err.), Kieden, 1926, Das Tierreich, Anura 11, p. 69. S n o u t short, scarcely prominent, b l u n t , once a n d three-quarters a s long a s t h e eye ; c a n t h u s rostralis n o t m a r k e d ; nostril m u c h nearer t h e t i p of t h e s n o u t t h a n t h e e y e ; interorbital space t h r e e t i m e s a s b r o a d a s t h e u p p e r eyelid ; t y m p a n u m hidden. F i r s t finger v e r y short. Toes depressed, three-fourths webbed, w i t h slight terminal dilatations ; n o distinct subarticular tubercles ; a scarcely prominent oval inner, b u t n o outer, metatarsal tubercle. Tarso-metatarsal articulation reaching t h e posterior corner of t h e eye. Skin smooth a b o v e a n d below ; a transverse occipital fold j u s t behind t h e eyes bending backwards o n t h e sides a n d t e r m i n a t i n g b e n e a t h t h e insertion of t h e arm. P a l e blackish brown above, w i t h some lighter d o t s a n d a yellowish vertebral line from t h e t i p of t h e s n o u t to t h e vent. Lower surfaces darker t h a n t h e upper, t h e line of demarcation pronounced ; thighs a n d tibiae almost black beneath ; throat, a b d o m e n a n d thighs w i t h numerous small yellowish white d o t s ; o u t e r side of tibiae wit.h some larger, irregular, cream-coloured spots ; groin, u p p e r surface of thigh and tibia w i t h a few white dots. L e n g t h : 43 mm. DISTRIBUTION : K n o w n only f r o m t h e t y p e locality, Sarare River, Colombia.

MICROH YLA, Tschudi Engys toma (part), Fitzinger, 1826, K. Class. Rept., p. 65 ; Gunther, 1858, Cat. Batr. Sal. Brit. Mus, p. 51 ; Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 194 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 160 ;Mir. Ribeiro, 1920, Rev. Mus. Paulista, XII, p. 281 ; idem, 1926, Arch. MUS.Nac. Rio de Janeiro, XXVII,p. 182 ; Nieden, 1926, Das Tierreich, Anura 11, p. 63 ; Dumhril and Bibron, 1841, Erpet. GBn., VIII, p. 738; Cope, 1889, Bull. U.S. Nat. Mus., xxxrv, p. 385. Engistonla, Peracca, 1904, Bol. Mus. Torino, xrx, 465, p. 23. Hysaplesia (part), Boie, 1826, in FBrussac, Bull. Sci. Nat., IX, p. 239. Hylaplesia (part), Boie, 1827 in Schlegel, Isis, xx, p. 294. Stenocephalus (non Latreille, 1825), Tschudi, 1838, Mem. Soc. Sci. Neuchiitel, 11, 1839, p. 86 (part). iMicrohyka, Tschudi, 1838, Mem. Soc. Sci. NeuchStel, 11, 1839, pp. 28 and 71 (Type species " Hylaplesia achatina Boie " (nom. md.)= ~Wicrohylaachatina, Tschudi) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 163 ; idem, 1890, Fauna Brit. India, Rept. and Batr., p. 491 ; Sarasin, 1910, Zool. Jahrb., Supp. XII, i, p. 18; Vogt, 1911, Sitzber. Ges. Natf..Fr. Berlin, p. 183 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 258 ; Smith, 1917, Journ. Nat. Hist. Soc. Siam, 11, p. 229 ;Rao, 1918, Rec. Ind. Mus., xv, p. 41 ;Prashad, 1918, Rec. Ind. Mus., xv, p. 101 ; Barbour, 1920, Occ. Papers Mus. Zool. Michigan, LXXVII,p. 2 ;Robinson and Kloss, 1920, Journ. F.M.S. Mus., vIn, p. 305 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 152 ; Nieden, 1926, Das Tierreich, Anura 11, p. 27 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 473; Smith, 1930, Bull. Raffles Illus., nI, p. 125 ;Pope, 1931, Bull. Amer. Mus. Nat. Hist., m r , p. 591 ; Noble, 1931, Biol. Amph., p. 537. Hicrhyh, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 121. Gastrophryne, Fitzinger, 1843, Syst. Rept., p. 33 (Type species Enyystoma carolinense, Holbrook) ; Stejneger, 1910, Proc. Biol. Soc. Washington, xxm, p. 166 ; Parker, 1927, Occ. Papers hlus. Zool. Michigan, CLXXXVII, p. 5 ; Noble, 1931, Biol. Amph., p. 537. Siphneus, Fitzinger, 1843, Syst. Rept., p. 33 (Type species Engystoma ornatum, Durn. and Bibr.) Dendromanes, Gistel, 1848, Naturg. Thierr., p. 11 (substitutemame for Microhyla). Diplopelma (part), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 50. Scapiophryne, Fitzinger, 1861, Sitzber. Ak. Wiss. Wien, XLII, p. 416 (Type species Xcaptophryne labyrinthica, Fitzinger =Engystoma pulchrum, Hallowell). Copea, Steindachner, 1864, Verh. Zoo1.-bot. Ges. Wien, p. 286 (Type species Copea fulva, Steindachner = Engystoma rubrum, Jerdon). CalEula (part), Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 192. Systoma (non Wagler), Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 193 (part). Ranina (non Lamarck), David, 1872, Nouv. Arch. Mus. Paris, VII, Bull. 1872, p. 416.

124

MONOGRAPH OF THE MICROHYLIDAE

Glossostoma, Giinther, 1900, Biol. Centr. Amer., Rept. and Batr., p. 210 (Type species Glossostoma aterrimum, Giinther) ; Nieden, 1926, Das Tierreich, Anura II, p. 68. Eupemphix (aon Steindachner), Boulenger, 1903, Ann. Mag. Nat. Hist., (7), XII, p. 552. Dasypops, Mir. Ribeiro, 1924, Bol. Mus. Nac. Rio de Janeiro, IV, p. 255 (Type species Dasypops schirchii, Mir. Ribeiro) ;Noble, 1931, Biol. Amph., p. 538. Prevomer divided, the post-choanal portion lost ; palatine present or absent (figs. 51 and 52). No clavicles, procoracoids or omosternum ; sternum cartilaginous. Vertebral column diplasiocoelous. Terminal phalanges simple, or T-shaped. Pupil circular. Tongue oval ; entire and free behind ; one or two dermal ridges across the pa1at.ein front of the pharynx, smooth or crenulate. Digits with or without terminal dilatations. Tadpoles with toes more fully webbed before than after metamorphosis.

Fig. 51. Anterior cranial elements of Microhyla berdmorei (seen from beneath)

Fig, 52. Anterior cranial elements of Microhyla inornata (seen from beneath)

As pointed out by Noble (1931), there is no character whereby Microhyla can be distinguished from Gnstrophryne, and there seems to be no plausible reason for retaining these two groups. The distribution of the combined genus is very similar to that of many other genera, as, for example, Trimereszcrus among snakes and the Tapirs among mammals, and there is no reason for believing that the group is other than monophyletic. The author (1928) in reviewing the Asiatic forms (Microhyla) has shown

MICROHYLA

125

how a series can be traced from the primitive species which retain their palatine, and have webbed feet and digital discs. to those, like inornaia, which lack all these characters. Among the American species the one with the greatest amount of digital webbing (aterri.ma)is also the most primitive in the retention of a palatine bone ; but digital discs are only retained by the most specialised burrowing species, M. microps. SYNOPSIS O F THE SPECIES I. Snout not more than twice as long as the eye. A. Toes webbed, with distinct discs.

+

webbed ; digital discs usually large. Palatine bone present. ( a ) Discs with a distinct median longitudinal groove (producing the appearance of two scut,es) on their upper surface. (i) Toes entirely webbed ; two metatarsal tubercles 1M.berdmorei, p. 127 (ii) Toes nearly webbed ; two metatarsal tubercles ; first finger much reduced . . . . . .M. borneensis, p. 128 (iii) Toes 3. to $ webbed ; an inner, but no outer, metatarsal tubercle ; first finger much reduced. ( a ) Toes webbed ; skin smooth . . M. anneetens, p. 129 (p) Toes 2 webbed ; skin rough . . M.annamensis, p. 130 (iv) Toes to 6 webbed ; two metatarsal tubercles ; a diagonal fold along the under surface of the tarsus from the inner metatarsal tubercle. . .M. butleri, p. 131 ( b ) Digital discs large, without any trace of a median cleft. Toes $ webbed. (i) No supraciliary spine (sometimes a tubercle) ; two outer fingers with discs; the inner measuring & to the length of the third . . . . . . M. palmipes, p. 132 (ii) A supraciliary spine ; fingers short, with scarcely perceptible discs, the inner measuring scarcely 4 the length of the third . . . . .1M. superciliaris, p. 133 (2) Toes not more than Q webbed ; digital discs small, but with a median longitudinal cleft above. Palatine bone absent. (a) Outer metacarpal tubercle divided ; a continuous dark lateral band from the end of the snout to the groin ; a ( )-shaped, small, dark spot on the centre of the back fW.heymonsi, p. 134 ( b ) Outer metacarpal tubercle single ; lateral band interrupted above the shoulder ; no ( )-shaped spot on the centre of the back . . . . . . . . . , N . achatina, p. 136 B. Toes with a t least a rudiment of web, but no discs (sometimes the digits end obtusely). (1) Two normal (not shovel-shaped) metatarsal tubercles. Palatine bone absent. (a) Toes more than webbed . . . . . . M. pulchm, p. 137 ( b ) Toes less than webbed. (i) Toes with or without traces of a groove on the upper surfaces of their tips. Tibio-tarsal articulation reaching the eye in females, or from the eye to beyond the tip of the snout in males. Toes of M. okinauensis, p. 1 38 females fringed . . . . . (ii) Toes without any trace of a groove on their tips ; tibio-tarsal articulation not extending beyond the eye ; toes not fringed . . . . . M. ornatn, p. 139 (1) Toes a t least

+

+

.

+

+

+

+

.

.

.

MONOGRAPH O F THE AIICROHYLIDAE (2) Two shovel-shaped metatarsal tubercles ; palatine bone absent.

( a ) Outermetacarpaltuberclesingle . . 1M. picta, p. 141 ( b ) Outer metacarpal tubercle divided . . . . . M. rtlbra, p. 142 (3) A single (inner) metatarsal tubercle. (a) Toes nearly half webbed ; palatine bone present .M. nterrima, p. 143 ( b ) Toes with only a rudiment of web ; no palatine bone. (i) A round inguinal spot not covered by the folded hindlimbs . . . . . . . . . . AT. elegans, p. 144 (ii) No marked inguinal spot . . . . LM. nequatorialis, p. 146 C. 'Foes quite free ; no digital discs. No palatine bone. (1) Snout not more than once and a half as long as the eve. (a) A single, not compressed, metatarsal tubercle (the outer is sornetimcs feebly-indicated). (i) Snout rounded, not longer than the diameter of the eye ; loreal region vertical ; hahitus rather slender ,IT. inosnata, p. 144 (ii) Snout pointed, longer than the diameter of the eye ; loreal region oblique ; habitus stout. (or) Eye longer than its distance from the nostril ; skin smooth or slightly pustular ; belly mottled and stippled with brown . &f. carolinensis, p. 146 (p) Skin areolated or pustular ; belly grey with lighter dots . . . . . . . . . M . areolata, p. 147 (y) Eye not longer than its distance from the nostril ; skin smooth or slightly pustular ; belly immaculate . . . . . . . . . M. texensis, p. 148 (6) Two compressed metatarsal tubercles . . . . .M. usta, p. 148 (2) Snout twice as long as the eye ; belly black, with large white spots . . . . . . . . . . . . M. pictiventris, p. I46 11. Snout more than twice as long as the eye. No palatine bone.

A. Toes not dilated. (1) Toes with a rudiment of web ; skin adherent to skull . M. schirchi, p. 149 (2) Toes quite free ; skin of head smooth, not adherent to skull M. subnigra, p. 150 B. Toes, except the inner, dilated to form small but distinct discs f i . microps, p. 150 SYNOPSIS O F THE TADPOLES I. Lower lip produced into a funnel which almost surrounds the mouth. A. A knob-like protuberance on each side of the mouth within the oral disc. . . . . . . . . . . . . . . . . M. heymonsi B. A boss with a central depression, sucker-like, on each side of the mouth within the oral disc . . . . . . . . . . M. achatina 11. No funnel-like projection of the lower lip. A. Head squarish anteriorly ; tail without terminal flagellum ,TI. palmipes, M. berdmorei, fM.carolinensis B. Head rounded anteriorly ; tail with a terminal flagellum (1) IIlouth terminal ; body broad anteriorly ; spiraculum opening

under a flap which extends posteriorly almost to the anterior edge of the anus ; latter large, slit-like, with pigmented edges ; . . . . . . . . M. butleri tail marked with scarlet in life

( 2 ) Mouth dorso-terminal ; body narrow anteriorly ; spiracular opening about the centre of the gut ; anus small, circular,

. .

il.1. borneensis, M. inornata, M. ornata, M. rubra, fM.pulchra, M. okinavensis If the relative proportions given by Wright for tadpoles of *TI. carolinensis can be relied upon, it is obvious that the tadpoles of this genus are enormously variable ; the tail of some specimens is reported to be twice as long as in others. Pope (1931) has also questioned the possibility of determining ~Microhylatadpoles by the relative lengths of their tails, and so it seems that the precise identification of species is a t present a matter of considerable difficulty and cannot be accomplished without an actual comparison with series of named specimens. unpigmented

Microhyla berdmorei (Blyth) Engystoma ( ? ) berdmorei, Blyth, 1866, Journ. Asiatic Soc. Bengal, xxrv, p. 720 (Type locality Pegu, Burma) ; Mason, 1860, Burma, ed. 2, p. 325. Diplopelma berdmorei, Giinther, 1868, Zool. Eec., rv, p. 146 ; Anderson, 1871, Proc. Zool. Soc. London, p. 163 ; Stoliczka, 1872, Proc. Asiatic Soc. Bengal, p. 109. illicrohyla berdmorii, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 166 ; idem, 1890, Fauna Brit. India, Rept. and Batr., p. 492 ;Sclater, 1892, Batr. Ind. Mus., p. 23 ; Plower, 1896, Proc. Zool. Soc. London, p. 908 ; idem, 1899, op. cit., p. 906 ;Werner, 1900, ZooI. Jahrb., Syst., XIII, p. 496 ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 390 ; van Kampen, 1905, Zool. Jahrb., Syst., XXII, p. 713 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 263; Smith, 1916, Journ. Nat. Hist. Soc. Siam, 11, p. 169 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 158 (part) ; Nieden, 1926, Das Tierreich, Anura 11, p. 33. ~iicrohylaberdmorei, Laidlaw, 1900, Proc. Zool. Soc. London, p. 888; Smith, 1924, Rec. Ind. Mus., xxvr, p. 141, PI. VII, fig. i ; Parker, 1928, Ann. Mag. Nat. Hht., (10). 11, p. 479 ; Smith, 1930, Bull. Raffles Mus., III, p. 127, fig. 13 right ; Cochran, 1930, Proc. U.S. Nat. Mus., LXXVII, 11, p. 7. Callula natatriz, Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 192 (Type locality Nr. Rangoon). Diplopelma pulchrum (part), Theobald, 1868, Cat. Rept. Mus. Asiatic Soc. Bengal, p. 83. ~Vicrohylamalcolmi, Cochran, 1927, Proc. Biol. Soc. Washington, x ~p., 182 (Type locality Pak Jong, Siam).

Habit slender. Snout obtusely pointed, scarcely prominent, a little longer or a little shorter than the diameter of the eye, canthus rostralis rounded ; loreal region nearly vertical ; interorbital space as broad as, or broader than, the upper eyelid ; tympanum hidden. Fingers scarcely dilated, but with a wide median groove on the upper surface of the tip separating two swellings ; first much shorter than the second, which is a little shorter than the fourth. Toes webbed to the very tips, with distinct discs, each with a median groove separating a pair of scale-like pads ; third and fifth subequal ; toes of newly metamorphosed specimens only webbed ; subarticular tubercles well developed ; two small, but prominent metatarsal tubercles ; sometimes sesamoid developments in the tendon of Achilles produce the semblance of shovelshaped tubercles a t the proximal end of the tarsus and a t the outer metatarsal tubercle. Tibio-tarsal articulation reaching well beyond the tip of the snout. Skin smooth or with small tubercles which may form longitudinal series, especially from the supra-scapular region to the groin ; a more or less distinct tympanic fold and another behind the angle of the mouth. Hinder part of belly and anal region granular. Olive to purplish brown or pink above ; an oblique dark line on the flanks and more or less distinct traces of a median spot which begins on the head between the eyelids and is truncate in front ; this narrows on the nape, broadens in the scapular region, narrows on the middle of the back, broadens on the sacral region and sends a branch postero-laterally towards the groin, narrows again and finally broadens and bifurcates, a branch running towards each leg. This marking may be well defined and white-edged, indistinct, broken a t the narrow regions, or entirely absent. Hind-limbs cross-barred ; a longitudinal dark streak is often present on the anterior aspect of the thigh, and a dark, triangular area surrounds the vent, Sides of head with indistinct dark vertical

+

128

MOXOGRAPH OF THE MICROHYLIDAE

bars. Yellowish white beneath, the throat and chin grey-marbled or dusted and almost black in males. Male with a vocal sac. Length : 45 mm. DISTRIBUTION : Burma, Siam, Cambodia, Malay Peninsula and Sumatra. A well-grown tadpole has the following characters : Body, 1 i to 1%times as long as broad ; tail about once and a half as long as the body, and about 3 times as long as deep. Head and body broad, depressed, almost square when seen from above. Mouth dorso-terminal ; eyes lateral ; nostrils midway between the mouth and a line connecting the eyes ; internarial space 4 thc interocular. Spiraculum median, below the centre of the gut. Tail pointed, but without terminal filament ; caudal crests subequal, shallower than the muscular portion a t its base. Grey-brown above, dusted with darker ; a dark patch between the eyes.

Q9

B.31. 68.4.3. 99-101 65.4.5. 5-6 1928.11.16. 8-19 1928.11.16. 53

?? juvs.

$

Pegu, Burma . Burma . . . . N. of Prae, N. Siam Pak Jong, N. Siam

. Theobald

. . . Theobald . . . M. A. Smith . M. A. Smith, TOPOTYPE of M. malcolmi

$

1928.11.16. 45 98.4.2. 54-56 97.10.8. 100 1916.3.27.46-48 1927.9.1. 4 1928.11.16. 47 1928.11.16. 63 1928.11.16. 54-55 1928.11.16. 46 92.4.18. 25 1928.11.16.48 1928.1 1.16. 49-50 1928.1 1.15 . 27-38

1928.11.16. 51-52

2.

Pak Maat, near Vien Chan, N. Siam . . M. A. Smith , Dong, Phya Fai, 700 ft., Siam Flower juv. Bortong, Siam . . Flower $ $, 9 Nakon Sri Tamarat, Siam . M. A. Smith cleared Nakon Sri Tamarat, Siam . M. A. Smith $ Kompong Sumbon, S.E. Siam M. A. Smith $ Hat Sanuk, S.E. Siam . . 81. A. Smith juvs. Klong Bang Lai, Penins. Siam RI. A. Smith Q Pulau Condor . . M. A. Smith ? Pera-Kan, Cambodia . . . Heron-Royer Bockor Plateau, Elephant Mts., Cambodia . . . . 31. A. Smith $, Q Sui Kat, Langbian Plateau, S. Annam . . . . . . M. A. Smith tadpoles Daban, Langbian Plateau, M A . Smith (series Annam . described by Smith) Q$? Dran, S. Annarn . . . AI. A. Smith

Microhy la borneensis, Parker

Xicrohyla annectens (part), Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 262 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 166 ; Smith, 1926, Journ. Sarawak Mus., 111, 8, p. 32 ;Nieden, 1926, Das Tierreich, Anura 11, p. 32. L71icrohylaborneense, Parker, 1926, Ann. Mag. Nat. Hist., (lo), 11, p. 473 (Type locality Bidi District, Sarawak).

Habit slender, snout pointed, slightly prominent, once and a half as long as the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space once and three-quarters to twice and a half as broad as the upper eyelid ; tympanum hidden. Fingers with distinct discs, with a median groove on their upper surfaces ; first finger very short ; second shorter than the fourth. Toes with discs similar to, but larger than, those of the fingers ; toes rather more than half webbed, the membrane midway between the third and fourth, extending half-way between the tubercles of the third toe ; fifth toe a ljttle shorter than the third ; subarticular tubercles well

'

developed ; an oval inner, and a conical outer, metatarsal tubercle. Tibio-tarsal articulation reaching a little beyond the tip of the snout. Skin smooth above and below ; a slight tympanic fold. Light brown above, with a darker area along the middle of the back ; from the head, where it connects the upper eyelids, it narrows behind the occiput, broadens and gives off a branch towards the fore-limbs or the fianks, narrows again and finally widens and becomes indistinct on the posterior part of the back ; this spot may be outlined in white ; a dark, oblique streak from above the shoulder to the middle of the flank ; hind-limbs indistinctly cross-barred ; a dark spot on the front of the knee ; a dark area about the vent. Upper lip with three dark brown spots and a light line from behind the eye to the fore-limb. Lower surfaces yellowish, mottled with light brown, especially on the throat and chest. Length : 23 mm. Larvae were collected by Hewitt in the pitcher of a Nepenthes in November. The youngest stage has the opercular fold just covering the gills, large, adhesive organs close to the angles of the small mouth and the intestine full of yolk ; length 6.5 mm. Development is not accompanied by any great increase in size ; examples 7 mm. long have the operculum and spiracle complete, the adhesive organs reduced to a brown patch on each side of the now functional mouth, and limb-buds developing ; but the intestine is still packed with yolk though commencing to develop its spiral form. At 10.5 Inm. the digits are visible on the hind-limbs and a t this length the nostrils break through and metamorphosis commences ; a newly metamorphosed example, with the tail resorbed, measures 4 mm. from snout to vent. A mature tadpole has the following characters : Body 1.8 times as long as broad ;tail about 1.7 times as long as the body and 2.5 times as long as deep. Head and body not depressed, as deep as broad ; snout rounded ; mouth dorso-terminal, the lower lip slightly prominent ; eyes lateral ; nostrils midway between the tip of the snout and a line connecting the anterior borders of the eyes ; internarial space one-third the interocular, and a little wider than the mouth. Spiraculum median, below the centre of the gut, its covering flap feebly emarginate. Tail with a short terminal filament ; caudal crests subequal, about equal to the muscular portion a t its base. Pale straw-colour above, feebly stippled with darker except on the caudal crests and posterior half of the tail. DISTRIBUTION : Sarawak.

B.M. 1911.1.30. 43 1933.6.20.15 1914.5.12. 61-78

9 9

Bidi Distr., Sarawak Mt. Dulit, Sarawak developmentaI Kuching . series

. .

. . Brooks, TYPE . . . Moore . . . Hewitt

3 . Microhyla annectens, Boulenger Microhyla annectens, Boulenger, 1900, Ann. Mag. Nat. Hist., (7), VI,p. 188(Type locality Larut Hills, Perak, 4,000 ft.) ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 389 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 262 (part); Smith, 1916, Journ. Nat. Hist. Soc. Siam,

11, p. 169 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 156 (part); Nieden, 1926, Das Tierreich, Anura 11, p. 32 (part); Parker, 1928, Ann. Mag. Nat. Hist., (lo),11, p. 482 ; Smith, 1930, Bull. Raffles Mus., III, p. 127. Habit slender. Snout rounded, slightly prominent, as long as, or a little longer than, the eye ; canthus rostralis rounded ; Ioreal region nearly vertical ; interorbital space once and three-quarters as broad as the upper eyelid ; tympanum hidden. Fingers with distinct discs divided by a median groove on their upper surfaces ; inner very short ; second much shorter than the fourth. Toes with similar discs to those of the fingers, about $ webbed, the membrane midway between the third and fourth reaching the level of the distal tubercle of t.he third toe, which is a little longer than 9

MONOGRAPH O F THE MICROHYLIDAE the fifth ; subarticular tubercles moderate ; an oval inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching well beyond the tip of the snout. Skin smooth above and beneath. Brown above, with an oblique black bar from above the shoulder, along the flanks, towards the groin ; a light streak, sometimes black-bordered posteriorly, from below the eye to the fore-limb ; a symmetrical dark marking on the back commencing between the eyes, where it connects the upper eyelids, narrowing behind the occiput, widening between the shoulders, narrowing again and then becoming indistinct ; an oblique dark cross-bar on the femur, tibia and tarsus ; a black spot on the anterior aspect of the knee and another around the vent. The head, from the level of the centre of the eyes to the tip of the snout may be green, with a triangular black spot on the lip in front of the eye. Lower surfaces lighter, closely marbled with brown. Sometimes a fine, white, dorsal line from snout to vent. Male with a vocal sac. Length : 20 mm. DISTRIBUTIOX : Federated Malay States and Peninsular Siam.

B.11. 1900.6.14. 34-35 1928.11.12.1

hgr. $

Larut Hills, Perak, 4,000 ft. Perak. . . . . . .

. . . .

Butler, COTYPES M.A.Smith

4. Microhyla annamensis, Smith illicrohyla annamensis, Smith, 1923, Journ. Nat. Hist. Soc. Siam, VI, p. 47, P1. v, fig. 2 (Type locality Sui Kat, Langbian Plateau, S. Annam) ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 483.

Habit moderately slender. Snout rounded, not prominent, as long as, or a little longer than, the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space once and a quarter as wide as the upper eyelid ; tympanum feebly indicated or hidden, half the diameter of the eye. Fingers with small, but distinct, discs, with a median longitudinal groove on their upper surfaces ; first very short, second shorter than the fourth. Toes with discs similar to, but slightly larger than, those of the fingers ; f webbed, the membrane midway between the third and fourth extending beyond the distal tubercle of the third toe, which is subequal to, or a little longer than, the fifth ; subarticular tubercles feeble ; a small inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tip of the snout, or a little beyond. Skin more or less warty or tubercular above ; smooth beneath. Grey-brown above, with a short, black streak above each shoulder ; a chevron-shaped dark spot on the scapular region and, sometimes, a few indistinct dark spots posteriorly ; usually there is a triangular dark spot between the eyes, and a light line fro111 beneath the eye to the fore-limb. Limbs with more or less distinct black cross-bars. Yellow beneath, more or less thickly brown dusted. Nale with a vocal sac and infuscate gular region. Length : 20 mm. DISTRIBUTION : Langbian Plateau, S. Annam.

B.M, 1923.5.14. 10

9

Sui Kat, Langbian Plateau, S. Annam, 1,000 m. . Rf. A. Smith, HOLOTYPE 1923.5.14. 11-13 $, $?9 Sui Kat, Langbian Plateau, S. Annam, 1,000 m. . M. A. Smith 1928.8.11. 2 cleared Sui Kat, Langbian Plateau, S. Annam, 1,000 m. . $1. A. Smith PA~ATyp"s 1928.11.16. 42-44 $ $, $? Dran, Langbian Plateau 31. A. Smith J

\

MICROHYLA

5 . Microhyla butleri, Boulenger Microhyla sp., Flower, 1899, Proc. Zool. Soc. London, p. 903, PI. LX, fig. 2 ; Kieden, 1926, Das Tierreich, Anura 11, p. 36, fig. BficrohyEa butleri, Boulenger, 1900, Ann. Mag. Kat. Hist., (7), VI, p. 188 (Type locality Larut Hills, Perak, 4,000 ft.) ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 388 ; Boulenger, 1912, Pauna Malay Penins., Rept. and Batr., p. 261 ; Smith 1917, Journ. Eat. Hist. Soc. Siam, 11, p. 268 ; idem, 1922, op. cit., rv, p. 214 ; idem, 1922. Journ. F. 31. S. Rlus., x, p. 281 ; idem, 1923, Journ. Nat. Hist. Soc. Siam, VI, p. 212 ; Nieden, 1926, Das Tierreich, Anura 11, p. 31 ; Parker, 1925, Snn. Mag. Nat. Hist., (lo), n, p. 483 ; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., IV, 2, p. 26 ; Pope, Bull. Amer. Nus., Nat. Hist., LXI, p. 592, fig. 37 ; Smith, 1930, Bull. Raffles Mus., 111, p. 127 ; Cochran, 1930, Proc. U.S. Nat. Mus., LXXVII,11, p. 6. Microhyla boulengeri, Vogt, 1913, Sitzber. Ges. Natf. Fr. Berlin, pp. 222, 227, 229 (Type IocaIity Hainan) ; idem, 1922, Arch. Naturg., LXXXVIIT, 10, p. 146 ;Mell, 1922, Arch. Katurg., LXXXVIII, 10, p. 130 ; Nieden, 1926, Das Tierreich, hnura 11, p. 31. ilficrohyla berdmorei ? (non Blyth), Annandale, 1917, Mem. Asiatic Soc. Bengal, VI, p. 151 ; Rao, 1918, Rec. Ind. Mus., xv, 4, p. 42 ; van ICampen, 1923, Amph. Indo-Austr. Arch., pp. 153 and 158. Microhyla latnstii, Boulenger, 1920, Ann. Mag. Nat. Hist., (9), VI, p. 107 (Type locality Saigon). Microhyla hainanensis (non Barbour), Mell, 1922, Arch. Naturg., LXXXVIII, 10, p. 131. Microhyla grahami, Stejneger, 1924, Occ. Papers Boston Soc. Nat. Hist., v, p. 119 (Type locality Sui-fu, Sze-chwan, China) ; idem, 1925, Proc. U.S. Nat. Mus., LXVI,25, p. 13. Microhyla sowerbyi, Stejneger, 1924, Occ. Papers Boston Soc. Nat. Hist., v, p. 119 (Type locality Nr. Yenping-fu, Fukien, China) ; idem, 1925, Proc. U.S. Nat. Mus., LXVI,25, p. 14.

Habit slender. Snout rounded, prominent, as long as, or a little longer than, the eye ; canthus rostralis rounded ; loreal region oblique ; interorbital space once and a quarter to once and three-quarters the width of the upper eyelid ; tympanum hidden. Fingers with more or less well-developed discs, with median longitudinal furrows above ; first shorter than the second, which is shorter than the fourth. Toes with discs similar to those of the fingers, and rather variable webbing ; usually 2 to webbed, but sometimes nearly half, with lateral fringes to all the toes and a distinct ridge on the outer side of the fifth toe continued along the outer metatarsal ; subarticular tubercles prominent ; two prominent metatarsal tubercles and a more or less distinct diagonal ridge from the inner towards the middle of the tarsus ; sometimes a very small tubercle near the tibio-tarsal articulation, which reaches the eye or the end of the snout when the hind-limb is laid forwards. Skin smooth or tubercular above ; a fold from the corner of the eye to the anterior side of the arm, and sometimes traces of an occipital fold. Lower surfaces smooth, the anal region granular. Grey or brown above, with darker markings consisting essentially of a single large marking which commences between the eyelids, connecting the two, narrows considerably behind the head, broadens abruptly between the fore-limbs, narrows slightly and then sends a branch postero-laterally towards the groin, narrows immediately, and finally broadens to send a diagonal branch on to each femur. This marking is often edged with a light border and may be broken up, the breaks occurring a t the narrow points on the nape and sacral region ; posteriorly the marking may be indistinct and anteriorly reduced to a triangular interorbital spot and a ) (-shaped figure or transverse bar on the back. A light streak is usually present from below the eye to the fore-limb, and dark spots are frequently present on the temporal region and above the fore-limb. The dermal tubercles, particularly in juveniles, are lighter (red in life). Limbs and digits with narrow dark cross-bars. Lower surfaces white. Male with a vocal sac and infuscate gular region. Length : 26 mrn. DISTRIBUTION : S. China (Fukien, Kwangtung and Sze-chwan), Tonkin, Hainan, Burma, Siam, Annam and the Malay Peninsula. Tadpoles only found in relatively permanent pools. Body 1.3 times as long as broad ; tail 1-6 to 1.8 times as long as the head and body and 2 . 2 times as long as

+

MONOGRAPH OF THE MICROHYLIDAE deep. Head and body broad, rounded anteriorly ; mouth terminal ; nostrils close together, a little nearer the tip of the snout than the line connecting the anterior border of the eyes ; internarial space about Q the interocular. Spiraculum median, opening under a transparent sheath which opens close to the anus. Anus an oval slit. Tail ending in a filament ; lower crest deeper than the upper and than the muscular portion. Transparent anteriorly ; posteriorly and on the tail some dark markings ; terminal filament unpigmented ; sides of tail red in life. Toes fully webbed. B.M. 1927.3.15. 1

d

SPECIMENS EXAMINED Suifu, Sze-chwan .

. . . (U.S.

Nat. Mus.), TOPOTYPE of M. grahami, Stejneger

Hummocks, nr. Hoi-hao, Hain a n . . . . . . . M. Smith . . Friihstorfer 1903.4.29. 97 9 Man-Son Mts., Tonkin 1923.5.14. 21 9 Den Chai, N. Siam . . . M. Smith 1917.5.14.40 9 Ta Rua, C. Siam . . . . M. Smith 1923 5.14.14-16 99 Ta Rua, C. Siam . . . . M. Smith 1928.11.15. 17-26 tadpoles Ta Rua, C. Siam . . . . M. Smith (series described bv M. Smith) 1928.11.16. 21 juv. Ta Rua, C. Siam . . M. Smith (reared from tadpole in captivity) Maa Yome, N. of Pra, Siam . M. Smith 1928.11.16. 20 hgr. 1928.11.16.24-25 39 Nong Khor, S.E. Siam . . M. Smith 1920.1.20.4054a 39 Saigon . . HolbC, COTWESof M . latastii, Roulenger 1923.5.14.17-20 33d,9 Paknam Chumpon, Penins. Siam . . M. Smith 1928.11.16. 26 3 Bukit Fraser, P.M.S. . . M. Smith 1928.8.11. 3 cleared Bukit Praser . . . . . M. Smith 1901.3.20. 5 9 Larut Hills, Perak, 4,000 ft. . Butler, TYPE 98.9.22. 124 . Flower d Kedah . Mus. Berlin 23334 2 ads. Hainan . . Mell, COTYPESof M. boulengeri, Yogt 1928.11.16. 22-23 h p .

6 . Microhyla palmipes, Boulenger Yicrohyh palmipes, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), XIX, p. 108 (Type locality Pengalengan, Java, 4,000 ft.) ; van Kampen, 1923, Amph. Indo-Austr. Arch, p. 157 ; Nieden, 1926, Das Tierreich, Anura II, p. 33 ; Mertens. 1927, Senckenbergiana, IX, 6, p. 242 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 485 ; Mertens, 1929, Senckenbergiana, XI, p. 24 ; idem, 1930, Abh. Senck. Nat. Ges., XLII,3, p. 237, P1. VI, fig. 5 ;Smith, 1930, BuU. Raffles Mus., m, p. 128, fig. 13 (left) ; Schijfsma, 1932, Treubia, xm,i, p. 48, figs. 3-5. Microhyla annectens (non Boulenger), van Kampen, 1907, Zool. Egebn. Ned. Ost. Ind., rv, 2, p. 404 ; Barbour, 1912, Mem. Mus. Comp. Zool. Harvard, xLrv, p. 71, PI. vrr, fig. 26 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 262 (part) ; Annandale, 1917, Journ. F. M. S. Mus., VII, p. 108 ; Boulenger, 1920, Journ. F.M.S. Mus., vm, p. 295; Robinson and Kloss, 1920, Journ. F.M.S. Mus., vm, p. 305 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 156 (part) ; Nieden, 1926, Das Tierreich, Anura n, p. 32 (part) ; Dunn, 1928, Amer. M u . Novitates, cccxv, p. 4. Microhyla niasensis, van Kampen, 1915, in Kleinweg de Zwaan, Die Insel Nias bei Sumatra, p. 279 (Type locality Gunung Sitoli, Nias). Microhyla berdmorei (non Blyth), Smith, 1922, Journ. F.M.S. Mus., x, p. 281.

MICROHYLA Habit slender, snout rounded, slightly prominent, as long as, or slightly longer than, the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical ; interorbital space nearly twice as broad as the upper eyelid ; tympanum hidden. Fingers with distinct discs, which lack any trace of a longitudinal cleft above ; first much shorter than the second, which is subequal to the fourth. Toes with discs similar to those of the fingers, to $ webbed, the membrane midway between the third and fourth reaching a little beyond the distal tubercle of the third toe, which is a little longer than the fifth ; subarticular tubercles well developed ; two metatarsal tubercles, the outer sometimes indistinct. Tibio-tarsal articulation reaching the tip of the snout, or slightly beyond. Skinsmooth or slightly tubercular above ; usually a very small tubercle on the posterior half of the upper eyelid, near its free edge ; a fold from the eye to the fore-limb and another, above this, from the eye to the middle of the flank. Smooth beneath. Purplish brown above, with a black streak, sometimes broken up or indistinct, below the upper lateral fold ; back with a darker marking commencing between the eyes and connecting the upper eyelids, narrowing behind the head, widening abruptly between the fore-limbs, narrowing slightly and then gradually widening and becoming more indistinct posteriorly ; frequently a black spot a t the extremity of each interscapular projection ; a light streak from below the eye to the fore-limb ; a dark bar across the fore-arm, femur, tibia and tarsus, and a black spot in front of the knee and heel ; anal r&on black; lower surfaces white, more or less closely marbled with brown, especially in males. Male with a vocal sac. 21-8 mm. ; 6 16 mm. Length : D I ~ T R I B ~ I:OMalay N Peninsula, Sumatra, Nias, Java and Bali. A full-grown tadpole measures about 28 mm. The body is 13 t o 12 times as long as broad ; tail about twice as long as the body and twice to twice and a half as long as deep. Mouth terminal ;nostrils close together on the upper surface, nearer the end of the snout than the eye ; eyes lateral, the interorbital space about 5 times the internarial. Spiraculum median, opening in a transparent sheath, which is bifid on its ventral edge ; anus median. Tail pointed, without terminal filament ; lower caudal membrane deeper than the upper. Back moderately, tail sparsely, dark pigmented. Toes fully webbed. (After Mertens.) SPECIMENS EXAMINED Gunong Tahan, Malay PeninB.M. 1928.11.16. 56 sula . . . . . . . M. Smith 1904.7.19.29-30 $2, hgr. Mouth of Batu Caves, Kuala Lumpur, Selangor . . . (F.M.S. Mus.) 1896.12.3.40 ? Pengalengen, Java, 4,000 ft. Fruhstorfer, TYPE 1928.9.25. 1 cleared Tjibodas, Java. . . . . M. Smith 1915.12.2. 21 ? Sungei Kumbang, Korinchi, Robinson and Sumatra, 4,700 ft. . Kloss

+

.

. .

7 . Microhyla superciliaris, Parker Microl~ylasuperciliaris,Parker, 1928,Ann. Mag. Nat. Hist.,(lO),II, p. 486 (Typelocality Batu Caves, Kuala Lumpur) ; Smith, 1930, Bull. Raffles Mus., III, p. 128. Habit slender. Snout rounded, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded; loreal region slightly oblique ; interorbital space nearly twice as broad as the upper eyelid ; tympanum hidden. Fingers very slightly dilated, the discs without any trace of a longitudinal cleft ; first very short ; third very long ; the second a little shorter than the fourth. Toes with distinct discs similar to those of the fingers, webbed to the discs of all except the fourth ; fifth a little longer than the third ; subarticular tubercles scarcely distinct ; two small metatarsal tubercles. Tibio-tarsal articulation reaching the tip of the snout. Skin smooth above and below ; a prominent spine on the upper eyelid.

134

MONOGRAPH O F THE MICROHYLIDAE

Yellowish brown above, with an irregular, interrupted dark streak from behind the eye, above the fore-limb to the middle of the flank ; indications of a median darker area commencing behind the head, expanding between the shoulders, narrowing and then broadening posteriorly; a light streak from behind the eye to the fore-limb ; upper lip with brown spots ; limbs cross-barred ; a black spot on the front of the knee ; anal region black ; lower surfaces yellowish, the throat and chin marbled with light brown. Length : 12 mrn. DISTRIBUTION : Batu Caves, F.M.S., and Deli, Sumatra. B.M. 19W.7.19. 31 Mus. Amsterdam

$2 $2

Batu Caves, Kuala Lumpur 'Deli, Sumatra . . . .

.

.

TYPE de Bussy, PARATYPE

8 . Microhyla heymonsi, Vogt Microhyla acchatim (mn Boie), Sclater, 1892, Proc. Zool. Soc. London, p. 347 ; idem, 1892, Cat. Batr. Ind. Mus., p. 23 ; Werner, 1893, Jahr. Ver. Magdeburg, p. 254 ;Flower, 1895, Proc. Zool. Soc. London, p. 908; idem, 1899, op. cit., p. 906; Laidlaw, 1899, Proc. Zool. Soc. London, p. 888, Werner, 1900, Zool. Jahrb., Syst., xm, pp. 496, 602 (part) ; Butler, 1904, Joum. Bombay Nat. Hist. Soc., XV, p. 389 ;Werner, 1905, Zool. Jahrb., Syst., XXII, p. 713 (part) ;van Kampen, 1909, Tidschr. Ned. Ind., LXIX, p. 45 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 261 (part) ;Barbour, 1912, Mem. Mus. Comp. Zool. Harvard, XLN, p. 17 ;Robinson and Kloss, 1916, Journ. F.M.S. Mus., v, p. 155 ; Smith, 1916, Journ. Nat. Hist. Soc. Siam, 11, p. 37, fig. ; idem, 1916, tom. cit., p. 169 ; Annandale, 1917, Mem. Asiatic Soc. Bengal, VI, p. 150, P1. VI, fig. 6 ; Hora, 1922, Journ. Asiatic Soc. Bengal, xvm, p. 12 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 154 (part), fig. 22 ; Smith, 1923, Journ. Nat. Hist. Soc. Siam, n, p. 212 :Lonnberg and Rendahl, 1925, Ark. Zool. Stockholm, XVII, A, p. 3 ; Smith, 1926, Proc. Zool. Soc. London, pp. 983-986, fig. 2 ; Nieden, 1926, Das Tierreich, Annra n, p. 32 (part) ; Cochran, 1930, Proc. U.S. Nat. Mus., L x x v n , 11, p. 6. Microhyh ap., van Kampen, 1907, Zool. Ergebn. Ned. Oat. Ind., N, 2, p. 405. Microhyh$esipes ( w n Boulenger, 1884), Boulenger, 1909, Ann. Mag. Nat. Hist., (8), IV, p. 495. Microhyla kymonsi, Vogt, 1911, Sitzber. Ges. Natf. W. Berlin, p. 181 (Type locality Kosempo, Formosa) ; Stejneger, 1925, Proc. U.S. Nat. Mus., LXVI, p. 13 ; Nieden, 1926, Das Tierreich, Anura n, p. 35 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 487 ; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., IV, 2, p. 26 ; Smith, 1930, Bull. Raffles Mus., m, p. 127 ; Pope, 1931, Bull. Amer. Mus. Nat. Hist., LXI, p. 593, fig. 38.

Habit slender. Snout obtusely pointed, prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space about once and a half the width of the upper eyelid ; tympanum hidden. Fingers with small terminal discs which usually have a median longitudinal cleft on their upper surface, though this may be reduced t o a mere notch terminally ; first much shorter than the second, which is a little shorter than the fourth ; outer metacarpal tubercle divided, or heart-shaped. Toes with discs similar to, but larger than, those of the fingers, webbed a t the base (not beyond the proximal tubercle of the third toe midway between the third and fourth) and with more or less distinct lateral fringes on each digit ; third distinctly longer than fifth ; subarticular tubercles well developed ; two small metatarsal tubercles. Tibio-tarsal articulation reaching - the eye or the end of the snout. Skin smooth above and below ; a slight glandular fold from the posterior corner of the eye to the fore-limb. Pink or grey above, with a black lateral band, extending from the tip of the snout t o the groin and entirely covering the sides of the head, sharply defined above but shading into the colour of the belly beneath ; traces of a narrow median dark area commencing between the eyes, with a branch on each eyelid, narrow on the nape, broadening between the scapulae and then becoming indistinct ; remainder of dorsum

MICROHYLA

135

frequently with fine brown dots arranged in linear series parallel with the edges of the median area ; generally a fine white line from snout to vent, and one, or sometimes two, small black, sharply-defined ( )-shaped spots at the middle of this white line ; anterior aspect of the thighs, anal region and lower surfaces of the'tarsus and foot, black ; limbs with numerous, narrow, oblique cross-bars. Lower surfaces white, the chin and chest brown stippled. Male with a vocal sac and infuscate gular region. Length : 26 rnm. ; 21.5 mm. : Formosa, S. China (Fukien), Hainan, Tonkin, Siam, Cochin China, DISTRIBUTION Malay Peninsula, Nias and Sumatra. Eggs laid in small, floating clumps. A full-grown tadpole measures about 22 mm. The body is about 13 times as long as broad and the tail about twice as long as the body. Snout bluntly rounded; mouth dorso-terminal, with the lower lip produced into a large, funnel-shaped structure which almost surrounds the mouth ;a knob-like prominence on each side of the mouth within the oral disc ; nostrils midway between the upper lip and the eyes ; eyes lateral, the interorbital space 4 to 7 times the internarial distance. Spiraculum opening in a transparent sheath below the centre of the g u t ; anus median. Tail ending in a fine point, about 4 times as long as deep ; lower caudal crest slightly deeper than the upper, and not quite as deep as the muscular pertion of the tail. Brown or black above with a metallic spot between the eyes and sometimes with gold or orange patches on the sides ; tail more or less dark dusted, sometimes with a clear, oblique, transverse bar. Toes fully webbed. The exact function of the oral funnel is in dispute, but it is almost certainly an aid to microphagy and possibly t o the gathering of food from the surface film of the water ; i t has been suggested that it is an adaptation to life in swift-flowing water, but though the species may be a mountain-dweller in some parts of its range (Pope), in others it is equally a t home in swampy lowlands.

B.M. 1909.10.29. 86-90

SPECIMENS EXAMINED 4 63, Kosempo, Formosa

. . . . Sauter. TOPOTYPES

Kanshirei, Formosa . . . . Sauter Tun Fao, Kachek River, Hainan. M. Smith Tun Fao, Kachek River, Hainan. M. Smith Man Son Mts., Tonkin, 3 4 , 0 0 0 ft. Fruhstorfer He-Ho Gorge, S. Shan States . M. Smith Pak Jong, N. Siam . . . . M. Smith Ban Me Yen, Me Wang, N. Siam . M. Smith Dong Phya Fai, Siam, 900 ft. . Flower Chantaboon, Siam . . . . . Flower Bangkok . . . . . . . M. Smith tadpoles, Bangkok . . . . . . . M. Smith Yg. tadpole Bangkok . . . . . . . M. Smith tadpoles Bangkok . . . . . . . M. Smith (described by M. Smith) Bockor, Elephant Mts., Cambodia M. Smith 1928.11.16. 32 6 1920.1.20.4054b. ? Saigon . . . . . . . . (Lataste) Q Penang, 2,200 ft. . . . . . Flower 98.9.22. 114 ? PenangHill . . . . . . Flower 97.1.28.4 tadpoles PenangHill . . . . . .Flower 96.6.25. 199-208 86.12.28. 37-38 Malacca . . . . . . . . Hervey d? 98.9.22. 115 Taiping, Perak . . . . . . Flower ? 89.11.12. 29 Deli, Sumatra . . . . . . Moesch ? Mus. Vienna 20 Kanshirei, Formosa . . . . Sauter and 99

d? ?? cleared ? hgr. ? ? ? 6 Q

MONOGRAPH OF THE MICROHYLIDAE

g. Microhyla achatina, Tschudi Hy8aplesia achdinu (n.n.), Kuhl and van Hasselt, 1826, in FBrussac, Bull. Sci. Nat., IX, p. 239 ; Hylapkaia acWina (n.n.), Schlegel, 1827, Isis, xx, p. 294. Microhyla achdiina, Tschudi, 1838, Mem. Soc. Sci. Neucutel, n, 1839, p. 71 (Type locality Java) ; DumBril and Bibron, 1841, Erpet. GBn., vm, p. 615 ; Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 193 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 166 ; Boettger, 1892, Ber. Off. Ver. Nat., p. 144 ; Werner, 1900, Zool. Jahrb., Syst., xm, p. 502 (part); van Kampen, 1905, Zool. Jahrb., Syst., x m , p. 713 (part) ; idem, 1907, Zool. Ergeb. Ned. Ost. Ind., IV, 2, p. 404 ;idem, 1909, Bull. Dep. Agr. Ind. Ned., XXV, p. 5 ;Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 261 ;Barbour, 1912, Mem. Mus. Comp. Zool. Harvard, xLrv, p. 71, P1. vn, fig. 27 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 154 (part), fig. ; Nieden, 1926. Das Tierreich, Anura 11,p. 32 (part) ;Dunn, 1928, Amer. Mus. Novitates, cccxv, p. 4 ;Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 489 ; Schijfsma, 1932, Treubia, XN, 1, p. 51. Dendromanes achatina, Gistel, 1848, Naturg. Thierr., p. 11. Micrhyh achatina, Giinther, 1858, Cat. Batr. Sal. Brit. Mua, p. 121. Diplopelm disciferum, Peters, 1867, Mon. Ak. Berlin, p. 36 (Type locality Java). Engystoma disciferum, Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 194.

Habit slender. Snout obtusely pointed, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical, interorbital space about once and a half the width of the upper eyelid ; tympanum hidden. Two or three outer fingers with small discs which usually have a median cleft on their upper surfaces ; first very short ; second shorter than the fourth ; outer metacarpal tubercle undivided. Toes with discs similar to, but larger than, those of the fingers, to ) webbed; the membrane midway between the third and fourth extending a little beyond the proximal tubercle of the third toe ; digits with lateral ridges ; third a little longer than the fifth ; subarticular tubercles not well developed ; two small metatarsal tubercles. Tibio-tarsal articulation reaching the nostril, or a little beyond the end of the snout. Skin smooth or feebly tubercular above ; smooth beneath ; anal region granular. Light pinkish-brown above, with an irregular dark-brown streak from above the shoulder almost t o the groin ; a light streak from beneath the eye t o the fore-limb, edged above with brown ; upper lip brown-spotted ; centre of the back with a narrow dark stripe, with undulating borders, from between the eyes to the end of the coccyx, or with irregular, indistinct, sinuous marblings ; fore and hinder side of thighs, and sometimes the lower surfaces of tarsus and foot, dark brown ; limbs indistinctly cross-barred. Lower surfaces white, the gular region brown-spotted. Male with a vocal sac and infuscate gular region. * Length : 23 mrn. DISTRIBUTION : Java, be!ow 1,600 m. Tadpole similar t o t h a t of M. heymonsi, but the expansion of the lower lip is slightly less, and the knob-like prominence on each side of the mouth has a central, sucker-like depression.

B.M. 44.2.22. 97 R.1928.6.30. 1 80.5.7. 5 91.1.30. 19-20 96.12. 32-35 96.12. 36-38 1928.11.17. 1-3 Mus. Vienna

. . . ? . . . . Posata, Bantam Java . . .

. . . . . (Mus. Leyden) COTYPE? ? . . . . ? . . . . . (Janson) ?? . . . . . Fruhstorfer 2 2 5, 2 $?$! Pengalengan, Java, 4,000 ft. . . Fruhstorfer 2, Pengalengan, Java, 4,000 ft. . . Fruhstorfer tadpoles Tjibodas, J a v a . . . . . . Dunn ?

Java

3 ads.

Pengalengan, Java, 4,500-5,000 ft. Fruhstorfer

MICROHYLA I o.

Microhyla pulchra (Hallowell)

Diplopelma ornutum (part), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 50. Engystoma pulchrum, Hallowell, 1860, Proc. Acad. Philadelphia, p. 506 (Type locality Between Hong Kong and Whampoa). Diplopelma pdchmm, Giinther, 1864, Rept. Brit. India, p. 417 ; Steindachner, 1867, Reise Novara, Amph., p. 36, P1. n, figs. 15-18 ; Theobald, 1868, Cat. Rept. Mus. Asiatic Soc. Bengal, p. 83 (part) ; Swinhoe, 1870, Proc. Zool. Soc. London, p. 412 ; Tirant, 1885, Rept. and Batr. Cochinchine et Cambodge, p. 96. Microhyla pulchra, Bodenger, 1882. Cat. Batr. Sal. Brit. Mus., ed. 2, p. 165 ; Boettger, 1885, Ber. Off. Ver. Nat.,p. 50 ; idem, 1888, op. cit., pp. 99 and 162 ; Flower, 1899, Proc. Zool. Soc. London, p. 905; Werner, 1903, Abh. Bayer. Akad. Wiss., (2), xxn, 2, p. 370 ; WolterstorIT, 1906, Abh. Mus. Magdeburg, I, 3, p. 131 ; Vogt, 1913, Sitzber. Ges. Natf. Fr. Berlin, pp. 222, 227, 229; Smith, 1916, Journ. Nat. Hist. Soc. Siam, n, pp. 39 and 169; Andersson, 1916, Vet. Ak. Handl. Stockholm, LV, 4, p. 17 ; Mell, 1922, Arch. Naturg., ~xxxvm,10, p. 131 ; Vogt, 1922, Arch. Naturg., LXXXVIII, 10, p. 145 ; Smith, 1923, Journ. Nat. Hist. Soc. Siam, VI, 2, p. 211 ; Parker, 1925, Ann. Mag. Nat. Hist., (9), XV, pp. 301, 304 ; Rao and Ramana, 1925, Proc. Zool. Soc. London, pp. 1445-1478 ; Nieden, 1926, Das Tierreich, Anura, n, p. 31, fig. ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 490 ; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., IV, 2, p. 26 ; Cochran, 1930, Proc. U.S. Nat,Mus., LXXVII, 11, p. 6 ;Pope, 1931, Bull. Amer. Mus. Nat. Hist., LXI, 8, p. 599, fig. 39. Scaptophryne labyrinthica, Fitzinger, 1861, Sitzber. Ak. Wiss. Wien, XLII, p. 416 (Type locality Hong Kong). Ranina symmetrica, David, 1872, Nouv. Arch. Mus. Bull., VII, p. 76 (Type locality " Hills near the Yangtsekiang, and becoming abundant towards Sze-chwan "). Microhyla hainanensis, Barbour, 1908, Bull. Mus. Comp. Zool. Harvard, LI, p. 322 (Type locality Mt. Wuchi, Central Hainan) ;idem, 1909, Proc. New England Zool. Club, rv, p. 57 ; idem, 1912, Mem. Mus. Comp. Zool. Harvard, XLIV, 1, P1. VII, fig. 2 8 ; Vogt, 1913, Sitzber. Ges. Natf. Fr. Berlin, pp. 227, 229; idem, 1922, Arch. Naturg., rxxxvm, 10, p. 145; Smith, 1923, J o u ~ ~ . Nat. Hist. Soc. Siam, VI, 2, p, 211 ; Nieden, 1926, Das Tierreich, Anura, n, p. 34. Microhyla melli, Vogt, 1914, Sitzber. Ges. Natf. Fr. Berlin, p. 101 (Type locality Environs of the city of Canton) ;Mell, 1922, Arch. Naturg., ~xxxvm,10, p. 130 ;Gee and Boring, 1929, Peking Soc. Nat. Hist. bull.,^^, 2,p. 26. Microhyla boulengeri (non Vogt), Mell, 1922, Arch. Naturg., L X X X V I ~ ,10, p. 130. Microhyla major, Ahl, 1930, Sitzber. Ges. Natf. Fr. Berlin, p. 317 (Type locality Yao-Shan Mts., Kwangsi, 1,500 m., 24" 5' N., 1loo E.).

Habit moderately stout. Snout obtusely pointed, slightly prominent, as long as, or slightly longer than, the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space as broad as, or a little broader than, the upper eyelid ; tympanum hidden. Digits not dilated, without median cleft above. First finger much shorter than the second, which is a little shorter than the fourth. Toes half webbed, or a little more or a little less ; membrane midway between the third and fourth not quite reaching the distal tubercle of the third toe ; toes more or less fringed, the outer much shorter than the third ; subarticular tubercles well developed ; two rather small, rounded, metatarsal tubercles. Tibio-tarsal articulation reaching the eye or beyond the tip of the snout. Skin smooth or feebly granular above, sometimes a transverse occipital fold connecting the posterior corners of the upper eyelids and continued obliquely backwards to the fore-limb. Smooth beneath ; anal region granular. Pale olive, grey, or pinkish brown above ; a narrow dark brown bar from eye to eye, an oblique lateral band from behind the eye to the middle of the flanks, and a A-shaped marking on the back, its apes between the scapulae and its extremities in the groins ; within this A are two or three collateral, smaller A S which are frequently incomplete or broken up into lines and elongate spots. Forehead and dorsum external to the A with alternately darker and lighter narrow lines, those of the forehead parallel with the interorbital bar, and those of the back parallel with the A . A dark canthal stripe. Limbs with numerous cross-bars, frequently very irregular. Yellowish-white beneath, the chin and- chest mottled with brown.

MONOGRAPH OF THE MICROHYLIDAE Male with a vocal sac and infuscate gular region. Length : 35 mm. DISTRIBUTION : S. China (Hong Kong, Kwantung, Hupeh), Hainan, Tonkin, Siam and Cochin-China. Eggs frequently laid in transitory puddles ; development very rapid. A full-grown tadpole measures about 28 mm. Body 1%times as long as broad ; tail slightly more than twice as long as the body. Nostrils close together, nearer the tip of the snout than the eye ; mouth dorso-terminal, the lips not produced ; eyes lateral, the interocular space about 5 times the internarial ; spiraculum opening into a transparent sheath about the middle of the gut ; anus median. Tail ending in a distinct filament ; 4 times as long as deep, with the lower crest a little deeper than the upper. Almost colourless and very transparent, sometimes faintly brown-stippled on the dorsum and muscular part of the tail. Toes fully webbed. SPECIMENS EXAMIXED B.M. 48.8.16. 13 56.11.17. 45 92.8.4. 11 51.12.27. 72.1.29. 19 86.6.30. 51 1928.11.16. 41 1924.9.19. 1 1924.12.9. 13 1903.4.295. 87-96 1928.11.15. 143

6 6 ? ? ? ? 6 ? 6

66, $2 tadpoles ? ? ? 6 ? hgr.

6 ? ?

Mus. Berlin

Ad.

I I.

Hong Kong . . . . . . Rowring Hong Kong . . . . . . Bowring Hong Kong . . . . . . Walker China . . . . . . . . . China . . . . . . . . Adams Macao . . . . . . . . Anderson . . . . M. Smith Hoi How, Hainan . . . . . Stevens Hanoi, Tonkin Thai-Nien, Tonkin . . . . Stevens Man-Son Mts., Tonkin, 3 4 , 0 0 0 f t . Fruhstorfer Nong Pling, nr. Paknampo, C. M. Smith (deSiam . scribed by M. Smith) . . . . . M. Smith Krabin, Siam Phrae, Siam . . . . . . M. Smith Dong Phya Fai, Siam, 700 ft. . Flower Bortong, Siam . . . . . Flower Hup Bon, S.E. Siam . . . . M. Smith Cambodia . . Mouhot . . . M. Smith Patiyu, Penins. Siam Praak Puttabat, Chumpon, Penins. Siam . . . . . M. Smith Yaoshan Mts., Kwangsi . . .,Shin, TYPEof M. major, Ah1 Hainan . . Mell, TYPE of M. melli, Vogt

Microhyla okinavensis, Stejneger

Microhylofiasipes(non Boulenger, 1884), Boulenger, 1887, Proc. Zool. Soc. London, p. 150 ;Boettger, 1888, Ber. Off. Ver. Nat., p. 162 (part) ; Okada, 1891, Cat. Vert. Jap., p. 66 ; Fritze, 1894, Zool. Jahrb., Syst., vn, p. 865; Boettger, 1895, Ber. Off. Ver. Nat., xxxvr, p. 106; Okada, 1931, TaiIless Batr. Jap. Emp. p. 71 (part), figs. 33,34, P1. vn, fig. 3, P1. x w , fig. 7, P1. xxm, fig. 6. Microhylo okimvensis, Stejneger, 1901, Proc. Biol. Soc. Washington, XIV, p. 189 (Type locality Okinawa Shirna, Riu Kiu Archipelago) ; idem, 1907, Bull. U.S. Nat. Mus., L=, p. 89 ; Barbour, 1909, Proc. New Eng. Zool. Club, rv, p. 58 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), 11, p. 492. Microhyla undulba, Brown, 1902, Proc. Acad. Philadelphia, p. 186 (Type locality Ooshima or Okinawa, Loo Choo Islands).

Habit moderately stout. Snout obtusely pointed, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical ;

MICROHYLA

139

interorbital space about once and a third the width of the upper eyelid ; tympanum hidden. Fingers not dilated, without cleft above ; first much shorter than the second, which is a little shorter than the fourth. Toes slightly spatulate distally, and sometimes with more or less distinct traces of a cleft on their upper surfaces ; toes with a rudiment of web (nearly 4)and with more or less distinct lateral frmges ; subarticular tubercles well developed ; two well-defined metatarsal tubercles. Tibio-tarsal articulation reaching the eye in females or from this point to beyond the tip of the snout in males. Skin smooth or slightly pustular above ; traces of an occipital fold and its continuation from the posterior conier of the eye to the fore-limb ; smooth beneath, the anal region granular. Grey or pinkish above, with a dark lateral stripe from the eye to the middle of the flanks or the groin, sometimes interrupted above the shoulder ; a dark, mid-dorsal marking, commencing between the eyes and connecting the upper eyelids, narrowing behind the occiput, broadening between the shoulders, narrowing, broadening again to give off a very short, backwardly directed process, then narrowing abruptly and bifurcating, the two branches extending towards the thighs ; the edges of this marking may be emphasised or the whole may be indistinct ; there are usually a few indistinct wavy bands parallel with the edges of the dorsal marking. Limbs cross-barred ; a dark patch on the anterior aspect of the knee and another enclosing the vent. White beneath, the gular region and chest clouded with pale brown. Male with a vocal sac and infuscate gular region. 2 5 mm. ; $2 3 1 mm. Length : DISTRIBUTION : Riu Kiu Archipelago. Eggs are laid in small masses adhering to water-plants. A well-grown tadpole measures 3 2 mm., and is essentially similar to that of M. pulch~a. Body 1 to 1 ) times as long as broad ; tail 2 to 2 4 times the length of the body. Nostrils nearer the tip of the snout than the eye; mouth dorso-terminal, the lower lip not expanded, but papillose ; eyes lateral, the interorbital space 3 & to 5 times the internarial. Tail 3 to 3 ) times as long as deep, with a terminal flagellum. Body and muscular portion of the tail with some black markings (after Okada).

B.M. 87.1.31.43-44 92.9.3.24-26 1902.5.19.21-22 1925.11.7. 8

12.

8 $2 $2 8

Loo Choo Islands . . . Motobu, Okinawa Shima . Ishigaki Shima . . . Ishigaki Shima . . .

. . . . . . .

.

. . .

. . Pryer

.

. Holst

'

. . Owston . . . Owston

Microhyla ornata (Durn. and Bibr.)

Engystoma ornatum, Dumbril and Bibron, 1841, Erpet. GBn., vnr, p. 745 (Type locality Malabar coast) ; Cope, 1867, Journ. Acad. Philadelphia, (2), VI, p. 194. Siphneus orlultum, Fitzinger, 1843, Syst. Rept., p. 33. Diplopelma ornatum, Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 50 (part); idem, 1864, Rept. Brit. India, p. 417 ; Theobald, 1868, Cat. Rept. Mus. Asiatic Soc. Bengal, p. 82 (part). Micmhyb ornata, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 165 ; Boettger, 1885, Ber. Off. Ver. Nat., pp. 48, 5 0 ; Miiller, 1887, Verh. Nat. Ges. Basel, V I ~2,, p. 251 ; Boettger, 1888, Ber. Off. Ver. Nat., pp. 98, 162 ; Anderson, 1889, Journ. Linn. Soc. London, xxr, p. 350 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 491 ; Boettger, 1892, Ber. Off. Ver. Nat., pp. 98,102 ;Sclater, 1892, Batr. Ind. Mus., p. 22 ;Flower, 1899, Proc. Zool. Soe. London, p. 901, P1. LX, fig. 1 ; Laidlaw, 1900, Proc. Zool. Soc. London, p. 887 ; Boettger, 1901, Ber. Senck. Nat. Gea., p. 52 ; Boulenger, 1903, Fasc. Malay. Zool., I, p. 133 ; Butler, 1904, Journ. Bombay Nat. Hist. Soc. xv, p. 387 ; Ferguson, 1904, Journ. Bombay Nat. Hist. Soc., xv, p. 506; Wolterstorff, 1906, Abh. Mus. Magdeburg, I, 3, p. 131 ; Sarasin; 1910, Zool. Jahrb., Syst., xu, pp. 131, 134 ; Boulenger, 1912, Bauna Malay Penins., Rept. and Batr., p. 260; Vogt, 1913, Sitzber. Ges. Natf. Pr. Berlin, pp. 222, 227; Smith, 1916, Journ. Nat. Hist. Soc. Siam, n, p. 169 ; idem, 1917, op. cit., 11, p. 268 ; Andersson, 1917, Vet. Ak.

MONOGRAPH O F T H E MICROHYLIDAE Handl. Stockholm, LV, 4, p, 1 7 ; Rao, 1917, Rec. Ind. Mus., XIU, p. 281 ; Rao, 1918, op. cit., p. 42 ;Mell, 1922, Arch. Naturg., LXXXYIII, 10, p. 130 ;Vogt, 1922, Arch. Naturg., L x x x v m , 10, p. 145 ; Smith, 1923, Journ. Nat. Hist. Soc. Siam, n, p. 212 ;Parker, 1925, Ann. Mag. Nat. Hist., (9), xv, p. 304 ; Rao and Ramana, 1925, Proc. Zool. Soc. London, pp. 1445-1478 ; iidem, 1925, Quart. Journ. Micr. Sci., LXIX, p. 735 ; Nieden, 1926, Das Tierreich, Anura 11, p. 30, fig. ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), n, p. 493; Gee and Boring, 1929, Peking Soc. Nat. Hist. Bull., rv, 2, p. 2 6 ; Cochran, 1930, Bull. U.S. Nat. Mus., ~ x x v n ,11, p. 7 ; Smith, 1930, Bull. Raffles Mus., m, p. 129 ; Pope, 1931, Bull. Amer. Mus. Nat. Hist., LEI, 8, p. 597 ;McCann, 1932, Journ. Bombay Nat. Hist. Soc., xxxv~,p. 176. Engyatoma carnaticum, Jerdon, 1854, Journ. Asiatic Soc. Bengal, XXII, p. 534 (Type locality the Carnatic) ; Mason, 1860, Burma, ed. 2, p. 325. Diplopelma carnaticum, Stoliczka, 1870, Journ. Asiatic Soc. Bengal, xxxrx, p. 154 ; Anderson, 1879, Zool. Yunnan, p. 841. ? Engyatoma malabaricum, Jerdon, 1854, Journ. Asiatic Soc. Bengal, XXII, p. 534 (Type locality Malabar). Microhyla$aaipe.s, Boulenger, 1884, Ann. Mag. Nat. Hist.. (5), xm, p. 397 (Type locality Taiwan-fu, S. Formosa); Boettger, 1887, Ber. Off. Ver. Nat., p. 162 (part) ; Werner, 1903, Abh. Bayer. Akad. Wiss., n, 22, p. 369 ; Stejneger, 1907, Bull. U.S. Nat. Mus., LVIII, p. 8 8 ; Vogt, 1914, Sitzber. Ges. Natf. Fr. Berlin, p. 101 ;Mell, 1922, Arch. Naturg., ~xxxvm,10, p. 130 ;Stejneger, 1925, Proc. U.S. Nat. Mus., ~xvr,p. 12 ; Nieden, 1926, Das Tierreich, Anura 11, p. 35 ; Okada, 1931, Tailless Batr. Jap. Emp. p. 71 (part). Microhyla eremita, Barbour, 1920, Occ. Papers. Mus. Zool. Michigan, LXXVI, p. 3 (Type locality Nanking) ; Stejneger, 1925, Proc. U.S. Nat. Mus., LXVI, p. 11 ; Schmidt, 1927, Bull. Amer. Mus. Nat. Hist., LIV, p. 562 ; Fang and Chang, 1931, Contr. Biol. Lab. Sci. Soc. China, Zool., VII, 2, p. 87. xv,

Habit moderately slender. Snout obtusely pointed, slightly prominent, a little longer than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space a little broader than the upper eyelid. Digits not dilated distally and without any trace of a median cleft above. First finger much shorter than the second, which is subequal to, or a little shorter than, the fourth. Toes with a rudiment of web, but without lateral fringes ; subarticular tubercles distinct ; two small, prominent metatarsal tubercles. Tibio-tarsal articulation reaching t o between the shoulder and the anterior border of the eye, or very slightly beyond the latter point. Skin smooth or slightly tubercular above, and often with small warts which sometimes form a longitudinal series from the eye to the groin ; sometimes an occipital fold connecting the posterior corners of the upper eyelids and descending obliquely to the fore-limb. Lower surfaces smooth ; anal region granular. Pinkish olive, grey, or brown above, with a more or less defined dark streak from behind the eye, above the shoulder to the flank ; a median dark spot commencing between the eyes, where it connects the upper eyelids, narrowing behind the occiput, broadening abruptly between the fore-limbs, narrowing slightly, broadening on the sacral region and sending a branch obliquely backwards to each groin, narrowing immediately and then broadening and bifurcating, each branch being directed obliquely backwards and continued across the thigh ; this area may be very ill-defined or broken up into spots, the breaks occurring a t the narrowed portions. Rest of the dorsum sometimes with undulating dusky lines ; a light streak from the eye to the insertion of the fore-limb. The warts and granules of the back are frequently lighter (red in life), particularly in young specimens. Limbs cross-barred ; anterior aspect of thighs and anal region darker. Lower surfaces whitish, the throat and chest closely stippled with brown. Male with a vocal sac which forms a moderately distinct fold across the chest. Length : 9 25 mm. ; $ 2 2 . 5 mm. : China (south of Sze-chwanand Kiangsu), Formosa, Tonkin, Haincn, DISTRIBUTION Burma, Assam, Siam, Annam, Cochin-China, Malay Peninsula, India and Ceylon.

Eggs frequently laid in transient pools ; development very rapid. Tadpole almost indistinguishable from that of M. pulchra, but slightly smaller, and often with a somewhat shorter tail.

MICROHYLA SPECIMENS EXAMINED

? d ?

B.M. 1925.11.7. 7

Nanking

.

. (Barbour) as M.

eremita, Barbour . . . . . . Ningpo Da-laen-saen, S.W. of Ningpo Walker hgr. Ning-kong-chow, S.W. of Ningpo . . . . . . Walker 94.11.14. 9-10 d? C.Formosa . .Hoist 84.3.11. 6 ? Taiwanfu, S. Formosa . . TYPE of M. fissipes, Boulenger 92.8.4. 9-10 . Walker ?? Hong Kong . 86.6.30. 50 . ? Macao Anderson 70.1.14. 2&25 . Swinhoe d? Sze-chwan . 1928.11.16. 35-36 Nr. Kachek, S.E. Hainan . M. Smith 1924.12.9. 114-115 ?? Thai Nien, Tonkin . Stevens 1903.4.29. 85-86 ?? Man-son Mts., Tonkin . . F ~ h s t o r f e r 1920.1.20. 3539 d Hue, Cochin-China . . , . Dorr 1928.5.16. 33-34 d? Tourcham, Annam . . . M. Smith 1927.3.20. 58-59 ?? Kontunl, Annam . Delacour and Lowe 61.8.12. 12 . ? Cambodia Mouhot 1928.11.16. 37 ? Pak Maat, nr. Vien Chan, N. Siam. . M. Smith 1916.3.27. 50 . M. Smith 6 Bangkok . 97.10.8. 85-91 5dd , 2 Bangkok . . Flower 97.10.8. 93-98 tadpoles Bangkok . . . Flower 97.10.8. 83-84 d? Bo=tong, Siam . . Flower 97.10.8. 7%82 9,hgr. Chantaboon, Siam . Flower 98.5.18. 3 7 4 0 2 3 8 , 2hgr. Ayuthia, Siam . Flower 1928.11.16. 38-39 d Paknam Chumpon, Penin. Siam . . . . . . M. Smith . . . . . Flower ?? Kedah , 98.9.22. 116-123 W Mabek, ~ a l o r Malay , Penin- Annandale and 1903.4.13. 123 sula. . Robinson 87.2.26. 25-31 38, Moulrnein, Burma Theobald hgr87.2.26. 3 2 4 1 , Pegu, B u m a . . Theobald hgr. 93.11.17. 18 ? Thayetmyo, Burma . . Watson 72.4.17. 231-232 ?? Assam . Jerdon 69.8.28. 4 6 4 7 India . . . . . . . 1928.6.30. 2 India . . . . . . . Belcher 46.11.22. 51-52 'W Madras . . Jerdon 80.11.10. 102 Nr. ~urnaserdian,Godaveri Blanford Valley 1908.1.29. 27-29 338 Godaveri Valley . Blanford 6 9,hgr. Palode, Travancore . . . Ferguson 97.2.8. 12-15 . . Sauter 92 6 d, 9, Kanshirei, Formosa Mus. Vienna

54.2.10. 28 93.4.20. 40 93.4.21. 1&13

.

-

.

Sd

m

9

I 3.

$Fp

.

Phuc-Son, Annam

.

.

. Fruhstorfer

Mimohyla picta, Schenkel

Yicrohyla picta, Schenkel, 1901, Abh. Nat. Ges. Basel, xm, p. 151, fig. (Typelocality " wahrscheinlich Cochinchina ") ; Smith, 1921, Proc. Zool. Soc. London, p. 437 ;Nieden, 1926, Das Tierreich, Anura 11, p. 30 ; Parker, 1928, AM. Mag. Nat,. Hist., (10),n, p. 494.

MONOGRAPH O F T H E MICROHYLIDAE Habit rather stout. Snout rounded, scarcely prominent, slightly shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region moderately oblique; interorbital space a little broader than the upper eyelid ; tympanum hidden. Digits not dilated, without median cleft above. First finger much shorter than the second, which is subequal to the fourth ; outer metacarpal tubercle entire. Toes about $ webbed, the outer a little shorter than the third ; subarticular tubercles distinct; two large, compressed, metatarsal tubercles. Tibio-tarsal articulation reaching to between the shoulder and the eye. Skin smooth or slightly warty above ; traces of an occipital fold connecting the posterior corners of the upper eyelids and continued obliquely backwards to the insertion of the fore-limb. Lower surfaces smooth ; the anal region granular. Reddish brown above, with a median dark spot which commences between the eyes, where it connects the upper eyelids, narrows on the nape, broadens between the shoulders, narrows slightly, then broadens and gives off a branch obliquely backwards to each groin, narrows abruptly and finally broadens and bifurcates, each branch being continued across the thigh. This marking may be finely edged with white and may be broken up into irregular spots. Remainder of dorsum and sides with dark insuliform spots or irregular marblings ; a light streak from behind the eye to the fore-limb ; canthus-rostralis darker. Limbs cross-barred. Lower surfaces yellowish, the gular region marbled with brown. Male with a vocal sac and infuscate gular region. Length : $ 29 rnm. ; 31 mm. DISTRIBUTION : Cochin-China, S. Annam.

?

B.M. 1921.4.1. 272 1921.4.1. 273-278 1927.9.1. 3

8$, 9$2 cleared

Mus. Vienna

I 4.

d

Tour Cham, S. Annam . . Cap St. Jacques, Cochin-China Cap St. Jacques, Cochin-China Annam . . . . . .

.

. .

M. Smith M. Smith M. Smith Fruhstorfer

Mcrohyla rubra (Jerdon)

Engystoma rubrum, Jerdon, 1854, Journ. Asiatic Soc. Bengal, XXII,p. 531 (Typelocality the Carnatic). i7licrohyla rubra, Boulenger, 1882, Cat. Batr. Sal. Brit. bfus., ed. 2, p. 164 ; Boettger, 1889, Ber. Senck. Nat, Ges., p, 298 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 491 ; Sclater, 1892, Batr, Ind. Mus., p. 22 ; Werner, 1893, Verh. Zoo1.-bot. Ges. Wien., XLIII,p. 353 ; Ferguson, 1904, Journ. Bombay, Nat. Hist. Soc., xv, p. 506 ; Annandale, 1009, Rec. Ind. Mus., III, p. 286 ; Rao, 1915, Rec. Ind. Mus., XI,p. 31 ; idem, 1917, op. cit., XIII,p. 282 ; idem, 1918, op cit., xv, p. 42 ; Prashad, 1918, Rec. Ind. Mus., xv, p. 101, P1. IX,fig. 10 ; Nieden, 1926, Das Tierreich, Anura 11, p. 29, fig. ; Schmidt, 1926, Publ. Field Mus. Chicago, Zool., XII,p. 168 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo),11, p. 496. Diplopelma ornatum (part),Giinther, 1858, Cat. Batr. Sal. Brit. Xus., p. 50 ; Theobald, 1868, Cat. Rep. Mus. Asiatic Soc. Bengal, p. 82. Copea fulva, Steindachner, 1864, Verh. Zoo1.-bot. Ges. Wien, p. 286, P1. XVII,fig. 5 (Type locality " Brazil ") ; Parker, 1932, Ann. Mag. Nat. Hist., (lo),X,p. 341. Habit stout. Snout rounded, scarcely prominent, a little shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space once and a quarter t o once and a half as broad as the upper eyelid ; tympanum hidden. Digits not dilated and without median cleft above. First finger much shorter than the second, which is very slightly shorter than the fourth; outer metacarpal tubercle completely divided or very distinctly notched distally. Toes 2 webbed, sometimes more, with traces of lateral fringes ; third distinctly longer than the fifth ; subarticular tubercles prominent ; two large, compressed, metatarsal tubercles. Tibio-tarsal articulation reaching to between the shoulder and the eye. Skin smooth, or slightly warty above ; sometimes a distinct row of warts from the

MICROHYLA SPECIMENS EXAMINED B.M. 1925.11.7. 7

?

Nanking

.

. (Barbour) as M.

eremita, Barbour . . . Ningpo . . Da-laen-saen, S.W. of Ningpo Walker h ~ . Ning-kong-chow, S.W. of Ningpo . . Walker 94.11.14. 9-10 89 C. Formosa . Holst 84.3.11. 6 ? Taiwanfu, S. Formosa . . TYPE of M. 3 s sipes, Boulenger 92.8.4. 9-10 . Walker ?? Hong Kong . 86.6.30. 50 . ? Macao Anderson 70.1.14. 24-25 89 Sze-chwan. . Swinhoe 1928.11.16. 35-36 89 Nr. Kachek, S.E. Hainan . M. Smith 1924.12.9. 114-115 ?? Thai Nien, Tonkin . . Stevens 1903.4.29. 85-86 $? Man-son Mts., Tonkin . . Fruhstorfer 1920.1.20. 3539 . Dorr 8 Hue, Cochin-China . 1928.5.16. 33-34 89 Tourcham, Annam . . ' . M. Smith 1927.6.20. 58-59 ?? Kontum, Annam . Delacour and Lowe 61.8.12. 12 Cambodia . . . . Mouhot 1928.11.16. 37 Pak Maat, nr. Vien ~ h a n , N. Siam. . M. Smith 1916.3.27. 50 . M. Smith 8 Bangkok . 97.10.8. 85-91 5 88,2 9 9 Bangkok . . Flower 97.10.8. 93-98 tadpoles Bangkok . . Flower 97.10.8. 83-84 89 Bortong, Siam . Flower 97.10.8. 79-82 2, hgr. Chantaboon, Siam . Flower 98.5.18. 3 7 4 0 2 88, 2hgr. Ayuthia, Siam . Flower 1928.11.16. 38-39 6.9 Paknam Chumpon, Penin. Siam . M. Smith 98.9.22. 116-123 8 , Kedah . Flower 1903.4.13. 123 hgr. Mabek, ~ a i o r'Malay , Penin- Annandale and Robinson sula. . 87.2.26. 25-31 8 , Moulmein, Burma . Theobald hgr. 87.2.26. 3 2 4 1 8, Pegu, Burma . . Theobald hgr. 93.11.17. 18 ? Thayetmyo, Burnla . . . Watson 72.4.17. 231-232 ?? Assam . Jerdon 69.8.28. 4 6 4 7 88 India . . . . . . . 1928.6.30. 2 ? India . . . . . . . Belcher 46.11.22. 51-52 hgr. Madras . . Jerdon 80.11.10. 102 h ~ Nr. ~umaserdian,Godaveri . Blanford Valley 1908.1.29. 27-29 888 Godaveri Valley . Blanford 97.2.8. 12-15 80, hgr. Palode, Travancore . . . Ferguson Mus. Vienna 92 8 8, 9, Kanshirei, Formosa . . Sauter Yg. 9 , Phuc-Son, Annam . Fruhstorfer

54.2.10. 28 93.4.20. 40 93.4.21. 10-13

8 ?

8

13. Microhyla picta, Schenkel Microhyla picta, Schenkel, 1901, Abh. Nat. Ges. Basel, xm, p. 151, fig. (Typelocality " wahracheinlich Cochinchina ") ; Smith, 1921, Proc. Zool. Soc. London, p. 437 ;Nieden, 1926, Das Tierreich, Anura U, p. 30 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo),11, p. 494.

MONOGRAPH O F T H E MICROHYLIDAE Habit rather stout. Snout rounded, scarcely prominent, slightly shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region moderately oblique; interorbital space a little broader than the upper eyelid ; tympanum hidden. Digits not dilated, without median cleft above. First finger much shorter than the second, which is subequal to the fourth; outer metacarpal tubercle entire. Toes about f webbed, the outer a little shorter than the third; subarticular tubercles distinct ; two large, compressed, metatarsal tubercles. Tibio-tarsal articulation reaching to between the shoulder and the eye. Skin smooth or slightly warty above ; traces of an occipital fold connecting the posterior corners of the upper eyelids and continued obliquely backwards to the insertion of the fore-limb. Lower surfaces smooth ; the anal region granular. Reddish brown above, with a median dark spot which commences between the eyes, where it connects the upper eyelids, narrows on the nape, broadens between the shoulders, narrows slightly, then broadens and gives off a branch obliquely backwards to each groin, narrows abruptly and finally broadens and bifurcates, each branch being continued across the thigh. This marking may be finely edged with white and may be broken up into irregular spots. Remainder of dorsum and sides with dark insuliform spots or irregular marblings; a light streak from behind the eye to the fore-limb ; canthus. rostralis darker. Limbs cross-barred. Lower surfaces yellowish, the gular region marbled with brown. Male with a vocal sac and infuscate gular region. Length : 8 29 mm. ; 31 mm. : Cochin-China, S. Annam. DISTRIBUTION

? B.M. 1921.4.1. 272 1921.4.1. 273-278 66, 1927.9.1. 3 cleared 6 Mus. Vienna

.

Tour Cham, S. Annam . . Cap St. Jacques, Cochin-China Cap St. Jacques, Cochin-China Amam . . . . . .

.

.

M. Smith M. Smith M. Smith Fruhstorfer

14. Microhyla rubra (Jerdon) Engystoma rubrum, Jerdon, 1854,Journ. Asiatic Soc. Bengal, XXII,p. 534 (Type locality the Carnatic). Mic~ohylaTubTa, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 164 ; Boettger, 1889, Ber: Senck. Nat. Ges., p. 298 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 491 ; Sclater, 1892, Batr. Ind. Mus., p. 22 ; Werner, 1893, Verh. Zoo1.-bot. Ges. Wien., xLm, p. 353; Ferguson, 1904, Journ. Bombay, Nat. Hist. Soc., xv, p. 506 ;Annandale, 1909, Re* Ind. Mus., m, p. 286 ; Rao, 1915, Rec. Ind. Mus., XI, p. 31 ; idem, 1917, op. cit., XIII,p. 282 ; idem, 1918, op cit., xv, p. 42; Prashad, 1918, Rec. Ind. Mu., XV,p. 101, P1. IX,fig. 10 ; Nieden, 1926, Das Tierreich, Anura 11, p. 29, fig. ; Schmidt, 1926, Publ. Field Mus. Chicago, Zool., XII,p. 168 ; Parker, 1928, Ann. Mag. Nat. Hist., (lo), n, p. 496. Diplopelma ovnatum (part), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 50 ; Theobald, 1868, Cat. Rep. Mus. Asiatic Soc. Bengal, p. 82. Gopea fulva, Steindachner, 1864, Verh. Zoo1.-bot. Ges. Wien, p. 286, P1. XVII,fig. 5 (Type locality " Brazil ") ; Parker, 1932, Ann. Mag. Nat. Hiat., (lo), x, p. 341.

Habit stout. Snout rounded, scarcely prominent, a little shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space once and a quarter to once and a half as broad as the upper eyelid ; tympanum hidden. Digits not dilated and without median cleft above. First finger much shorter than the second, which is very slightly shorter than the fourth ; outer metacarpal tubercle completely divided or very distinctly notched distally. Toes webbed, sometimes more, with traces of lateral fringes ; third distinctly longer than the fifth ; subarticular tubercles prominent ; two large, compressed, metatarsal tubercles. Tibio-tarsal articulation reaching to between the shoulder and the eye. Skin smooth, or slightly warty above ; sometimes a distinct row of warts from the

a

MICROHYLA

143

posterior corner of the eye to the flanks ;a fold from the eye to the fore-limb. Smooth beneath, the anal region granular. Reddish brown above, with a dark lateral band from the tip of the snout through the eye to the groin ; an elongate dark brown lumbar spot, continued across the thigh ; mid-dorsal region with faint traces of a dark mark similar to that of M. picta, but often broken up ; a dark streak along the anterior aspect of the knee ; hinder side of the thighs and anal region dark marbled. Limbs indistinctly cross-barred. White beneath, the throat and chest clouded with light brown. Male with a vocal sac and infuscate gular region. Length : 29 mm. DISTRIBUTION : Ceylon, Southern and Central India, Assam. Eggs laid in floating masses. A well-grown tadpole measures 36 mm. and has the following characters : Body about 1%times as long as broad ; tail 2; times as long as the body. Mouth dorso-terminal ; nostrils close together, nearer a line connecting the anterior borders of the eyes than the end of the snout ; eyes lateral, the interorbital space about 6 times the internarial, twice and a half as broad as the mouth. Spiraculum opening under a transparent sheath near the middle of the gut ; anal tube reflected forwards and opening close behind the spiraculum. Tail with a terminal filament, 4 times as lon as deep ;the lower caudal crest twice as deep as the upper. Transparent, with some dafk spots above and on the caudal membranes posteriorly.

B.M. 87.2.26. 24 72.4.17. 233-234 46.11.22. 60-64 46.11.22. 60-64 72.4.17. 196196 95.12.30. 50 1903.9.26. 29-33 97.2.8. 6-9 74.4.29. 268 76.3.21. 48 1908.7.2. 12-13

Mus. Vienna

15.

? ?? 33, $29,

.

hgr skeleton

$29 ?

tadpoles and yg.

8, ?? & $29

3, ,

yg.

. Theobald . . Jerdon . . Jerdon . . . . . Jerdon . . . . . Jerdon (as

Moulmein, Burma Assam . . . . . . Madras Madras S. India

.

typical of Engystoma TUb u m , Jerdon) Trevandrum, Travancore Ferguson Trevandrum, Travancore Ferguson

Palode, Travancore . Ceylon . . . . ~e$lon . . . . Trincomalee, Ceylon " Brazil " . . .

. . . .

.

Ferguson Beddome Ferguson Green " Natterer," CoTYPES of Copea fulva, Steindachner

Microhyla aterrima (Giinther)

Qbssostoma atenimum, Giinther, 1902, Biol. Centr. Amer., Rept. and Batr., p. 210, PI. LXIV, fig. A (Type locality Costa Rica). Boulenger, 1913, Proc. Zool. Soc. London, p. 1025; Nieden, 1926, Das Tierreich, Anura n, p. 68, fig. 45. Gastrophryne aterrimum, Parker, 1927, Occ. Papera. Mus. Zool. Michigan, CLXXXVII, p. 5.

Habit moderately stout. Snout rounded, prominent, about once and two-thirds as long as the eye ; canthus rostralis rounded ; loreal region oblique, slightly concave ; nostril much nearer the tip of the snout than the eye ; interorbital space twice as broad as the upper eyelid. Digits not dilated and not cleft above. Fingers short, tapering, the first shorter than the second, which is shorter than the fourth. Toes nearly half webbed ; subarticular tubercles flat ; a flat, oval, inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching to between the shoulder and the tympanic region.

MONOGRAPH O F THE MICROHYLIDAE Skin thick, smooth above and beneath ; a strong fold connecting the posterior comers of the upper eyelids and descending to the angle of the lower jaw. Edge of lower jaw, upper surfaces of arms, tibiae, tarsi and soles of feet, edges of the digits, and the anal region beset with more or less numerous, minute spinose pustules. Dark purplish brown or blackish above ; lighter beneath, the throat and limbs more nearly approaching the colour of the upper surfaces. Length : 5 5 mm. : Costa Rica, Colombia and N.W. Ecuador. DISTRIBUTION B.M. 1902.5.12.6 1913.1 1.12. 9 5

Costa Rica . . Underwood, TYPE PeAa Lisa, Condoto, Choco, ColomSpurrel bia ( 3 0 0 ft.) Rio Durango, N.W. Ecuador

d

.

1902.5.27.20 I

. .

6 . Microhyla elegans (Boulenger)

Engystoma carolinense (part), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 51. Engystoma elegans, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 162 (Type locality Cordoba, Mexico) ; Giinther, 1902, Biol. Centr. Amer.; Rept. and Batr., p. 210 ; Nieden, 1926, Das Tierreich, Anura 11, p. 66. Gmtrophryne elegans, Stejneger, 1910, Proc. Biol. Soc. Washington, x x m , p. 166; Kellogg, 1932, Bull, U.S. Nat. Mus., CLX, p. 187. Engystoma ustum (non Cope), Gadow, 1905, Proc. Zool. Soc. London, p. 194.

Dermal ridges of the roof of the mouth curved, more or less crenulate. Habit stout. Snout pointed, p~ominent,nearly twice as long as the eye ; canthus rostralis rounded ; loreal region oblique, not concave ; nostril much nearer the tip of the snout than the eye; interorbital space twice as broad as the upper eyelid. Digits not dilated and without cleft above. Fingers moderate, ending obtusely, the first much shorter than the second, which is shorter than the fourth. Toes with a rudiment of web ; subarticular tubercles flat ; a small oval inner, but no outer, metatarsal tubercle. Tibiotarsal articulation reaching the tympanic region. Skin smooth above and beneath ; a strong fold connecting the posterior corners of the upper eyelids and extending downwards to the angles of the mouth. Brown above ; a dark lateral streak from the tip of the snout to the groin, covering the loreal and temporal regions ; an indistinct dark blotch on the back, commencing between the eyelids, narrowing on the nape, then broadening and becoming indistinct ; a well-defined, dark, circular spot in the inguinal region, not covered by the folded hind-limbs. A single, broad, dark bar across the femur, tibia and tarsus ; these bars sometimes paler in the middle and so becoming partially divided. ~ o & e surfaces r yellowish, with a bold, very regular and well-marked, dark reticulum, with smaller meshes beneath the throat and on the limbs ; a triangular dark spot enclosing the vent ; lower surfaces of tarsus and foot black. Male with a vocal sac ; throat infuscate ; limbs, upper and lower lips, and to a less extent the dorsum, beset with scattered, almost microscopic, spinules. Length from snout to vent : 3 1 mm. DISTRIBUTION : S.E. Mexico. SPECIMENS EXAMINED

Q

B.M. 56.3.17. 2 7 1903.9.30.234-235

3

. . . . . . . . .

Cordoba Presidio, Vera Cruz

. . . . . .

Sall6, TYPE Gadow

17. Mimohyla inornata, Boulenger Microhyla inornata, Boulenger, 1890, Proc. Zool. Soc. London, p. 37 (Type locality Deli, Sumatra): idem, 1893, Ann. Mus. Genova, (2), x m , p. 342 ; Boettger, 1893, Zool. Anz., p. 430 ; Flower, 1899, Proc. Zool. Soc. London, p. 905 ;Laidlaw, 1900, Proc. Zool. Soc. London, p. 887 ;Werner,

MICROHYLA 1900, Zool. Jahrb., Syst., x m , p. 502 ; Butler, 1904, Journ. Bombay Nat. Hist. Soc., xv,p. 388 ; van Kampen, 1905, Zool. Jahrb., Syst., xn, p. 713 ; Boulenger, 1912, Fauna Malay Penins., Rept. and Batr., p. 259 ; Smith, 1916, Journ. Nat. Hist. Soc. Siam, II, p. 109; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 153; Smith, 1924, Rec. Ind. Mus., xxvr, p. 141 ; Nieden, 1926, Das Tierreich, Anura n, p. 33; Sambon, 1928, Ann. Trop. Med. Parasit., m, p. 85 ; Smith, 1930, Bull. Raffles Mus., m, p. 129. ? Microhyla steinegeri, Boulenger, 1909, Ann. Mag. Nat. Hist., (8), IV, p. 494 (Tgpe locality Kanshirei, Formosa) ; Stejneger, 1910, Proc. U.S. Nat. Mus., xxxvm, pp. 92, 95 ; Gee and Boring, 1929, Peking, Soc. Nat. Hist. Bull., N, 2, p. 27. M. stejnegeri, Nieden, 1926, Das Tierreich, Anura n, p. 935 ;Okada, 1931, Tailless Batr. Jap. Emp., p. 74.

Habit moderately stout. Snout blunt, slightly prominent, as long as, or slightly shorter than, the diameter of the eye ; canthus rostralis rounded; loreal region vertical ; interorbital space about once and a third the width of the upper eyelid ; tympanum hidden. Digits without discs, or clefts above. Fingers ending obtusely, the first shorter than the second, which is slightly shorter than the fourth. Toes free, swollen distally, the fifth shorter than the third ; subarticular tubercles strongly developed ; a rounded, prominent, inner metatarsal tubercle, but the outer absent or minute. Tibio-tarsal art~culationreaching to between the shoulder and the eye. Skin smooth, or slightly shagreened above ; a feeble fold from the posterior corner of the eye t o the fore-limb ; smooth beneath ; the anal region feebly granular. Purplish grey to light brown above, immaculate or, usually, spotted with darker brown. The spots may be smaller and scattered or larger, irregular or arranged in longitudinal series ; sometimes the spots run together t o form an irregular reticulum ; usually a dark streak from the tip of the snout, through the eye and along the flanks, is more or less indicated ; upper lip with a light line or series of light spots ; limbs spotted like the dorsum, sometimes pinkish. Lower surfaces white, more or less spotted or marbled with brown. Male with a vocal sac and infuscate gular region. Length : 30 mm. DISTRIRUTION : Formosa (?), Burma, Siam, Cochin-China, Malay Peninsula and Sumatra. Tadpole almost indistinguishable from that of M. ornata.

B.M. 1909.10.29. 92-96

88,

99 ?

? d? 8 , $? $? d

cleared

d?

?

8 d d?

Kanshirei, Formosa

.

. Sauter,

COTYPES of M. steinegeri, Boulenger Palon, Pegu, Burma . . Fea Ban Yan, Me Wang, N. Siam . . M. Smith Paknampo, Siam . . . M. Smith Lat Bua Kao, Siam . . . M. Smith Klong Bang Lai, Siam . . M. Smith Chantaboon, Siam . . Flower . . . . . M. Smith Bangkok Nong Khor, S.E. Siam . . M. Smith (taken in copula, Sept. 1922) Nong Khor, S.E. Siam . . M. Smith Saigon . . . . . (Lataste) Khao Koo, near ~ a k o n Sritamarat, Penin. Siam . M. Smith . . . . . .Werner Perak Deli, Sumatra . . . . Moesch, COTYPES

MONOGRAPH O F THE MICROHYLIDAE I8.

Microhyla aequatorialis (Peracca)

Engistom oquatoriole, Peracca, 1904, Bol. Mus. Torino, XIx, 465, p. 23 (Type locality Cuenca, Ecuador). Engyatoma aequatoridia, Nieden, 1926, Das Tierreich, Anura 11, p. 67. Gaatrophryne aquatorialia, Stejneger, 1910, Proc. Biol. Soc. Waehington, xxm, p. 166.

Snout subacuminate, slightly prominent, about once and a quarter as long as the diameter of the eye ; eye a little longer than its distance from the nostril ; interorbital space broader than the upper eyelid. Fingers rounded terminally, the first a little shorter than the second. Toes not dilated, with a rudiment of web ; subarticular tubercles well developed ; an oval inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder in males, but not quite so far in females. Skin smooth in females, but more or less rough and with a glandular appearance in males ; usually a transverse fold behind the upper eyelids descending to the shoulders. Upper surfaces covered with small grey and light brown dots, together with some lighter grey spots and larger, irregular, anastomosing, dark brown spots ; a dark brown, above light-margined, lateral streak from the nostril, through the eye above the shoulder to the groin, where it merges with the anastomosing dorsal spots. Lower surfaces with small, ramifying, dark brown flecks which anastonlose to a greater or less extent, especially towards the anus. Male with a vocal sac. Length from snout to vent : 37 mm. DISTRIBUTION : Cuenca, Ecuador.

19. Microhyla pictiventris (Cope) Engy8toma pictivenire, Cope, 1886, Proc. Amer. Phil. Soc., 1885, p. 272 (Type locality Nicaragua) ; Giinther, 1900, Biol. Centr. Amer., Rept. and Batr., p. 209. Engyatomu pictiventria, Nieden, 1926, Das Tierreich, Anura n, p. 66. Q a s l r o p h r y ~pictiventria, Stejneger, 1910, Proc. Biol. Soc. Washington, xxm, p. 166. Snout long, prominent, twice as long as the diameter of the eye ; nostrils much

nearer the end of the snout than the eye. First finger shorter than the second. Toes free, the first very short ; third1 toe much longer than the fifth ; an inner, but no outer, metatarsal tubercle. Tarso-metatarsal articulation reaching the end of the snout. Skin smooth above and beneath ; no transverse occipital fold. Olive brown above ; a black band, narrowly light-edged above, from the tip of the snout to the groin ; no inguinal spot ; black beneath, with large white spots on the abdomen, smaller ones on the hind-limbs, and still smaller ones on the t?roat and chest. Length : 22.5 mm. ~ ~ S T F L I B U T I O N : Nicaragua.

20.

Microhyla carolinensis (Holbrook)

Engystoma carolinenae, Holbrook, 1836, N. Amer. Herpet., ed. 1, I, p. 83, P1. 11 (Type locality Charleston, S. Carolina) ; Dum. and Bibr., 1841, Erpet. GBn., v n ~p. , 743 ; Holbrook, 1842, N. Amer. Herp., ed. 2, v, p. 23, P1. VI; Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 51 ; Brocchi, 1882, Miss. Sci. Mex., Batr., p. 93 (part) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 162 ; Cope, 1889, BuU. U.S. Nat. Mus., xxxrv, p. 385, fig. 97 ; Boulenger, 1891, Ann. Mag. Nat. Hist., (6), vIn, p. 453; Garman, 1892, Bull. Illinois Lab., 111, p. 331 ; Dickerson, 1906, Frog Book, New York, p. 166, figs. 189-192, col. P1. n, fig. 11; Strecker, 1908, Proc. Biol. Soc. Washington, XXI, p. 57, P1. I ; idem, 1909, op. cit., xxn, p. 116 ; E. carolinensis, Nieden, 1926, Das Tierreich, Anura 11, p. 64. amtrophryne caroliwnsia, Stejneger, 1910, Proc. Biol. Soc. Washington, XXIII, p. 166; Stejneger and Barbour, 1917, Check List N. Amer. Amph. and Rept., ed. 1, p. 4 0 ; Iidem 1923, op.cit. 2, p. 3 8 ; Wright and Wright, 1923, Anat. Rec., xxm, , 336, 337, P1. I, fig. 24 ; Strecker and Bkierson, p. 406 ; Iidem, 1924, Amer. Nat., ~ v m pp. 1 The original description mys " second," but this is probably an error.

MICROHYLA 1926, Contr. Baylor Univ. Mua., v, p. 5 ; Strecker, 1926, Contr. Baylor Univ. Mus., ~ I I ,p. 5 ; Wright, 1932, Frogs of Okehokee Swamp, p. 452 ; (7. carolinense, Strecker, 1915, Baylor Bull., xvm, 4, p. 46. ? Engystoma rugosum, Durn. and Bibr., 1841, Erpet. Gbn., vm,p. 744 (Type locality Southern part8 of N. America) ; Brocchi, 1882, Miss. Sci. Mex., Batr., p. 94. Gastrophryne rugoeum, Fitzinger, 1843, Syst. Rept., p. 33. Engystoma olivaceum, Hallowell, 1856, Proc. Acad. Philadelphia, p. 252 (Type locality not cited). Engystoma, Byder, 1891, Amer. Nat., p. 838. Habit stout. Snout pointed, prominent, once and a quarter as long as the eye ; eye a little longer than its distance from the nostril ; canthus rostralis not distinct ; loreal region oblique, not concave ; interorbital space once and a half to once and

three-quarters as broad as the upper eyelid, tympanum hidden. digit^ not dilated, without median cleft above ; first finger much shorter than the second, which is shorter than the fourth. Toes free ; subarticular tubercles well developed ; a prominent inner, b l ~ no t outer, metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder or a little beyond. Skin smooth above, or slightly pustular on the posterior portion of the body ; smooth beneath ; a dermal fold across the head behind the upper eyelids, descending to the angles of the lower jaw. Black, brqwn or grey above, uniform or with more or less distinct traces of the following colour pattern : A dark lirteral streak from the tip of the snout to the sides of the body; a dark dorsal blotch commencingbetween the eyes and gradually widening posteriorly, its edges reaching the groins ; an oblique, broad, dark bar across the femur, sometimes divided into two. Lower surfaces yellowish, mottled and stippled with brown, or brown with yellowish stippling. Male with a vocal sac and infuscate throat. Length : 38 mm. DISTRIBUTION : Virginia to Florida, westwards to Texas and northwards to Indiana. Eggs laid May to August in strings coiled to form large floating masses ; larvae hatch in about 3 days ; metamorphosis reached in 20 to 60 days from oviposition ; full-grown tadpoles measure about 26 to 37 mm. ; newly metamorphosed frogs 8.5 to 12 mm. The mature tadpole is characterised as follows : Body 1.2to 1.6 times as long as broad ; tail 1-4to 2-8times the length of the body, 2.2 to 5.4 times as long as deep. Head and body broad, depressed ; snout somewhat truncate ; nostrils close to the mouth, which is terminal, without mandibles or teeth ; a distinct canthus rostralis ; eyes lateral. Spiraculum median, close in front of the anus. Tail bluntly rounded terminally with subequal crests, which are not as deep as the muscular portion. Width of mouth contained 1.6 to 2.3 times in the interorbital distance. Black above, finely stippled with lighter ; lower surfaces grey with a yellow transverse stripe, a light stripe on each side of the belly, and another in the gill region, or regularly spotted with yellow ; muscular region of tail black with a light stripe or series of dots ; tip of tail blackish (after Wright, 1932). SPECIMENS EXAMINED N. America . B.M. R.1931.2.13. 1 6 45.11.9.32 6 New Orleans. . . 45.11.9. 33-34 99 New Orleans. . . 87.5.14. 54-55 69 Pensacola, Florida . 88.2.13.64-66 6 9 yg. Key West, Florida . 1920.1.20. 3161 9 Florida . . . . 1925.11.13.4 cleared Florida . . . . 2I .

.

. Doublerlay

. . . . . . . . . .

. Bollman . . . . . Gamier . . . . . (Lataste)

. (M.C.Z.)

Microhyla areolata (Strecker)

Engystoma areokzta, Strecker, 1909, Proc. Biol. Soc. Washington, xxn, p. 118 (Type locality Guadalupe River Bottom, Victoria, Texas) ; idem, 1910, op. cit., m,p. 117, P1. n, fig. 1 ; Nieden, 1926, Dm Tiemich, Anura rr, p. 65. Gmtrophryne arwlata, Stajneger, 1910, Roc. Biol. Soc.

'148

MONOGRAPH O F T H E MICROHYLIDAE

Washington, X ~ I p., 166 ;Strecker, 1915, Baylor Bull., xvnI,4, p. 47. Stejneger and Barbour, 1917, Check List N. Amer. Amph. and Rept., ed. 1, p. 40 ; Iidem, 1923, op. cit., ed. 2, p. 37. Similar t o M. carolinensis, hut the snout shorter ; skin areolated or pustular ; hind-limbs and feet shorter ; metatarsal tubercle larger. Light or dark grey above, with heavy dark marblings or blotches ; sometimes a

dark line from eye t o eye around the tip of the snout, and a small, dark, broadly V-shaped spot between the upper eyelids. Light grey beneath, closely spotted with lighter. Length : 22 mm. DISTRIBUTION : S.E. Texas. If this species is really distinct from M. tecensis, perhaps the name M. rugosum (Dum. and Bibr.), a t present placed as a synonym of M. carolinensis, ought to be used for it. 22.

Microhyla texensis (Girard)

Engystoma tezense, Girard, 1859, Proc. Acad. Philadelphia, p. 169 (Type locality Rio Seco, Texas) ; Dickerson, 1906, Frog Book, New York, p. 168, fig. 193; Strecker, 1908, Proc. Biol. Soc. Washington, XXI, p. 5 7 ; idem, 1909, op. cit., XXII, p. 117. Gastrophryne texana, Stejneger, 1910, Proc. Biol. Soc. Washington, xxm, p. 166. E. texensis, Nieden, 1926, Daa Tierreich, Anura n, p. 65. Gastrophryne texensis, Strecker, 1915, Baylor Bull., xvm, 4, p. 47 ; Stejneger and Barbour, 1917, Check List N. Amer. Amph. and Rept., ed. 1, p. 40 ;Iidem, 1923, op. cit., ed. 2, p. 38 ; Strecker, 1926, Contr. Baylor Univ. Mus., v n ~ (I), p. 12 ; Strecker and Williams, 1927, Contr. Baylor Univ. Mus., XII, p. 8 ; Allen, 1933, Occ. Papers. Mus. Zool. Michigan, CCLIX, p. 3. Engystoma carolinense (part), Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 162 ; Cope, 1889, Bull. U.S. Nat. Mus., xxx~v, p. 385. Habit stout. Snout rounded, prominent, once and a half as long as the eye ; eye as long as its distance from the nostril ; canthus rostralis not distinct ; loreal region oblique, not concave ; interorbital space about once and three-quarters as broad as the upper eyelid ; tympanum hidden. Digits not dilated and without median cleft

above. Fingers moderate, the first much shorter than the second, which is a little shorter than the fourth. Toes free ; subarticular tubercles well developed ; a distinct inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the region of the shoulder. Skin smooth (or slightly pustular in breeding males) above and beneath ; a feeble transverse fold across the head behind the upper eyelids, descending t o the angles of the lower jaw. Grey, greenish or light brown above, uniform or with some darker markings in the form of scattered, irregular spots tending to be arranged in linear seriew femur and tibia sometimes with an ill-defined cross-bar or two narrow ones. Lower surfaces uniformly whitish, except the throat, which may be faintly mottled with brown. Male with a vocal sac ; walls of the sac dark brown or black, showing by transparency through the skin beneath the throat. Digits of males flattened a t the breeding season, with feeble fleshy lateral fringes, especially on the inner sides of the second and third fingers. Length : 29 mm. : Texas, N. Mexico. DISTRIBUTION SPECIMENS EXAMINED 3 3,$2 $2 Duval Co., Texas . . . . . . Taylor B.M. 87.1.4.41 92.10.29. 4 5 4 8 1931.12.18. 7-8 33 2 miles S. of Dallas, Texas . . Burt

)

.

23.

Microhyla usta (Cope)

Engystoma carolinense (part), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 51 ; Brocchi, 1882, Miss. Sci. Mex., Batr., p. 94.

MICROHYLA

149

Engystoma rugosum (non Durn. and Bibr.), Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 52. Engystoma ustum, Cope, 1866, Proc. Acad. Philadelphia, p. 131 (Type locality Guadalajara, Jalisco, Mexico) ; idem, 1871, op. cit., p. 216 ; Brocchi, 1877, Bull. Soc. Philom. Paris, (7), I, p. 193; idem, 1882, Miss. Sci. RIex., Batr., p. 93, PI. x, fig. 1 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 161 ; Giinther, 1902, Biol. Centr. Amer., Rept. and Batr., p. 209. Systoma ustum, Cope, 1867, Journ. Acad. Philadelphia, (2), VI,p. 194. Gastrophryne usta, Stejneger, 1910, Proc. Biol. Soc. Washington, x x m , p. 166. Engystoma usta, Nieden, 1926, Das Tierreich, , 187. Anura 11, p. 65 ; Kellogg, 1932, Bull. U.S. Nat. Mus., c ~ xp. Engystoma mexicanurn, Peters, 1869, Mon. Ak.Berlin, p. 881 (Type locality State of Puebla, Mexico). Eupemphix gadovii, Boulenger, 1903, Ann. Mag. Nat. Hist., (7), XII, p. 552 (Type locality San Mateo del Mar, Tehuantepec).

Habit stout. Snout pointed, prominent, once and a quarter as long as the eye ; eye a little longer than its dist,ance from the nostril ; canthus rostralis rounded ; loreal region slightly oblique, not concave ; interorbital space nearly twice as broad as the upper eyelid ; tympanum hidden, digits not dilated and without median cleft above. Fingers moderate, the first much shorter than the second, which is a little shorter than the fourth. Toes free, subarticular tubercles well developed ; two well-developed, compressed, metatarsal tubercles. Tibio-tarsal articulation reaching the shoulder or the tympanic region. Skin smooth or slightly shagreened above ; tarsus sometimes slightly pustular. Smooth beneath, the anal region slightly granular ; a transverse dernial fold on the head behind the upper eyelids, descending to the angles of the lower jaw. Brown or greyish above, uniform or with more or less distinct traces of the following colour pattern : A dark lateral band from the eye, above the forearm, along the flanks ; a dark mid-dorsal area, commencing on the snout, with a transverse branch to each upper eyelid, and gradually widening caudad ; a branch from this area arises in the sacral region, crosses the groin, and is continued as a broad diagonal bar on the femur and tarsus. A fine white line from snout to vent, meeting another which runs from heel to heel along the hinder side of the legs, may be present. Yellowish-white beneath, stippled with pale brown or with faint darker reticulations. Male with a vocal sac ; throat dark brown or black ; edge of lower jaw with a few spinules. Length : 30 mrn. DISTRIBUTION : Mexico, as far south as Tehuantepec.

Q

3f. 57.10.28. 38 60.6.17. 26 47.12.28. 43 83.4.5. 1-2 56.3.17. 30-31 1903.9.30.259-261

hgr. $ $! hgr. juvs.

R. 1931.2.13. 2-3

8$?

Mexico . . . . . . . . Sall6 Mexico. . . . . . . . Mexico . . . , . . . . Presidio de Mazatlan, W. Mexico . Forrer Cordova . . . . . . . . Salle S. Mateo del Mar, Tehuantepec . Gadow, TYPES of Eupernphix gadovii, Boulenger Tehuantepec . . . , . . (Brussels Mus.)

24. Microhyla schirchi

.

(Mir.Ribeiro)

Dasypops schirchi, Mir. Ribeiro, 1924, Bol. Mus. Nac. Rio de Janeiro, IV, p. 266 (Type localit,y Rio Rlutum, Espirito Santo, Brazil); idem, 1926, Arch. Mus. Nac. Rio de Janeiro, x x v ~pp. , 185 and 225, PI. XXI (col.). Habit stout. Snout long, prominent, obliquely truncate, 24 times as long as the diameter of the eye ; canthus rostralis rounded ; loreal region oblique ; nostril nearer the tip of the snout than the eye ; tympanum almost hidden. Fingers short, depressed; not dilated distally, with rudimentary lateral fringes ; first, second and fourth subequal,

MONOGRAPH O F THE MICROHYLIDAE toes short, bufoniform, without discs, and with a rudiment of web ; subartic~llar tubercles feebly developed ; an inner, but no outer, metatarsal tubercle. Skin smooth on the body, with feebly marked mid-dorsal and post-femoral furrows ; head rugose, the skin adherent to the skull ; a transverse occipital groove. Dark yellow above, with two series of darker spots meeting on the nape and diverging behind ; a blotch on the temples and shoulders, and two smaller ones on the anterior part of the flanks ; loins marbled. Head marbled with dark brown, wit11 a darker loreal blotch. Lower surfaces pale yellow. Limbs cross-barred. Length : circa 39 mm. DISTRIBUTION : Rio Blutum, Espirito Santo, Brazil.

25.

Microhyla subnigra (Mir . Ribeiro)

Engystoma subnigrun~,Mir. Ribeiro, 1920, Rev. Mus. Paulista, xIr, p. 285, P1. (Type locality Serra pp. 183 de MacahB, State of Rio, Brazil) ; idem, 1926, drch. Nus. Nac. Rio de Janeiro, XXVII, and 225. Habit stout. Snout very long, pointed, strongly projecting, slightly hooked, and 2 or 3 times as long as the eye ; canthus rostralis rounded ; loreal region nearly vertical, very slightly concave ; nostril t ~ i i c eas far from the eye as from the tip of the snout ; interorbital space 3 times as broad as the upper eyelid ; tympanum hidden. Digits not dilated, without median cleft above. Fingers short, depressed, not dilated ; first much shorter than the second, which is subequal to, or a little shorter than, the fourth ; second and third (at least in males) with a fleshy fringe along their inner margins. Toes short, free ; subarticular tubercles feebly developed ; a very indistinct rudiment of an inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder. Skin smooth above and beneath; a transverse occipital fold descending to the angles of the mouth, and a tympanic fold from the eye to the fore-limb. Dark brown to black above ; lower surfaces brown, regularly stippled with yellow ; hind-limbs sometimes with transverse black bands. Male with a vocal sac. Length : 37 mm. DISTRIBUTION : State of Rio de .Janeiro, Brazil. SPECIMENS EXAMINED Mus.Vienna Petropolis,Riode Janeiro . . . . .Mehaelis

. .

Microhy la microps (Durn. and Bibr .) Eqystomn microps, Dum. and Bibr., 1841. Erpet. GBn., VIIT, p. 744 (Type locality Brazil) ; Giinther, 1858, Cat. Batr. Sal. Brit. illus., p. 52 ; Steindachner, 1864, Verh. Zoo].-bot. Ges. Wien, p. 285, PI. xv, fig. 3 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 163 ; Baumann, 1912, Zool. Jahrb., Syst., XXXIII, (2), pp. 122, 148, 161 ; Nieden, 1926, Das Tierreich, Anura 11, p. 67. Gastrophryne microps, Stejneger, 1910, Proc. Biol. Soc. Washington, XXIXI, p. 166 ; Crawford, 1931, Ann. Carnegie RIus., xsr, (l),p. 38. Engystoma dumerili, Mir. Ribeiro, 1926, Arch. Mus. Nac. Rio de Janeiro, XXVII, pp. 183, 82.5 (substitute name for E. microps, Dum. and Bibr.). Habit stout. Dermal ridges of the roof of the mouth slightly crenulate. Snout acutely pointed, very prominent, slightly hooked in profile, 3 times as long as the eye in adults, but scarcely twice as long in the young ; canthus rostralis rounded ; loreal region nearly vertical, distinctly concave ; nostril nearly twice as far from the eye as from the tip of the snout ; interorbital space more than twice as broad as the upper eyelid ; tympanum hidden. Fingers short, depressed, tapering, without discs ; first shorter than the second, which is subequal to, or shorter than, the fourth. Toes free, the four outer with small terminal dilatations, which are but little wider than the penultimate phalanges, but have a horseshoe-shaped groove separating their upper and lower surfaces, though without any trace of a median cleft above ; inner digit very short, without a disc ; subarticular tubercles flat, indistinct ; an indistinct,

rudimentary inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching to between the shoulder and the tympanic region. Skin smooth above and beneath ;a very feeble fold connecting the posterior corners of the upper eyelids and descending to the angles of the lower jaw. Pale brown above, uniformly flecked with yellow ; sometimes a light streak along the canthus rostralis and the edge of the upper eyelid, continuing straight backwards almost to the level of the fore-limb. Yellow beneath, the throat darker, with a few yellow flecks. Male without a vocal sac. Length : 32 mm. DISTI~IBUTION : Guianas, Brazil. SPECIRIENS EXAMINED

B.M. 72.10.16. 46-49 1'305.11.l. 31-33

J5,92

$9,yg

Demerara Falls, British Guiana. Tung District., British Giuana, 3,000 ft. Bovallius

SPHENOPHRYNINAE, Noble Eleutherognathinae (part),RI6hely, 1901, Termds. Fiizetek, x x ~ vp. , 189. Sphenophryninae and Cacopinae (part), Noble, 1931, Biology Amph., pp. 551 and 532. Maxillary teeth absent, or rarely present as minute vestiges ; vomerine teeth absent. Maxillae not overlapping the premaxillae ; prevomer large, entire, almost surrounding the choana, with a mesial wing-like expansion which is in contact with its fellow ; post-choanal portion of the vomer sometimes bearing small odontoids ; ethmoid divided ; ear fully developed; vertebral column procoelous. No free-swimming tadpole, the metamorphosis taking place within the egg ; the larva lacks external gills, has no open gill clefts, internal gills, operculum, spiracle, or teeth ; respiration is performed through a non-muscular, vascular tail. This subfamily appears to be the procoelous, eleutherognathine analogue of the Asterophryinae, and a further differential character is provided by the tongue. A large subcircular tongue, entirely adherent and often with a median furrow and posterior pouch, has been claimed to be a variant which has been evolved many times. But this type of tongue is constantly present throughout the Dyscophinae and the Asterophryinae, and persists in some of the more primitive members of the Microhylinae. All members of the Sphenophryninae, like the majority of those of another procoelous subfamily, the Cophylinae, have an oval tongue which is half free behind and without any trace of median furrow or pouch. SYNOPSIS OF THE GENERA I. Clavicles and procoracoids present. A. Clavicle extending from the mid-line of the girdle to the scapula Sphenophrynt., p. 151 R. Clavicle not reaching the scapula . . . Oreophryne, p. 159 11. Procoracoids, but no clavicles, present . . . . iicrobatrachus, p. 170 111. No clavicles or procoracoids. A. Sternum present . . . . . . . . . . . Cophizalus, p. 170 B. Sternum vestigial . . . . . . , . . . .Aphnntophryne. p. 178

.

. .

. .

SPHENOPHR YNE, Peters and Doria Cornufer (part),Tschudi, 1838, Riem. Soc. Sci. Neuchiitel, TI, 1839, pp. 28, 71. Sphenophryne, Peters and Doria, 1878, ,4nn. Mus. Genova, XIII,p. 430 (Type species Sphenophryne cornuta, Peters and Doria) ; Boulenger, 1882, Cat. Batr. Sal. Brit. RIus., ed. 2, p. 167 ; van Kampen, 1919, Bijdr. Dierk,, xxx, p. 53 ; idem, 1923, Amph. Indo-Austr. Arch., p. 105 f art) ; Nieden, 1926, Das Tierreich, Anura TI, p. 43 (part) ; Noble, 1931, Biol. Amph., p. 531.

152

MONOGRAPH OF THE MICRQHYLIDAE

Liophryne, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), XIX, p. 11 (Type species Liophryne r h b dactyla, Boulenger) ; Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, pp. 429,432 ;van Kampen, 1923, Amph. 1ndo.-Austr. Arch., p. 100 (part) ;Nieden, 1926, Das Tierreich, Anura 11,p. 50 (part) ; Noble, 1931, Biol. Amph., p. 531. Chuperim (non Mocquard), MBhely, 1901, TermBs. Ezetek, XXIV, pp. 206,256 ; Vogt, 1911, Sitzber. Ges. Nat. Fr. Berlin, pp. 429, 432. Austrochqerim, Fry, 1912, Rec. Austr. Mus., IX, 1, p. 87 (Type species Austrochaperim robusta, Fry). Ozydactyla, van Kampen, 1913, Nova Guinea, IX,Zool., 3, p. 464 (Type species Ozydactyla brevicrus, van Kampen) ; idem, 1923, Amph. Indo-Austr. Arch., p. 122 ; Nieden, 1926, Das Tierreich, Anura n, p. 49 ;Noble, 1931, Biol. Amph., p. 531. Hylophorbus ? (mMacleay), Smith, 1927, Proc. Zool. Soc. London, p. 215.

Vestiges of maxillary teeth rarely present. Prevomer large, undivided, in contact with its fellow mesially (fig. 53) ; the post-choanal portion overlying the palatine region and bearing a more or less strongly raised ridge on which a few small odontoids may be present. Clavicles and procoracoids present, well developed, reaching the mid-line of the shoulder girdle and the scapulae ; no omosternum ; sternum cartilaginous (fig. 54). Vertebral column procoelous. Terminal phalanges usually T-shaped.

Fig. 53. Anterior cranial elements of Sphenophryne cornuta (seen from beneath)

Fig. 54. Ventral elements of the shoulder-girdle of Sphenophryne cornuta

SPHENOPHRYNE Pupil horizontal. Tongue oval, entire and half free behind. A denticulated dermal ridge across the palate in front of the pharynx, sonletimes preceded by a shorter smooth one. Digits free, with, or without, terminal dilatations. SYNOPSIS OF THE SPECIES I. Snout longer than the eye, or, if only as long as the eye, the canthus rostralis angular and the loreal region nearly vertical. A. Discs of fingers larger than those of the toes ; upper eyelid with a S. cornuta, p. 153 spine-like tubercle . . . . . . . . . B. Discs of fingers smaller than those of the toes ; no supraciliary spines. ( 1 ) Canthus rostralis angular ; tibio-tarsal articulation reaching to between the eye and the tip of the snout S. schlaginhaufeni, p. 154 ( 2 ) Canthus rostralis rounded ; tibio-tarsal articulation reaching . . . . . . . . . . . S. gracilipes, p. 1 55 the eye 11. Snout not longer than the eye ; canthus rostralis rounded. A. Digital disc of third finger at least once and a half as broad as the . S. macrorhyncha, p. 155 penultimate phalanx B. Digital disc of third finger scarcely broader than the penultimate phalanx. ( 1 ) Tibio-tarsal articulation reaching the eye. ( a ) Tibia equal to about half the distance from snout to vent. Fingers moderately long, the fourth as long as, or longer than, the internarial space. Length from snout to vent circa 48 mm.. . . . . . S. rhododactyla, p. 156 ( b ) Tibia equal to about half the distance from snout to vent. Fingers short, the internarial space half as long again as the fourth. Length from snout to vent circa 28 mm. S. mehelyi, p. 156 ( c ) Tibia distinctly less than half the distance from snout to vent. Fingers very short, the internarial space twice as long as Ihe fourth. Length from snout to vent drca 28 mm. . . . . . . . . . . . S. robusta, p. 157 ( 2 ) Tibio-tarsal articulation reaching the shoulder . .S. polysticta, p. 157 C. Fingers without discs. ( 1 ) Toes slightly dilated ; terminal phalanges T-shaped S. brevipes, p. 158 ( 2 ) Toes pointed ;terminal phalanges club-shaped . . S brevicrus, p. 158 I.

Sphenophrym cornuta, Peters and Doria

Cornufer unicolor (part), Tschudi, 1838, Mem. Soc. Sci. Neucbltel, n. 1839, pp. 28, 71. Sphenophryne co~nutcs,Peters and Doria, 1878, Ann. Mus. Genova, XIII, p. 430, PI. VII, fig. 4 (Type locality near the Wa Samson River, N. New Guinea) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 157 ;idem, 1898, Ann. Mus. Genova, (2), xvm, p. 707 ;Wandolleck, 1911, Abh. Mus. Dresden, XIII,6, p. 4, figs. 1-9 ; Boulenger, 1914, Trans. Zool. Soc. London, xx, 5, p. 251 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 105, fig. 13; Nieden, 1926, Dm Tierreich Anura 11, p. 47, fig. 33. Chaperina ceratophthalmufi,van Kampen, 1009, Nova Guinea, IX,Zool., 1, p. 43, PI. m, fig. 8 (Type localities Noord River, near Geitenkamp and Resi Peak, Dutch New Guinea); idem, 1913 op. cit., IX,Zool., 3, p. 464 ; idem, 1914, Zool. Jahrb., Syst., x x x w , p. 378 ; Idem, 1915, Nova, Guinea, xm, Zool., 1, p. 41.

Minute vestiges of maxillary and rem maxillary teeth sometimes present. Denticulated ridge in front of the pharynx preceded by a median swelling. Snout prominent, once and a third as long as the eye ; canthus rostralis angular ; loreal region vertical, not concave ;nostril much nearer the tip of the snout than the eye ; interorbital space much broader than the upper eyelid ; tympanum distinct, 4 to $ the diameter of the eye. First finger very short, with a small disc ; remaining fingers with large discs ;

MONOGRAPH OF TIIE MICROHYLIDAE second a little shorter than the fifth. Toes with discs similar to, but smaller than, those of the fingers ; third longer than the fifth ; subarticular tubercles indistinct ; an illdefined elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching to between the eye and the end of the snout. Skin smooth or very slightly granular above ; usually a spine-like tubercle on each eyelid and a smaller one on each heel. Smooth beneath ; sometimes a row of small tubercles along the outer side of the tibia and forearm. Greyish brown above, with indistinct darker and lighter spots or reticulations ; greyish white beneath, with scattered white spots and brown patches. A fine, white mid-dorsal line from snout to vent, continued along the hinder side of the thighs and the outer edge of the tibiae ; a similar white line on the mid-line of the belly, intersected by a transverse one across the chest and on the outer side of the arm. ,Male with a vocal sac. Ovarian eggs large, 2.5 mm. in diameter. Length : 41 mm. DISTRIBWION : New Guinea. SPECIMENS EXAMINED $' Foot of Charles Louis Mts., B.M. 1897.3.23. 5 New Guinea . . . . v. Renesse 1913.11.1. 136-139 3 8 QQ Launch Camp, Setekwa River, Dutch New Guinea Wollaston Exp.

. . . .

2.

Sphenophryne schlaginhaufeni,Wandolleck

Sphenophryne schlaginhaufeni, Wandolleck, 1911, Abh. Mus. Dresden, XIII, 6, p. 5, figs. 10-17 (Type locality " Oberlauf des Rienjamur, 650-700 m.," Toricelli Mts.). Sphenophryne kloaai, Boulenger, 1914, Trans. 2001. Soc. London, xx, 5, p. 251, PI. XXVII, figs. 3-3b (Type locality Launch Camp, Setekwa River, Dutch Now Guinea) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 107 ; Nieden, 1926, Das Tierreich, Anura 11, p. 46 ; Mertens, 1930, Zool. Mededeel Leiden, XIII,p. 145.

Denticulated ridge in front of the pharynx, preceded by a median swelling. Snout slightly prominent, as long as, or a little longer than, t h e diameter of the eye ; canthus rostralis angular : loreal region slightly oblique, concave ; nostril nearer the tip of the snout than the eye ; interorbital space broader than the upper eyelid ; tympanum distinct, Q the diameter of the eye. Fingers with small discs, the first shorter than the second, which is as long as, or a little longer than, the fourth. Toes with small discs, the third longer than the fifth ; subarticular tubercles indistinct ;an indefinite elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching to between the eye and the tip of the snout. Skin smooth or very slightly tubercular above and on the flanks ; straight fold from the posterior corner of the eye above the tympanum t o the flank; a slight ) (-shaped fold between the shoulders. Lower surfaces smooth. Yellowish brown above ; sides of head and tympanic region below the glandular fold, dark brown : indications of darker bars between the eyes, between the ) (-shaped folds and across the sacral region ; a black, white-tipped wart on either side of the coccyx. Anal region, anterior aspect of the knee and lower surfaces of the tarsus and foot, black. Limbs with narrow, dark, diagonal cross-bars. White beneath, the throat and limbs with a brown wash. Male with a vocal sac. Length : 42 mm. DISTRIBUTION : New Guinea. SPECIMENS EXAMINED B.M. 1913.11.1. 140-141 8 8 Launch Camp, Setekwa Wollaston Exp., River, Dutch New Guinea COTYPESof S. klossi, Boulenger. Mus. Dresden ad. Rienjamur, 650-700 metres. Schlaginhaufens, Toricelli Mts. COTYPE

SPHENOPHRYNE

3. Sphenophryne gracilipes (Fry) Austrochaperina gracilipes, F r y , 1912, Rec. Austr. Mus., IX,1, p. 93, fig. 39, PI. vIn, figs. 1-lb (Type locality Somerset, Cape York, N. Australia) ; Nieden, 1926, Das Tierreich, Anura 11,p. 48.

Denticulated ridge in front of the pharynx preceded by a median swelling. Snout depressed, prominent, longer than the eye ; canthus rostralis rounded ; loreal region oblique, concave ; nostril slightly nearer the tip of the snout than the eye ; interorbital space much broader than the upper eyelid ; tympanum hidden. Fingers slender, with scarcely enlarged discs, the first very short. Toes with very small discs, the fifth a little shorter than the third ; no metatarsal tubercles. Tibio-tarsal articulation reaching the anterior border of the eye. Skin smooth above and below. Pinkish brown above, with sparsely distributed dark brown spots on the legs and sides ;an indistinct inguinal spot present ;a dark brown streak from the nostril through the eye to the shoulder. Lower surfaces pale brown, with faint brown reticulations and spots. Length : 1 7 mm. DISTRIB$TION

: Cape York Peninsula.

4. Sphenophryne mamorhyncha (van Kampen) Chaperina mmrorhyncha, van Kampen, 1906, Nova Guinea, v, p. 168, fig. 3 (Type looality Manikion District, Dutch New Guinea); idem, 1909, op. cit., rx, Zool., 1, p. 43. Sphenophryne nuzerorhyncha, van Kampen. 1923, Amph. Indo-Austr. Arch., p. 107 ; Nieden, 1926, Das Tierreich, Anura n, p. 48 ; Burt, 1932, Bull. Amer. Mus. Nat. Hist., Lxm, p. 484. Chaperina bmipalmata, van Kampen, 1906, Nova Guinea, v, p. 169, figs. 4 , 5 (Type localities Tawarin and Timena River, Dutch New Guinea) ; idem, 1913, op. eit., IX,Zool., 3, p. 464 ; idem, 1914, Zool. Jahrb., Syst., xxxvn, p. 377 ; Boulenger, 1914, Trans. Zool. Soc. London, xx, 5, p. 251 ; Sphenophryne basipalmata, Nieden, 1926, Das Tierreich, Anura n, p. 46. Chaperina quatu07lobata, Wandolleck, 1911, Abh. Mus. Dresden, XIII, 6, p. 9, figs. 3 6 4 6 (Type locality Toricelli Mts.). Chaperina punctata, van Kampen, 1913, Nova Guinea, rx, Zool., 3, p. 463, P1. XI, fig. 7 (Type localities Went Mts., 800-1,050 m. and HeUwig Mts., 2,500 m.): idem, 1914, Zool. Jahrb., Syst., x x x w , p. 378.

Denticulated ridge in front of the pharynx preceded by an indistinct smooth one, or three swellings. Snout slightly prominent, rounded, as long as, or slightly shorter than, the eye ; canthus rostralis rounded ; loreal region oblique, almost flat ; nostril equidistant from the eye and the tip of the snout, or a little nearer the latter ; interorbital space as broad as the upper eyelid ; tympanum rather indistinct, $ to # the diameter of the eye. Fingers with well-developed discs; the first shorter than the second, which is a little shorter than the fourth. Toes with small discs, slightly webbed a t the base and sometimes with indistinct, fleshy lateral fringes ; subarticular tubercles indistinct ; an elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the eye. Skin smooth above and below ; a cumed supratympanic fold from the eye to the shoulder. Brown or grey above, uniform or with darker marblings ; there is often a dark streak along the loreal region, and the limbs are usually cross-barred or spotted. Lower surfaces white, the throat and limbs washed with pale brown. Ovarian eggs large, 2.5 mm. in diameter. Length : 35 mm. DISTRIBUTION : New Guinea.

MONOGRAPH O F T H E MICROHYLIDAE $2

B.M. 1928.2.10. 3 1913.10.31. 248

Mus. Dresden

$2 $2

Went Mts., 800 m.

. . . . .

COTYPEof Chaperina basipalmata, van Kampen Mirnika River, Dutch New Guinea Wollaston Exp. Toricelli Mts. . . . . . . COTYPEof Chaperina quatuorlobata, Wandolleck

5. Sphenophryne rhododactyla (Boulenger) Liophryne rhododactyla, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), XIX, p. 11, PI. 2, fig. 2 (Type locality Mt. Victoria, Owen Stanley Range, New Guinea) ; van Kampen, 1923, Amph. IndoAustr. Arch., p. 100 ;Nieden, 1926, Das Tierreich, Anura. n, p. 50. Denticulated ridge in front of the pharynx preceded by a smooth one ; vomerine

odontoids sometimes present and better developed in females than in males. Snout short, scarcely prominent, as long as the diameter of the eye ;canthus rostralis angular, curved ; loreal region oblique, rather concave ; interorbital space broader than the upper eyelid ; tympanum 3 the diameter of the eye. Fingers with small discs, the first shorter than the second, which is slightly shorter than the fourth ;latter about as long as the internarial space. Toes with small dilatations which are scarcely wider than the rest of the toe ; subarticular tubercles indistinct: an inner, but no outer, metetarsal tubercle. Tibio-tarsal articulation reaching to between the eye and the end of the snout. Skin smooth above and below ; a slight fold from the posterior corner of the eye, bifurcating above the tympanum, the upper branch continuing along the flank and the lower curving around the tympanum to the shoulder. Brown above, finely speckled with lighter; supratympanic fold darker ; a round white spot above the vent ; lower surfaces brown, with numerous light spots ; upper surfaces of fingers and toes yellow (red in life). Male with a vocal sac. Ovarian eggs 6 mm. in diameter. Length : 6 0 mm. : Owen Stanley Range, New Guinea. DISTRIBUTION B.M. 1896.10.31. 28-30

3, $2,

yg.

Mt. Victoria, Owen Stanley Range, New Guinea

v

'

Anthony, COTYPES

6. Sphenophryne nzdhelyi sp. n. Chuperina fusca (non Mocquard), MBhely, 1901, TermBs. Fiizetek, x m , pp. 207 and 257, P1. XII. fig. 3 ; Werner, 1901, Verh. Zoo1.-bot. Ges. Wien, LI, p. 613; Vogt, 1911, Sitzber. Ges. Xatf. Fr. Berlin, p. 431. Sphemphryne fusea, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 109. Type $2 : Denticulated ridge in front of the pharynx preceded by a median swelling. Snout short, rounded, as long as the diameter of the eye, canthus rostralis rounded ; loreal region oblique ; nostril nearer the tip of the snout than the eye ; interorbital space twice as wide as the upper eyelid ; tympanum distinct, rather less than half the diameter of the eye. Fingers short, with small discs ;the first shorter than the second, which is shorter than the fourth ; latter measuring about 2 the internarial distance.

Toes with larger discs than the fingers. Tibio-tarsal articulation reaching the eye. Skin smooth above and below ; a fold from the posterior corner of the eye to the shoulder. Uniformly brown above, the flanks with scattered lighter dots ;lower surfaces dirty white, the throat and chest washed with pale brown and irregularly spotted with white.

SPHENOPHRYNE

157

Length : 20 mm. ; fore-limb 10 mm. ; hind-limb 29 mm. The paratype, a male from the same locality as the type, differs only in having the lower surfaces of the limbs dusted with light brown. : Sattelberg, New Guinea. DISTRIBUTION SPECIMENS EXAMINED Mus. Hung. Amph. 2414111. Sattelberg, New Guinea 2414111

3

Sattelberg, New Guinea

Bir6, TYPE (lacking shoulder-girdle) Bir6, PARATYPE (skull and shoulder-girdle presemed separately)

7 . Sphenophryne robustsa (Fry) Austrochaperina robusta, Fry, 1912, Rec. Austr. Mus., rx, 1, p. 89, figs. 35, 35a, and 37, P1. vm, fig. 2a, P1. IX (Typelocality Russel River, N.E. Queensland). Sphenophryne robusta, Andersson, 1916, Vet. Ak. Handl. Stockholm, L I ~ ,9, p. 7 ; Nieden, 1926, Das Tierreich, Anura 11, p. 48.

Denticulated dermal ridge in front of the pharynx preceded by a curved smooth or feebly denticulate one. Snout slightly prominent, rounded, shorter than the eye; canthus rostkalis rounded ; loreal region slightly oblique, not concave ; nostril nearer the tip of the snout than the eye ; interorbital space broader than the upper eyelid ; tympanum indistinct, about half the diameter of the eye. Fingers short, with small discs ;first much shorter than the second ; fourth only about half as long as the internarial space. Toes with feeble fleshy fringes and moderate sized discs ; a small oval inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the eye. Skin smooth above and below. Uniform brown above, with or without a few brownish speckles and a fine white dorsal line from snout to vent which intersects another running from the vent to the heel ; loreal region dark brown ; a dark brown band from behind the eye, above the tympanum t o the shoulder. Lighter beneath, uniform or faintly speckled with darker brown. Male with a vocal sac. Length : 28.5 mm. DISTRIBUTION : N.E. Queensland. B.M. 1929.2.4. 1

3

SPECIMENS EXAMINED Malanda, N. Queensland

. . . .

Mjoberg

8 . Sphenophryne polysticta (Mkhely) Chaperinapolysticta, MBhely, 1901, Termgs. Fiizetek, XXW, pp. 208, 258, PI. xIr, fig. 4 (Typelocality Sattelberg, New Guinea) ; Werner, 1901, Verh. Zoo1.-bot. Ges. Wien, LI, p. 613. Sphenqhryne polysticta, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 108 ; Nieden, 1926, Das Tierreich, Anura n, p. 47. A~strocha~erina robusta (part), Fry, 1912, Rec. Austr. Mus., IX, 1, p. 89, P1. mu, figs. 2 and 2b. Austrochaperinu brevipes, Fry, 1915, Proc. R. Soc. Q~eensland~xxm~, 4, p. 61 (Typelocality Bloom-

field River, near Cooktown, Queensland). Denticulated dermal ridge in front of the pharynx preceded by a smooth one which may be reduced t o a median tubercle. Snout rounded, slightly prominent, shorter than the eye ; canthus rostralis rounded ; loreal region oblique and slightly concave ; nostril midway between the orbit and the tip of the snout or nearer the latter ; interorbital space broader than the upper eyelid ; tympanum distinct, about Q the diameter of the eye. Fingers with very small discs, that of the first smallest ; first much shorter than the second. Toes with discs not quite half as wide as the tympanum ; third longer than the fifth. Tibio-tarsal adiculation reaching the tympanum. Skin smooth above and below.

MONOGRAPH OF T H E MICROHYLIDAE Yellow or pale brown above ; back and limbs irregularly spotted with dark brown ; a dark streak from the tip of the snout along the loreal region, through the eye, above the tympanum and shoulder to the middle of the flanks. Lower surfaces yellowish, the throat, chest, and hind-limbs faintly dusted with brown. Length : 16.5 mm. DISTRIB~IO : Sattelberg, N New Guinea, and Cooktown, Queensland.

Mus Hung. Amph. 2414112.

ad.

Sattelberg, New Guinea

Bir6, TYPE(Skull, part of shoulder-girdle and humerus preserved separately.)

9 . Sphenophryne brevipes (Boulenger) Liophryne brevipes, Boulenger, 1892, Ann. Mag. Nat. Hist., (6), xrx, p. 11 (Type locality Mt. Victoria, Owen Stanley Range, New Guinea) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 101 ;Nieden, 1926, Das Tierreich, Anura n, p. 51.

Denticulated ridge in front of the pharynx preceded by a smooth one. Snout rounded, not prominent, shorter than the diameter of the eye ; canthus rostralis obtusely angular, curved ; loreal region oblique and slightly concave ; nostril slightly nearer the end of the snout than the eye ;tympanum 8 the diameter of the eye. Fingers not dilated, tfie first shorter than the second. Toes slightly dilated distally, the third a little longer than the fifth ; subarticular tubercles indistinct ; a weak inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation not quite reaching the tympanum. Skin smooth above and below ; indications of a )(-shaped pair of folds between the shoulders, and a straight fold from the posterior corner of the eye, above the -tympanum to the flanks. " Grey b r o p above, p-inkish on the sides of the back ; sides of head dark brown ; belly greyish, marbled wlth brown, with a fine white median line ; throat vinaceous red." Length : 22 mm. DISTRIB~ION : Owen Stanley Range, New Guinea.

B.M. 1896.10.31.31 juv.

I o.

Mt. Victoria, Owen Stanley Range, New Guinea

Anthony, TYPE (colour faded)

Sphenophryne brevicrus (van Kampen)

Ozydactyb brevicms, van ICampen, 1913, Nova Guinea, IX, Zool. 3, p. 465, P1. XI, fig. 8 (Type localities Hellwig Mts., 2,500 m., and W i c h m a ~ Mts., 3,000 m., Dutch New Guinea); idem, 1923, Amph. Indo-Austr. Arch., p. 122 ; Nieden, 1926, Das Tierreich, Anura n, p. 49, fig. 34.

Denticulated dennal ridge in front of the pharynx preceded by a smooth one. Snout rounded, feebly prominent, shorter than the eye, about as deep as long ; canthus rostralis rounded ; loreal region oblique, not concave ; nostril a little nearer the tip of the snout than the eye ; interorbital space broader than the upper eyelid ; tympanum scarcely, or not, distinct in adults, more distinct, and about half the diameter of the eye in young specimens. Fingers not dilated, the first shorter than the second, which is shorter than the fourth. Toes not dilated, the third a little longer than the fifth ; subarticular tubercles very feeble ; a feebly developed inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the axilIa or the shoulder, and the tarso-metatarsal joint the tympanum or the posterior border of the eye. Skin smooth above and below. Violet above, marbled or spotted with darker ; sometimes a dark band along the

OREOPHRYNE

159

sides of the head and body; yellowish beneath, usually marbled with brown, especially on the throat and limbs. Length : 30 mm. DISTRIBUTION : New Guinea (IdenberglMts., Hellwig Mts., Wichrnann Mts.).

? Sphenophryne sp. Hewitt, in November 1907, collected from the pitcher of a Nepenthes at Kuching two large eggs containing embryos of the type peculiar to the Asterophryinae and Sphenophryninae. The embryo has clavicles which reach the scapulae and the mid-line of the girdle, no teeth, two denticulated ridges across the palate, digits with large discs and T-shaped terminal phalanges, and a Sphenophryne-like habitus. The fingers have a rudiment of web, the toes are $ webbed and the outer extends beyond the third. B.M. 1933.6.23.1

SPECIMENS EXAMINED embryo Kuching . . . .

.

. . . . .

Hewitt

OREOPHR YNE, Boettger Microhyla (non Tschudi), Peters and Doria, 1878, Ann. Mus. Genova, xm, p. 428. ? Callula (non Gray), Horst., 1882, Notes Leyden Mus., v, p. 243. Orwphryne, Boettger, 1895, Zool. Anz., p. 135 (Type species Orwphryne senckenbergianu,Boettger = Microhykz achatinu, war. mluecensis, Peters and Doria) ; MBhely, 1901, TermBs. Fuzetek, xxm, p. 254 ; van Kampen, 1918, Bijdr. Dierk., xxr, p. 54 ; i&m, 1923, Amph. Indo-Austr. Arch., p. 110 ;Nieden, Das Tierreich, Anura 11, p. 38 ; Noble, 1931, Biol. Amph., p. 532. Sphenophryne (non Peters and Doria), MBhely, 1901, Termb. Fiizetek, xxm, p. 251 ; Vogt, 1911, Sitzber. Ges. Natf. Fr. Berlin, pp. 425-432 (part). i Phrynizalue (non Boettger), Stejneger, 1908, Proc. U.S. Nat. Mus., xxxnI, p. 573. dlehelyia, Wandolleck, 1911, Abh. Mus. Dresden, xm, 6, p. 7 (Type species Mehelyia k'neata, Wandolleck = Sphenophryne biroi, MBhely). Cophidus (non Boettger), van Kampen, 1913, Nova Guinea, IX,Zool., 3, p. 462. Chapmenu(part), Taylor, 1920, Philippine Journ. Sci., XVI,3, p. 332.

No trace of teeth. Prevomer large, undivided, in contact with its fellow mesially (fig. 55) ; the post-choanal portion overlying the palatine and sometimes raised to form a distinct ridge and, rarely, bearing one or more very small odontoids. Clavicles and procoracoids present, but reduced ; the former never reach the scapulae, and are inclined a t an angle to the coracoids ; procoracoids extending to the scapulae, or not extending greatly beyond the clavicles (figs. 56 and 57) ; no omosternum ; sternum large, cartilaginous. Vertebral column procoelous. Terminal phalanges T-shaped.

Fig. 55. Anterior cranial elements of Orwphryne variabilis (seen from beneath)

160

MONOGRAPH O F T H E MICROHYLIDAE

Pupil horizontal. Tongue oval, entire or slightly notched, and half free behind. A denticulated dermal fold across the palate in front of the pharynx, preceded as a rule by a shorter, curved, smooth one. Digits with distinct discs.

Fig. 56. Ventral elements of the shoulder-girdle of Oreophryne variabilis

[AfterMkhely Fig. 57. Ventral elements of the shoulder-girdle of Oreophryne biroi

SYNOPSIS O F THE SPECIES

s

I. Fifth toe shorter than the third. The procoracoid reaching the scap"la (fig. 56). A. Toes free. (1) Tibio-tarsal articulation reaching the middle of the eye or 0. cebbensis, p. beyond ; discs of fingers large. . . . . (2) Tibio-tarsal articulation reaching the tympanum or the eye ; discs of fingers small. (a) Tympanum less than half the diameter of the eye ; no dorso-lateral line of glandular warts. . 0. monticola, p. (b) Tympanum 8 the diameter of the eye; a dorso-lateral line of elongate warts . . . . . . 0. jeflersoniana, p. (3) Tibio-tarsal articulation reaching the shoulder or the tympanum ; discs of fingers moderately large. (a) Tympanum 4 to 8 the diameter of the eye, or hidden. (i) Belly of adult with glandular yellow patches O.uariabilis, p. . . . . 0. zimmeri, p. (ii) Belly uniform yellow (b) Tympanum about $ the diameter of the eye ; belly without glandular yellow patches . . . . . 0. rookmaakeri, p. B. Toes 3 webbed . . . . . . . . . . 0. kampeni, p.

.

.

161

162 163

.

163 164

.

164 164

OREOPHRYNE 11. Third and fifth toes subequal. Toes webbed. A. The procoracoid reaches the scapula. (1) Interorbital space much broader than the upper eyelid 0. moluccensis, p. . . 0. cruczjera, p. (2) Interorbital space as broad as the upper eyelid B. The procoracoid connected to the scapula by ligament (fig. 57) 0. albopunctata, p. 111. Fifth toe longer than the third. A. Toes free ; the procoracoid reaches the scapula. (1) Snout as long as, or longer than, the eye ; belly infuscate 0. ann~llata,p. (2) Snout shorter than the eye ; belly immaculate 0.jlava, p. B. Toes webbed; the procoracoid connected to the scapula only by ligament. (1) Tympanum about half the diameter of the eye ; palatine odontoids present, a t least in adult females . . 0. anthonyi, p. (2) Tympanum, if visible, about 4 the diameter of the eye; no palatine odontoids . . . . . . . 0. biroi, p.

. . .

.

I.

.

165 166 166

167 168

168 269

Oreophryne celebensis (Miiller)

9phenophrynecelebensis, Miiller, 1894, Verh. Nat. Ges. Basel, X, p. 841, fig. (Type localities Boelawa Mts., 1,200 m., and Totoiya Valley, 800 m., summit of the Gunung Sudara, Celebes); Boulenger, 1897, Roc. Zool. Soc. London, p. 235, P1. XVI, fig. 4 ; Schenkel, 1901, Verh. Nat. Ges. Basel, xrn, pp. 144, 151 ; Roux, 1904, Verh. Nat. Ges. Baael, XV, p. 432. Orwphryne cdebed8, MBhely, 1901, Term&. Fiizetek, XXTV, p. 256, P1. x, fig. 1 ; van Kampen, 1923, Amph. IndoAustr. Arch., p. 112 ; Nieden, 1926, Das Tierreich, Anura 11, p. 40 ;Ahl. 1933. Mitt. Zool. Mm. Berlin, xrx, p. 582. ?Orwphryneverrucosa (non Boulenger), van Kampen, 1913, Nova Guinea, IX, Zool., 3, p. 463 ;idem, 1923, Amph. Indo-Austr. Arch., p. 113 (part).

Two dermal ridges in front of the pharynx. Snout not prominent, rounded, slightly shorter than the diameter of the eye, canthus rostralis obtuse : loreal region nearly vertical, flat ; interorbital space broader than the upper eyelid ; tympanum indistinct, ) to 4 the diameter of the eye. Fingers moderately long, with large discs, that of the third as large as the tympanum ; Arst finger shorter than the second, which is subequal to the fourth. Toes moderate, free, the fifth distinctly shorter than the third ; discs well developed, but smaller than those of the fmgers ; no subarticular tubercles ; indications of an elongate inner metatam31 tubercle, but no outer. Tibio-tarsal articulation reaching the centre of the eye, or the centre of the loreal region. Skin smooth or slightly tubercular above, slightly granular on the belly ; an indefinite curved fold from the posterior corner of the eye above the tympanum to the shoulder. Reddish, yellowish, pink or brownish grey above, uniform or with darker spots ; sometimes there are indications of a light transverse bar between the eyes and an indefinite W-shaped dark mark between the shoulders; sometimes a dark lateral stripe along the loreal region, through the tympanum above the shoulder to the groin ; this stripe being sharply defined above, but merging into the colour of the lower surfaces beneath. The tubercles of the back sometimes picked out in white and sometimes a narrow light streak from snout to vent. Lower surfaces dirty white, uniform or dusted with brown beneath the throat. An indefinite triangular, dark area, lightbordered above, enclosing the vent. Two brown rings, separated by a light interspace, on the wrist. Males without vocal sacs. Length : 32 mrn. The procoracoid reaches the scapula. DJSTRIBUTION : Celebes, New Guinea.

MONOGRAPH OF THE MICROHYLIDAE SPECIMENS EXAMINED

9,hgr.

B.M. 94. 9. 28. 4-5 96.12.9. 101-102 96.4.29. 69-73

31)

d d, y 9

Mus. Amsterdam A 18l

2.

Totoiya Valley, 800 m., Celebes Sarasin, COTYPES Emponglar Volcano, Masarang, N. Celebes . . . . . Sarasin Indmlaman, 2,000 ft. S . Celebes Everett Hellwig Mts., 2,600 m., New Det. van Kampen Guinea as 0. zwucosa (Boul.)

Oreophryne monticola (Boulenger)

Sphenophryne manticola, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), XIX, p. 508 (Type locality Lombok) ; van Kampen, 1907, in Weber, Zool. Ergeb. Reise 0.Ind., N, p. 413 ; Roux, 1911, Zool. Jahrb., Syst., xxx, p. 506; Roux, 1912, in Elbert, Sunda Exped., n, p. 305; Barbonr, 1912, Mem. M u . Comp. Zool. Harvard, XLW, pp. 30, 173 ; Nieden, 1926, Das Tierreich, Anura n, p. 44 ; Barbour and Loveridge, 1929, Mem. Mus. Camp. Zool. Harvard, LXIX, p. 345. Oreophryne monticola, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 113 ;Mertens, 1927, Senckenbergiana, rx, pp. 237,242 (part);Dunn, 1928, Amer. M u . Novitates, cccxv, p. 3 ;Mertens, 1928, Zool. Anz., ~xxvm,p. 80; idem, 1930, Abh. Senck. Natf. Ges., x ~ n3,, p. 233, P1. VI, figs. 2 and 3. Callula mdesta (nom. nud.), Boettger, 1898, Ber. Senck. Nat. Ges., p. LW. Callula n. sp., Boettger, 1900, Abh. Senck. Nat. Ges., xxv, p. 338, footnote.

Two dermal ridges in front of the pharynx. Snout very slightly prominent, rounded, slightly longer than the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique, flat ; interorbital space as broad as, or broader than, the upper eyelid ; tympanum scarcely distinguishable, less than half the diameter of the eye. Fingers moderately long with small discs, that of the third being but little more than half the size of the tympanum ; first finger shorter than the second, which is slightly shorter than the fourth. Toes free, the fifth shorter than the third ; discs smaller than those of the fingers ; no subarticular tubercles ; an indistinct inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum or the posterior border of the eye. Skin smooth or slightly tubercular above ; a curved fold from the posterior corner of the eye above the tympanum to the shoulder, and indications of a pair of )(-shaped interscapular folds ; belly indistinctly granular. Magenta, purplish-brown, brown or grey above, uniform or with darker and lighter markings ; sometimes the back is irregularly stippled with darker, and sometimes there are traces of a darkw-shaped mark between the shoulders, with or without a dark interorbital spot ; flanks darker, sometimes a well-defined dorso-lateral streak of yellowish white from the posterior comer of the eye to the groin ; occasionally a fine white dorsal line from snout t o vent which bifurcates and runs along the hinder side of the legs to the tarso-metatarsal joint ; snout sometimes lighter, and there may be traces of a dark, white-bordered, ocellar, lumbar spot. Lower surfaces whitish, immaculate or faintly dusted with darker ; the throat and chest infuscate. Concealed surfaces of hind-limbs usually bright pink in life. Male with a vocal sac. Length : 29 mm. The procoracoid reaches the scapula. : Lombok, Bali. DISTRIBUTION

B.M. 97.6.21. 92-98

d d , 91)

Lombok

. . . .

.

Everett, COTYYES

This specimen differs from typical examples from Celebes in a slightly shorter first finger and longer fifth toe, but whether these relatively trivial differences indicate the existence of a distinct race or species i t is impossible t o decide on the basis of s single individual.

OREOPHRYNE

3. Oreophryne jeffersoniana, Dunn Orwphryne montiwla (part), Mertens, 1927, Senckenbergianarx, 6, p. 236. Oreophryne jepersoniana, Dunn, 1928, Amer. Mus. Novitates, cccxv, p. 3 (Type locality Komodo, 2,500 ft.) ; Mertens, 1930, Abh. Senck. Natf. Ges., XLII, 3, 1930, p. 232, PI. m, fig. 4.

Two dermal ridges in front of the pharynx. Snout short, rounded ;canthus rostralis rounded ;interorbital space broader than the upper eyelid ;tympanum feebly distinct, ) the diameter of the eye. Fingers with feebly developed discs; f i s t much shorter than the second. Toes free, discs very small ; subarticular tubercles absent ; a feebly developed inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the eye. Skin with a few scattered small warts above and a line of dongate ones from the eye more than half-way towards the groin ; belly finely granular. Tan above, the warts marked or outlined in black ; a fine mid-dorsal line and an X-shaped mark on the scapular region ; sides, below the dorso-lateral glandular line, darker. Thighs indistinctly barred. Length : 18.5 mm. DISTRIBUT~N : Komodo and Sumbawa.

4 . Oreophryne variabilis (Boulenger) aphemphryne uariabilis, Boulenger, 1896, Ann. Mag. Nat. Hist., (6), xvm, p. 64 (Type locality Mt. Bonthain, 6,500 ft., S. Celebes); idem, 1897, Proc. Zool. Soc. London, p. 235, P1. xm, fig. 5 ; Schenkel, 1901, Verh. Nat. Ges. Basel, XIII,pp. 144, 151 ; Nieden, 1926, Das Tierreich, Anura 11, p. 45. Orwphryne uariabilis, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 112 ; Smith, 1927, Proc. Zool. Soc. London, p. 214.

Two dermal ridges in front of the pharynx, the anterior sometimes reduced to three tubercles. Snout not prominent, as long as the diameter of the eye ; canthus rostralis blunt ; loreal region slightly oblique, flat ; interorbital space broader than the upper eyelid ; tympanum scarcely distinguishable, about 4 to 3 the diameter of the eye. Fingers moderately long with large discs, that of the third finger being about as large as the tympanum ;first finger shorter than thc second, which is subequal to, or slightly shorter than, the fourth. Toes moderate, free, the fourth shorter than the third ; discs smaller than those of the fingers ; no subarticular tubercles ; an indistinct, elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder or the tympanum. Skin smooth or very slightly tubercular above ; a curved fold from the posterior corner of the eye, above the tympanum, to the shoulder, and indications of a pair of )(-shaped folds between the scapulae ; belly indistinctly granular. Grey, brown, purple, pink or crimson above, uniform or with darker or lighter markings ; the loreal and tympanic regions beneath the tympanic fold usually darker, sometimes edged with lighter above ; there may be an indefinite transverse, dark bar between the upper eyelids and indications of a W-shaped dark mark between the shoulders ; sometimes a dark, light-bordered, ocellar lumbar spot ; lighter markings may also be present in the form of a pair of broad, yellow dorso-lateral stripes from the upper eyelid to the groin, a fine white line from snout to vent, or an oval yellow spot entirely covering the upper surface of the snout. Lower surfaces brownish, grey, or dirty white, with, in the adult, numerous glandular, white or yellow patches ; in halfgrown and juvenile specimens the belly is either immaculate or dusted with brown. Male with a vocal sac. Length : 28 mm. The procoracoid reaching the scapula. DISTRIBUTION : Mt. Bonthain, S. Celebes, from 1,500 to 2,000 metres.

MONOGRAPH O F THE MICROHYLIDAE B.M. 96.4.29.75-84

8 8, $! $!

8 $! 96.12. 99-100 1927.3.7. 5W54 8 8, 99 1929.5.18.1 cleared

Bonthain Peak, S. Celebes, 6,500ft.

Everett, COTYPES Bonthain Peak, S. Celebes, 5,000ft. Sarasin Bonthain Peak, S. Celebes, 6,000ft. M. Smith Bonthain Peak, S. Celebes . . . M. Smith

5 . Oreophryne ximmeri, Ah1 Orwphryne zirnmeri, Ahl, 1933, Mitt. Zool. Mus. Berlin, XIX,p. 482 (Type locality Tanke Salokko, Mengkoka Mts., 2,000 m., S.E. Celebes). Snout short, rounded, a little prominent, as long as the eye ; canthus rostralis distinct ; loreal region almost vertical, concave ; nostril much nearer the tip of the snout than the eye ; interorbital space twice as broad as the upper eyelid ; tympanum hidden. Fingers long, with discs measuring about half the diameter of the eye ; first

much shorter than the second, which is subequal to the fourth. Toes free, with lateral fringes, the 6fth shorter than the third ; discs smaller than those of the fingers ; no subarticular tubercles. Tibio-tarsal articulation reaching the tympanic region. Skin with only a few, small, scattered warts above ; an indefinite, narrow fold from the eye to the shoulder. Lower surfaces finely wrinkled and granular. Grey, with a yellowish sheen above ; back with scattered, irregular, black flecks ; a broad black stripe along the loreal and temporal regions below the canthus rostralis and supratympanic fold. Sides of the body and limbs lemon yellow with darker marbling. Lower surfaces uniform orange-yellow. Length : 37 mm. DISTRIB~TION : Tanke Salokko, Mengkoka Mts., 2,000 m., S.E. Celebes.

6 . Oreophryne rookmaaki, Mertens Oreophryne rooknuuakeri, Mertens, 1927, Senckenbergiana, Ix, 6, p. 236 (Type locality Rana Mese, W. Flores, 1,200 m.) ; idem, 1930, Abh. Senek. Natf. Ges., mu, 3, p. 230, PI. vr, fig. 1.

Two dermal ridges in front of the pharynx. Snout short, not prominent, with rounded, indistinct, canthus rostralis ; interorbital space broader than the upper eyelid ; tympanum distinct, about $ the diameter of the eye. Fingers moderate, with large digital discs which are about equal t o half the diameter of the eye ; first shorter than the second. Toes moderate, free; discs smaller than those of the fingers ; subarticular tubercles very indistinct ; an ill-defined; elongate inner, but no outer, meta9 tarsal tubercle. Tibio-tarsal articulation reaching the shoulder. Skin smooth or slightly warty. Chocolate brown above, clay-yellow beneath ; rarely a stippling of darker spots or a fine yellow line from snout to vent and along the hinder side of the legs. Length : 32.5 mm. : Rana Mese, W. Flores, 1,200 m. DISTRIBUTION

7 . Oreophryne kampeni, sp. nov. Sphenophryne loriae (in part), Boulenger, 1898, Ann. Mus. Genova, (2), xvm, p. 707.

Holotype a $2, Number 97.12.10. 143 in the British Museum, from Moroka, British New Guinea. A denticulated dermal ridge across the palate in front of the pharynx, preceded by an indefinite meaian tubercle. Snout truncate, not prominent, slightly shorter than the diameter of the eye ; canthus rostralis rounded, curved ; loreal region oblique, slightly concave ; interorbital space broader than the upper eyelid ; tympanum rather indistinct, the diameter of the eye. Fingers moderate, with large discs, that of the third finger larger than the tympanum ; first finger shorter than the second, which is

+

OREOPHRYNE shorter than the fourth. Toes short, ) webbed, the fifth distinctly shorter than the third ; discs much smaller than those of the fingers ; subarticular tubercles very indistinct ; an ill-defined elongate inner, but no outer, metatarsal tubercle. Tibiotarsal articulation reaching the posterior border of the eye. Skin smooth above ; a curved supratympanic fold ; belly slightly granular posteriorly. Pale brown above, spotted and washed with darker, a pale band from eye to eye extending forwards over the snout ; two broad paler stripes from the upper eyelids towards the groin, converging slightly between the shoulders ; two dark brown bands on the wrist, separated by a light interspace. Lower surfaces yellowish brown, washed with darker beneath the throat. Ovarian eggs large, 2 mm. in diameter. Length from snout to vent, 23 mm. Fore-limb, 13 mrn. Hind-limb, 31 mm. The thirteen paratypes from the same locality show little variation. The snout may be as long as, or slightly longer than, the eye, and the tibio-tarsal articulation may reach the centre of the eye. The two dorso-lateral light streaks may be entirely absent, but more frequently, though present, they are invaded by spots of dark brown ; there is almost constantly a dark rectangular spot on the anterior part of the back, commencing between the eyelids, where i t connects the two, and bounded by the dorsolateral streaks ; posteriorly this dark area merges gradually into the lighter ground colour ; sometimes a h e silvery stripe from snout to vent. Male with a vocal sac opening by a slit on each side of the tongue. These specimens formed part of the series from which Boulenger described 0. loriae. The figure of this species, however, shows quite definitely a frog with the second finger as long as, or longer than, the fourth, and the fifth toe longer than the third; consequently the name loriae must be reserved for that form, although comparison reveals it to be identical with bird Mkhely. The form with the short fifth toe, which is here described as new, differs also in the possession of a procoracoid which reaches the scapula, and thus closely resembles 0.moluccensis and 0. d f e r a ; it is, however, distinguished from either of these by its much shorter fingers and toes and their relative proportions. SPECIMENS EXAMINED ? Moroka, British New Guinea Loria, TYPE B.M. 97.12.10. 143 97.12.10. 144-145 3, yg. Moroko, British New Guinea Loria, PARAMus. Genova

11

3 3, 9 9, yg.

TYPES

Moroko, British New Guinea Loria, PARATYPES

8. Oreophryne moluccensis (Peters and Doria) Microhyla achatino scar. moluccenais, Peters and Doria, 1878, Ann. Mus. Genova, xu~, p. 428 (Type localities " Acqui Conom," Ternate, and Batjan). 9 Ccallulafrontifcasci&, Horst, 1883, Notes Leyden Mu.,v, p. 243 (Type localities Salawatti, Morotai and Halmaheira). Oreophryne senckenbergicano,Boettger, 1895, Zool. Anz., xvm, p. 135 (Type localities Supu, N. Halma heira ; Dodinga, W. Halmaheira ;and Patani, E. Halmaheira) ; idern, 1901, Abh. Senck. Natf. Ges., XXV, p. 376, P1. XVI,figs. lla-c ; Mbhely, 1901, Termbs. Fiizetek, xxrv, p. 255 ; Mertem, 1922, Senckenbergiana, rv, p. 165 ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 115; Nieden, 1926, Das Tierreich, Anura 11, p. 39.

Two dermal ridges in front of the pharynx. Snout obtuse, not prominent, slightly longer than the diameter of the eye ; canthus rostralis rounded, curved ; loreal region oblique ; interorbital space broader than the upper eyelid ; tympanum indistinct, less than half the diameter of the eye. Fingers short, with large discs, that of the third being larger than the tympanum ; first finger shorter than the second, which is longer than the fourth. Toes webbed, the third and fifth subequal ; discs slightly smaller than those of the fingers ; subarticular tubercles indistinct ; an ill-defined, elongate

+

166

MONOGRAPH OF THE MICROHYLIDAE

inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum or the posterior border of the eye. Smooth above ; a curved supratympanic fold; belly and lower surfaces of the thighs granular. Whitish, marbled with grey above ; a white cross-bar between the eyes ; a black temporal streak, bordered above and below with white ; sides of body spotted with black and white ; a large black, white-bordered lumbar ocellus ; brownish yellow beneath, immaculate or spotted and marbled with white. Male with a vocal sac. Length : 26 mm. DISTRIBL~ION : Halmaheira, Ternate, Batjan, Salawatti ? and Morotai ?

. . .

Mus. Genova

ad.

Batjan

Mus. Senckenberg

$

Halmaheira

.

Beccari, COTYPE of Microhyla achatina var. mluccensis, Peters and Doria Kiikenthal, COTYPEof Oreophryne senckenbergiana, Boettger

9 . Oreophryne crucifera (van Kampen) Cophizalwr cmifer, van Kampen, 1913, Nova Guinea, IX, 3, p. 462, P1. XI, fig. 6 (Type locality Went Mts., 800 and 1,050 m.). Orwphryne crucifera, van Kampen, 1919, Bijdr. Dierk., XXI, p. 54 ; idem, 1923, Amph. Indo-Austr. Arch., p. 116 ; Nieden, 1926, Das Tierreich, Anura n, p. 41.

A denticulated dermal ridge across the palate in front of the pharynx, sometimes preceded by a feebly developed smooth one. Snout truncate, not prominent, shorter than the diameter of the eye ; canthus rostralis angular, cumed ; loreal region oblique, slight3y concave ; interorbital space as broad as the upper eyelid ; tympanum hidden or distinct, about 3 the diameter of the eye. Fingers moderate, with large discs, that of the third finger larger than the tympanum ; first finger shorter than the second, which is longer than the fourth. Toes webbed at the base, the third and fifth subequal ; discs a little smaller than those of the fingers ; subarticular tubercles indistinct ; an indistinct inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaches the eye. Skin smooth above, granular beneath. Reddish brown above ;an hour-glass-shaped figure on the back and a round spot in the lumbar region, yellowish ; a yellow crossbar between the eyes. Lower surfaces whitish, immaculate or finely dusted with brown. Length : 24 mm. DISTRIB~IO :N New Guinea (Went Mts.). 10.

Oreophryne albopumtata (van Kampen)

SpLnophryne dbopunetata, van Kampen, 1909, Nova Guinea, Ix, 1, p. 42, P1. n, fig. 7 (Type locality North River [= Lorentz River], near Sabang, Dutch New Guinea). Oreqphryne a.%opnetata, van Kampen, 1918, Bijdr. Dierk, xxr, p. 54; i h , 1923, h p h . Indo-Austr. Arch., p. 119; Nieden, 1926, Das Tierreich, Anura n, p. 41.

Two dermal ridges across the palate in front of the pharynx, the posterior denticulate and the anterior smooth. Snout about as long as the diameter of the eye, with rounded canthus rostralis and nearly vertical, slightly concave, loreal region ; interorbital space broader than the upper eyelid ; tympanum indistinct, about 3 the diameter of the eye. Fingers moderate, with large discs, that of the third fmger being more than half the diameter of the eye ;first finger shorter than the second, which is subequd to, or very slightly shorter than, the fourth. Toes slightly webbed, the third and fifth

OREOPHRYNE

167

subequal ; discs smaller than those of fingers ; subarticular tubercles indistinct ;traces of an elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum or the posterior border of the eye. Skin smooth or slightly tubercular above ; a curved supratympanic fold ; belly granular. Brownish above, with darker and lighter markings ;a dark bar between the posterior halves of the upper eyelids, a W-shaped marking between the shoulders, and an indehite lumbar ocellus ; a white dot in the middle of the tibia ; anal region darker. Lower surfaces whitish, dusted with darker. Length : 25 rnrn. DISTRIBUTION : Lorentz River, Dutch New Guinea.

II .

Oreophryne annulata (Stejneger)

Phrynixalus onulatus, Stejneger, 1908, Proc. U.S. Nat. Mus., xxxnr, p. 573 (Type locality Davao, Mindanao). P . onnulatwr, Taylor, 1920, Philippine Journ. Sci., XVI, 3, p. 336 ; idem, 1921, Amph. and Turtles Philippine Islands, p. 137. Chperino Pris yo, Taylor, 1920, Philippine Journ. Sci., xv, 3, p. 335, P1. IX,fig. 3 (Type locality Biliran I L d ) ;idem, 1921, Amph. and Turtles Philippine Islands, p. 136, Pi. IX,fig. 3.

Two transverse dermal ridges across the palate in front of the pharynx. --

Snout very slightly prominent, as long as, or a little longer than, the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical, slightly concave ;interorbital space much broader than the upper eyelid ; tympanum moderately distinct, about half the diameter of the eye. Fingers short, with large discs, that of the third finger quite as large as the tympanum ; first finger shorter than the second, which is subequal to, or very slightly shorter than, the fourth. Toes free, the fifth longer than the third ; discs scarcely, if a t all, smaller than those of the fingers ; subarticular tubercles indefinite ; an indistinct, elongate inner, but no outer, metatarsal tubercle. Tibiotarsal articulation reaching the tympanum or the posterior corner of the eye. Skin smooth or slightly tubercular above, granular on the belly and beneath the thighs ; an indefinite supratympanic fold. Brownish above, shaded with darker and lighter; a dark cross-bar between the posterior halves of the upper eyelids, preceded by a lighter transverse bar ; sometimes a W-shaped dark marking between the shoulders and traces of ocellar spots in the groin ; a pale, dark-edged streak from eye to shoulder, a light, dark-edged annulus proximal to each finger disc ; sometimes a h e white line from snout to vent. Lower surfaces whitish, more or less profusely dusted with brown. Male with a vocal sac. Females with large ovarian eggs (1.5 mm.). Procoracoid reaching the scapula. Length : 19.5 mm. DISTRIBUTION : Mindanao and Biliran, Philippine Islands. The statement that the paratype is " apparently just through its transformation " was, no doubt, prompted by the presence of a conical excrescence, just above the vent, which recalls the vanishing tail of a methorphosing frog ; this prominence is, however, more probably due to the pressure of the urostyle against the skin, caused by the dislocation of the ilia from the sacrum.

U.S. Nat. Mus.,

juv.

B.M. 1929.6.1.1-5

,

Todaya, Mt. Apo, 4,000 ft. Mindanao . . . . Milbok to Santo Cotabato, Mindanao

Mearns,P~~~m~ Taylor (identified by Taylor as Chaperina vis a ~ ~ )

MONOGRAPH OF THE MICROHYLIDAE I 2.

Oreophryne JEava sp. n.

Orwphryne anthnyi (non Boulenger), van Kampen, 1915, Nova Guinea, XIII, 1, p. 40 ; idem, 1923, Amph. Indo-Austr. Arch., p. 117 (part). ? Orwphryne vemoaa (non Boulenger), van Kampen, 1923, Amph. Indo-Austr. Arch., p. 113 (part).

Holotype a $2, A.15 in the Zoological Museum, Amsterdam, from the Lorentz River (Kloofbivak), New Guinea. Two dermal ridges in front of the pharynx, the posterior denticulate, the anterior smooth and divided into three lobes. Snout rounded, not prominent, shorter than the diameter of the eye ; canthus rostralis rounded, straight ; loreal region nearly vertical, slightly concave ; interorbital space a little broader than the upper eyelid ; tympanum distinct, nearly half the diameter of the eye. Fingers moderate, with large discs, that of the third finger about as large as the tympanum; first finger shorter than the second, which is subequal to the fourth. Toes moderately long, free, the fifth longer than the third ; discs smaller than those of the fingers ; subarticular tubercles indistinct ; an indefinite elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum. Skin indistinctly tubercular above ; belly granular. Yellowish, mottled with brown above ; a darker cross-bar between the posterior halves of the upper eyelids, preceded by a lighter band ; an indefinite W-shaped mark between the shoulders, and a short dark streak from the posterior corner of the eye, above the tympanum ; flanks more mottled than the back, the spots tending to form short longitudinal streaks ; a light, dark-bordered, annulus proximal to each finger disc and on the wrist. A fine white line from snout to vent. Lower surfaces yellowishwhite, the throat faintly dusted with darker. Length 21 mm. Fore-limb, 12 mm. Hind-limb, 29 mm. The procoracoid reaches the scapula. The specimens found breeding near the Idenburg River by van Heurn and referred by van Kampen to " 0.verrucosa " may also belong to this species. They have the same type of shoulder-girdle, and free toes of similar proportions, but the only specimens available (A.16 and 17 of the Amsterdam Museum) are, unfortunately, so shrivelled as to render their accurate determination almost impossible ; one of them possesses one or two small palatine odontoids on each side, a condition only found elsewhere in the genus in 0. anthonyi. The large (5 mm.) eggs, ten to twenty in number, were arranged in a group hanging by a muailaginous cord from the roof of a cavity in the tuber of an epiphytic plant of the genus Hydnophytum ; development was " direct." Mus. Amsterdam A.15 A.16-17

I 3.

SPECIMENS EXAMINED Lorentz River (Kloofbivak) Near Idenburg River (2,4002,900 m.) .

$2

.

.

. . . .

9

Versteeg, TYPE van Heurn

Oreophryne anthonyi (Boulenger)

Sphnophryne anthunyi, Boulenger, 1897, Ann. Mag. Nat. Hist., (6), xrx, p. 10, P1. n, fig. 1 (Type locality Mt. Victoria, Owen Stanley Range, New Guinea). Orwphryne anthohyi, van Kampen, 1919, Bijdr. Dierk., xxr, p. 64 ; idem, 1923, Amph. Indo-Austr. Arch., p. 117 (part) ; Nieden, 1926, Das Tierreich, A n m n, p. 42.

A denticulated dermal ridge across the palate in front of the pharynx, preceded by a smooth one which is reduced to three tubercles ; prevomers forming a pair of strong, posteriorly convergent, ridges behind the choanae ; these ridges, smooth in males and juveniles but beset with a few small, rounded, tooth-like prominences, which are best developed mesially, in adult females. Snout rounded, slightly shorter than the diameter of the eye, canthus rostralis obtuse, curved ; loreal region somewhat oblique, slightly concave ; tympanum indistinct, half the diameter of the eye. Fingers long,

OREOPHRYNE

169

with large triangular discs, that of the third finger the diameter of the eye ; first shorter than the second, which is shorter than the fou=th. Toes webbed at the base, the fourth slightly longer than the third ; discs distinctly smaller than those of the fingers; subarticular tubercles feebly developed ; an indefinite, elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanum. Skin smooth or faintly tubercular above ; a curved supratympanic fold and a pair of )(-shaped folds between the shoulders ; belly granular. Pale brown, grey or reddish above, with irregular darker spottings and marblings ; the supra-scapular folds usually outlined with darker. Lower surfaces, flanks and concealed portions of the hind-limbs brown, with large white spots, or white with a brown reticulum ; in adult males this pattern is entirely lost or only rudimentary, but a fine median white line is developed on the chin and throat. Sometimes a fine median dorsal line from snout to vent. Males with a vocal sac. Length : 45 mm. The procoracoid is connected to the scapula by a ligament only (fig. 57). : British New Guinea. DISTRIBUTION B.M. 96.10.31. 43-49

SPECIMENS EXAMINED $3, 99,juvs. Mt. Victoria, Owen Stanley Range

Anthony, COTYPES

14. Oreophryne biroi (Mdhely) Sphenophrym bbiroi, MBhely, 1897, TerrnBs. Fiizetek, xx, pp. 400, 411, P1. x, figs. 3-6 (Type locality Friedrich Wibelmshaven, New Guinea) ; Boulenger, 1898, Ann. Mus, Genova, (2), x w , p. 709 ; MBhely, 1901, Term&. Fiizetek, xxrv, p. 252, P1. x, fig. 2, P1. xr, fig. 6 ; van Kampen, 1913, Nova Guinea, IX,Zool., 3, p. 462. Orwphryne biroi, van Kampen, 1918, Bijdr. Dierk., XXI, p. 54 ; idem. 1913, Amph. Indo-Austr. Arch., p. 118 ; Nieden, 1926, Das Tierreich, Anura 11, p. 40. r9phemphryne lo&, Boulenger, 1898, Ann. Mus. Genova, (2), xvm, p. 707, P1. vnr, fig. 3 (Type locality Moroka, British New Guinea) ; van Kampen, 1909, Nova Guinea, rx, 2001. I, p. 41. Orwphryne loriae, van Kampen, 1918, Bijdr. Dierk., XXI,p. 54 ; idem, 1923, Amph. IndoAustr. Arch., p. 117 ;Nieden, 1926, Das Tierreich, Anura n, p. 41. ? Sphemphryne mertoni, Roux, 1910, Abh. Senck. Natf. Ges., xIn, p. 227 (Type locality Samang, Wokam, Aru Islands). Oreophryne mertoni, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 120. Mehelyia line&, Wandolleck, 1911, Abh. Mus. Dresden, x m , 6, p. 7, figs. 18-26 (Type locality " Sacksack-hutte," Torricelli Mts., New Guinea). Mehelyia a&&, Wandolleck, 1911, Abh. Mus. Dresden, xnI, 6, p. 8, figs. 27-35 (Type locality Torricelli Mts., New Guinea). Cophizalus crucifer ( m n van Kampen), Boulenger, 1914, Trans. Zool. Soc., London, xx, 5, p. 251.

A denticulated dermal ridge in front of the pharynx, preceded by a row of three transversely elongate tubercles. Snout truncate, about as long as the eye ; canthus rostralis blunt ; loreal region slightly oblique, concave ; tympanum scarcely distinct, very small, f to the diameter of the eye. Fingers with well-developed discs, that of the third finger being nearly half the diameter of the eye ; first finger shorter than the second, which is as long as, or slightly longer than, the fourth. Toes webbed a t the base, the fifth slightly longer than the third and with a larger disc ; discs but very slightly smaller than those of the fingers ; subarticular tubercles indistinct ; an indefinite elongate inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation rearhing the eye. Skin smooth above ; a curved supratympanic fold ; belly granular. Grey, brown or yellowish above with dark dots and indefinite longitudinal streaks, with or without a fine white line from snout to vent ; sometimes there are traces of a brown blotch on the occiput, preceded by a light bar between the anterior halves of the upper eyelids, and of aW-shaped marking behind the head, or a faint lumbar

+

170

MONOGRAPH OF THE MICROHYLIDAE;

ocellus. Sometimes a light, dark-bordered annulus proximal to each finger disc and on the mist. Lower surfaces whitish, uniform or dusted with brown. Male with a vocal sac. Eggs large (4 mm.) Length : 30 mm. DISTRIBUTION : New Guinea and Ferguson Island. The procoracoid is connected to the scapula by ligament only. Van Kampen has suggested that the two species described by Wandolleck as Mehelyia afinis and Mehelyia lineatu are both conspec&?with 0. biroi ;after examining cotypes of these two species I am able t o conflm van Kampen's suggestion. The differences in the shoulder-girdle which Wandolleck stresses are probably due to individual, or perhaps sexual, variation. The angles of incidence of the axis of the clavicle with a line joining the point of contact of the clavicle with the coracoid and the anterior mesial extremity of the coracoid, were approximately 30' and 44' respectively in a 8 and 0. biroi from Sattelberg ; these two angles are approximately the same as those shown in Wandolleck's figures of M . afinis and M . lineata. SPECIMENS EXAMINED B.M. 83.10.23. 2 ? Katow, New Guinea . Doria 97.12.10. 141-142 9,hgr. Moroka, British New Loria, COTYPESof Guinea . . . . Sphenophryne loriae, Boulenger 1903.4.30. 21 ? Milne Bay, British New Guinea . . . . Meek 190P.ll.l. 62-65 8 8 , ?? Ferguson Island British New Guinea Meek 1904.11.1. 70 juv. Ferguson Island, British New Guinea Meek 1909.10.30. 23 8 Wendessi, Dutch New . Guinea . . . . Pratt 1913.11.1. 142 juv. Setekwa River, Dutch New Guinea . . . Kloss 1929.5.19. 1 8 Torricelli Mts., New Schlaginhaufens, COGuinea . . . . TYPE of Mehelyia lineata, Wandolleck 1929.5.19. 2-3 8? Torricelli Mts., New Schlaginhaufens, COGuinea . . . . TYPE of Mehelyia aflnis, Wandolleck 1934.1.5. 7-9 99,juv. Kokoda (1,200 ft.), British New Guinea Cheeswn 1934~1.5.10 ? Orrori (3,000 ft.), British New Guinea . . Cheesman Mus. Hung. Amph. 2126 B 13 ad. and Friedrich Wilhelmsjuv. hafen . . . . Biro, COTYPES Amph. 2414110 2 9 8 8 , $29,Sattelberg . . . . Biro juvs. and 1 skull 3 Adults Moroka, British New Loria, COTYPES of Mus. Genova Guinea Sphenophlyne loriae, Boulenger

COPHIXAL US, Boettger C o p h i d u s , Boettger, 1892, Kat. Batr. Senck. Nat. Ges., p. 24 (Type species Cophizalw geislerorum, Boettger) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 146 (part) ; Nieden, 1926, Das Tierreich, Anura n,p. 56 ;Noble, 1931, Biol. Amph., p. 533. Phrynizalus, Boettger. 1895, 2001. h. wm, , p. 132 (Type species Phrynizdua nwnkznua, Boettger) ; MBhely, 1901, TermBs. Fiizetek, xxw, pp. 195, 245 ; Boettger, 1903. Abh. Senck. Natf. Ges., xxv, p. 368.

COPHIXALUS Cq'uka, MBhely, 1901, TermBs. Fiizetek, xxrv, pp. 193, 242 (Type species Phrynizalua ozyrhinus, Boulenger) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 134 ;Nieden, 1926, Das Tierreich, Anura u, p. 56 ;Noble, 1931, Biol. Amph., p. 533. Awtrochzperina (part), Fry, 1912, Rec. Austr. Mus., 11,p. 91. Choerophryne, van Kampen, 1915, Zool. Jahrb., Syst., xxxvn, p. 376 (Type species Chmrophryne poboscidea, van Kampen = Copiu2a (?) roetellifer, Wandolleck) ; i&m, 1923, Amph. IndoAustr. Arch., p. 136 ; Nieden, 1926, Das Tierreich, Anura n, p. 62 ;Noble, 1931, Biol. Amph., p. 534. Hylopbrbus (part), van Kampen, 1923, Amph. Indo-Auetr. Arch., p. 137; Nieden, 1926, Das Tierreich, Anura u, p. 57. Sphenqpryne (non Peters and Doria), Bonlenger, 1898, Ann. Mus. Genova, (2). X-, p. 707.

No teeth. Prevomer large, undivided, in contact with its fellow mesially, the postchoanal portion overlying the palatine region, without odontoids (fig. 58), sometimes with a distinct raised ridge and sometimes (C. rostellifer) fused with the ethmoids. Clavicles, procoracoids and omosternum absent; sternum cartilaginous. Vertebral column procoelous. Terminal phalanges T-shaped. Pupil horizontal. Tongue oval, entire and half free behind ; a denticulated dermal ridge across the palate, sometimes preceded by a curved smooth one. Digits with terminal dilftations.

fig. 58. Anterior cranial elements of Cophimka uerrueosua (seen from beneath)

SYNOPSIS OF THE SPECIES

I. Digital discs large, those of the fingers larger than those of the toes. A. Toes webbed a t the base . . C. geislerorum, p. 172 B. Toes free. (1) Third toe extending beyond the fifth. Inner fmger very short () or less the length of the second), C. dteles, p. 172 without a disc. . . . . . . . . . Inner finger well developed (about half the length of the second), with a small disc. (i) Snout shorter than the eye. (a) Tibio-tarsal articulation reaching the tympanum or the eye . . . . . . . . C. ornutw, p. 173 (p) Tibio-tarsal articulation reaching t o between the eye and the tip of the snout. . C. montanrcs, p. 174

.

. . . . . .

MONOGRAPH OF THE MICROHYLIDAE (ii) Snout longer than the eye. (a) Tibio-tarsal articulation reaching the shoulder C. biroi, p. (p) Tibio-tarsal articulation reaching to between the eye and the end of the snout. * Snout rounded, scarcely prominent ; anterior and posterior surfaces of the thighs dark, . C. venucosus, p. with circular white spots . ** Snout acutely pointed, prominent ; anterior and posterior surfaces of the thighs uniform yellow or red . . C. cheesmanae, p. (2) Fifth toe extending beyond the third. (a) Snout not more than once and a half as long as the eye C. van'egatus, p. (6) Snout a t least twice as long as the diameter of the eye C. rostellifer, p. 11. Digital discs small, those of the toes larger than those of the fingers C. oxyrhinus, p. I.

174

175 175 176 177 178

Cophixalus geisbrorum, Boettger

Cophizalus gei~lerorum,Boettger, 1892, Kat. Batr. Senck. Nat. Ges.,p. 24 (Type locality " Kaiserwilhelmsland," New Guinea) ;Vogt., 1911, Sitzber. Gea. Natf. Fr. Berlin, p. 431 ;van Kampen, 1923, Arnph. Indo-Austr. Arch., p. 146 ;Nieden, 1926, Das Tierreich, Anura rr, p. 56.

A denticulated dermal ridge in front of the pharynx. Snout truncate, shorter than the diameter of the eye ; canthus rostralis feebly developed ; loreal region slightly concave ; interorbital space broader than the upper eyelid; tympanum distinct. Fingers free, with large discs, first shorter than the second. Toes distinctly webbed a t the base with discs slightly smaller than those of the fingers ; subarticular tubercles feebly developed ; an indefinite, compressed inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching to between the shoulder and the tympanum. Skin smooth above ; belly and lower surfaces of the thighs granular. Blackish brown above, the sides and limbs dotted with yellow; a distinct light marking in the lumbar region ; a curved white streak from the eye to the shoulder ; lower surfaces yellowish white, the throat and hinder side of the thighs finely dusted with brown. Length 26 mm. DISTRIBWION : N.E. New Guinea (known only from the type locality.) 2.

Cophixalus ateles (Boulenger)

Sphenophryne ateles, Boulenger, 1898, Ann. Mus. Genova, (2), x w , p. 708, fig. (Type locality Moroka, New Guinea, 700 m.) ; Vogt., 1911, Sitzber. Ges. Natf. Fr. Berlin, p. 431 ; Orwphryne atelm, van Kampen, 1923, Amph. Indo-Austr. Arch, p. 115.

A denticulated dermal ridge in front of the pharynx, preceded by a curved smooth one. Snout rounded, slightly prominent, a little shorter than the diameter of the eye; canthus rostralis blunt ; loreal region slightly oblique and concave ; interorbital space broader than the upper eyelid ; tympanum distinct, a little less than half th- diameter of the eye. Fingers short, all except the first with large discs, which are truncate distally ; first finger extremely short, not more than the length of the second,l which is a little shorter than the fourth. Toes free, with small but distinct discs ; the third much longer than the fifth. Subarticular and inner metatarsal tubercles present, but very feebly developed. Tibio-tarsal articulation reaching the eye. Skin slightly warty above, especially on the posterior part of the back and the *flanks; smooth beneath. Grey or pale brown above, the flanks and upper surfaces of the snout yellowish ;

+

1

The figure given with the original description exaggerates the length of this digit.

the colour of the flanks forms a more or less regular streak from the eye to the groin which may be edged above by darker ; a light cross-bar connecting the upper eyelids bordered behind by darker. A faint dark W-shaped mark may be present on the anterior part of the back and a light vertebral stripe may be present. Lower surfaces lighter, more or less profusely dusted with dark brown ; forearm with a distinct brown annulus ; anal region blackish. Length : 15 mrn. Male with a vocal sac. DISTRIBUTION : Moroka, New Guinea (700 m.) SPECIMENS EXAMINED 3, hgr. Moroka, British New Guinea B.M. 97.12.10. 146-147 (700 m.) . . . . . . Loria, COTYPES

3. Cophixalus ornatus (Fry) Sphenophryne loriae (part), Boulenger, 1898, Ann. Mus. Genova, (2), x m , p. 707. Austrochaperina ornata, Fry, 1912, Rec. Austr. Mus., IX, 1, p. 91, fig. 38 (Type localities Russell Nieden, 1926, Das Tierreich, River and Cairns, N.E. Queenaland). Sphenophryne OW, Anura IIQ. 48. Phrynimlus regime, Andersson, 1917, Svenska Vet. Ak. Handl. Stockholm, m, 9, p. 4, figs. (Type locality Malanda, Queensland). Sphenophryne variabilia (non Boulenger), Procter, 1923, Proc. Zool. Soc. London, p. 1071. Hylophorbus sp. ?, Smith, 1927, Proc. Zool. Soc. London. p. 215.

A denticulated dermal ridge in front of the pharynx, preceded by a curved smooth one, which may be broken up into three lobules. Snout truncate, or rounded, scarcely prominent, a little shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region almost vertical, not concave ; interorbital space once and a half as broad as the upper eyelid ; tympanum hidden or slightly distinct, about half the diameter of the eye. Fingers with discs, those of the three outer large, truncate, that of the inner smaller ; first about half the length of the second, which is very slightly shorter than the fourth. Toes with smaller but distinct discs ; the third much longer than the fifth ; subarticular and inner metatarsal tubercles present. Tibio-tarsal articulation reaching the tympanum on the hinder border of the eye. Skin smooth or slightly tubercular above ; belly and hinder side of thighs more or less granular. Coloration variable. Brown or blackish above, uniform or marked with darker and lighter. The following markings may, or may not, be all present ; a light streak connecting the upper eyelids bordered behind by darker ; irregular black spots on the back ; a narrow black line from the posterior corner of the eye to the middle of the flank, interrupted above the shoulder ; a narrow black streak from beneath the eye to the corner of the mouth ; an oval black spot, margined anteriorly by white in the groin, and traces of a similar ocellus above each arm ; a fine white streak from snout to vent. Limbs dotted with dark brown ; a transverse black bar on the outer side of the forearm ; anal region blackish ; the heel lighter. Lower surfaces yellowish-white, the throat and chest dusted with brown. Ovarian eggs large (2 rnrn.). Length : 26 mm. DISTRIBUTION : Queensland, S.E. New Guinea. SPECIMENS EXAMINED B.M. 1929.5.17.1 3 Malanda, N. Queensland . Mjoberg, COTYPE of Phrynixulus regime, Andersson 1923.10.13.12 Ravenshoe, N. Queensland . Sherrin Mus. Genova ad. Moroka, British New Guinea. Loria, CCyrnrp~ of Sphmphryne loriue, Boulenger.

.

MONOGRAPH OF THE MICROHYLIDAE

4 . Cophixalus montanzls (Boettger) P h r p i z d w r montanus, Boettger, 1895, Zool. Anz., xvm, p. 133 (Type locality N. Halmaheira, 2,200-2,500 ft.) ; idem, 1901, Abh. Senck. Natf. Ges., xxv, p. 368, P1. XVI,figs. 9-9b ;MQhely, 1901, Term&. Ffizetek, XXN, pp. 201,251. Hylophrbua mmkanua, van Kampen, 1923, Amph. Indo-Anstr. Arch., p. 139 ;Nieden, 1926, Das Tierreich, An- 11, p. 62 ; Burt, 1932, Bull. Amer. Mus. Nat. Hist., LXIII, p. 484.

Prevomer with a strongly raised edge which produces a prominent curved fold behind each choana ;a denticulated dermal ridge in front of the pharynx, preceded by a curved smooth one. Snout obtusely pointed, shorter than the eye ; canthus rostralis rounded ; loreal region slightly concave; interorbital space broader than the upper eyelid; tympanum moderately distinct, a little more or a little less, than half the diameter of the eye. Fingers with discs, those,of the three outer very large, truncate distally ; first about half the length of the second, which is a little shorter than the fourth. Toes free, with small, but definite, discs ;the fifth much shorter than the third ; subarticular and inner metatarsal tubercles present, but feebly developed. Tibio-tarsal articulation reaching to between the eye and the tip of the snout. Skin smooth above or slightly tubercular posteriorly and on the flanks; lower surfaces entirely smooth, or the posterior part of the belly slightly granular. Reddish brown (green in life), spotted and marbled with blackish ; a dark triangular spot between the eyes and a broad W-shaped figure of the same colour on the anterior part of the back ; sometimes a broad, white vertebral line ; limbs dark spotted ; fingers and toes yellow, with blackish rings. Brownish yellow beneath, throat, breast and thighs powdered or spotted with blackish. Length : 26 mrn. DISTRIBUTION :' Halrnaheira (2,200-2,500 ft.), ? New Guinea. SPECIMEN EXAMINED Mus. Senckenberg ad. N. Halmaheira (2,200-2,500 ft.) . Kiikenthal, COTYPE

5 . Cophixalus biroi (Mthely) Phrynizalus biroi, MQhely,1901, TermQs.Fiizetek, X ~ I Vpp. , 197,247, P1. IX, figs. 1-5, P1. XII, fig. 3 (Type locality Sattelberg, New Guinea) ; Vogt, 1911, Sitzber. Ges. Natf. Fr. Berlin, p. 430. Hylophorbua birooi, van Kampen, 1918, Bijdr. Dierk., XXI, p. 54; idem, 1923, Amph. IndoAustr. Arch., p. 138 ; Nieden, 1926, Das Tierreich, Anura 11, p. 59, figs. 40, 41.

Prevomer with an upturned edge which forms a strong, curved ridge behind eaTh choana ; a denticulated dermal ridge in front of the pharynx preceded by a feebly developed smooth one. Snout obtusely pointed, somewhat longer thamthe diameter of the eye ;canthus rostralis well marked ; loreal region almost vertical ; interorbital space twice as broad as the upper eyelid ; tympanum indistinct, about the diameter of the eye. Fingers with discs, those of the three outer large, truncate distally ; first finger about half the length of the second, which is about as long as the fourth. Toes free, with smaller discs than the fingers ; third much longer than the fifth. Subarticular and inner metatarsal tubercles present, but feebly developed. Tibio-tarsal articulation reaching the shoulder or the tympanum. Skin smooth above, a fine fold from the tip of the snout to the anus. Lower surfaces smooth, except the hinder part of the belly, which is feebly granular. Dirty pink above, finely and densely spotted or marbled with reddish brown ; a dark streak along the sides of the head which may be continued along the sides of the body, with, sometimes, another dark streak along each side of the back. m i t e or yellowish beneath, the throat and chest sometimes dusted with dark brown. Eggs large, connected by a gelatinous cord ; deposited in water (?). Development direct, with no free larval stage ; the developing embryo lacks gills, respiration taking place by means of an expanded leaf-like, non-muscular, vascular tail. Length : 33 mm. DISTRIBUTI,ON : New Guinea (Tor River, Sattelberg).

+

COPHIXALUS B.M. 1901.3.9. 2 Mus. Hung. Amph. 241419. Amph. 240911

juv. in egg

Sattelberg

1 ad., 2 hgr., 43 juvs. (2

Sattelberg

skulls ; anat. prep.) Eggs, 4 juvs.

Sattelberg

. . .

BW Bir6 Bir6

6. Cophixalus verrucosus (Boulenger) 8phenophryne verruwsa, Boulenger, 1898, Ann. MUR. Genova, (2), xvm, p. 707, PI. vm, fig. 2 (Type locality Mt. Victoria, Owen Stanley Range, New Guinea) ; Vogt, 1911, Sitzber. Ges. Natf. Fr. Berlin, p. 431. Oreaphryne verrucosa, Nieden, 1926, Dss Tierreich, Anura rr, p. 43 (part).

Prevomer with an upturned edge which forms a distinct ridge posterior to each choana ; a denticulated dermal ridge in front of the pharynx preceded by a rather indistinct smooth one which may be broken up into three lobes. Snout truncate, scarcely prominent, a little longer than the diameter of the eye; canthus rostralis angular ; loreal region vertical ; interorbital space a little broader than the upper eyelid ; tyrn anum moderately distinct, about half the diameter of the eye. Fingers with discs, t lfose of the three outer large, truncate distally ; first finger about half as long as the second, which is slightly shorter than the fourth. Toes free, with distinct discs which are slightly smaller than those of the fingers ; the third much longer than the fifth. Subarticular and inner metatarsal tubercles present, but indistinct. Tibiotarsal articulation reaching the anterior border of the eye or the tip of the snout. Skin irregularly warty above, the warts sometimes confluent into ridges, of which the most constant are a )(-shaped pair on the anterior part of the back ; sometimes a moderately prominent tubercle about the middle of the edge of the upper eyelid. Smooth beneath, or the hinder part of the belly slightly granular. Brown or olive above, with indistinct darker and lighter spots and rnarblings ; sometimes a W-shaped dark marking on the anterior part of the back, a light cross-bar from eye to eye and a crescentic white spot in front of the groin ; anterior and posterior sides of thighs and posterior surface of tibia dark brown or black, with large, circular, white spots ; a dark transverse bar across the forearm. Lower surfaces dirty white, more or less profusely dusted and spotted with dark brown; belly sometimes with tiny white glandular spots. Length : 33 mm. Eggs large ; male with a vocal sac. : S. New Guinea (Mt. Victoria, Milne Bay), Ferguson Island. DISTRIBUTION B.M. 96.10.31. 37-42 1903.4.30. 22 1904.11.1.67-69

SPECIMENS EXAMINED Mt. Victoria, Owen Stanley Range, Brit. New Guinea Anthony, COTYPES juv. Milne Bay, British New Guinea. . . . . . Meek 9,hgr. Ferguson Island, British Meek New Guinea

66, $29

7. Cophixalus cheesmanae sp. n. Holotype a male, number 1934.1.5.5 in the BritishMuseum from Kokoda (1,200 ft.), British New Guinea ; collected by Miss L. E. Cheesman. Prevomer with an upturned edge which forms a slight ridge behind each choana ; a denticulated dermal ridge in front of the pharynx, preceded by an indefinite, curved, smooth one. Snout acutely pointed, prominent, once and a third as long as the eye ; canthus rostralis rectangular ; loreal region vertical, not concave ; interorbital Space once and three-quarters as broad as the upper eyelid ; tympanum moderately distinct, about half the diameter of the eye. Fingers depressed, the three outer with large,

MONOGRAPH O F THE MICROHYLIDAE truncate discs ; first finger about half as long as the second, with a smaller, rounded disc ; second finger shorter than the fourth. Toes depressed, free, the third much longer than the fifth, with slightly smaller discs than the fingers; subarticular and inner metatarsal tubercles present, but indistinct. Tibio-tarsal articulation reaching midway between the eye and the tip of the snout. Skin feebly shagreened above ; a straight fold from the posterior corner of the eye above the tympanum and arm to the middle of the flanks ; an indistinct tubercle on the middle of the free edge of the upper eyelid and a small papilla on the heel. Posterior half of the belly granular. Brownish yellow above, heavily dusted with black ; sides of head below the canthus rostralis, and temporal region below the supratympanic fold, black. Fore and hinder sides of the thighs and lower surfaces of the tibiae uniform pinkish yellow. Lower surfaces yellow, the chin, chest, and anal region stippled with brown. A fine white line from snout to vent. , A vocal sac present. Length :27 rnm. The paratype is a female from the same locality and differs only in details from the type. The tympanum is more distinct and measures nearly the diameter of the eye ; the tibio-tarsal articulation only reaches the anterior border of the eye, and the sides of the thighs, lower surfaces of the tibiae and inner edge of the tarsus and foot are bright red. Ovarian eggs 2.5 mm. in diameter. Length : 34 mm. SPECIMENS EXAMINED B.M. 1934.1.5. 5 Kokoda, 1,200 ft., British New Guinea Cheesman, TYPE 1934.1.5.6 Kokoda, 1,200 ft., British New Guinea Cheesman, PARATYPE

.

.

8. Cophixalus variegatus (van Kampen) Sphenophryne loriue (part), Boulenger, 1898, Ann. Mus. Genova, (2), xvnr, p. 707. Hylophorbua vuriegalus, van Kampen, 1923, Amph. Indo-Austr. Arch., p. 138 (Type Iocality Digul River, New Guinea).

A denticulated dermal ridge in front of the pharynx, preceded by a shorter, smooth one. Snout obtusely pointed, slightly prominent, as long as, or a trifle longer than, the eye ; canthus rostralis rounded ; loreal region almost vertical ; interorbital space broader than the upper eyelid ; tympanum distinct, nearly ) the diameter of the eye. Fingers, with large discs rather truncate distally ; the first much shorter than the second, which is about as long as the fourth. Toes short, free, with smller discs than the fingers ; the fifth very distinctly longer than the third. Subarticular tubercles very feebly developed; no metatarsal tubercles, or the inner merely indicated. Tibio-tarsal articulation reaching the eye. Skin smooth, or with a few small tubercles on the upper eyelid. Brown above with darker and lighter markings; a fine light line connecting the upper eyelids ; sometimes a yellowish lateral streak from the upper eyelid to the groin giving off dorsally a short, blunt btanch in front of the sacral prominences ; sometimes a W-shaped dark mark on the anterior part of the back, bordered anteriorly by lighter ; a dark spot, bordered anteriorly by lighter, may be present in the groin. Limbs lighter ; a single broad cross-bar on the femur, tibia, and tarsus ; anal region, knee and heel darker. Lower surfaces whitish, more or less profusely dusted with brown. Length : 18 mm. DISTRIBUTION : New Guinea (Digul River, Moroka). SPECIMENS EXAMINED ..

Mus. Genova

ad.

Mus. Amsterdam

ad.

Moroka, British New Guinea . Loria, COTYPEof Sphenoph7yne Zoriae, Boulenger Digul River, New Guinea . TYPE

.

COPHIXALUS

177

9 . Cophixalus rostellifer (Wandolleck) Cop'wla ( 1 ) rostellijer, Wandolleck, 1910, Abh. Mus. Dresden, xm,6, p. 11, figs. 47-49 (Type locality Torricelli Mts., New Guinea) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 135 ; Nieden, 1926, Das Tierreich, Anura 11, p. 57. Choerophryne poboaeidea, van Kampen, 1914, Zool. Jahrb., Syst.. xxxvn, p. 377 (Type locality Njao, Dutch New Guinea) ; idem, 1923, Amph. Indo-Auatr. Arch., p. 136 ; Nieden, 1926, Das Tierreich, Anura n, p. 63.

A denticulated dermal ridge in front of the pharynx, preceded by a curved smooth one.* Snout very long, pointed and projecting far in front of the mouth, twice to twice and a thud as long as the eye, the nostrils close to the tip ; canthus rostralis rounded ; loreal region nearly vertical ; interorbital space twice and a half the width of the upper eyelid ; tympanum rather indistinct, about $ the diameter of the eye. Fingers with large discs ; first shorter than the second, which is a little shorter than the fourth. Toes short, free, with discs scarcely smaller than those of the fingers ; fifth a little longer than the third. Subarticular tubercles and inner metatarsal tubercles present, but feebly developed. Tibio-tarsal articulation reaching the tympanum. Skin slightly warty above, smooth beneath, except the hinder part of the belly and lower surfaces of the thighs, which are distinctly granular. Brown abbve, with darker spots and reticulations, the largest of these having an hour-glass shape on the middle of the back ; a white spot bordered with darker behind, in the lumbar region and a light transverse bar connecting the anterior borders of the upper eyelids. Lower surfaces pale brown with heavy dark vermiculations. Length : 19.5 mm. DISTRIBUTION : New Guinea (Torricelli Mts., Tor River, Njao). Mus. Amsterdam

SPECIMEN EXAMINED ad. (bad state) Tor River, Dutch New Guinea. I o.

Cophixalus oxyrhinus (Boulenger)

Mantophryne lateralis (part), Boulenger, 1897, Ann. Mag. Nat. Hist., (6), XIX, p. 12. Phrynizalus ozyrhinus, Boulenger, 1898, Proc. Zool. Soc. London, p. 480, P1. xxxvm, fig. 3 (Type locality St. Aignan Island, British New Guinea). Copiula ozyrhina, MBhely, 1901, TermBs. Fuzetek, xxrv, pp. 194, 243, P1. v m ; van Kampen, 1906, Nova Guinea, v, Zool. 1, p. 168; idem, 1909, op. cit., rx, Zool, 1, p. 41 ;Vogt. 1911, Sitzber. Ges. Natf. Fr. Berlin, p. 431 ; van Kampen, 1913, Nova Guinea, rx, Zool., 3, p. 462 ; idem, 1914, Zool. Jahrb., Syst., xxxvn, p. 376 ; idem, 1923, Amph. Indo-Austr. Arch., p. 135, fig. 19 ; Nieden, 1926, Das Tierreich, Anura 11,p. 57 ; Burt, 1932, Bull. Amer. Mus. Nat. Hist., ~ x m p. , 483. Prevomer forming a distinct ridge behind each choana ; a denticulated dermal

ridge in front of the pharynx. Snout moderately prominent, bluntly pointed, a little longer than the diameter of the eye ; canthus rostralis obtuse, rounded ; loreal region slightly oblique, concave ; interorbital space about twice as broad as the upper eyelid; tympanum moderately distinct, 8 to 9 the diameter of the eye. Fingers with small (scarcely wider than the penultimate phalanges) discs ; the first shorter than the second which is a trifle shorter than, or subequal to, the fourth. Toes free, with slightly larger discs than the fingers ; the third longer than the fifth ; subarticular and inner metatarsal tubercles present, but poorly developed. Tibio-tarsal articulation reaching to between the eye and the end of the snout. Skin smooth, or finely granular, above ; smooth beneath. Pale brown above, with, or without, small darker spots, which are largest and most numerous on the sides ; sides of the head with a dark brown streak, which may be continued above the tympanum and on the anterior part of the flanks ; groin and Van Kampen states " two denticulated transverse dermal ridges," but in the specimen examined the anterior appears quite smooth. 12

178

MONOGRAPH OF THE MICROHYLIDAE

hinder side of thighs sometimes lighter, with dark brown spots or rnarblings. Lower surfaces white, immaculate or with the throat brown-spotted. Male without vocal sac ; ovarian eggs 3 mrn. in diameter. Length : 37 rnm. DISTRIBUTION : St. Aignan Island, New Guinea. B.M. 98.3.31.12-16

$, $ $

96.10.31. 36

juv.

1901.11. 27.22

$

97.12.10.148,165

$, juv. ad. 2 juvs. ad.

Mus. Amsterdam

St. Aignan Island, S. of Ferguson Island, British New Guinea . . . . . . Meek, COTWES Mt. Victoria, Owen Stanley Anthony, COTYPE Range, New Guinea . . . of Mantophryne lateralis, Boul. Albert Edward Ranges, New Guinea, 6,000 ft. . . . . Rohu Moroka, British New Guinea . Loria Hellwig Mts., New Guinea . . Lorenz Went Mts., New Guinea . . Lorenz Zoutbron (3" 1' 13"S., 140°57' 30" E.), New Guinea . . . van Kampen

APHANTOPHR YNE, Fry Aphanlophryne, Fry, 1917, Proc. Linn. Soc. N. S. Wales, 1916, p. 770 (Type species Aphanlophryne pama, Fry) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 145 ; Noble, 1931, BioL Amph., p. 534.

No teeth. Prevomer large, undivided, in contact with its ,fellow mesially, the post-choanal portion overlying the palatine regi0n.l Clavicles, procoracoids, omosternum and sternum absent (fig. 59). Terminal phalanges T-shaped. Pupil horizontal. Tongue oval, entire and free behind ; a denticulated dermal ridge across the palate in front of the pharynx, preceded by a short, smooth or slightly papillose one. Digits free, without terminal dilatations.

Fig. 59. Ventral elements of the shoulder-girdleof Aphanlophryne pama I.

Aphantophryne pansa, Fry

Aphantophrynepansa, Fry, 1917, Proc. Linn. Soc. N. S. Wales, 1916, p. 772, Pls. LIV, LV, fig. 2(Type locality Mt. Scratchley, Owen Stanley Range, New Guinea, 12,200 ft.) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 145. Inferred from footnote on p. 773 of original description.

MICROBATRACHUS Snout rounded, scarcely prominent, as long as, or slightly shorter than, the diameter of the eye ; canthus rostralis rounded ; loreal region slightly oblique, concave ; interorbital space broader than the upper eyelid ; tympanum slightly distinct, about half the diameter of the eye. Fingers without discs, the first shorter than the second, which is a little shorter than the fourth. Toes short, free, without discs; the fifth a little shorter than the third. Subarticular tubercles not developed ; a very weak inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the axilla. Skin smooth above and below. Uniformly dark brown above ; lower surfaces dark brown, or white, clouded and speckled with dark brown ; anterior part of forearm sometimes yellowish. Length : 27 rnm. DISTRIB~ION : Mt. Scratchley, Owen Stanley Range, New Guinea.

MICROBATRACH US, Roux Microbalrachus, Roux, 1910, Abh. Senck. Natf. Ges., xxxm, p. 228 (Type species Microbatrachus pusillus, Roux) ; van Kampen, 1923, Amph. Indo-Austr. Arch., p. 121 ; Nieden, 1926, Das Tie~~eich,~Anura 11, p. 49.

No teeth. Clavicles and omosternum wanting, procoracoids slender, reaching the scapulae and the mid-line of the shoulder-girdle; sternum cartilaginous. Pupil horizontal. Tongue entire and free behind. A denticulated dermal ridge across the palate in front of the pharynx. Digits free, the toes slightly dilated distally. This genus may, as has been suggested, prove to have been founded on an immature O~eophyneor Sphenophyne, more probably the latter.

I.

Microbatrachus pusillus, Roux

Microbatrachus pmillus, Roux, 1910, Abh. Senck, Nat. Ges., xxxm, p. 228, P1. xrv, figs 6-6b (Type locality Pobdjetur, Terangan, Aru Islands); van Kampen, 1923, Amph. Indo-Austr. Arch., p. 121; Nieden, 1926, Das Tierreich, Anura 11, p. 50.

Snout short, rounded, as long as the diameter of the eye ; canthus rostralis rounded ; nostril midway between the eye and the end of the snout ; interorbital space once and two-thirds as broad as the upper eyelid ; tympanum hidden. Fingers very short, obtuse, the first shorter than the second, which is shorter than the fourth. Toes free, with distinct discs; fifth shorter than the third ; no subarticular or metatarsal tubercles. Tibio-tarsal articulation reaching the hinder border of the eye. Skin smooth above and below. Brown above, the head and anterior part of the back darker; arms pale grey; legs yellowish brown. Yellowish white beneath ; the chin darker, with small lighter flecks. Length : 7 mm. DISTRIBUTION : Known only from the type locality, Pobdjetur, Terangan, AN Islands.

BREVICIPITINAE, Cope Brevicipitidae, Cope, 1867, Journ. Acad. Philadelphia, (2), W,p. 191. Brevicipitinae + Dyscophinae (part), Noble, 1931, Biol. Amph., pp. 528, 539.

Maxillary and vomerine teeth absent ; prevomer large, with a median expansion, but no post-choanal portion (fig. 60) ; palatine absent ; ethmoid absent ; ear fully developed ; vertebral column diplasiocoelous. No free-swimming tadpole, metamorphosis occurring on land ; the lawa lacks adhesive organs, external gills, gill clefts (?),

180

MONOGRAPH O F T H E MICROHYLIDAE

spiracle and teeth ; an operculum is present ; the tail is muscular and the larva leaves the egg before metamorphosis. This exclusively African group of genera differs from all the other members of the family in the condition of the prevomer, which, though reduced behind the choanae, still retains a large median expansion. All the genera are closely allied, but do not appear to be particularly closely related to any other existing genera, and may be considered as an independent Ethiopian line of descent from some primitive, toothless, Dyscophine ancestor. The specialised breeding habits resemble, but do not exactly parallel, those of the Asterophryinae and Sphenophryninae. SYNOPSIS OF THE GENERA

I. Coccyx articulating by two condyles. A. Terminal phalanges T-shaped . B. Terminal phalanges simple . .

. . . . . . . Callulina p. 180 . . . . . . Spelaeophryne, p. 182

11. Coccyx and sacrum fused. A. A series of pharyngeal folds, but no glandular area on the palate Probreviceps, p. 183 B. A large glandular area on the palate in front of the pharyngeal folds B ~ ~ c e pp.s 186 ,

CALLULINA, Nieden Callulina. Nieden, 1910, Sitzber. Ges. Natf. Fr. Berlin, x, p. 449 (Type species Calldim &#ti, Nieden) ;Noble, 1924, Bull. Amer. Mus. Nat. Hist., XLIX, p. 277 ;Nieden, 1926, Das Tierreich, Anura n, p. 7 ;Noble, 1931, Biol. Amph., p. 529.

Prevomer a thin expanded plate, nearly in contact with its fellow mesially, but not bordering the choana posteriorly (fig. 60). Clavicles and procoracoids present, welldeveloped, straight. reaching the mid-line of the shoulder girdle and the scapulae; coracoids moderately dilated; omosternum small, cartilaginous ; sternum cartila$nous (fig. 61). Vertebral column diplasiocoelous ; coccyx articulating by two condyles. Terminal phalanges T-shaped. Pupil horizontal. Tongue large, subcircular, entire and very slightly free behind ; three strong ridges across the palate in front of the pharynx, the posterior straight and denticulate, the middle curved, smooth, and the anterior chevron-shaped, with the apex in front, and smooth. Digits free, with small, but distinct, triangular terminal dilatations ; inner two toes opposable to the remainder.

Fig. 60. Anterior cranial elements of Cdlulina krefli (seen from beneath)

Fig. 61. Ventralklements of the shoulder-girdleof Cdlulino krefli

I.

Callulina kreffti, Nieden

Cdlulina krefli, Nieden, 1910, Sitzber. Ges. Natf. Fr. Berlin, x, p. 449 (Type localities Amani and Tanga, E. Africa); idem, 1915, Mitt. Zool. Mns. Berlin, vn, p. 380; Noble, 1924, Bull. Amer. Mus. Nat. Hist., XLIX, p. 345 ; Nieden, 1926,Das Tierreich, Anura n,p. 7, fig. 7 ;Barbour and Loveridge, 1928,Mem. Mus. Comp. Zool. Harvard, L, 2, p. 241 ; Loveridge, 1930, Proc. Zool. Soc. London, (I),p. 16. Snout short, rounded, not prominent, about as long as the eye ; canthus rostralis rounded ; loreal region rather oblique ; interorbital space as broad as (juveniles), or

broader than (adults), the upper eyelid. Tympanum hidden or very indistinct. Fingers moderate, with distinct, triangular dilatations ; the first shorter than the second, which is shorter than the fourth. Toes dilated distally, the discs of the inner two toes about as large as those of the fingers, those of the outer three distinctly smaller ; first and second toes subequal, opposable to the remainder ; fifth extending well beyond the third. Subarticular tubercles well developed ; two flat, oval, metatarsal tubercles. Tibio-tarsal articulation reaching to the middle of the flank (juveniles) or to the tympanic region. Skin thick, very rough and warty above ; granular beneath. Grey-brown to blackish above, irregularly mottled with darker and lighter. Sometimes a lighter area from behind the eye to the angle of the mouth; yellowish or whitish beneath, indistinctly washed with grey ; throat more or less infuscate. Ovarian eggs large, 3 rnrn. ; male with a vocal sac. Length : $! 47 mm. ; $ much smaller. DISTRIBUTION : Usambara and Uluguru Mts., E. Africa (Tanganyika Territory). B.M. 1929.6.1.8 1929.6.1. 7 Mus. Berlin 21778

$! $!

9

SPECIMENS EXAMINED Vituri, Uluguru Mts., E. Africa Amani, E. Africa . . . . Amani, E. Africa . . . .

. . . . . . . . .

Loveridge Loveridge Kreflt, TYPE

MONOGRAPH O F T H E MICROHYLIDAE

SPELAEOPHRYNE, Ah1 Spelueophryne, Ahl, 1924, Zool. Anz., LXI, p. 99 (Type species Spelaeophrym methneri, Ahl) ; Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Harvard, L, 2, p. 243 ;Noble, 1931, Biol. Amph. p. 539.

Prevomer a thin expanded plate, almost in contact with its fellow mesially, but not bordering the choana posteriorly (fig. 62). Clavicles and procoracoids present, well developed, straight, reaching the mid-line of the shoulder girdle and the scapulae ; coracoids moderately dilated ; omosternum rudimentary ; sternum cartilaginous. (fig. 63). Vertebral column diplasiocoelous ; coccyx articulating by two condyles. Terminal phalanges simple.

Fig. 62. Anterior cranial elements of Spelaeophryne methnevi (seen from beneath)

Fig. 63. Ventral elements of the shoulder-girdle of Spelueophryne methneri

PROBREVICEPS Pupil horizontal. Tongue oval, scarcely free behind. A denticulated dermal ridge across the palate in front of the pharynx, preceded by a curved smooth one. A dermal papilla between the choanae and a similar structure beneath each Harderian gland. Digits free, not dilated distally. The swellings beneath the Harderian glands are, apparently, homologous with the anterior palatal fold of Callulina.

I.

Spelaeophryne methneri, Ah1

8pelaeqphryne methneri, Ahl, 1924, 2001. Anz., LXI, p. 99 (Type locality Nangoma Cave, Matumbi, E. Africa) ; Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Harvard, L, 2, p. 243, PI. n, fig. 6 ; Loveridge, 1930, Proc. Zool. Soc. London, (I), p. 14.

Snout subconical, slightly prominent, once and a half as long as the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical ; interorbital space once and a half as broad as the upper eyelid ; tympanum distinct, 3 the diameter of the eye. Fingers well developed, the first a little shorter than the second, which is subequal to the fourth. Toes well developed, the outer much shorter than the third but extendiflg beyond the second. Subarticular tubercles prominent; a prominent oval inner, and a smaller, rounded outer, metatarsal tubercle. Tibio-tarsal articulation reaching the elbow. Skin smooth, but very thick and glandular, above; smooth beneath. Purplish black above ; a broad, white (red in life) band extends from the tip of the snout (where it meets its fellow) above the canthus rostralis, along the upper eyelid to above the tympanum or the level of the shoulder ; brownish beneath, the edges of the lower jaw darker. Male with a vocal sac. Length : 53 mm. ; 52 mm. DISTRIBUTION : East Africa (Tanganyika Territory).

B.M. 1928.5.1.5 Mus. Berlin 2756

Nyange, Uluguru Mts., E. Africa . Nangoma Cave, Matumbi, E. Africa

. . Loveridge . . Methner, TYPE

PROBREVICEPS, Parker Breviceps (non Memem), Tornier, 1906, Kriechth. Deutsch-0. Afrika, III, 4, p. 160 ; Nieden, 1926, Das Tiemeich, Anura 11, p. 4 (part) ; Barbour and Loveridge, 1928, Mem. Mus. Comp. 2001. Harvard, L, 2, p. 246 (part) ; Loveridge, 1930, Proc. Zool. Soc. London, (I), p. 15. Probreviceps, Parker, 1931, Ann. Mag. Nat. Hist., (lo), vm,p. 262 (Type species Breviceps macrodactylus, Nieden) ; de Villiers, 1933, h a t . Anz. u x v , p. 257.

Prevomer a thin expanded plate in contact with its fellow mesially, but not bordering the choana posteriorly. Clavicles and procoracoids present, well developed, straight, reaching the mid-line of the shoulder girdle and the scapulae ; coracoids moderately dilated ; omosternum small, cartilaginous ; sternum cartilaginous. Vertebral column diplasiocoelous ; coccyx and sacrum fused. Terminal phalanges simple. Pupil horizontal. Tongue large, subcircular, scarcely free behind, with a median furrow. A denticulated dermal ridge across the palate in front of the pharynx, preceded by a similar, narrow, strongly curved one ; a dermal papilla between the choanae. Digits free, not dilated distally. In all the known species the first finger is a little shorter than the second, which is approximately equal to the fourth ; outer toe shorter than the third, but extending well beyond the second.

MONOGRAPH O F T H E MICROHYLIDAE SYNOPSIS OF THE SPECIES

I. Tympanum visible ; tibio-tarsal articulation reaching the shoulder or beyond. A. Tympanum as large as the eye . . . P. macrodactylus runpensis, p. B. Tympanum Q the diameter of the eye. P. macrodactylus macrodactylus, p. C. Tympanum half the diameter of the eye . P. macrodactylus loveridgei, p. 11. Tympanum hidden ; tibio-tarsal articulation reaching the elbow or the shoulder . . . . . . . . . . . . . . P. uluguruensis, p. I.

184 184 185 185

Probreviceps marnodactylus rungwensis, Loveridge

Probreviceps nacroddylus ruqwenais, Loveridge, 1932, Bull. Mus. Comp. Zool. Harvard, LXXII, 10, p. 387 (Type locality Nkuka Forest, Rungwe Mt., E. Africa). Snout prominent, pointed, a shade longer than the diameter of the eye ; canthus rostralis angular ; loreal region vertical, flat ; interorbital space about 13 times as

broad as the upper eyelid; tympanum distinct but less well marked in adult than in juvenile specimens, its diameter equal to, or a little greater than, that of the eye. Fingers short. Toes moderate, the fourth very long; subarticular tubercles well developed ; a flat oval inner, and an indistinct outer, metatarsal tubercle. Tibio-tarsal articulation reaching the shoulder or the tympanum. Skin beset with numerous small conical warts above ; granular beneath, the granules becoming smaller but more prominent on the chin and thighs. Uniform purplish brown above ; pale brown with darker spots and marblings beneath ; throat uniform purplish brown. Ovarian eggs large, 3.5 mm. ; male with a vocal sac. Length : 58 mm. : Nkuka Forest, Rungwe Mt., Tanganyika Territory. DISTRIBUTION SPECIMENS EXAMINED Nkuka Forest, Rungwe Mt., E. Africa . . . . . Loveridge, TYPE 16462-333 Nkuka Forest, Rungwe ~ t . , E. Africa. . . . . . Loveridge, PARATYPES B.M. 1931.12.13. 1 $! Nkuka Forest, Rungwe Mt., . . Loveridge, PARATYPE E. Africa. . . M.C.Z. Haward 16461 $!

.

2.

Probreviceps macrodactylus macrodactylus (Nieden)

Breviceps vewvcosus ( m n Rapp), Tornier, 1896, Kriechth. Deutsch-0. Afrika, p. 160 ; Nieden, 1910, Sitzber. Ges. Natf. Berlin, p. 449 ; idem, 1915, Mitt. Zool. Mus. Berlin, p. 380. Brewicepa macrodactylus, Nieden, 1926, Das Tierreich, Anura n, p. 6 (Type locality Usambara,, E. Africa). Probreviceps macrodactylus macrodactylus, Parker, 1931, Ann. Mag. Nat. Hist., (10). w,p. 262. Breviceps usambaricus, Barbour and Loveridge, 1928, Mem. Mus. Comp. 2001. Harvard, L, 2, p. 251 (Type locality Amani, Usambara Mts., E. Africa) ; Loveridge. 1930, Proc. Zool. Soc. London, p. 16. Snout moderately pointed, slightly prominent, as long as the diameter of the eye ; canthus rostralis rounded ; loreal region nearly vertical, very slightly concave ; interorbital space about once and a third as broad as the upper eyelid ; tympanum

distinct, though sometimes ill-defined, about f the length of the eye. Fingers moderate. Toes moderate, the fourth very long ; subarticular tubercles well developed ; a large oval inner, and an ill-defined flat outer, metatarsal tubercle. Tibio-tarsal articulation reaching to between the shoulder and the tympanum. Skin rugose or granular above ; smooth or slightly granular beneath.

PROBREVICEPS Uifnorm;dark, purplish brown above ; pale brown beneath, with numerous small round, whitish spots ; throat coloured like the upper surfaces, but light-spotted like the belly. Male with a vocal sac ; ovarian eggs large, 4-5 rnm. Length : 65 rnrn. DISTRIBUTION : Usambara Mountains, Tanganyika Territory, from ~ , W , 0 0 0 ft. B.M. 1928.5.1. 7 1930.10.3. 2

M.C.Z. Haward 15709-15718 Mus. Berlin 11362

I

$2

Amani, Usambara Mts., E. Loveridge Africa . . . . . PARATYPES hgr. 6 Amani, Usambara Mts., E. of B~ezn~ceps Africa . . . . . Loveridge usambaricus, Barbour and 6 6, $2 9 Amani, Usambara Mts., E. Loveridge Africa . . . . . Loveridge 2 ads. Amani . . . . . . Krefft, COTYPES

3. Probreviceps macrodactylus loveridgei, Parker b

Breviceps m q o m (non Power), Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Haward, L, 2, p. 249 ;Loveridge, 1930, Proc. Zool. Soc. London, (I), p. 16. Probrevicepa ntacrodaetyha loveridgei, Parker, 1931, AM. Mag. Nat. Hist., (lo), vm, p. 263 (Type locality Bagilo, Uluguru Mts., E. Africa). Snout moderately pointed, slightly prominent, as long as the diameter of the eye ; canthus rostralis distinct ; loreal region nearly vertical, concave ; interorbital space about as broad as the upper eyelid ; tympanum distinct, its diameter about half the

length of the eye. Fingers moderate. Toes moderate, the fourth very long; subarticular tubercles well developed ; flat oval inner, and a rounded outer, metatarsal tubercle ; the inner metatarsal tubercle is relatively smaller in juveniles. Tibio-tarsal articulation reaching the tympanum or the eye. Skin beset with numerous small, subconical warts above ; uniformly granular beneath. Uniform purplish brown above ; brown beneath, with a faint darker reticulum. Male with a vocal sac. Eggs large (6 mm., including the capsules), laid in burrows on land. 46 mm. ; 6 34 mrn. Length : DISTRIBUTION : Bagilo, Vituri, and Nyange, Uluguru Mts., E. Africa (Tanganyika Territory). B.M. 1928.5.1. 8 1930.10.3. 1 M.C.Z. Haward 15701-15708 Mus. Kimberley

? 6 6d,? ?

. . Loveridge, TYPE . . Loveridge Vituri, Uluguru Mts. . . Loveridge PARATYPES Bagilo, Uluguru Mts. . . Loveridge) Bagilo, Uluguru Mts. Vituri, Uluguru Mts.

1

4 . Probrevkeps ulquruensis (Loveridge) Brewicepa uluguruensis, Loveridge, 1925, Proc. Zool. Soc. London, p. 789, P1. I, fig. 3 (Type locality Bagilo, Ulugaru Mts.) ; Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Harvard, L, 2, p. 247 ; Loveridge, 1930, Proc. Zool. Soc. London, p. 16. Probreviceps dugurwensie, Parker, p. 257. 1931, Ann. Mag. Nat. Hist., (lo), vm,p. 264 ;de Villiers, 1933, Anat. Am., ~ x x v ,

Snout short, moderately pointed and very slightly prominent, a little shorter than the eye ; canthus rostralis rounded ; loreal region slightly oblique ; interorbital space as broad as the upper-eyelid ; chin swollen. Fingers moderate, with a distinct fleshy ridge along the inner side of the second and third. Fourth toe very long ; subarticular

186

MONOGRAPH O F THE MICROHYLIDAE

tubercles moderately well developed ; a flat, oval inner, and an indistinct outer, metatarsal tubercle. Tibio-tarsal articulation reaching the elbow or the shoulder. Skin of the upper surfaces beset with small warts ; lower surfaces granular. Cream to terra-cotta coloured above, sometimes washed with purple ; juveniles with traces of a light dorso-lateral band on each side. Flanks and lower surfaces purplish brown, with some indistinct lighter blotches. The eggs are large (5.5 mm., including the capsules) and are laid in burrows on land. Length : $2 45 mm. ; 3 30 rnrn. DISTRIBUTION : Bagilo, and Nyingwa, Uluguru Mts. (6,00&8,000 R.), E. Africa (Tanganyika Territory). SPECIMENS EXAMINED Nyingwa, Uluguru Mts. Loveridge B.M. 1931.6.6. 1-5 4 9 9, juv. M.C.Z. Haward 1 0 3 d,? 9, h g ~ . Nyingwa, Uluguru Mts. Loveridge Mus.Kimberley $2 Nyingwa, Uluguru Mts. Loveridge

BRE VICEPS, Merrem R a m (part), Linn., 1758, Syst. Nat., ed. 10, p. 211. Bufo (part), Laurenti, 1768, Syn. Rept., p. 25 ; Schneider, 1799, Hist. Amph., I, p. 111 ; Daudin, 1803, Hist. Rainettes, p. 71. Breviceps, Merrem, 1820, Syst. Amph., p. 177 (Type species R a m gibbosa, Linn.) ; Gravenhorst, 1829, Delic. Mus. Vratis., p. 69 ; Dumbd and Bibron, 1841, Erpet. Gbn., vm,p. 752 ; Giinther, 1858, Cat. Batr. Sal. Brit. Mu., p. 52 ; Steindachner, 1869, Reise Novara, I, Amph., p. 38 ; Peters, 1882, Reise. n. Mossambique, p. 175 ;Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 176 ;i L m , 1910, Ann. S. Afr. Mus., v, 9. p. 534 ;Hewitt, 1925, Ann. Natal. Mus., v, 2, p. 189 ; Power, 1926, Ann. S. Afr. Mus., xx, 6, p. 451 ;Nieden, 1926, Das Tierreich, Anura n, p. 4 (part). Engystoma, Fitzinger, 1826, Class. Rept., pp. 39, 65 (Type species Rana gibbosa, Linn.) ; Stejneger, 1910, Proc. Biol. Soc. Washington, xxIn, p. 166. Systoma, Wagler, 1830, Syst. Amph., p. 205 (Type species Breviceps gibbosua, Memm =R a m gibbosa, Linn.);Tschudi, 1838, Mem. Soc. Sci. Neuchktel, 1839, n, p. 86 (part) ;Fitzinger, 1843, Syst. Rept., p. 33 (part).

Prevomer a thin expanded plate in contact with, or narrowly separated from, its fellow mesially, but not bordering the choana posteriorly (fig. 64). Clavicles and procoracoids present, well developed, straight, reaching the mid-line of the shouldergirdle and the scapulae ; coracoids strongly dilated ; omosternurn vestigial ; sternum cartilaginous (fig. 65). Vertebral column diplasiocoelous ; seven presacral vertebrae ; coccyx and sacrum fused. Tenninal phalanges simple. Pupil horizontal. Tongue large, oval, entire and scarcely free behind. a denticulated dermal ridge across the palate in front of the pharynx, preceded by a large glandular area occupying the centre of the roof of the mouth ; a dermal papilla between the choanae and sometimes a pair of glandular swellings behind each choana. Digits free, not dilated.

Fig. 64. Anterior cranial elements of Brevicep a h p e r m (seen from beneath)

Fig. 65. Ventral elements of the shoulder-girdleof Brevieeps adsperm

SYNOPSIS OF THE SPECIES

I. Outer toe extending beyond the level of the union of the third and fourth. Length of eye contained more than ten times in the length from snout to vent. Outer finger extending as far as the distal subarticular tubercle of the third. A. Tympanum more or less visible ; outer toe extending slightly beyond the second. . . . . . . . . . . . . B. tympanifer, p. B. Tympanum hidden ; outer toe not extending beyond the second. (1) Dorsal surfaces beset with numerous small, conical, almost spinose tubercles ; uniformly brown above. . B. fusczcs, p. (2) Dorsal surfaces smooth, or coarsely granular ; upper surfaces with dark, or light and dark, markings. (a) Skin granular; outer finger about 4 the length of the second. (i) First finger shorter than the second . . B. gibboswr, p. (ii) First fmger a little longer than the second B. sylvestris, p. ( b ) Skin almost smooth ; outer fmger the length of the second B. rosei, p.

+

188

189

190 189 191

11. Outer toe not reaching beyond the level of the union of the third and fourth. Length of eye contained less than 9 times in the length from snout to vent. A. Outer finger reaching the level of the distal subarticular tubercle of the third, and measuring or more the length of the second. ( 1 ) Outer toe moderately well developed, with a subarticular tubercle ; length of eye contained less than 7 times in the total length. . . . . . . . . . B. namaquensis, p. 191 ( 2 ) Outer toe rudimentary, without a subarticular tubercle ; length of eye contained more than 7 times in the total length. (a) Belly smooth. B. penthen', p. 192 ( b ) Belly densely granular B. montanus, p. 192

. . . . . . . . . . . . . . . .

MONOGRAPH OF THE MICROHYLIDAE B. Outer finger reaching midway between the subarticular tubercles of the third finger, or not so far, and measuring less than 8 the length of the second. (1) Fingers and toes all distinct, not bound together ; second finger more than half as long as the eye. (a) Outer finger measuring to Q of the second ; outer toe about as long as the inner. (i) Skin warty above ; back with a row of light spots on each side of the vertebral line and traces of a lateral row of similar spots ; throat with two longitudinal dark bars. . . . . . . . . B. adspersus, (ii) Skin pitted, but not warty, above ; no light spots on the back, but perhaps some dark streaks ; throat uniformly infuscate . . . . . B. mossmlricw, (b) Outer finger measuring of the second ; outer toe minute B. pmeri, (2) Two inner toes bound together; second finger measuring less than half the diameter of the eye . . . . . B. macrops,

p. 193 p. 194 p. 195 p. 196

Breviceps tympanifer, Hewitt

I.

Brevicepa tppanijer, Hewitt, 1925, Ann. Natal Mus., v, 2, p. 190, PI. x, figs. 1-2 (Type locality Pirie, near Kingwilliamstown, S. Africa) ; Power, 1926, Ann. S. Afr. Mas.,xx, 6, p. 463, Pls. xxxrx, fig. 5, XL, fig. j, XLI, fig. E, XLU, fig. 7 ; Essex, 1928, Ann. Mag. Nat. Hist., (lo), I, pp.

265-268 ;Hewitt, 1932, Ann. Natal Mus., vn, 1, p. 106. Breviceps ncgom, Power, 1926, Ann. S. Afr. Mus., xx, 6, p. 467, Ple. x x m , fig. 6, xr, fig. h, x m , 6gs. 8, 9 (Type locality Mariannhill, Natal) ; Loveridge, 1930, Proc. 2001. Soc. London, (l),

P. 16 (part). Snout short, rounded, not prominent, about 8 the diameter of the eye ; interorbital space a little broader than the upper eyelid ; tympanum more or less distinct, about Q the diameter of the eye. Fingers short, the first shorter than the second ; fourth reaching the level of the distal subarticular tubercle of the third and measuring Q to the length of the second. Toes short, the outer relatively well developed, with a subarticular tubercle, and extending beyond the second. Subarticular tubercles well developed. A large, oval, inner, and a flat, rounded, outer, metatarsal 9bercle. Skin granular and porous above, granular beneath. Dark brown above ; snout, from the middle of each upper eyelid, paler ; a broad mid-dorsal area anteriorly yellowish, mottled and spotted with darker ; sometimes also a light line posteriorly, which is joined above the vent by another light line that passes along the posterior side of the hind-limb t o the base of the outer toe. A dark streak from beneath the eye to the anterior face of the fore-limb, bordered in front and behind by broad lighter patches ; sometimes a narrow, pale, dorso-lateral streak. Yellowish white beneath, sometimes mottled or streaked with pale brown on the throat and towards the flanks. Length : 51 mrn. DISTRIBWION : S.E. Cape Province to Natal.

B.M. 1903.4.8.24 1903.6.19. 5 Mus. Kimberley

9 $ ad.

.

. . Cregoe . Stenning . . . . Felix, TYPEof B~evi-

Durban . . . . . Pirie Bush, Kingwilliamstown Mariannhill, Natal. .

ceps rugoszrs, Power

BREVICEPS *

2.

Breviceps fuscus, Hewitt

Breviceps veerrumsus (non Rapp), Boulenger, 1907, Roc. Zool. Soc. London, p. 480. Breviceps gibbosus (part), Boulenger, 1910, Ann. S. Afr. M u . , v, 9, p. 534. Brevieeps fuscus, Hewitt, 1925, Ann. Natal Mus., v, 2, p. 191, P1. x, fig. 5 (Type locality Knysna, S. Africa); Power, 1926, Ann. S. Bfr. Mus., xx, 6, p. 462, Pls. xxx~x,XL, jig. d, =I, fig. D ; Essex, 1928, Ann. Mag. Nat. Hist., (lo), I, pp. 265, 268 ; FitzSimons, 1930, Ann. Transvaal Mus., XIV, 1, p. 48 ; de Villiers, 1931, Anat. Anz., L x x n , pp. 164-178.

Snout short, rounded, not prominent, about $ the diameter of the eye ; interorbital space broader than the upper eyelid; tympanum hidden. Fingers short, the first shorter than the second ; fourth reaching the level of the distal subarticular tubercle of the third and measuring f to $ of the second. Toes short, the outer relatively well developed, with a subarticular tubercle, extending to the level of the proximal tubercle of the fourth, but not as far as the tip of the second ; subarticular tubercles well developed; inner metatarsal tubercle large, oval, oblique ; outer smaller, flat, scarcely separated from the inner. Skin uniformly thickly beset with small, regular, conical tubercles above ; similar tubercles on the lower surfaces, but less numerous except on the chin and throat. Uniform blackish brown above ; lower surfaces paler except the chin and throat, which are infuscate. Length : 57 mm. DISTRIBUTION : Districts of Knysna and George, S.W. Cape Province. B.M. 92.6.27. 2

juv.

1907.4.9. 60-61 $, 1910.7.28. 5 $ Mus. Kimberley 578 9 569 $

9

.

Cape of Good Hope " . . Knysna, Cape Province, S. Africa Cape Province, S. Africa . . Knysna, Cape Province, S. Africa Knysna, Cape Province, S. Africa "

. (Christiania Mus.) . Grant . (S. Afr. .

Mus.)

.

3 . Breviceps sylvestris, FitzSimons Breviceps sylv,estris, FitzSimons, 1930, Ann. Transvaal Mus., locality Woodbush, Pietersburg District, Transvaal).

XIV,

pp. 46 and 48, figs. 27129 (Type

Snout short, rounded, not prominent ; interorbital space considerably broader than the upper eyelid ; tympanum hidden. Fingers short, the first a little longer than the second ; fourth reaching the level of the distal subarticular tubercle of the third and measuring $ of the second. Toes short, the outer reaching the proximal subarticular tubercle of the fourth, but not extending as far as the second ; subarticular tubercles well developed, prominent. Inner metatarsal tubercle large, oblique, compressed, with a digging edge ; outer large, subcircular, flat, distinct from the inner. Dorsal surfaces porously granular ; a distinct granular ridge or fold on either side of the middle line from the posterior angle of the eye-cleft to about the middle of the back ; granular beneath, the granules smaller and more closely set anteriorly. Dark chocolate brown above, with very faint, irregular splashes of dirty yellowish white ; the space between the granular ridges well defined by uniform darker colour along its outer edges. Flanks with large, irregular yellowish spots. Lower surfaces dirty buff, with dense argus-brown vermiculations on the belly which become dense anteriorly, the chin and throat being uniformly brown. A wide, dark-brown band from beneath the eye to the breast. Eggs laid in a subterranean chamber in which metamorphosis takes place. Length : 4.4 mm. DISTRIBUTION : Forested regions of Pietersburg and Zoutpansberg Districts, N. Transvaal.

MONOGRAPH OF THE MICROHYLIDAE

4 . Breviceps gibbosus (Linn.) [Bufo acephulus, Klein, 1751, Quadrup. Dispos., p. 121.1 Rana gibbosa, Linn., 1758, Syst. Nat., ed. 10, p. 21 1 (Type locality ?). Bufo gibbosus, Laurenti, 1768, Syn. Rept., p. 27 ; Daudin, 1803, Hist. Nat. Rainettes, p. 80, P1. x m x , fig. 2 ; idem, 1803, Hist. Nat., w, p. 158. Breviceps gibbosus, Merrem, 1820, S p t . Amph., p. 178 ; Gravenhorst, 1829, Delic. Mus. Vratialav, p. 69 ; Dum6ril and Bibron, 1841, Erpet. Gn., vm, p. 754 ; Smith, 1849, Zool. S. Afr., I U , Appesdix, p. 27; Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 53 ; Peters, 1882, Reise n. Mossambique, Zool., Amph., p. 177 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 176, fig. ; idem, 1884, Ann. Mag. Nat. Hist., (5), m,p. 388 ; idem, 1910, Ann. S. Mr. Mm., v, 9, p. 534 (part); Beddard, 1911, Proc. Zool. Soc. London, p. 404; Noble, 1924, Bull. Amer. Mus. Nat. Hist., X L I X ,p. 344 (part) ; Nieden, 1926, Das Tierreich, Anura n, p. 5 (part) ; Power, 1926, Ann. S. Afr. Mus., x x , 6, p. 460, Pls. x x x x , fig. 4, z, fig. b ; X L I ,fig. c, X L ~ fig. i ; Essex, 1928, Ann. Mag. Nat. Hist., (lo), I , pp. 265, 268; Loveridge, 1930, Proc. Zool. Soc. London, (I), p. 16 (part). Engystoma gibbosa, Fitzinger, 1826, Class. Rept., p. 65 ; Henle, 1839, Verg. An. Beschr. Kehlk. Rept., p. ? (not seen) ; Sys2oma gibbosum, Wagler, 1830, Syst. Amph., p. 205 ; Fitzinger, 1843, Syst. Rept., p. 33 ;Parker, 1868, Mon. Shoulder Girdle, p. 69, P1. vn, fig. 9. R a m brewiceps, Schneider, 1799, Hist. Amph., I , p. 140 (substitute name for R a m gibbosa, Linn.) ; Shaw, 1802, Gen. Zool., nI, p. 170, PI. LU. Systoma breviceps, Tschudi, 1838, Mem. Soc. Sci. NeuchLtel, n, 1839, p. 86. Engystoma dorsalum, Cuvier, 1829, RAgne Animal, ed. 2, n, p. 112 (substitute name for Bufo gibbosus, Daud.). ? Breviceps verrumsus, Rapp, 1842, Arch. f. Naturg., vm, 1, p. 291, P1. v r , fig. 5 (Type locality Natal) ;Steindachner, 1869, Reise Novara, Zool. I , Amph., p. 38 ;Peters, 1882, Reise n. Mossambique, Zool., Amph., p. 177 ; Werner, 1910, Jena Denksch., xvr, p. 293 (part).

Snout very short, vertically truncate, not prominent; interorbital space broader than the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second ; fourth reaching the level of the distal subarticular tubercle of the third and measuring 8 to $ of the second. Toes short, the outer reaching the proximal subarticular tubercle of the fourth, but not extending as far as the second; subarticular tubercles well developed ; inner metatarsal tubercle large, oblique, compressed, with a digging edge ; outer flat, scarcely distinct from the inner. Dorsal surfaces porous and warty ; uniformly granular beneath. Brown above with darker and lighter markings; usually a curved light bar connecting the anterior halves of the upper eyelids ; light spots forming an indefinite longitudinal series on each side of the vertebral line, and sometimes fusing to form a broad light median area with festooned lateral margins ; another series of light spots arranged in a longitudinal row dorso-laterally may also be present ; an-oblique dark bar from beneath the eye to the anterior side of the fore-limb constantly present and bordered anteriorly and posteriorly by a broad lighter area. Lower surfaces whitish, irregularly spotted with brown. Male with a vocal sac. Length : 74 rnm. DISTRIBUTION : Cape Peninsula. The status of B. vewucosus, Rapp., from "Natal" is still somewhat uncertain. Prof L. Miiller has very courteously examined the type, now preserved in Stuttgart, and from the information supplied by him, and from the descriptions, it is evident that we have to do with a species falling into section IB of the preceding key ; the general characters and colour agree well with B. gibbosus (Linn.), but this species is not known to extend into Natal. SPECIMENS EXAMINED B.M. R.1931.1.1. 1 9 ? . . . . . . . . . . (Damon) 56.11.28.11 hgr. " S. Africa . . . . . . . . Brandt 90.5.20. 6 ? " Cape of Good Hope " . . . . . Fisk 1920.6.3. 1-6 8 8 , ? 9 '' Cape Colony " . . . . . . . (Zool. Soc.) Mus. Kimberley 2 ads. Cape Peninsula . . .. . . . .

,

5 . Breviceps rosei, Power Breviceps rosei, Power, 1926, Ann. S. Afr. Mus., xx, 6, p. 466, Pls. XXXIX, fig. 10, XL,fig. i, XLII, figs. 1, 2, x m , fig. 2 (Type locality Lakeside, Cape Penins., S. Africa).

Snout short, obliquely truncate, slightly prominent ; interorbital space about equal to the width of the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second ; fourth reaching the level of the distal subarticular tubercle of the third, and measuring 8 of the second. Toes short, the outer reaching the level of the proximal tubercle of the fourth, but not extending as far as the second. Subarticular tubercles well developed. Inner metatarsal tubercle large, oblique and compressed, with a digging edge ; outer flat, subcircular. Upper surfaces almost smooth, with some flat, indistinct warts; forehead, lips, sides of head and lower surfaces finely granular. Dark above, with indistinct brown markings; a brown patch on the forehead between the eyes, and another on each side of the head behind the eye ; whitish beneath, the belly speckled with black, and the chin and throat with heavy black verrniculations which form a pair of indefinite, broad, longitudinal stripes somewhat divergent posteriorly. Length : 36-5 mm. : Lakeside, Cape Peninsula. DISTRIBUTION Possibly this species is based on,an aberrant, smooth-skinned B. gibbosus. Mus. Kimberley

hgr. (?)

SPECIMENEXAMINED Lakeside, Cape Peninsula

. . . . . Rose, TYPE

6 . Breviceps namaquensis, Power Brevieeps ahperms (nonPeters), Boulenger, 1910, Ann. S. Afr. Mus., V, 9, p. 534 ; Noble, 1924, Bull. p. 344. Amer. Mus. Nat. Hiat., XLIX, ? Breviceps aerrueosus (part),Werner, 1910. Jena Denksch., XVI,p. 293. Breviceps mssambicua (part),Nieden, 1926, Das Tierreich, Anura n, p. 5. Brevieeps namaquensia, Power, 1926, Ann. S. Afr. Mus., xx, 6, p. 465, Pls. xxx~x,fig. 2, XL,fig. g, XLII,fig. 3 (Type locality Port Nolloth, Namaqualand).

Snout short, obliquely truncate, slightly prominent ; eye large ; interorbital space narrower than the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second ; fourth reaching the distal subarticular tubercle of the third and measuring 2 t o of the second. Toes short, the outer not quite reaching the proximal tubercle of the fourth. Subarticular tubercles very prominent, often double ; supernumerary double tubercles between the subarticulars often present. Inner metatarsal tubercle large, compressed, with a digging edge ; outer smaller, subcircular. Dorsal surfaces porous and with a few small, scattered warts ; finely granular or almost smooth beneath. Brown above with lighter markings in the form of a curved bar between the eyes, a row of spots on each side of the vertebral line, of which the first pair are elongately oval, and another dorso-lateral row ; these light spots may be more or less fused both transversely and longitudinally ; an oblique dark bar beneath the eye. Lower surfaces whitish or yellowish. Length : 5 0 mm. DISTRIBUTION : Namaqualand.

B.M. 46.6.18.30 1911.3.18. 1

9 9

. .

Narnaqualand Garris, Namaqualand

. . . . . . . . . . . . . . . .

Broom

192

MONOGRAPH OF T H E MICROHYLIDAE

7 . Breviceps pentheri pentheri, Werner Brevicepspentheri. Werner, 1899, Zool. Anz., XXII, p. 116 (Type locality S. Africa, probably Grahamstown). 7Breviceps vernreosus (non Rapp), Beddard, 1908, Proc. Zool. Soc. London, p. 11. Breiceps parvus, Hewitt, 1925, Ann. Natal. Mus., v, 2, p. 192, figs. 6-8 (Type locality Grahamstown, S. &ca) ; Power, Ann. S. Afr. Mus., xx, 6, p. 464, Pls. XXXIX, fig. 7, XL, fig. a, XLI, fig. B ; Essex, 1928, Ann. Mag. Nat. Hist., (lo),I, pp. 265-268 ; de Villiers, Ann. Transvaal Mus., XIII, 3, pp. 142-151, 8 figs. ; WtzSimons and van Dam, 1929, Ann. Transvaal Mus., XIU, 3, p. 152 ; PitzSimom, 1930, Ann. Transvaal Mus., xrv, 1, p. 48 ; Hewitt, 1932, Ann. Natal Mus., VTI, 1, p. 108. Breviceps gibboszls (part), Nieden, 1926, Das Tierreich, Anura XI,p. 5. Snout short, rounded, slightly prominent ; interorbital space a little broader than the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second ;

fourth just reaching the level of the distal subarticular tubercle of the third, and measuring 8 to Q the length of the second. Toes short, the outer not, or scarcely extending beyond the first. Subarticular tubercles well developed. Inner metatarsal tubercle large, compressed, with a digging edge ; outer small, rounded, scarcely distinct from the inner. Upper surfaces porous and more or less warty ; smooth beneath. Dark brown or greyish above, with traces of lighter markings in the form of a bar from eye to eye, a row of indefinite, paired spots on each side of the vertebral line, of which the anterior pair are elongately oval, and, sometimes, a dorso-lateral row of similar spots ; a more or less distinct, oblique, dark bar from beneath the eye, bordered by definite lighter areas anteriorly and posteriorly. Lower surfaces whitish, heavily reticulated and spotted with brown. Eggs laid in holes on land ; no free larval stage. Developing embryo without gills, spiraculum, or mouth armature ; tail large. Male with a vocal sac. Length : 36.5 mrn. DISTRIBUTION : S.E. Cape Province (Zululand, Portuguese E. Africa and Transvaal ?) It is probable that this species is only a southern form of B. adspe~sus,Peters, and the Transvaal, Zululand and Portuguese East African records may be based on immature examples of that species. SPECIMENS EXAMINED B.M. 78.10.12. 19-20 $2, juv. Kaffraria . . . . . . . Weale Lester 89.12.28. 2 juv. Port Elizabeth. . . . . . 90.7.16. 23 JUV. Port Elizabeth. . . . . . . Spencer hgr. Coldspring, nr. Grahamstown . . Hewitt 1925.7.1. 1

.

.

.

. .

8. Breviceps pentheri cafler, Hewitt Breviceps p a w caffer, Hewitt, 1932, Ann. Natal Mus., Province, S. Africa).

VII,

1, p. 109 (Type locality Kei Road, Cape

Snout more prominent and obliquely truncate and nostrils more closely approximated than in the typical form; skin corrugated, roughened or warty. Dorsurn and throat uniformly infuscate ; no dark oblique streak from the eye to the fore-limb. Length : 28 mm. DISTRIBUTION : Kei Road, S.E. Cape Province.

9. Breviceps montanus, Power Brevicepe modanus, Power, 1926, Ann. S. Afr. Mus., xx, 6, p. 466, Pls. xxx~x,fig. 9, x L I n , figs. 3,4. (Type locality Table Mountain, S. Africa).

XL,

fig. c,

BREVICEPS

193

Snout short, obliquely truncate, slightly prominent ; interorbital space about equal to the width of the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second ; fourth reaching the level of the distal subarticular tubercle of the third and measuring Q the length of the second. Toes short, the outer not reaching the proximal subarticular tubercle of the fourth. Inner metatarsal tubercle large, oval, and prominent, but without a digging edge ; outer rounded, more or less distinct from the inner. Skin densely granular both above and beneath. Black above with indistinct brownish grey markings ; yellowish or grey beneath, spotted and vermiculated with darker, especially on the chin and throat. Length : 30.5 mm. DISTRIBUTION : Table Mountain.

10.

Breviceps adspersus, Peters

Breviceps verrwosus (non Rapp), Smith, 1849, Zool. S. Afr., m, Appendix, p. 27 ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus., p. 52 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 177 ; Odhner, 1908, Ark. Zool. Stockholm, rv, 18, p. 7 ; Sternfeld, 1911, Fauna Deutach Kolon., IV, p. 57, fig. ;Noble, 1924, Bull. Amer. MUE.Nat. Hist., XLIX, p. 345. Breviceps adspersus, Peters, 1882, Reise n. Mossambique, Zool., Amph., p. 177 (Type localities Damaraland and Transvaal) ;Hewitt, 1932, Ann. Natal Mus., VII, 1, p. 107, P1. VI, fig. 1. Breviceps mossambicus var. occidentalis, Werner, 1903, Abh. Bayer. Akad., xxn, 2, p. 383 (Type locality S. W. Africa). Breviceps mossambicus (nm Peters), Boulenger, 1907, Proc. Zool. Soc. London, p. 480 ;idem, 1910, Ann. S. Afr. Mus., v, 9, p. 534 (part) ; Sternfeld, 1911, Fauna Deutsch Kolon., rv, 2, p. 57 ; Werner, 1915, in Michaelsen, Beitriige Fauna Deutsch S. W. Afrika, m, p. 372 ; Nieden, 1926, Das Tierreich, Anura n, p. 5 (part) ; Power, 1926, Ann. S. Afr. Mus., xx,6, p. 461 (part) ;Essex, 1928, Ann. Mag. Nat. Hist., (lo), I, pp. 265-268 ;FitzSimons, 1930, AM. T r a n s v d Mus., XIV, 1, p. 48 (part). Brevicepsgibbosw (non Linn.), Boulenger, 1910, Ann. S. Afr. Mus., V, 9, p. 534 (part) ; Nieden, 1926. Das Tierreich, Anura n, p. 5 (part) ; Barbour and Loveridge, 1928, Mem. Mns. Comp. Zool. Harvard, L, 2, p. 246, ? Brevieeps pretoriensis, FitzSimons, 1930, Ann. Transvaal Mus. XIV, 1, p. 45, figs. 24-26 (Type locality Jericho, Pretoria District, Transvaal).

Snout short, rounded, scarcely prominent; interorbital space about as broad as the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second; fourth only reaching to midway between the subarticular tubercles of the third and measuring g to Q the length of the second. Toes short, the outer not reaching the level of the union of the third and fourth. Subarticular tubercles moderately well developed. A large, compressed, oblique, inner metatarsal tubercle, with a digging edge ; inner small, compressed, almost continuous with the outer. Skin porous and granular above ; smooth beneath, except the throat, which, in juveniles, may be granular. Brown above with lighter markings in the form of a cross-bar between the anterior halves of the eyelids, a row of oval or subcircular spots on each side of the vertebral line, the anterior pair largest and longitudinally oval, and a dorso-lateral row of similar but smaller spots. These markings may be more or less obscured, or there may be partial fusion of the spots to form stripes ; in addition there may be irregularly scattered light verrniculations. An oblique dark streak from beneath the eye to the fore-limb, bordered before and behind by a broad lighter area, is constantly present. Yellowish white beneath, the chin and throat mottled with darker, the mottlings forming a pair of more or less well-defined, posteriorly divergent, longitudinal stripes. Sometimes a light streak from heel to heel, above the vent. Length : 52 mm. DISTRIBUTION : Damaraland, Bechuanalsnd, S. Rhodesia, Transvsal, N.E. Cape Province, Zululand and Natal. 13

MONOGRAPH OF THE MICROHYLIDAE SPECIMENS EXAMINED B.M. 1908.2.14. 13 $2 Gnamayaga River, 150 miles N. of . Chubb Bulawayo . 1909.4.30. 60 Barberton, Transvaal . . (Rosenberg) 1907.4.9. 62-63 $2 9 Klein Letaba, Transvaal . . . Grant 1907.4.9. 64 skel. Klein Letaba, Transvaal . . . Grant 1903.3.27. 30 hgr. Zoutpansberg, Transvaal . . Cregoe 1906.11.19.4 JUV. Mseleni, Zululand . . Warren 1906.11.19. 5 9 E. Zululand . . Warren 1907.4.17. 86 & Pietermaritzburg . .Warren 97.11.16.12-14 2 hg., juv. Lower Unkomaas River, Natal . Marshall . . . . Sir A. Smith 58.11.25. 7 juv. Cape of Good Hope 46.1.18.29 skel. South Africa. . . . . . . 46.6.18. 31 juv. South Africa. . . . . . . Mus. Berlin 6294 ad. Damaraland . . . . . . . TYPE Mus. Kimberley ad. Eldorado, S. Rhodesia . . . . 2 Marandellas S. Rhodesia . . . . ad. Pietersburg, Transvaal . . . . 2 Mariamhill, Natal . . . . . 2 Kimberley . . . . . . .

I I.

Breviceps mossambicus, Peters

Engyetoma granosum (mm. nud.), Cuvier, 1829, R h p e Animal, ed. 2, 11, p. 112. Syatoma granosum, Parker, 1868, Mon. Shoulder girdle, p. 69, PI. VII. fig. 10. Brevtkeps nwssambicue, Peters, 1854, Mon. Ak. Berlin, p. 628 (Type localities Island of Mozambique, Sena) ; idem, 1855, Arch. f. Naturg., x x , 1, p. 58 ;Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 177 ; Peters, 1882, Reiae n. Mossambique, Zool., Amph., p. 176, PL. xxv, fig. 2, x x v ~ fig. , 11 ; Boettger, 1892, Kat. Batr. Senck. Nat. Ges., p. 25 ; Tornier, 1906, Deutech-0. Afrika, III, 4, p. 160; Chubb, 1909, Proc. Zool. Soc. London, p. 591 ; Boulenger, 1910, Ann. S. Afr. Mus., v, 9, p. 534 (part) ; Nieden, 1915, Mitt. Zool. Mus. Berlin, vn, p. 381 ; Procter, 1920, Proc. Zool. Soc. London, p. 420; Noble, 1924, Bull. Amer. Mus. Nat. Hist.. XLIX, p. 345; Loveridge, 1925, Proc. Zool. Soc. London, p. 790 ; Nieden, 1926, Das Tiemich, Anura II, p. 5 (part); Power, 1926, h. S. Afr. Mus. XX,6, 1926, p. 461, PL. xxrx, fig. 8, XL, fig. f, fig. A ;Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Harvard, L, 2, p. 245 ; Loveridge, 1930, Proc. Zool. Soc. London, (I), p. 16 ; FitzSimons, 1930, Ann. Transvaal Mue., XN, 1, p. 48 ; de Witte, 1932, Bull. Cercle Zool. Congolais, ~I, 3, p. 86, figs. 4-5.

=I,

Snout short, rounded, scarcely prominent ; interorbital space a little broader than the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second; fourth not reaching the distal subarticular tubercle of the third and measuring to f the length of the second. Toes short, the outer not reaching the level of the union of the third and fourth. Subarticular tubercles well developed. A large, strongly compressed, oblique, inner metatarsal tubercle, with a digging edge ; outer small, compressed, almost continuous with the inner. Skin smooth but porous above ; smooth beneath. Dark brown to pinkish above, immaculate or with some irregular dark markings and sometimes a narrow light dorsal streak ; an indistinct dark lateral streak sometimes present. A broad, dark, oblique streak from beneath the eye to the fore-limb, bordered in front and behind by a lighter region, which is continuous round the tip of the snout and also borders the lower jaw. Lower surfaces whitish, immaculate or with irregular dark reticulations ; chin and throat more or less uniformly infuscste. Length : 48 mm.

5

e

D~STRIBV~ION : Tanganyika Territory, Portuguese E. Africa, Zululand, Nyasaland, Katanga, Rhodesia and Angola.

BREVICEPS B.M. 1920.5.10.2628

SPECIMENS EXAMINED Morogoro, Tanganyika Terri99,juv. tory . . . . . . . Loveridge . . . Nutt juv. Lake Tanganyika d LakeNyasa . . . . . Shirk Highlands, Nyasaland . Johnston 9 juv. Fwambo, Nyasaland. . . Carson Fort Johnston, Nyasaland . Johnston Masaka Mt., N. Nyasaland, juv. 6,000-7,000 ft. . . . . Johnston 9 Fort Hill, Nyasaland, 4,000ft. Johnston $2, hgr., juv. Zomba, Nyasaland . . . Lennon d Mozambique . . . . . Peters, COTYPE ~UV. Delagoa Bay . . . . . (Monteiro) JUV. Delagoa Bay . . . . . (S. Afr. Mus.) d Zambesi River . . . . (Stevens) hgr. Mazoe, Rhodesia . . . . Darling hgr. Mseleni, N. Zululand . . . Warren hgr. and juv. Interior of Benguella, 4,0005,000 ft. . . . Wellman h ~ . Chiyaka ~ i s t r i c tAngola , . Wellman R. Donda, 300 miles inland 6 from St. Paul de Loanda . (Higgins) d Guissanga, Angola . . . Bocage Bagilo, Uulguru Mts. . . ad. Tanganyika Territory . . Loveridge

?

87.3.23. 5 Mus. Kirnberley

I 2.

Breviceps poweri, sp. n.

Breviceps m s s o m b i c w (nan Peters), Boulenger, 1907, Mem. and Proc. Mancheater Litt. Phil. Soc., LI, 3, p. 5. Holotype a female, number 1932.9.9. 1 in the British Museum, from Broken Hill,

N. Rhodesia (3,900 ft.). Snout short, rounded, scarcely prominent ; interorbital space a little broader than the upper eyelid ; tympanum hidden. Fingers short, the first shorter than the second ; fourth only reaching the level of the proximal subarticular tubercle of the third, and measuring less than 4 of the second. Toes short, the outer reduced to a mere tubercle. Subarticular tubercles well developed. A large, strongly compressed, oblique, inner metatarsal tubercle with a digging edge ; outer metatarsal tubercle continuous with the inner. Skin smooth but porous above ; smooth beneath. Uniform dark grey above, lighter towards the flanks, a lateral row of 3 or 4 yellow spots (orange in life) and a similar circular spot above the vent ; an oblique dark streak from beneath the eye to the fore-limb, bordered anteriorly by a white area which extends round the tip of the snout. White beneath, the chin and throat uniformly infuscate. Length : 46 mm. Paratype an immature female, number 1906.11.22. 36 from Petauke, Mterize River, E. Loangwa District, N.E. Rhodesia. This specimen is essentially similar to the type, even to its coloration and the distribution of the yellow spots ; length 29 mrn. DISTRIBUTION : N. Rhodesia. SPECIMENS EXAMINED B.M 932.9.9. 1 9 Broken Hill, N. Rhodesia, 3,900 ft. Pitman, TYPE 11906.11.22. 36 hgr. $2 Petauke, Mterize River, E. Loangwa District, N. Rhodesia . . Neave, PARATYPE

MONOGRAPH O F T H E MICROHYLIDAE I 3.

Breviceps macrops, Boulenger

Breviceps macrops, Boulenger, 1907, Ann. Mag. Nat. Hist., (7), XX, p. 46, P1. n (Type locality Namaqualand) ; i h m , 1910, Ann. S. Afr. Mus., v, 9, p. 534 ;Noble, 1924, Bull. Amer. Mus. Nat. Hist., XLIX,p. 345 ;Nieden, 1926, Das Tierreich, Anura n, p. 6, figs. 3,4 ; Power, 1926, Ann. S. Afr. Mus., xx, 6, p. 462, Pls. xxx~x, fig. 1, XL, fig. e, x m , fig 4 ; Essex, 1928, Ann. Mag. Nat. Hist., (lo), I, pp. 265-268 ; FitzSimons,l930, Ann. Transvaal Mus., xrv, 1, p. 48. Snout short, rounded, not prominent ; interorbital space narrower than the upper eyelid ; eye very large ; tympanum hidden. Fingers very short, the outer reaching

midway between the subarticular tubercles of the third and measuring 3 to 4 the length of the second. Toes very short, the two inner bound together and scarcely distinguishable ; the outer not reaching the level of the union of the third and fourth. Subarticular tubercles feebly developed. Inner metatarsal tubercle oblique, compressed, but only feebly developed; outer small, oval, flat. Skin porous, but smooth above, sometimes granular towards the flanks ; smooth beneath. Pale brown to yellowish above, with some black or reddish brown blotches or streaks, and some scattered, small light dots ; a dark band along the outer edge of the upper eyelid continued across the forehead in front of the eyes to meet its fellow ; a broad, irregular dark band connecting the posterior halves of the upper eyelids. Lower surfaces immaculate white. Length : 48 mm. DISTRIBUTION : Namaqualand. B.M. 1907.6.12.1

Namaqualand .

. . . . . . Peringuey, TYPE

MELANOBATRACHINAE, Noble Melanobalrachinae + Hoplophryniw, Noble, 1931, Biol. Amph., pp. 538-539. Maxillary and vomerine teeth absent ; prevomer small, confined to the antero-mesial border of the choana ; palatine absent ; ethmoid fused with the parasphenoid ; ear

without tympanum, cavum tympani, annulus tympanicus, columella auris or Eustachian tube ; vertebral column procoelous. No free-swimming tadpole, metamorphosis taking place on land, but not in the egg ; the lama lacks external gills, has only a single open branchial arch with very short gill-filaments, but has adhesive organs, an operculum and a median spiracle ; there is a pair of peculiar flaps over each branchial region. The three genera of this subfamily appear t o be closely related, a d d i f f e r from all the other members of the family in their incomplete auditory apparatus; as has already been pointed out, their true affinities may lie elsewhere. SYNOPSIS OF THE GENERA I. Clavicle and procoracoid present ; well developed. A. Omosternum calcified. Toes webbed. . . . . .Melanobatrachus, p. 196 B. Omosternum rudimentary, cartilaginous. Toes free Parhoplophryne, p. 198 11. Clavicle reduced to a small nodule or absent . . . . Hoplophryne, p. 198

.

MELANOBATRACHUS, Beddome Melanobatrachus, Beddome, 1878, Proc. Zool. Soc. London, p. 722 (Type species Melanobalrachus inalicus, Beddome) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 156 ; i h m , 1890, Fauna Brit. India, Rept. and Batr., p. 489 ; Nieden, 1926, Das Tierreich, Anura n, p. 18; Noble, 1931, Biol. Amph., p. 538.

Prevomer confined to the antero-mesial border of the choana, without any postchoanal portion; palatine absent; ethmoid and parasphenoid fused, the former

MELANOBATRACHUS

197

entirely surrounding the posterior portions of the nasal capsules and bordering the choanae posteriorly. Clavicle and procoracoid present, straight, well developed, reaching the mid-line of the girdle and the scapula ; ornosternum small, calcified ; sternum large, cartilaginous (fig. 66). No colurnella auris or annulus tympanicus ; operculurn present, normal, with the usual attachment of the pars opmcularis of the m. levatur scapulae superior. Vertebral column procoelous. Terminal phalanges simple. Pupil circular. Tongue oval, entire and half free behind. No defined palatal ridges. No tympanum, cavum tympani or Eustachian tubes. Tips of digits not dilated ; toes webbed.

Fig. 66. Ventral elements of the shoulder-girdleof Melanobatrachua indicus

I.

Melanobatrachus indicus, Beddome

Melanabdrachua indicw, Beddome, 1878, Roc. Zool. Soc. London, p. 722 (Type locality Anamallai Hills, S. India, 4,000 ft.) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2, p. 157 ;Thurston, 1888, Cat. Batr. Sal. and Apoda S. India, p. 40 ; Boulenger, 1890, Fauna Brit. India, Rept. and Batr., p. 489 ; Sclater, 1892, Cat. Batr. Ind. Mus., p. 22 ; Nieden, 1926, Das Tierreich, Anura 11, p. 18. Snout vertically truncate, as long as the diameter of the eye ; canthus rostralis blunt ; loreal region vertical ; nostril nearer the tip of the snout than the eye ; inter-

orbital space broader than the upper eyelid. Fingers short, depressed, tapering and ending obtusely ; first much shorter than the second, which is subequal to the fourth. Toes webbed a t the base, depressed, tapering and ending obtusely ; fifth much shorter than the third, but longer than the second ; subarticular tubercles indistinct; an indefinite inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching to midway between the shoulder and the eye. Skin with small white pustules set on small prominences and larger pustule-bearing warts above ; smooth beneath ; anal region granular. Black or dark brown above, uniform ; lower surfaces brown, dotted with greyish ; a large subcircular white (red in life ?) spot on the lower and anterior surfaces of esch thigh.

MONOGRAPH OF THE MICROHYLIDAE Male without a vocal sac ; a diffuse nuptial pad on the base of the first and the dorsal surfaces of the first and second fingers. Ovarian eggs, 3 rnm. in diameter. Length : 34 mm. DISTRIBUTION : Hills of Travancore and Cochin, S. India. B.M. 78.9.3. 1 78.1.11. 10 82.2.10.70-73 Mus. Vienna

8 $2, hgr. 2

$29

Anarnallai Hills, 4,000 ft. . N. Travancore . . . . . S.W. India, 3,00&4,500 ft. S. India . . . . . .

. Beddome, COTYPE . Beddome . . Beddome

PARHOPLOPHRYNE, Barbour and Loveridge

.

Parhapbphryne, Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. H m a r d , L, 2, p. 260 (Type species Parhoplophryne wrombaricua, Barbour and Loveridge) ; Noble, 1931, Biol. Amph., p. 539.

Prevomer confined to the antero-mesial border of the choana, without any postchoanal portion. Clavicle and procoracoid present, well developed, reaching the midline of the girdle and the scapula ; omosternum rudimentary ; sternum cartilaginous. Terminal phalanges simple. Pupil circular. Tongue oval, entire and free behind. No defined palatal ridges. Tips of digits not dilated ; toes free. I . Parhoplophryne usambaricus, Barbour and Loveridge Parbpbphryne w a m b a r i c u ~Barbour and Loveridge, 1928. Mem. Mus. Comp. Zool. Harvard, L, 2, p. 260 (Type locality Amani, Tanganyika Territory) ;Loveridge, 1930, Proc. Zool. Soc. London, (11, P. 15. Snout short, acuminate ; mouth small, only extending backwards slightly beyond the centre of the eye ; interorbital space twice as broad as the upper eyelid. Fingers

moderate, the first shorter than the second, which is slightly shorter than the fourth. Toes well developed, the first short ;third longer than the fifth. Subarticular tubercles well developed ;a strong inner, and an almost equally strong outer, metatarsal tubercle. Tibio-tarsal articulation barely reaching the axilla. Skin smooth, but beset with many short, rugose folds, above and beneath. Purplish grey above, darker on the flanks; limbs and digits with dark purplish blotches on a grey ground ; a white bar from the eye to the base of the fore-limb. White beneath, blotched and spotted with dark brown. v Length : 23 rnm. DISTRIBUTION : Amani, Usambara Mts., Tanganyika Territory.

HOPLOPHRYNE, Barbour and Loveridge Hoplophryne, Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Harvard, L, 2, p. 253 (Type species Hoplophryne uluguruemia, Barbour and Loveridge) ;Noble, 1931, Biol. Amph., p. 539.

Prevomer wnfined to the antero-mesial border of the choana, without any postchoanal portion ; palatine absent ; ethmoid and parasphenoid fused, the former expanded to embrace the hinder part of the nasal capsule and bordering the choana posteriorly. Clavicle reduced to a small nodule distally, or absent ; no omosternum ; sternum cartilaginous (fig. 67). No columella auris or ~ M ~ Utympanicus S ;operculum present with normal attachment of the pars opercularis of the m. bvatw scapulae superiw. Vertebral column prowelous. Terminal phalanges T-shaped. Pupil round (or horizontal). Tongue oval, entire and free behind. No defined palatal ridges. No tympanum, cavum tympani or Eustachian tubes. Tips of digits slightly dilated, but without distinct discs ;toes free.

HOPLOPHRYNE

Rg.67. Ventral elements of the shoulder-girdleof Hoploph~ynauluguruensia (3) 6

SYNOPSIS OF THE SPECIES I. Vestiges of clavicle and procoracoid present. Snout shorter than the eye ; fingers very short, the first represented by a mere swelling; belly uniform or slightly speckled with silver . . . . . H. u l u g u r u e ~p. , 199 11. No vestiges of clavicle or procoracoid. Snout longer than the eye ; fingers moderately well developed, the first represented by a distinct stump ; belly boldly vermiculated with brown and white . H. rogersi, p. 200 I.

Hoplophryne uluguruensis, Barbour and Loveridge

Hoplophryne ulugu~uemis,Barbour and Loveridge, 1928, Mem. Mus. Comp. 2001. Harvard, L, 2, p. 254, P1. n, figs. 3 4 (Type locality Mt. Mbova, Nyange, Tanganyika Territory) ; Noble, 1929, Bull. Amer. Mus. Nat. Hist., LvnI, 7, p. 292, figs. 1-3 ; Loveridge, 1930, Proc. Zool. Soc. London, (1). p. 14. Vestiges of clavicle and procoracoid present. Head depressed ; snout rounded, shorter than the diameter of the eye ;canthus rostralis indistinct ;loreal region oblique, convex ;nostril equidistant from the eye and the end of the snout ; interorbital space

more than twice as broad as the upper eyelid. Fingers very short, free, spatulate distally ; first represented by a swelling ; second very short-shorter than the fourth. Toes moderate, free, spatulate distally; fifth and third subequal. Subarticular tubercles developed ; an indefinite inner, but no outer, metatarsal tubercle. Tibiotarsal articulation reaching the shoulder or the eye. Skin beset with minute papillary tubercles on the back, on the upper and lower lips and on the throat and chest ;belly and thighs smooth. Brownish black above, with indistinct light dots, which sometimes are concentrated to form a fine vertebral line and an irregular lateral line from the eye to the knee ; a dark bar, bordered by lighter, across the femur and tibia ;grey-brown beneath, faintly white-stippled on the throat. Male without a vocal sac but with well-marked nuptial excrescences in the form of a group of 3 or 4 spines on the rudiment of the first finger, a similar group of 5 or 6 spines on the back of the second finger, a marnmiform rosette of 9 to 15 similar spines on each side of the chest and a number of smaller scattered spines on the tibia and foot. Length : 25' mrn. Eggs large, laid in the axils of banana trees or in the hollows of bamboos ; developing embryo without external gills a t any stage, and with only a single branchial arch with very short gill maments. Tadpole hatching a t a late stage.

MONOGRAPH O F T H E MICROHYLIDAE Description of Tadpole. Body twice as long as broad. Eyes on upper surface of head; the interocular distance twice as great as the internarial. Mouth terminal, without teeth or mandibles ; upper lip arcuate, lower folded, with a median groove and two lateral flaps ; a prominent locomotor (?) flap over each branchial region. Spiraculurn median, nearer the snout than the anus. Anus median. Tail bluntly pointed, 5 times as long as the head and body and 34 times as long as deep. Length (fully developed), 15 mm. DISTRIBUTION : Ulugum Mts., Tanganyika Territory, 3,000-7,500 ft. B.M. 1928.5.1.4 2.

3

Nyange, Uluguru Mts.

. .

.

. Loveridge, PARATYPE

Hoplophryne rogersi, Barbour and Loveridge

Hoplophryne rogersi, Barbour and Loveridge, 1928, Mem. Mus. Comp. Zool. Harvard, L, 2, p. 258, PI. n, fig. 5 (Type locality Mt. Bomoli, Amani, Tanganyika Territory) ;Noble, 1929, Bull.Amer. Mus. Nat. Hist., LVIII, 7, p. 300, figa. 6-9 ; Loveridge 1930, Proc. Zool. Soc. London, (I),p. 14. Clavicle and procoracoid absent. Head depressed ; snout rounded, not prominent, once and one-third as long as the diameter of the eye ; canthus rostralis rounded ;

loreal region nearly vertical, ⁢ interorbital space twice the width of the upper eyelid. Fingers short, the first a short stump, the remainder spatulate distally ; fourth much longer than the second. Toes moderate, free, spatulate distally, the fifth distinctly shorter than the third. Subarticular tubercles well developed ; a small inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the posterior corner of the eye. Skin smooth above and below except for a few scattered minute spines on the back, lips, throat and limbs. Brownish black above, with lighter and darker patches on the flanks and fore-limbs ; hind-limbs definitely cross-barred with black on a light ground colour ; beneath reticulated with brown and white. Male without a vocal sac but with well-marked nuptial excrescences thus : a group of large spines about the base of the first digit and on the inner side of the forearm ; some smaller spines on the dorsal surfaces of the other fingers ; a small circular patch of spines on the ventral surface of the upper arm close to its insertion, and a larger patch of similar spines on the chest, close to, but separated from, its fellow of the opposite side. A few large spines on the knee may also prove to be a secondary sexual development of the male. Length : 26 mm. Breeding habits probably similar to those of H. ulugumensis. Tadpole with a single branchial arch and very short gill filaments. v Description of Tadpole. Body once and one-third as long as broad. Mouth terminal, without teeth or mandibles ; upper lip arcuate, lower folded. A prominent locomotor ( 1 ) flap over each branchial region. Spiraculum median, midway between the snout and the median anus. Tail twice as long as head and body, bluntly pointed terminally. Integument of snout and lips beset with prominent papillae. Toes fully webbed. Length : 28 mm. DISTRIBUTION : Eastern range of the Usambara Mts., Tanganyika Territory. SPECIMEN EXAMINED B.M. 1928.5.1.3 8 Amani, Usambara Mts. . . . Loveridge, PARATYPE

.

INDEX acephdua. Bufo, 190 ac?dinu, Dendromanea, 136 ac?dina. Hybpleaia, 123, 136 ac?dina. Micrhyla, 136 ac?dinu. Microhyb. 134, 136, 165 acutiroatris. Calophruynua. 97 acutiroatria, Kalophrynua, 97 Adhesive organs. 7 adaperatca, Bretticepa, 191, 193 Aelzcrophryne. 15 aquntoriale. Engiatoma. 14f3 aepuatorialis. Engyatoma, 146 aequetorialk, GLastrophryne, 146 aquutorialia, Microhyb, 146 afinia, Mehdyia. 169 africana, Rana, 1 albopunddat Chiamwdeia, 118. 119 albopundato, Oaatrophryne, 119 albopuetata, Oreophryne. 166 albopunetata, Sphenophryne, 166 albopunclatum, Engy&nna, 116, 118 alluadi, Dyacqphw. 45 alluaudi, Plethadontohyb, 45 amboinenais, Aaterophrya, 61 amboinenaia, Hylophorbua, 61 angulifera, PlethodmItohyla. 35 angulifems, Mantipua. 35 annamensie, Microhyb, 130 anneetens, Microhyla, 128, 129, 132 annzllata. Orwphryne, 167 annuhtw, Phrynizalua, 167 Anodonthyb. 48 Anodontohyb, 48 anthunyi. Oreophryne, I68 anthonyi, Sphenophryn'e. 168 antongili, Dyacophua. 24 anulatua. Phrynizalua, 167 Aphuntophryne, 178 areolata, Engyatoma, 147 areohto, Castrophryne, 147 aredata. Mkrohyla, 147 Aaterophrydidae. 19, 52 Aaterophryinae. 10.52 Asterophrya. 58 d m . Cophiza2ua. 172 deles, Oreophryne. 172 alelea, Sp7mwphryne. 172 Atelopalidae. 8 Atelopus. 1 alerrima, Mkrohyb, 143 aterrimum, Castrophryne, 143 derrimum, (%aaostoma, 124, 143 dra, Mehpoatira. 70 Auatrochuperinu, 152, 171

baleutua. Hyluductylua. 88 balecrtua, Hylaedactylua, 88 baaipdmata. Chuperina, 155 bmipalmata. Sphenoph.ryne, 155 beloensis. Dyacophua. 26 B e r d w e a . 95 b e r d w e i , Diplopelma, 127 berdwei. Engyatoma. 127 berdmorei, Microhyla. 127, 131, 132 berdmwii, Microhyb, 127 beyeri, Chperina, 103 beyeri, Sphenophryne, 103 bicqoi, Chiamocleis, 118 bicolor, Elachistocleis, 121 bicdor, Engystoma, 121 bicdor, Oxyrhynchua, 121 bidena, Xenobatrachua, 57 bidena, Xenorhina, 57 biporcatua, Bufo, 88 biroi, Cophixalua, I74 biroi, Hylophorbua, 174 biroi, Oreophryne, 169 biroi, Phrynixalua, 174 biroi, L?phenuphryne, 169 bivillatua, Hylaedactylua, 85 boettgeri, Asterophrya, 61 boettgeri, Onathophryne. 61 boettgeri, Hylophorbu-8, 61 boliviana, C h i m o d e i s . 119 Bo7n6inator. 58, 73. 77 Bombinatorea, 19 Bombinatorina, 19 Born binatoroidea, 19 borealis, Cacopoides. 77, 82 borealis, Kaloula, 82 borneensis, Engystoma, 104 borneensis, Gaatrophryne, 105 borneemis, Microhyla, 128 borneenae, Engyatonuz, 104 borneenae, Oaatrophryne, 105 borneenae, Oaatrophrynoides, 104 borneeme, Microhyb. 128 boulengeri, Anodonthyb, 48, 49 bdengeri, Kaloula, 106 boulengeri, Microhyla, 131, 137 bouwensi, Asterophrys. 66 bouwensi, Pseudengystm, 59, 66 Brachycephalidae, 8 Brachycephdua, 1 Brachymeridae. 19 Branchiae. 7 Branchial Clefts. 7 Brevicepa, 183. 186 brewicepa, Rana, 190 brevicep8, Syatoma, 75. 190 Brevicipitidae, 1, 19, 179 Brevicipitinae, 2, 10, 179 breva'cm, Oxydactyb, 152, 158

baleata, Callzlla, 88 bale&, Kaloukr, 88 balecrtw, Bombinator, 77, 88

4

204

MONOGRAPH O F T H E MICROHYLIDAE

brevicrua, Sphenophryne, 158 brevipea, Auatrmhaperina, 157 brevipea, Liophryne, 158 brevipea, Plethdmtohyla, 45 brevipes, Sphenophryne, 158 brookai, Colpogloaatca, 30, 31 Bufo, 95, 120, 186 Bufonea. 19 bufonia, Rana, 121 Bufonidae, 19 Bufoniformes, 19 Bufonoidea, 19 bungurana, Microhyla, 100 bunguranum, Diplopelma, 100 bunguranus, Kalophrynua, 100 butleri, Microhyla, 131 Cacopinae, 2, 52, 71, 151 Cacopoidea, 77 Cacopua, 73 Cacosteminae, 2, 3 caeaarii, Engystoma, 121 caffer, Breviceps, 192 Calliglutwr, 30 Calluella, 27 Call&, 42, 76, 90, 107, 123, 159 Callulina, 180 Callzllops, 58 Calohyla, 42, 76, 107 Calophrynus, 95 carnaticum, Diplopelma, 140 carnaticum, Engystoma, 140 carnaticua, Hylaed.uctylua, 93 carolinense, Engystoma, 123, 144, 146, 148 carolinensis, Qastrophryne, 146 carolinensis, Microhyla, 146 celebemis, Calohyla, 88 celebensis, Hylaedactylus, 88 celebensis, Oreophryne, 63, 161 celebenais, Sphenophryne, 161 Ceratophyidae, 3 Ceratophrydes, 19 ceratophthalmus, Chaperina, 153 Chaperina, 101, 152, 159 cleamanae, Cophizalua, 175 Chiasmocleis, 116 Chanacantha, 54 Cherophryne, 171 Co~pogloasus,30 concolor, Ef~gystoma,121 conjuneta, Callula, 89 conjuncta, K a h d a , 14. 89 coniunctua. Holonectes. 77. 89 m3unctua; ~~laedactylus,' 89 Copea, 123 Cophizalua, 159, 170 Cophyla, 38, 41, 42 Co~hvlidae.19. 32 ~ o i h i l i n a e ;10; 32 Copiula, 171 Cornufer, 151 cornzlta, Sphenophryne, 151, 153 cowani, Platypelis, 39 crucifer, Cophizalus, 166, 169 cmifera, Orwphryne, 166

Ctenophryne, 122 cuneua, Hypopachua, 114 DWPOP, 124 Dendromanea, 123 Dentary, 5 Dermatonotua, 110 Didynamipua, 3 Diplopelma, 95, 123 discgerum, Diplopelma, 136 disciferum, Engystoma, 136 Discoglossidae, 19 Distribution, 11 doriae, Aeterophrya. 65 doriae, Callulops, 58, 65 dorsatum, Engyatoma, 190 dubia, Aaterophrys, 63 dubia, Gnathphryne, 63 dubia, Xenorhina, 63 dubiua, Hylophrbus, 63 d u m e d i , Engystoma, 150 Dyscophidae, 1, 19, 22 Dyscophina, 27 Dyscophinae, 2, 10, 22, 179 Dyacophua, 22, 42 Ear, 4 Eggs, 7 Egg-tooth, 7 Elachiatocleis. 120 elegans, Engyatoma, 144 degans, Gastrophryne, 144 elegans, Microhyla, 144 Eleutherognathinae, 1, 52, 151 Emydops, 115 Engiatoma, 123 Engystoma, 73, 95, 104, 110, 116, 120, 123, 186 Engystomatidae, 1, 19 Engystornidae, 19 eremda, Microhyla, 140 Ethmoid, 4 Eupemphiz, 124

$mipea, Microhyla, 134, 138, 140 Flagellum, 7 Jlava, Oreophryne, 168 frontifaaciata, Callula, 165 fulva, Copea, 123, 142 fwca, Asterophrya, 63 fusca, Chaperina, 101, 103, 156 fusca, Phrynomantis, 58, 63 fusca, Sphenophryne, 103, 156 fwcua, Breviceps, 189 gadovii, Eupemphiz, 149 Gastrophrynae, 16, 19, 71 Gdrophryne, 104, 116, 120, 123 Castrophrynoidea, 104 geagi, Ctenophryne, 123 geayi, Ctenophryne, 122, 123 geislerorum, Cophizalus, 170, 172 Cenyophryne, 52 Genyophrynidae, 1, 19,52 Genyophryninae, 52 gibbosa, Engystoma, 190

gibbosa, R a m , 1, 186, 190 gibbow, Breviceps, 186, 189, 190, 192, 193 gibbosus, Bufo, 190 gibbomm, Systoma, 190 gigantea, Xenorhina, 57 giganteus, Xenobatrachus, 57 Gas, 7 globulow, Cacopus, 75, 76 globulomm, Systoma, 76 globuloszmm, Uperoda, 76 Glossostoma, 124 Glyphoglosatls, 72 W h o p h r y n e , 58 gracilipes, Austrochaperina, 155 gracilipea, Sphenophryne, 155 grahami, Microhyla, 131 gmndidieri, Dyscophus, 25 grandis, Platyhyla, 37 granosum, Engystoma, 194 guineti, Dyscophus, 24 guineti, Kaloula, 24 guttulata, Calkella. 28 gutkhta, Cailuh, 28 gultulata, Megalophrys, 27, 28 Habits, 14 hainanemis, Microhyla, 131, 137 Hemisinae, 2, 3 heterochims, Calophrynus, 100 heymomi, Microhyla, 134 hildebrandti, Mantipua, 33, 35 Holonecta, 77 Hoplophyne, 198 Hoplophryninae, 2, 196 Huladactvla. 77 ~jlzadactjlz~S, 77 Hylae, 19 Hylaedactyli, 16, 19 Hylaedactylidae, 19 Hyla~dactylus,77, 90 Hvlaeformes. 19 ~ i l a ~ z e ~i23 ia, Hylophorbus, 58, 68, 152, 171 Hyoid, 5 Hypercdun, 75 hypomeha, EmyJops, 115, 116 hypomeha, Ribeirina, 116 Hyppachua, 110, 120 Hysapleaia, 123 incrassatus, Hypopachua, 111 incrassatus, SteTeocyclops, 110 indicus, Meladatrachus, 196, 191 inguinalis, Engyslmna, 112 inguinalis, Hypopachus, 112 inguinalis, Mantipua, 34 inguinalis, Plethodontohyla, 34 inwnata, Microhy!a, 1% i w h i s , Dyswphus, 22, 24,25,26 interlineah, Berdmorea, 99 interline&, Calophy n u s , 99 interlineatwr, Kalophynua, 99 interlineatum, Engyatoma, 95, 99 jeffersoniana, Oreuphryne, 163

kalingensis, Kaloula, 84 Kulluella, 27 Kalophrynina, 71 Kalopbryninae, 2 Kalophynua, 95 Kaloula, 22, 76, 90, 105, 107 Kaloulinae, 2, 71 kampeni, Liophryne, 64 kampeni, Oreophryne, 164 klossi, Sphenophryne, 154 kopsteini, Hylophorbus, 70 kreai, callulina, 180, 181 labyrinthica, Scaptophryne, 123, 137 laeve, Phrynocara, 46 laeve, Plethodontohyla, 46 laevis, Plethdontohyla, 46 laevipes, Mantiphrys, 33, 35 laevipes, Yantophys, 35 Larvae, 7 Larynx, 5 latclstii, Microh,yla, 131 lateralis, Manthophryne, 62 lateralis, Muntophryne, 58, 62, 177 leopoldi, Bsterophrys, 66 l~achena,ulti,Systoma, 75 leucosticta, Chiasmocleis, 118 leucosticta, Gastrophryna, 118 leucostictum, Engystoma, 118 leucostdgma, Yicrohyh, 103 leucostigma, Sphenophyne, 103 lineata, Engystoma, 121 lineata, MehpJyia, 159, 169 Liophyne, 58, 152 lividus, Hylaeda~tylus,88 loriae, Oreophryne, 169 loriae, Sphenophryne, 164, 169, 173, 176 louisiadensis, Aeterophrys, 62 loven'dgei, Probreviceps, 185 macra, Metopostira, 69 nurcrdactyla, Callula, 85 macrodactylus, Brevicep8, 183, 184 macrodactylus, Probreviceps, 184, 185 macrops, Breviceps, 196 macrops, Xenobatrachus, 56 macrops, Xenorhina, 56 macrorhyncha, Chaperina, 155 macrorhyncha, Sphenophryne, 155 madagascariensis, Stumpfia, 52 major, Microhyla, 137 malabaricum, Engystoma, 140 malcolmi, Yicrohyla, 127 manehurienais, Kaloula, 83 ManteUa, 48 Mantiphrys, 33 Manlipus, 33 Mantophyne, 58 Mantophrys, 33 mannwatum, Engystoma, 73, 76 mannoratum, Hypercdun, 75 mamoratum, Bysknna, 75 marmosatum, Uperodon, 76 Maxillae, 3 medidineata, Callzcla, 81

%

206

MONOGRAPH OF THE MICROHYLIDAE

mecliolinenta, Kaloula, 81 Hehelgia. 159 mehelyi, Choancacantha, 54 m a y i , Sphemphyne, 156 Melanobatrachinae, 2. 10, 196 Mdadatrachua, I96 melli, Mbmhyla. 137 Mento-meckelian bones, 5 mertoni, Oreophryne, 169 mertoni. Sphenophyne, 169 Metamorphosis, 7 Metaph~ynella,107 methneri, Spelaeophryne, 182, 183 Methods, 16 Metopoetira, 68 Micrhylidae, 19, 71 Microbairachw, 179 Microhgla, 95, 102,123, 159 Microhylidae, 9, 19, 16 Microhylinae, 2, 10, 71 Mbrqps, 120 micropa, Engystoma, 150 microps, Gnstrophyne, 150 microp. Microhyla, 150 microps, S t e m e p h l w r , 120, 121 microtia, Asterophys, 64 microtia, Mantophyne, 64 minima, Aaterophy8,67 mod&, Callula, 162 molosm~a,Glyphogloam~a,72 mdplcEenma, Microhyla, 166 moluceaaia, Oreophyne, 165 mentam, Anodonthylu, 50 montana, R a m a n e k , 91 montanw, Brevicepa, 192 montanw, Cophixalus, 174 montanua, H y ~ t y l u a 91 , monfanus, Hylophorbw, 174 montanua, Phrynixdus, 61, 170, 174 monticola, Orwphyne, 162, 163 monticola, S p h a o p h y n e , 162 rnossambicua, Breviceps,, 191, 193, 194, 195 rnottae, Engystoma, 121 miilleri, Derntatonotus, 111 miilleri, Engyetoma, 110, 111 mZilleri, Gaatrophyne, 111 mltlleri, Hypopachw, 111 namuquenais, Breviceps, 191 Nares, 7 W r i x , Calluh, 127 Nectodactylus, 116 Nectoph~yne,5, 102, 107 * negrosenak, Kaloula, 89 n e u b w s i , Hylophorbw, 65 n e u k w s i , Mantophyne, 65 n i a a e k , Microhyla, 132 Nomenclature, 15 notodicta, Callula, 42, 43 notosticta, Cdohyla, 43 notnsticta, Cophglu, 43 notosticta, Pldhdontohyla. 39, 43 obmura, Callulu, 91, 92, 93 obscura. Ramanella, 92

oecidentalia, Breviceps, 193 m d h t a , Mdopostira, 68, 69 mdlata, Pldhodontohyla, 44 odlata, Xenorhina, 56, 67 d l a t u a , Hyluphorbus, 69, 10 meUatua, Mantipurr, 44 ocdlatua, Xenobatrachus, 56 okinavensis, Microhyla, 138 olivacea, C~llulu,'93 olivacea, Gaatrophyne, 201 olivacea, Microhyla, 201 olivaceum, Engyetoma, 147, 201 Operculum, 7 ophiodon, Xenobatrachua, 54,58 OreophFyne, 159 Oreophryndla, 1 m i a , Austrmhaperina, 173 m t a , Mkrohyla, 139 onata, Sphenophyne, 173 m t u m , Diplupelma, 137, 139, 142 m a t u m , Engystma, 123, 139 omaturn, Biphneua, 139 ernatus, Cophizalw, 173 Orthogenesis, 15 Otophyne, 108 ovale, Ehhktocleia, 121 wale, Engyatoma, 121 m l k , Bufo, 121 ovalis, Ehhiatocleia, 121 ovolis, Gaetrophryne, 121 ocalk, R a m , 120, 121 ozycephala, Aderophys, 67 oxycephala, X e w h i n a , 67,70 oxycephalw, Bombinator, 58,67 Oxydactyla, 152 oxyrhinu, Copizlla, 177 oxyrhinus, Cophizalua, 177 oxyrhinus, ffypopachus, 113 ~xyrhinus,Phynixalua, 171, 177 Omjrhynchua, 120

.

,

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Pachybat7achu8, 74 Palatine, 4 pallidus, ~ s e o p h w24 , pdmaia, RamaneUa,93 , palmipea, Microhyla, 132 pama; Aphntophryne, 178 Parhoplophryne, 198 parvw, Breviceps, 192 pearsei, Hypopachus, 121 Pedostibes, 5 pentheri, Breviceps, 192 petersi, Pachybatrachua, 74, 75 Phalanges, 6 Pharyngeal Folds, 6 P h y n e k , 105, 107 Phryniacidae, 19 Phrynimlw, 159, 170 Phrynacara, 22, 42 Phrynomantis, 58, 68 Phrynomeridae, 9 Phrynomerinae, 2 phyllodactyla, Cophyla, 41 pictiventre, Engystom, 146 pictiventrie, Gaatrophryne, 146

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mr

MONOGRAPH OF THE MICROHYLIDAE 9

unicolor, Comufer, 153 uniwkw, Microps, 120, 121 Uperodon, 73 Urostyle, 6 usambaricus, Brevicepe, 184 usambaricus, Parhoplophryne, 198 usta, Cfastrophryne,149 &, Microhyla, 148 wrtecm, Engystoma, 144, 149 zcstum, S y s t m , 149

surinamenais, Bufo, 121 sylveatris, Brevieepa, 189 aymbioitica, Ramanella, 90, 93 symmetrica, Ranina, 137 Symphygnathinae, 2, 52 Systoma, 73, 110, 123, 186 syatoma, Bombinator, 75 aystoma, Cacopus, 75 qstoma, R a m , 75 systoma, Uperodon, 75 Systomata, 19

.

valvifera, Asterophrys, 64 valvifera, Pomatops, 58, 64 variabilis, Orwphryne, 163 variabilis, Sphenophryne, 163, 173 variegata, Callula, 93 variegnta, Kaloula, 93 variegata, Ramanella, 93 variegatus, Ccphixalus, 176 van'egaius, Hylophbzcs. 176 variolosum, Engystoma, 112 variolomcm, Systoma, 112 variolosus, Hypopachua, 112,113, 114 v m o s a , Callula, 80, 82 , v e m a a a , Kaloula, 80 verrucosa, Orwphryne, 161, 168, 175 verrucosa, Platyhyla, 37 v e m o s a , Sphenophryne, 175 verrucom, Breviceps, 184, 189, 190, 191, 192,

.Tadpoles, 7 taprobank, Kaluula, 88 Terminal Phalanges, 6 testudo, Rhombopjlryne, 47, 48 texana, W r o p h r y n e , 148 texense, Engy8toma. 148 texensis, k t r o p h r y n e , 148 texensis, Microhyla, 148 Thigh Muscles, 6 thomsoni, Genyophryne, 52, 54 Tongue, 6 Tmierelk, 8 t m i e r i , Cacopoides, 82 t a i e r i , C a U h , 82 t a i e r i , Kaloula, 82 Trends, 15 triangularis, Callula, 94 triangularis, KdozJa, 94 triangularis, Ramanella, M tuberata, Plethodontohyla, 46 tuberatum, Phrynocma, 42, 46 tuberculata, Cophyla, 40 tuberculata, Platypelis, 40 tuberculosus, Pedostibea, 5 tuberifera, Platyhyla. 39 tuberifera, Platypelis, 39 tuberifera, Plethodontohyla, 39 turpieola, Asterophrys, 66 turpicola, Ceratophyne, 66 turpicola, Ceratophrys, 58, 66 turpicula, Asterophrys, 66 tympanifer, Breviceps, 188

-

193

v e w u c o k , Cophixalua, 175 Vertebral Column, 6 visaya, C h p w i n a , 167 voeltzkaui, Platyhyla, 37 volzi, CallueUa, 30 volzi, Dyscophina, 27, 30 r

Kaloula, 82 - wdcwstorji, woltersturfi, Kaloula, 82 Xenobairachus, 54 Xenorhina, 54.68.68 Xenorhinidae, 19, 53

e

uluguruensis, Breviceps, 185 duguruensis, Hoplophryne, 198,199 uluguruensia, Probreviceps, 185 zcndzclata, 4brohyki. 138

yunnhnenais, ~ n l l u e ~29 a, yunmnensis, Kalluella; 29 zimmeri, O r q h r y n e , 164 Zoogeography, 11

*