LS ait th feta Re aT SOU Pann ea Uae peed le : hel \ phan y YENI ENT YS ort a Pern TALL Pan ANRC IR i K due eat USI he ) sia Coin vt Pa GEN Te if, nn ; a ips fehee) sit My prt “ air ay feast eee Shiai it ; pte HP ei rn Steere UY sot : er te Mone ci mi a Mae aoee 1" Nat fe seve roti eRe i! een Ue ete: f cnr Agree ena ‘ Mis Pari ee CORNELL UNIVERSITY LIBRARY BOUGHT WITH THE INCOME OF THE SAGE ENDOWMENT FUND GIVEN IN 1891 BY HENRY WILLIAMS SAGE Gmina? UREA Kescary ie maon The tailless batrachi 7 | i i THE AY SOO 1 ET Y. INSTITUTED MDCCCXLIV. This volume is issued to the Subscribers to the Ray Society for the Year 1896. LONDON: MDCCCXCTIE, THE TAILLESS BATRACHIANS or KUROPE. BY G. A. BOULENGER, F.R.S. PAR £, LONDON: PRINTED FOR THE RAY SOCIETY. MDCCCXCVII. YRINTED BY ADLARD AND SON, BARTHOLOMEW CLOSE, EC., AND 20 HANOVER SQUARE, W. PREFACE. NotwirnstanpvineG the great progress attained in our knowledge of European Batrachians within the last twenty years with regard to the distinction of species and varieties, the study of their anatomy, life-histories, and distribution, there exists at the present day no work dealing with them as a whole from these different aspects. For twenty-five years a close student and collector of these animals, which have always exercised an extraordinary fascination on my mind, I have often wished for an opportunity of supplying such a de- sideratum, by utilising the enormous material which had gradually accumulated in the literature, my own notes, and the unrivalled collection in the British Museum. Thanks to the Council of the Ray Society, who, on the special recommendation of Sir W. H. Flower, acceded to my request, I have at last the satisfaction of seeing my hopes realised, and, owing to the talent of my artistic collaborators, Messrs. P. J. Smit and J. Green, in a manner which fully satisfies my aspira- tions. May those, few as they still are, who share my fondness for this group of animals endorse my opinion of the beautiful and accurate illustrations which are one of the principal features of this work, il PREFACE. and derive from the perusal of these pages one-tenth of the pleasure it has given me to write them; I shall feel amply rewarded for the trouble I have taken. I would also express a hope that a little book of this kind, embodying in a concise form the result of much study, at present scattered in a multitude of publications in some ten different languages, may have the effect of stimulating interest to a subject that has been too much neglected, and in the cultiva- tion of which new workers will find much to repay their efforts, especially if applied in other regions of the globe, which, though much richer in Batrachians, have as yet yielded little or nothing to our knowledge of the life-histories. This work has been planned with the view of assisting the beginner as well as of affording new information to the advanced student, and the synoptic treatment has therefore been frequently resorted to in the Introduction; whilst all descriptions in the sys- tematic part are strictly comparative. Technical terms have not been avoided, but the numerous figures in the text should render them easily intelligible to the beginner. At the desire of the Council of the Ray Society this volume is issued in two parts. The second part, continuously paged and with bibliographical and alpha- betical indexes, is to follow in a few months. The plates, twenty-four in number, have been exe- cuted by Mr. P. J. Smit and printed by Messrs. Mintern Brothers. With two exceptions, all the coloured figures have been taken from living speci- mens. Five of the plates (I, Il, III, XVI, XVII) have already appeared in the ‘ Proceedings of the PREFACE. ii Zoological Society’ (1884, 1885, 1891), to illustrate papers of mine, and have been copied with the sanc- tion of the Committee of Publication of that Society. The figures in the text have been drawn by Mr. J. Green, and reproduced by the Typographic-Etching Company. Most of them are original ; if copied from other works, the source has in every case been referred to. A few cuts illustrating a paper by me on Tadpoles in the ‘ Proceedings of the Zoological Society ’ for 1891, and another by Dr. Ridewood on the development of the hyoid, published in 1897, have been reproduced by permission, for which I beg to tender my acknow- ledgments to the Committee of Publication of the Zoological Society. I also wish to publicly express my thanks to Sir W. H. Flower, K.C.B., and to the Rev. Professor Wiltshire for the interest they have shown in the publication of this work; and to the various friends and correspondents who have assisted me with notes or material, especially Dr. W. G. Ridewood, to whom I am indebted for many hints and preparations; M. F. Lataste, who bas placed his private collection at my disposal; Dr. J. de Bedriaga, Count M. Peracca, Mr. F. A. Moller, Dr. F. Werner, and Herr W. Wolter- storff, who have supplied me with an abundance of living specimens from France, Italy, Portugal, Austria, and Germany ; and Messrs. Chevreux and Doumergue, through whose kindness I have been able to make observations on a large number of living specimens of _ Discoglossus from Algeria. G. A. B. British Museum (NATURAL History) ; October 28th, 1897, THE TAILLESS BATRACHIANS OF HUROPE. INTRODUCTION. . CLASSIFICATION II. II. IV. . SKELETON VI. Vil. VIII. IX. . SPERMATOZOA XI. XII. XITI. XIV. XV. EXTERNAL CHARACTERS INTEGUMENT DERMAL SECRETION VISCERA Hasits . VOICE PAIRING AND OVIPOSITION Haas DEVELOPMENT AND METAMORPHOSIS TADPOLES HYBRIDS GEOGRAPHICAL DISTRIBUTION A PAGE 21 30 33 49 57 61 64 75 78 85 99 112 115 2 INTRODUCTION. I. CuasstricaTion. For many years the Tailless Batrachians (Hcaupata s. AnurA) were classified in a very unsatisfactory manner. The genera Bufo and Hyla were, it is true, always regarded as the types of distinct groups ; but Pelodytes, Pelobates, Discoglossus, Bombinator, and Alytes were placed with Rana, or in separate families in most unnatural associations. Domerit and Bipron, in the eighth volume of their standard work, ‘ Erpétologie générale,’ published 1841, after dividing the order into two sub-orders, Phanéro- glosses and Phrynaglosses (= Aglossa, the two exotic genera Pipa and Xenopus), a primary division first introduced by Wagler in 1830 and which has stood the test of time, combined the Phaneroglossal genera into three families—Raniformes, Hyleformes, Bufoni- formes ; the two latter containing each a single Euro- pean genus, whilst under the former all others were arranged in a series without further subdivision. So unnatural an arrangement evoked criticism from all who were acquainted with the life-histories of the European forms; and an excellent French observer of these animals, A. T'Homas, in a paper published in the ‘Annales des Sciences naturelles,’ 1854, dwelt upon the correlation which exists between the shape of the pupiland the mode of amplexation during parturition, and proposed to group together on the one hand those genera that have a horizontal pupil and an axillary embrace, on the other those that combine a vertical or triangular pupil with a lumbar embrace. These divisions were later provided with names by Bruc# in 1863 (Plagioglena, Orthoglena), and by pr wIste in 1877 (Alamplewes, Inguinampleces). How exaggerated the importance attached to this correlation, which, besides, holds good only for the Huropean forms, is now apparent to all. Yet the ar- rangement proposed by Thomas was a decided advance OLASSIFIOATION, 3. on that of the ‘ Erpétologie générale,’ and on the views of J. Mizner (1832), who overrated the taxonomic importance of the structure of the ear, and inspired the classification followed by Srannivus in the ‘ Zoo- tomie der Amphibien,’ published in 1854. The faults of the latter classification were further developed by GinrHsr in 1858 (‘Cat. Batr. Sal.’), whose avowedly artificial scheme has been followed by Favio in 1872 and by De Burra in 1874, Cors’s epoch-making classification in 1865 (‘ Nat. Hist. Review’), based on a correct appreciation of the osteological characters, placed the matter on a sound footing, and his groupings, slightly modified in 1867, received confirmation, so far as European genera are concerned, from Larastu’s study of the larval characters in 1878 and 1879. In revising the classification in 1882 (‘Cat. Batr. Ecaud.’) I could introduce but slight improvements to Cope’s scheme, as may be seen from the arrangement followed in this work, in which the various groups are placed in ascending order. Principal Schemes of Classification. 1. Dumerit and Bisron, 1841. Tongue absent: PHRYNAGLOSSES. Rana. Discoglossus. without disks: Raniformes Peleuates, T Alytes. ongue present: (toothed; Puantrociosses.| digits BOS Unperdaw 8 Bombinator. Pper Jaw with disks: Hyleformes . Hyla. toothless . ‘ . Bufoniformes Bufo. 2. Sranntius, 1854. = (absent: AGLOssA. a absent. ; ‘ F , Pelobatoidea | pe neres . & | present : scintten on inator. PHANERO-4 | .ocent: oe bein ;fabsent: Bufonina cue Bt Osek. present ; | manubrium ‘ (Alytes. ry digits sterni present: Ranina . Rana Ene dilated at the end . Hyloidea . Hyla. INTRODUCTION. 8. GunruER, 1858. ‘epwuofng 91} WIA poqan aie aepiuhsydowmyy ayy pus ‘pouopuvge mou st ‘auhsydourya SUES UVOTXOTY eT} UO posed “WOISIATP SITE, % oh epy hy ‘ofig - apuosng DULUO{NT a } aaprwoqwurgquog DULLOPDULQULO T POP TL : podopaaap Ajgoasaedurt Aone 4 epyfyy :yueseid BIQOWIOA noid Jonsvatona ssn) PME ih pODSHCED . : “pULwory muy pry POFETIP FOU"; padojaaap Apqoasaad y VSSOTDOURLOU | WOT Ul vey ‘punjhp ‘wpigowphanrg + poyettp quesqe a T99.L sy8tq! rea S Different forms of pupils. a. Bombinator pachypus. B. Alytes obstetricans. c. Pelobatesfuscus. p. Ranaarvalis. E. Bufo vulgaris. in Rana (Fig. 2, Dd), and Bufo viridis and calamita, with an upper and lower angle in Hyla and Bufo vulgaris (Fig. 2,8). That of Discoglossus and Bombinator may be described as roundish or subtriangular according to specimens, the lower angle being always distinct, but the upper border sometimes convex, when the EXTERNAL CHARAOTERS. 11 pupil appears nearly round, or straight when the subtriangular form obtains (Fig. 2, a). In some specimens of Bombinator the upper border is even emarginate, so as to produce the shape of a heart; and when extremely contracted the pupil divides into three branches, affecting the shape of a y (Fig. 2, a) This type of pupil approaches the vertical, and in some specimens of Discoglossus the much-contracted pupil is in fact a little deeper than broad. In Alytes, Pelodytes, and Pelobates (Fig. 2, 8B, c) we find a regular cat’s pupil, vertically elliptical or club-shaped, linear when fully contracted. ‘ The tympanum, or drum of the ear, is absent in Bombinator and Pelobates. When present it may be concealed under the skin, as in some specimens of Discoglossus, Pelodytes, and Bufo vulgaris, or appear on the temple behind the eye as a round or oval disk covered with thin skin (Fig. 1, a, p. 9). The mouth is large, and cleft to beyond the eyes. The jaws are edentulous in Bufo; the upper is armed with numerous closely-set teeth in the other genera— ten to fifteen in each premaxillary, forty to fifty in each maxillary. ‘The teeth have sharply-pointed, slightly-hooked crowns with long shafts applied to the inner side of the premaxillary and maxillary bones. The palate (Fig. 3, p. 12) is pierced by two ° pairs of orifices, the choane, or inner openings of the nostrils anteriorly, and the openings of the Eusta- chian or auditory tubes, situated near the commis- sures of the jaws; the Hustachian tubes are extremely fine, or sometimes even indistinct, in Bombinator. In all the genera except Bufo there are two groups or series of teeth, each implanted on an eminence of the vomerine bones; these vomerine teeth are situated between or behind the choanz, as shown on p. 12, Fig. 8, A, ©, D. The tongue, thick and papillose, flat, is attached in front and in the middle, and the posterior portion, by being bent over and rapidly thrust out of the mouth, 12 INTRODUCTION. acts as an organ of prehension in most tailless Batra- chians; in the Discoglosside, however, as in the newts and salamanders, the tongue is entirely or nearly entirely adherent to the floor of the mouth, and the prey is seized by the jaws. The tongue serves also as an organ of taste, for although frogs seize almost any moving object, they will reject before deglutition anything that is noxious to them, as may be witnessed on offering a frog a brandling or manure-worm (Allolobophora fetida), a ladybird beetle, or a young Bombinator. Before being swal- lowed, the food, if it be a small mollusc, crustacean, or hard beetle, is crushed between the tongue, the depressed eyeballs, and the vomerine teeth; if the latter be absent, as in Bufo, the sharp, sometimes Fig, 3. Open mouths of—a. Discoglossus pictus. B. Bufo calamita. oc. Hyla arborea. D. Rana temporaria, Showing the shape of the tongue ae disposition of the choane, Eustachian tubes, and vomerine teeth. serrated edge of the palatine bones supplies their function. ‘The tongue is circular and entire in the EXTERNAL CHARACTERS. 13 Discoglosside (Fig. 8, a); circular and entire or feebly nicked behind in the Pelobatide and Hyla (Fig. 3, c), resembling a mushroom when thrust out; elliptical or pyriform and entire in Bufo (Fig. 8, 8); oval and forked behind in Rana (Fig. 3, p). In the two latter genera the extensibility of the tongue is so great that a large specimen is able to seize a prey two inches distant from it. Owing to the nature of the skin and the absence of true ribs, the shape of the body varies enormously according to the degree of moisture of the sub- cutaneous sacs, the inflation of the lungs, or the condition of the ovaries, and all measurements that are not based on the bony framework are useless for systematic purposes. The frogs of the genus Rana appear hump-backed in a squatting posture, an angle being produced at the articulation of the cylindrical sacral diapophyses with the ilia; the other genera, with flattened and dilated sacral diapophyses, have a rounded back when at rest, as may be seen on com- paring a true frog with a toad or tree-frog. The vent, or cloacal opening, is a small rounded orifice at the very posterior extremity of the body, above and between the thighs. The fore limb is divided into a brachiwm or arm, an antebrachium or forearm, and a manus or hand with four functional fingers, of which the third is the longest. The length of the first finger as compared to the second is often used as a specific character. In order to preclude misunderstandings, it is well to remark that when the first finger is stated to extend as far as or beyond the second, the two are taken to meet halfway. A radimentary pollex is often indicated externally by a tubercle at the base of the inner finger (Fig. 4, 4). The hind limb is longer, usually very much longer, than the fore limb, and divided into four distinct segments—the femur or thigh, the crus or tibia, or leg, the tarsus, and the pes or foot. Anatomically, 14 INTRODUCTION. the tarsus is, of course, part of the foot proper; but as it here forms a distinct segment, the term foot is used in a restricted sense. The foot is measured from the base of the metatarsal tubercle; it has five elon- gate toes, gradually increasing in length to the fourth, the fifth being again shorter. ‘There is in addition a rudimentary sixth toe, the so-called pre- hallue, conspicuous externally in the form of a tubercle or spur at the base of the inner toe. In Pelobates (Fig. 4, 8) this preehallux or inner metatarsal tubercle acquires a very great development, is covered by a horny sheath with sharp cutting edge, and serves as a shovel for digging in the soil. ‘here is often another tubercle on the sole, at the base of the fourth toe; this so-called outer metatarsal tubercle (Fig. 4, c) is merely a thickening of the integument. Fie. 4. A. Hand of Alytes obstetricans. B. Foot of Pelobates fuscus. c. Foot of Bufo calamita (lower views). Other dermal tubercles are usually present, more or less developed, single (Fig. 6, a, p. 16) or paired (Fig. 4, 0), under the digits at the articulations between the phalanges; they are called subarticular tubercles. The fingers are free except in Hyla arborea, in which they are provided with a rudimentary web. The toes are more or less webbed or bordered by membranes in all our Batrachians, but the extent of the web varies greatly according to the species, and is usually more developed in males than in females, especially during the breeding season. The web is usually smooth and transparent; it is thicker, and often warty, especially towards the margin, in Bufo. EXTERNAL CHARACTERS. 15 In our single representative of the arboreal type, the tree-froe—Hyla arborea—the extremities of the fingers and toes are expanded into adhesive disks (Fig. 6, B), which assist the animal in climbing on vertical smooth surfaces. These disks do not act as suckers, as was once believed, but adhere by rapid and intense pressure of the distal phalanx and special muscles upon the lower surface, which is at the same time provided with numerous glands producing a viscous secretion. The upward rotation of the terminal phalanx, the swollen base of which is hinged upon an interarticular cartilage situated under the extremity of the penulti- mate phalanx, and can be raised or lowered like the claw of a cat, is easily to be observed on a living specimen. When the disk does not adhere (Fig. 5), its upper surface shows a short ridge produced by the claw- like terminal phalanx, and the lower surface is convex with some longitudinal grooves. During adhesion, on the other hand, the claw-like phalanx is no longer visible on the upper surface, but is indicated by a groove, whilst the lower surface is flat and expanded. Section through the extremity of the third toe of Hyla arborea, the bones being indicated by oblique bars. ph?, ph3. Second and third phalanges. ia. Interarticular cartilage. d. Ad- hesive disk. sé, Subarticular tubercle. The adhesion of tree-frogs to smooth vertical sur- faces is, however, not effected entirely through the above-mentioned organs, their function being supple- mented by the suctorial action of the belly, such as is developed, though to a lesser degree, in the young of most Batrachians which, devoid of digital expan- 16 INTRODUCTION. sions, are nevertheless able to slowly ascend a pane of glass. In most of our genera the fourth and fifth toes are cleft almost to the base, and the metatarsal portions of them are separated by the web. In two genera, Bufo and Hyla, this is not the case; the metatarsals of the two outer toes are bound together by the common integument, the angular divergence of the fifth toe proceeding only from the basal phalanx, as may be seen from the following figures (Fig. 6) representing the foot of the common frog and that of the tree- frog. Fia. 6. Lower mo of foot of Rana temporaria (a) and Hyla arborea (B). The tree-frog differs from other Batrachians in having the articulations of the limbs, especially the wrist, marked above by a strong fold, reminding one of the arms and legs of a jointed papier-maché doll. The proportions of the hind limbs to the body are best appreciated by bending the limb forwards against the sige, and ascertaining the position of the tibio- tarsal or tarso-metatarsal articulation ; a certain allow- ance must, however, be made in cases when the abdo- men of a female is much distended with ripe ova. A convenient method of appreciating the length of the tibia as compared to the femur is to fold the limb so as to place the former at a right angle to the axis of the body, as shown in the accompanying figure (Fig. 7) ; in which position the heels (tibio-tarsal articulation) meet when the tibia is nearly equal to the thigh, fail to meet when shorter, or overlap when longer. EXTERNAL CHARACTERS. L Fria. 7. Limbs of Hyla arborea, showing variations in the proportion of the tibia. The dotted line indicates the middle line of the body. The great development of the hind limbs enables these Batrachians to proceed by leaps ; there is, how- ever, one exception among the European forms, viz. Bufo calamitu, in which the limbs are so short as to render hopping impossible, and this toad progresses at a running pace when pursued. The other extreme is furnished by Rana agilis, which owing to its extremely long hind limbs is able to cover a distance of six feet at one leap. The aquatic habits of a Batrachian can, as a rule, be gauged by the extent of the web between the toes: thus Bufo calamita and Alytes, which have only rudimentary webs, are very bad swimmers, and do not venture far from the shore; whilst Ieana esculenta and Bombinator, with their broadly-webbed toes, spend nearly the whole of the active period in the water. But there are exceptions: Pelobates, the toes of which are completely webbed, sojourns but a very short time in the water; it is true that in this case the web is probably of use for pushing aside the soil in the process of digging. The following synopsis of the species is based entively on external characters, and applies to both SeXes. I. Tongue circular, entire, adherent, or slightly free behind ; vomerine teeth behind the level of the choane; first finger shorter than second (DiscocLossip#). r B 18 INTRODUCTION. A. Pupil round, subtriangular, or cordiform. — 1. Vomerine teeth in long transverse series ; tympanum distinct or concealed under the skin. . . . 1. Discoglossus pictus. 2. Vomerine teeth in two groups; no tym- panum. Tibia shorter than the foot 2. Bombinator iqneus. Tibia as long as or a little longer than the foot . . . . 38. Boimbinator pachypus. 8. Pupil vertical; vomerine teeth in short trans- verse or slightly oblique series; tympanum distinct. Three palmar tubercles ; fourth finger as long as or slightly shorter than second 4. Alytes obstetricans. Two palmar tubercles; fourth finger much shorter than second 5. Alytes cisternasit. II. Tongue circular, entire or slightly notched, and free behind; vomerine teeth between the choane ; pupil vertical (PELopatips). A. Tympanum distinct or concealed under the skin; toes webbed at the base and bordered ; inner metatarsal tubercle small, soft 6. Pelodytes punctatus. pn. No tympanum; toes nearly entirely webbed ; inner metatarsal tubercle large, hard, com- pressed, sharp-edged. Interorbital space and occiput convex ; meta- tarsal tubercle yellowish or pale brown 7. Pelobates fuscus. Interorbital space and ccciput flat; metatarsal tubercle black =. 8. Pelobates cultripes. IIL. Tongue elhptical or pyriform, entire, free behind; no teeth ; pupil horizontal (Buronipm). toes at least nearly half webbed, with paired subarticular tubercles; no tarsal fold; inter- EXTERNAL CHARACTERS. 19 orbital space at least as broad as the upper eyelid . . . . . 9. Bufo vulgaris. Toes at least nearly half webbed, with single subarticular tubercles; a tarsal fold; fingers very obtuse ; interorbital space narrower than the upper eyelid . 10. Bufo viridis. Toes webbed at the base, with paired subarticular tubercles ; a tarsal fold usually present; inter- orbital space narrower than the upper eyelid; hind limb not, or but slightly, longer than head and body . . . . 11. Bufo calamita. IV. Tongue cordiform, free behind; fingers and toes dilated at the end into distinct disks; fingers webbed at the base; vomerine teeth between the choane ; pupil horizontal (Hyripz). 12. Hyla arborea. V. Tongue forked and free behind; pupil horizontal (Rantpm). a. Vomerine teeth between or extending slightly behind the level of the choanz; interorbital space not more than half as broad as the upper eyelid . . 13. Rana esculenta. B. Vomerine teeth extending behind the level of the choane ; interorbital space at least one- half the width of the upper eyelid. 1. Tibio-tarsal articulation rarely reaching the tip of the snout, and never beyond; distance between the dorso-lateral folds five to seven times in the length from snout to vent. Inner metatarsal tubercle large, hard, compressed, measuring one-half to two-thirds its distance from the tip of the inner toe; first finger extending beyond second; tibia shorter than the forelimb. . . 14. Rana arvalis. Inner metatarsal tubercle small, soft, oval; first and second fingers equal, or first extending 20 INTRODUCTION. slightly beyond second; tibia as long as or shehtly shorter than the fore limb 15. Rana camerani. Inner metatarsal tubercle small, soft, oval; first finger extending beyond second; tibia con- sider ably shorter than the fore limb 16. Rana temporaria. 2. Tibio-tarsal articulation reaching the tip of the snout or beyond ; tibia nearly as long as the fore limb; distance between the dorso-lateral folds contained four to five and a half times in the length from snout to vent. a. Tympanum not more than two-thirds the diameter of the eye, distant from it. Distance between the nostrils greater than the interorbital width; first and second fingers equal, or first extending slightly beyond second; inner metatarsal tubercle nearly half the length of the inner toe, as long as the diameter of the tympanum . . . 17. Rana greca. Distance between the nostrils greater than the interorbital width; first and second fingers equal, or first extending slightly beyond second ; inner metatarsal tubercle one-third the length of the inner toe, shorter than the diameter of the tympanum . . 18. Rana iberica. Distance between the nostrils not greater than the interorbital width ; first finger extending beyond second; inner metatarsal tubercle one- third the length of the mner toe, shorter than the diameter of the tympanum 19. Rana latastii. hb, 'Tympanum two-thirds to five-sixths the diameter of the eye, and close to it ; first finger extending beyond second ; inner metatarsal tubercle very promi- nent, but moderately large. 20. Rana agilis. INTEGUMEN?. 21 III. Inrecumenr, The integument of the Ecaudata differs from that of other Batrachians in the slight attachment to the Fia. 8. Rana temporaria, showing the dermal attachments and the dis- position of the lymph-sacs, upper and lower views. The position of the lymph-hearts is indicated by asterisks. a, Abdominal sac. | g. Gular sac. | sb. Subbrachial _ sac. b. Brachial » | ¥ Interfemoral ,, sbp. Subplantar 5 ce. Crural » | Lateral a sf. Supra-femoral ,, d. Dorsal » |p. Pectoral ss sp. Supra-plantar_,, f. Femoral 55 muscles, thus rendering the skinning of a frog a most easy operation. But the body is not enclosed in a loose sac, as is often stated in popular descriptions ; there are definite lines of attachment (Fig. 8) by means of connective tissue, which form the demarca- 22 INTRODUCTION. tions between the lymph-sacs that exist on both the dorsal and ventral surfaces immediately beneath the integument. The lymph, fed by the moisture absorbed through the skin, is pumped into the veins by two pairs of lymph-hearts, contractile muscular sacs, the anterior of which are situated in the muscular tissue close to the transverse processes of the third vertebra, the posterior on the sides of the urostyle near its extre- mity; their pulsations may be observed on the back of the living frog. The number and extent of the lymph-sacs varies in the different genera. In Rana and Pelodytes there are 22, 4 unpaired and 9 pairs. ‘The dorsal sac extends from the tip of the snout to above the vent, and is limited by the line of attachment which corre- sponds to the canthus rostralis, the supra-orbital border, the supra-temporal fold, and the glandular dorso-lateral fold. The lateral sac is limited below by a band on each side of the belly, which may be marked externally by a slight groove; the abdominal surface of the head and body is divided into three sacs, gular, pectoral, and abdominal, with partitions across the preecoracoid and coracoid regions. The further sacs (brachial, subbrachial, femoral, supra-femoral, inter- femoral, crural, supra- and sub-plantar) belong to the limbs. In Lyla there is besides a close areolar attach- ment of the abdominal integument. The Discoglos- side differ in the absence of coracoid attachment, or if such exists, as in Discoglossus, it is widely interrupted on the middle line; there is, therefore, no distinet pectoral lymph-sac. In Pelobates the skin of the belly is attached in its posterior half. In Bufo the ventral integument is more or less broadly attached, together with the whole of the dorsal integument. One of the most remarkable peculiarities connected with the integument of Batrachians is the cutaneous respiration, or the power which the surface of the INTEGUMENT, 23 skin possesses of effecting those changes in the blood which are usually performed by the lungs or gills, and by which frogs may remain active for long periods under water. Results of various experiments have proved that pulmonary respiration alone is not sufficient to support life without the aid of that of the cutaneous surface; whilst some of the tailed Batra- chians have lately been shown to be deprived of both lungs and gills, so that the contrary does not hold good for them. But in order to carry out this im- portant function it is of course necessary that the surface be kept in a moist state, and this is effected by a secretion of fluid from the skin itself. An immense number of close-set glandular czaca open upon the surface of the skin, the slime-cells, in addition to which a greater or lesser number of larger poison-secreting glands are scattered or disposed in very conspicuous prominent aggregations, such as the dorso-lateral fold of Rana, the large dorsal warts of Bufo and Bomlbinator, and especially the so-called parotoids, situated above the ear in Bufo and Alytes, and a similar gland on the calf of Bufo calaimita (Fig. 9, p. 24). These glands are pierced with large pores, visible to the naked eye or with the aid of a low magnifier. A round gland, the frontal or pineal gland, the homologue of the so-called parietal eye of reptiles, is more or less discernible under the skin of the forehead in front of and between the eyes; this gland, in the early stages, was connected with the brain. ‘The presence of lime concretions in the skin of old specimens of Bufo vulgaris has been first pointed out by Leydig. The same have been found in B. calamita by O. Seeck. A character which does not appear to have been noticed before is the presence of a filiform line or raphe, on which the skin is much thinner, extending along the middle line of the back from the snout to the extremity of the coccygeal region in all specimens of Bufo. This raphe, which may “pest be observed on 9A, INTRODUCTION. a stripped skin held against the light, gives rise to the yellow vertebral line which is normal in Bufo calamita, and only exceptional in B. viridis and vulgaris, inde- pendently of another light vertebral streak which is sometimes also present. The independence of the two is most conspicuous in cases of deviation of the former, as often happens in Bufo calamita, which appears to be caused by the presence of large glands in the course of the raphe necessitating a winding (Fig. 9). Bufo calamita, showing yellow raphe, independent of light vertebral streak, and its deviation from the median line in the middle of the back. The epidermis is remarkable for the formation of horny cells, as the spines on the warts of some toads and Bombinator (Fig. 10, p. 29), or the sheaths on the tips of the digits in Bufo and Pelobates, or on the metatarsal tubercles when these are much developed. There are also, in some Batrachians, special small, pearl-like warts, mostly pigmentless, the “ tact-spots ” INTEGUMENT. 25 of Merkel, formed by the grouping together of terminal ganglion-cells. These sense-orgaus are found principally on the limbs and flanks of Rana esculenta, more developed in males than in females; the small warts on the fore limbs of Pelobates, which have by some been compared with the nuptial horny excres- cences, seem to be of the same nature. Very similar productions appear during the breeding season in the females of Rana temporaria and It. arvalis, but are of a merely temporary character, and will be noticed in the chapter on Pairing and Oviposition, together with the horny excrescences which arm the digits or other parts in the males at the same period, and assist them In maintaining their hold whilst pairing. Pigment, usually confined to the cutis, sometimes occurs in small quantity in the epidermis. As Leydig has shown, five kinds of special cells may be pre- sent in the cutis—black, yellow, red, white, and metallic or iridescent, the last being the so-called guanin-cells or iridocytes. The yellow and black pig- ments combine to produce the bright green colour, as in normal specimens of Rana esculenta typica and Hyla arborea : if the yellow pigment be absent, as in the German specimens of Rana esculenta, var. ridibunda, the black and gold produce the dull green or olive hue; if both yellow and gold pigments are absent, the black below the cloudy medium of connective tissue and epidermis produces blue, as in the sky-blue specimens of Rana esculenta and Hyla arborea. In the play of the special pigment-cells or chromatophores, which contract or expand and radiate, we find the explana- tion of the changes of colour which some specimens undergo with so great rapidity. A tree-frog will turn from yellow to green and black in less than an hour, yellow when the black pigment contracts, black in the - opposite process. ‘hat these rapid changes of colour harmonise, within certain limits, with the surroundings is well known. The researches of Dutartre on Rana escu- 26 INTRODUCTION. lenta have shown that the changes are controlled by the animal, the contraction or expansion of the chromatophores not being due entirely to the direct action of light and moisture upon the skin, but also to a reflex action of the sympathetic nervous system brought about by the visual impression. Comparative experiments conducted on frogs possessed or deprived of the sense of sight, and placed i in different conditions of light, show the blind individuals to change colour much less rapidly. The changes which we observe in Hyla, Rana, and Bufo ave not limited to the ground colour, but also extend to the markings. ‘The tree-frog, normally uniform, may become covered with lighter or darker dots or spots, sometimes disposed with great regu- larity and even assuming the form of cross-bars on the limbs. An edible frog, of a fine grass-green with few black spots above and with a pure white belly when captured, may shortly after, when transferred to an aquarium, appear largely marbled with black both above aud beneath. IF, Werner has proposed to group the Huropean Batrachians, with respect to this peculiarity, into such as actually change colour (.!/ytes, Hyla, Bufo, Rint), and such as only hghten or darken their shade (Misvoglossus, Pelodytes, LPelobutes). He oe Bombinator as representing a third category, n changes of any kind taking place; but I cannot agree with him in this, as I have seen B. pachypus pass from yellowish to a very dark brown, and also vary in the presence, absence, or distinctness of the light and dark markings. In a recent work on Hast African Reptiles and Batrachians G. Tornier endeavours to show that the yellow and red pigments do not exist as chemical entities, but are only degenerations of the brown pigment or “melanin.” And as a first “ simple and sure” proof of his contention he adduces the example of Pelobates fuscus larve, which according to him are INTEGUMENT. Oe “quite black,” and during metamorphosis assume the light yellowish-grey ground colour through the greater part of the pigment-cells turning pale. A more glaring misrepresentation could not be imagined. Far from being black, the tadpole of Pelobates Fusens is normally pale brown, with darker spots and marb- lings, as may be seen from the figure in this work. And the tadpole of Pelobates cultripes, a species which has the same dark spots as P. fuscus after transfor- mation, is usually very pale reddish yellow. A proof that the yellow and red pigments are chemically dis- tinct from the brown or black is to be found in the different action of alcohol on them. ‘Thus yellow pigment is rapidly destroyed; therefore green frogs turn olive or blue in spirit. Certain crimson pig- ments, not unfrequent in frogs, resist the action of alcohol and light better than the brown, which by dis- coloration may turn to reddish brown or rufous, these two reds strongly contrasting in specimens that have long been in spirit. ‘l'ornier goes even so far as to doubt the | presence of any but the brown pigment in reptiles ; surely the green of certain tree-snakes of the genera Dryophis and Lachesis, which colour the spirit in which they are preserved to the extent of resembling green Chartreuse, is due to the presence of a chemically distinct colouring matter, and has no relation whatever to the green of frogs, which, produced, as we have seen, by a combination of black and yellow, disappears in spirit when the latter pigment is destroyed. Another instance of the same author’s carelessness in the selection of examples is to be found a little further in the same work (p. 130), where he declares that Batrachian larve which develop in the light are, with few exceptions, uniformly dark, or more “usuall deep black, and retain that coloration until close to the metamorphosis. I believe the exception is just the other way. The embryos of Bombinator and Alytes are yellowish or pale brown striped with blackish, those of Hyla uniform yellowish, those of Lana 28 INTRODUCTION. esculenta feebly pigmented; and of black tadpoles, or of a dark brown approaching black, I only know Bufo and Raw temporaria; all the other Kuropean species are only exceptionally, not normally very dark. That the light cannot have a great effect in producing the pigment in Batrachian larve is shown by Alytes and Bombinator, both of which develop the same amount of pigmentation, the one under paternal care never being exposed to daylight, the other reared in pools or puddles exposed to the full action of the sun’s rays. Cases of melanism, or better, absence of iridocytes and all pigment but the brown or black, which may be very abundantly and exclusively present, have been observed by Héron-Royer in Alytes obstetricans, by Alfred Dugés, and more recently by Héron-Royer and by Vaillant, in Rana esculenta. I found a similar young specimen of the latter species in a marsh near the Belgian coast in 1884; the back was brown, nearly black, the hinder side of the thighs and the ventral surface pigmentless, transparent flesh-colour, the iris black without any gold. A male specimen of ana greea from Italy, showing the same abnormality in the absence of all but the brown colour, was sent to me last spring by Count Peracea, and is figured on Plate XXII of this work. Albinism, or absence of the brown pigment, on the other hand, has been observed in a young /ombinator pachypus by Fatio, in larvae of Alytes obstetricuns by Lataste and Héron-Royer, who succeeded in rearing them through the metamorphosis, in a larva of Disco- ylossus pictus by myself, in Lane esculenta by Pavesi, in larve and young of Bufo viridis by Born, Camerano, and myself, in larvae of Rana temporaria by Lessona, Fischer-Sigwart, and Camerano; an adult female of the latter species found in England was exhibited alive by Mr. Rowland Ward at the Linnean Society in 1891, and another, from Wiltshive, presented by Mr. W. Hannaford, is now preserved in the British INTEGUMENT. 29 Museum. These albinos are of a uniform yellowish flesh-colour, with silvery or golden iris and cherry-red pupil; the yellow ventral blotches are preserved in Bombinator. A figure of a young Alytes albino reared by Lataste is given on Plate VII. Albino specimens of Alytes obstetvicans obtained from tadpoles by Héron- Royer paired in confinement, and produced offspring that were likewise affected with complete albinism. But for the difficulty in getting these Batrachians to breed in captivity, a race of albinos could no doubt be fixed by artificial selection, as in the case of the axolotl, the numerous “ white’? examples of which are said to be all descended from an albino male which paired with normal females in the Jardin des Plantes at Paris some thirty years ago. Batrachians often ‘“‘change skin,’—about once a month during the spring and summer. The thin, trans- parent outer layer of the epidermis becomes detached about the mouth, and, lubricated by an abundant secre- tion of the slime glands, is pulled off by movements of both fore and hind limbs, often in a single piece turned inside out, especially when the operation is performed in the water, and finally swallowed by its owner. The casting of the epidermis on land is a slower process, accompanied by ludicrous contortions ; in the common toad it may last nearly an hour. Fra. 10. Piece of dorsal integument of male Bombinator pachypus (x 8), showing warts and epidermal horny asperities. 30 INTRODUCTION. TV. Durmat SECRETION. The skin of the dorsal surface of all Batrachians is, as we have seen, studded with glands and follicles. In various species, of which our Dufones are good examples, some of the larger glands appear as very prominent warts pierced with wide pores, especially a large mass situated on each side of the back of the head behind the eyes, which has received the mis- nomer of parotid, a name now generally changed to parotoid gland. These glands secrete, more or less abundantly according to the species, a viscid or milky more or less and variously odorous flud, which is exuded, when the animal is disturbed, by means of the contraction of small muscles disposed in the skin around the glands. If the larger glands be pressed or subjected to an electric current the secretion is squirted out with great force, and sometimes to a considerable distance. Kobert recommends, as the best means of obtaining an abundant supply of toad- poison, the subcutaneous injection of a small dose of chloride of barium ; after a few minutes the animal is so covered with its secretion as to appear as if coated with whitewash. The object of the secretion is to guard Batrachians from the attacks of various enemies. A dog will not often repeat the experiment of seizing a toad; the signs of pain which it evinces after the poisonous fluid has come into contact with its mouth is familiar to most of those who have kept young dogs in the country. That excellent Hungarian naturalist, Prof. v. Méhcly, relates the serious effects of the poison of Bufo viridis on a small terrier which accompanied him on a collecting excursion. " | Bufo viridis ~ Rana yreea Tine iberica . tian agilis. —. 10. Bufo calamitu . 11. Rana esculenta 12. Hylu arborea ~ f-Alyles obstetricans 4:, 6 ‘ : 7. Bombinator pachypus 8 : 0 INTRODUCTION. Feet. 6500 6500 5800 5900 4500 42.00 4,000 3500 3300 The other species do not ee to 3000 feet. * The following table shows the distribution in the different countries of Kurope: 1 2. . Discoglossus pictus er . Bombinator igneus i puchypis : Alytes obstetricans » clstermesls . a cultripes . » viridis . mo Ge » calanita + > «rvales. camercite ae 4 temporaria ey bse se Guiven. ees » tberica. yy lttastie, nn agilis Di ++++4ti ti: 1. Great Britain and Iveland. 2. Scandinavia and Denmark. 3. France. 4. Belgium und Holland. 5 Germany and Switzerland. 6. Spanish Peninsula. 7. Italy. 3. D++ettee4: tH: of! 4. D+i Di ttt+i tit: 8. Austria-Hungary and Roumania. 9. Dalmatia, Bosnia, and countries south of the Danube. 10. Russia. P +44: 9 Di f++Httti t: +44. t4¢i tttii +e +) D+: ++ +4: +i 43 Ptti tt +4 8. 9. + ous + + + + + + + + + + + + AE ast + + ado Ate + + 10. + t+4++44+4+4+4+ 4+ * 15,000 feet in the Himalayas. GEOGRAPHICAL DISTRIBUTION. 119 It may also not be without interest, in order to bring out their associations, to enumerate the species occurring in the environs (7. e. a radius of about fifteen miles) of some of the principal towns the Batrachian fauna of which has been thoroughly searched by com- petent workers. It will be seen, then, that no district can boast of possessing more than ten out of the twenty recognised species, and that, in spite of their northern position, Paris and Bonn yield to no other and are richer than many more southern localities. 1. London (three species). Bufo vulgaris, B. calamita, Kana temporaria. 2. Copenhagen (nine species). Bombinator igneus, Pelobates fuscus, Bufo vul- garis, B. viridis, B. calumita, Hyla arborea, Rana esculenta, It. arvalis, R. temporaria. 3. Paris (ten species). Bombinator pachypus, Alytes obstetricans, Pelo- dytes punctatus, Pelobates fuseus, Bufo vulgaris, B. culamita, Hyla urborea, Rana esculenta, It. temporaria, Et. agilis. 4, Bordeaux (nine species). Bombinator pachypus, Alytes obstetricans, Pelo- dytes punctatus, Pelobates cultripes, Bufo vulgaris, B. calanita, Hyla arborea, Rana esculenta, Lt. agilis. Brussels (five species). Bufo vulgaris, B. culamita, Hyla arborea, Rana esculenta, It. temporaria. 6. Berlin (nine species). Bombinator igneus, Pelobates fuscus, Bufo vul- garis, B. vividis, B. calumita, Hyla arborea, Rana esculenta, Lt. arvalis, R. temporaria. 7. Bonn (ten species). Bombinator pachypus, Alytes obstetricans, Pelo- bates fuscus, Bufo vulgaris, B. viridis, B. calamitu, Hyla arborea, Rana esculenta, R. arvalis, R. temporaria. rt 120 INTRODUCTION. 8. Geneva (eight species). : Bonbinalor pachypus, Alyles obstetricaus, Bufo vulgaris, B. calumita, Hyla arborea, Runa esculenta, Re. temporaria, Le, agilts. 9. Coimbra (ten species). Discoglossus pictus, Alytes obstelrivans, 1. cister- nusti, Pelodyles yraetatus, Pelobates cul- tripes, Bufo vulgaris, B. calamita, Hyla arlurea, Rana esculenta, R. ‘berica. 10. Turin (seven species). Pelobutes fuscus, Bufo culyavis, Be vividis, Hyla arborea, Lana esculenta, R. latastit, De. ayilis, 11. Palermo (five species). Discoglossus pictus, Bufo vulgaris, B. viridis, Hyla arborea, Rana esculenta. 12. Vienna (ten species). Bombinutor iynens, Be pachypus, Pelobates Juscus, Bufo culgaris, Bo viridss, Hyla arborea, Lana esculenta, I. arvalis, 2. temporaria, R. agilts. 13. St. Petersburg (three species). Bufo vulyaris, Rane urcalis, R. temporaria. 14. Moscow (eight species). Bombinator iqneus, Pelobates fuscus, Bufo vul- garis, DB. viridis, Hyla arborea, Luna esculenta, Lt. arvalis, Lt. temporaria. And finally, in order to complete the sketch of the Geographical Distribution, a list is appended of all the tailless Batrachians known from other parts of the Palearctic region in its widest sense, viz. North Africa and Asia north of the 30th parallel. Column A stands for North-western Africa, B for North-eastern Africa, C for South-western Asia, D for North- western and Central Asia, E for Hastern Asia with Tibet, and F for Japan. The names of species occur- ring also in Hurope are prefixed with an asterisk. GEOGRAPHICAL DISTRIBUTION. DiscocLossipH&. *Discoglossus pictus, Otth . Bombinator orientulis, Blgy. PELOBATIDA, Pelodytes caucasicus, Blyy. *Pelobates fuscus, Laur. 55 syriacus, Bttgr. . BuFonips. *Bufo vulgaris, Laur. » formosus, Bler. » viridis, Laur. » vraddit, Strauch > maur itunicus, Sehleg. » regularis, Reuss HYLips. *Hyla arborea, L. » stepheni, Blgr. . RANID a. Rana boulengeri, Gthr. » Umnocharis, Boie » rugosa, Schleg. * |, esculenta, L. » plancyi, Lat. * 4, arvalis, Nilss. . * 4, camerant, Bigr. >» macrocnemtis, Blgr. . * ,, temporaria, L. » amurensis, Blgr. a martenstt, Blgr. » japonica, Blgr. * 4, ayilis, Thom. » mascar eniensis, D.& B. schmackert, Bttgr. Rhacophorus buergert, “Schleg. 3 davidi, Sauv. Ss schlegelti, Gthr. A. + +: B. +++ 2 Spe ee boskeps +4+i4¢+4+ 44+ ++: +4+++ ++ Pi + 121 ++ +++ [Aé page 123. 70 » PACHYPUS, ALYTES OBSTETRICANS, “ CISTERNASII. 50 DISTRIBUTION OF EUROPEAN DISCOGLOSSID A. DISCOGLOSSIDA. 123 Order ECAUDATA., Four limbs and no tail. Radius and wna, and tibia and fibula confluent; tarsus (astragalus and caleaneum) elongate, forming an additional segment in the hind limb. Frontal bones confluent with parietals. Sub-order PHANHROGLOSSA. Hustachian tubes separated ; tongue present. Series AW—ARCIFERA. Pectoral arch with the opposite halves moveable, the coracoids and precoracoids connected by an arched cartilage (the epicoracoid), that of the one side over- lapping that of the other. Family 1.—DISCOGLOSSID.A. Vertebree opisthoccelous ; short ribs articulated to the anterior diapophyses; diapophyses of sacral vertebra dilated. Upper jaw toothed. The genera combined under this family constitute ‘a most interesting and perfectly natural group, as 1s abundantly evidenced by the bony structure, the larval characters, &c. That they occupy the most lowly position among the Heaudata, and show the nearest approximation to the Caudata, is another point on which there can be no question. The opisthoccelous vertebree with distinct ribs, the increased number of carpal and tarsal ele- ments, the non-extrusible tongue, the presence of the azygos (posterior cardinal) vein discovered by Hoch- stetter in Bombinator, and since shown by Howes to be 124 DISCOGLOSSIDE. fairly distinctive of the whole family, the structure of the urogenital apparatus, together with other cha- racters, give ample foundation to this proposition. This family comprises only four genera. Three are confined to the Palwarctic region and represented in HKurope; the fourth, Liopelma, Fitzinger, closely allied to Alytes, is the only representative of the Batrachians in New Zealand. The range of the species dealt with in this work is shown on the accompanying map. The European genera are distinguished as follows : Tympanum present, distinct or hidden; pupil roundish or triangular; diapophyses of sacral vertebra moderately dilated . 1. Discoglossus. ‘'ympanum absent; pupil roundish or triangular ; diapophyses of sacral vertebra very strongly dilated. . . . . . . . 2, Bombinator. Tympanum distinct ; pupil vertical; diapophyses of sacral vertebra strongly dilated. 3. Alytes. The relationships of these three genera cannot be well expressed in a linear arrangement LDiscoglossus is unquestionably the most generalised, and Bombinator and Alytes are almost equally related to it, the latter being, on the whole, more affine to Discoglossus than to Bonbinator, especially with regard to the osteological characters. 1. DiscoaLossus. Otth, Neue Denkschr. allgem. Schweiz. Naturf. Ges., i, 1837, p. 6. Pupil roundish or triangular. Vomerine teeth in long transverse series behind the choana. Tongue circular, entire, scarcely free behind. Tympanum more or less distinct or concealed under the skin. Fingers free, toes webbed ; outer metatarsals separated by web. Diapophyses of sacral vertebra moderately dilated. Urostyle articulated to two condyles. A single species, confined to the western parts of the Palearctic region. DISCOGLOSSUS. 125 The pupil is described as roundish or triangular ; in its contracted condition it varies considerably according to individuals. It is never absolutely round, for it forms an angle below, whence a subtri- angular form results, I have seen specimens in which the vertical diameter somewhat exceeds the horizontal, so that the fully contracted pupil might be termed vertically pear-shaped, or, the upper border forming an open angle, kite-shaped. 1. Discocossus PIoTus. (Plate IV.) Cetti, Anf. e Pesce. Sard., iii, p. 38 (1777). Rana temporaria (non L.), Rozet, Voy. Alg., i, p. 230 (1833). Discoglossus pictus, Otth, N. Denkschr. allgem. Schweiz. Nat. Ges., i, 1837, p. 6, figs.; Tschudi, Class. Batr., p. 80 (1838); Bonaparte, Icon. Faun. Ital., Rett. Anf. (1838); Duméril & Bibron, Erp. Gén., viii, p. 425 (1841); Ginther, Cat. Batr. Sal., p. 35 (1858); Strauch, Erp. Alg., p. 77 (1862) ; De Betta, Atti Ist. Venet. (3), xiii, 1868, p. 77, and Faun. Ital., Rett. Anf., p. 67 (1874); Schreiber, Herp. Eur., p. 112 (1875); Camerano, Atti Acc. Torin., xiv, 1879, p. 448, pl. —, figs. 6—8; Lataste, Act. Soc. Linn. Bord., xxxiii, 1879, p. 275, pls. ii—v; Bedriaga, Bull. Soc. Nat. Mosc., 1881, p. 292; Boulenger, Cat. Batr. Ecaud., p. 445 (1882); Camerano, Mem. Ace. Torin. (2), xxxv, 1883, p. 204; Bedriaga, Arch. f. Nat., 1883, p. 254; Héron-Royer, Bull. Soc. Zool. France, 1885, p. 565, pl. xiv; F. E. Schulze, Sitzb. Ges. Naturf. Fr. Berlin, 1886, pp. 5 and 31; Héron-Royer & Van Bambeke, Arch. Biol., ix, 1889, p. 280; Bedriaga, Bull. Soc. Nat. Mosc, 1889, p.545, and Amph. Rept. Portug., p. 22 (1889); Héron- Royer, Bull. Soc. Et. Sc. Angers (2), xix, 1889, p. 160, pls. i and ii; Boulenger, Trans. Zool. Soc., xiii, 1891, p. 160, and Proc. Zool. Soc., 1891, p. 620, pl. xlvii, fig. 3; Anderson, Proc. Zool, Soc., 1892, p. 24; Bedriaga, Suppl. Amph. Port., p. 13 (1893); Mina-Palumbo, Nat. Sicil., xu, 1893, p. 262. Discoglossus sardus, Tschudi, in Otth, 1. ¢., and Class. Batr., . 80; Bonaparte, 1. c.; Camerano, Atti Acc. Torin., xiv, 1879, p. 437, pl. —, figs. 4 and 5. Pseudis sardoa, Gené, Syn. Rept. Sard., p. 24, pl. v, figs. 1—3 & 6 (1839). ' Rana picta, Schlegel, in Wagner, Reisen Alg., iii, p. 134 (1841). Colodactylus czerulescens, Tschudi, Faun. Per., Herp., p. 68, pl. xi, fig. 2 (1845); Peters, Mon. Berl. Ac., 1873, p. 414, pl. iii, fig. 1. Discoglossus scovazzit, Camerano, Atti Acc. Torin., xiii, 1878, p. 548, and xiv, 1879, p. 447, pl. —, figs. 1—8. Discoglossus auritus, Hévon-Royer, Bull. Soc. Zool. France, 1888, pp. 205 and 220, and Bull. Soc. Et. Sc. Angers (2), xix, 1889, p. 177, pls. i and ii. 126 DISCOGLOSSID. Vomerine teeth in a long, straight or slightly curved series, narrowly interrupted in the middle, behind the choanw, and extending outwards to the Fra. 48. vertical of their outer borders or even a little beyond. Tongue moderately large, circular, thick, adherent, scarcely free behind. Head much depressed, a little broader than long; snout rounded or obtusely pointed, projecting considerably be- yond the mouth, as long as or a little longer than the diameter of the orbit ; no canthus rostralis; dorsal region slightly grooved ; nostril a little nearer the tip of the snout than the eye; eye moderate; interorbital space as broad as or a little narrower than the upper eyelid, and equal to the distance between the nostrils; tympanum measur- ing three-fifths to two-thirds the diameter of the eye, sometimes feebly distinct, often completely hidden. Open mouth. Fie. 49. Heads of males, upper views. A. Corunna. gs. Oran. ¢. Tlemsen. p. Sardinia. DISCOGLOSSUS. 127 Fingers rather short, obtusely pointed, first shortest, third longest, second and fourth equal; no sub- articular tubercles ; three palmar tubercles, the inner (rudiment of pollex) largest and very prominent, the two others flat and close together, at the base of the third and fourth fingers. Hind limb rather long; the tibio-tarsal articulation reaches the tympanum, the eye, or the nostril; tibia longer than the femur, the heels overlapping when the legs are folded at right angles to the rhachis. Foot a little shorter than the tibia ; toes rather slender, one-fourth or one-third webbed in females and young, three-fourths or nearly entirely in adult males; no subarticular tubercles ; no tarsal fold ; asmall rounded inner metatarsal tubercle, the length of which equals one-third to one-half the length of the inner toe. Skin remarkably shiny in life, smooth or with small warts or short glandular folds on the back ; a more or less developed glandular lateral fold from the eye to above the shoulder, often prolonged on the side of the body to the inguinal region; a fold of the skin may be present across the occiput behind the eyes. Lower parts smooth except the thighs, which are granulate near the vent; throat and belly sometimes with isolated small granules. Coloration very variable. Pale brown, grey, greyish olive, yellowish or red above, uniform or with dark, often light-edged spots; some specimens with a broad yellow vertebral stripe, which may be bordered on each side by one or two dark stripes; the lateral glandular fold light, often reddish or golden; a dark streak on each side from the tip of the snout to the eye, and a dark temporal blotch or streak ; limbs with transverse dark spots, sometimes forming regular cross-bands; a more or less distinct light triangular or heart-shaped spot often present on the middle of the back between the fore limbs, and another, more in the form of a streak, on the coccygeal region. The dorsal spots sometimes small and irregular, some- 128 DISCOGLOSSIDA. times forming handsome symmetrical markings; alarge triangular, heart-shaped, or chevron-shaped marking is usually present between the eyes. Lower parts ivory-white, uniform or closely speckled with brown, yellowish or more or less carneous under the limbs. Iris golden in its upper portion, dark bronzy in its lower two-thirds, the dark lateral streak of the head passing through the eye ; sometimes entirely bronze- brown, as in Bonbinutor, with a fine golden border to . the pupil, interrupted at the lower angle. Fia. 50. Lower view of male with nuptial excrescences. Male distinguished from the female by much stronger muscular fore limbs, more devcloped inner carpal tubercle, a great enlargement and flattening of the inner finger, and more fully webbed toes, the web even extending to the inner metatarsal tubercle. Vocal sacs are in a rudimentary condition, and do not com- DISCOGLOSSUS. 129 municate by any openings with the floor of the mouth. Horny blackish excrescences are largely developed and distributed as isolated minute spines over the ventral surfaces and on the hind limbs; they form large groups on the inner palmar tubercle and the two inner fingers, a band round the chin, and often border the web of the foot. These excrescences usually persist longer than in most other Batrachians, being found, more or less developed, in adult specimens all the year round. GEOGRAPHICAL Variations.—The polymorphism of this species has given rise to the establishment of various species and sub-species, which were believed to be restricted to certain parts of the habitat of the genus. Thus a Discoglossus sardus, Tschudi, was stated to be peculiar to Sardinia and Corsica, and a D. scovazzii, Camerano, was described from Morocco. Lataste, in his monograph of 1879, had, it seemed, disposed once for all of these supposed species by showing the inconstancy of the characters adduced for their separation ; and, for my part, after examin- ing avery large material from almost every part of the habitat, I fully agree with him. But, since 1879, the D. sardus has been maintained, no longer as a species, but as a sub-species, by Camerano, and the D. scovazzii, or a form very closely agree- ing with it, has been revived by Héron-Royer under the name of D. auritus; it may therefore be well to explain in a few words why I take no notice of them even as varieties. According to Camerano’s latest definition, J. sardus, from Sardinia, Corsica, and neighbouring small islands, differs from the typical D. pictus from Sicily, Malta, and the Spanish Peninsula in having a less acuminate snout, more robust limbs, and the length of the tibia contained twice, or a little over twice, in the length of head and body. The two former characters are really too slight to be easily appreciable ; and besides, I find Maltese, Algerian, and Portuguese specimens with the snout I 130 DISCOGLOSSIDE. quite as short and broadly rounded as in the Sardinian form. As already observed by Lataste, this difference is no greater than that between specimens of Rana temporaria, and is certainly not more constant. With regard to the third character, I need simply refer to my tables of measurement and to Camerano’s own table, where Sardinian as well as Sicilian specimens are shown to have the tibia less than half the length of head and body. Bedriaga also is inclined to retain D. sardus as a distinct form, although admitting that the existence of annectant specimens renders a precise definition impossible. On the whole, he finds the body shorter in proportion to the head and limbs in Corsican and Sardinian specimens than in those from Algeria and Portugal. ‘That there is nothing constant in that supposed difference may be seen from the following measurements (in millimetres) of four adult males, the heads of which are figured above (p. 126),—a from Corunna, 6 from Oran, ¢ from Tlemsen, prov. Oran, and d from Luras, Sardinia : a b c. d. From snout to vent : BS ... 62 «. Fl «x 60 Length of head ‘ i ... 19... 20 .. 18 Width of head at angles of mouth TO: 22: OE ne BB vB is 5, below eyes . » 16 .. 17d. 20 .. J8 From end of snout to fore limb OF tee DOr yen BO gas “DO Fore limb 5 ‘ . 84... 85... 850, 85 Hind limb. : B83 my 95 acs 92 a, 91 Tibia . ‘ ‘ » B28 wn SF we 932 ae 30 It will be noticed that the Spanish and Algerian specimens a and J have the head a little longer in proportion to the body than in the Algerian ¢ and the Sardinian d; that the width of the head below the eyes is as great in proportion in the Algerian ¢ as in the Sardinian d; and that the hind limb is shorter in pro- portion in the Algerian c than in the Sardinian d. Héron-Royer’s D. awritus is founded on Algerian specimens (which had been previously referred by Camerano to D. scovazati), supposed to differ from the Europeans in having the tympanum distinct, the tem- DISCOGLOSSUS. 181 poral spot larger, and the inner metatarsal tubercle smaller. Buta male specimen from Algiers, collected by Mr. Sclater, has the tympanum completely con- cealed ; and, on the other hand, the organ is perfectly distinct in a male from Corunna and in another col- lected by M. Boscd on the Sierra Morena; whilst a number of specimens from Europe and Africa are in- termediate between the two extremes. I have now before me a large number of living specimens from Oran. In some the tympanum is very apparent, whilst in others its presence can hardly be detected. The shape and extent of the dark temporal band and the size of the metatarsal tubercle are subject to variation in specimens from the same locality, and I have failed to find any constancy in the other very trivial distinc- tive characters pointed out by Héron-Royer. Dr. Anderson, who has collected specimens in Algeria, some of which show no trace of a tympanum exter- nally, concurs with me in rejecting D. auritus even as a local form. Specimens from Sardinia, Corsica, Montecristo, and Giglio, it has been observed, never show the striped form so frequent in Spain and Africa; but neither do the Maltese specimens, which are referred by Camerano to D. pictus. Measvrements (in millimetres). é 3 1. 2. 3. 4, 5. 6. 7. 8. From snout to vent 58 ...55 ...60 ... 73 ...55...54...43 1.76 Length of head 18 ...16 ...18 ... 21 ...16...15...14 ...19 Width of head : 19 ...18 ...21 ... 25 ...19...16...15 ...23 Diameter of eye Be sc Bee BD eee Cay Bis thee Blom 6 Interorbital width . Sb. B Dena A ace Ae Baw Baw BD we D From eye to nostril 2b ..4 450. Fb 4 4 4a 8 _ 3 endofsnout . 8 ... 75... 85... 10 0. 7... 7... 6 2.9 Fore limb . i 2 84 ...29 1.85 0. 40 ...27...26...24 ...85 Hind limb . 83 ...80 2.91 1.105 ...84...77...64 2.95 Tibia ; 28 ...26 ...80 ... 85 ...26...25...20 ...33 Foot . ; ‘ 26 2.25 ...26 ... 82 ...24...22...18 ...80 1, 5. Corunna: Seoane. 4. Oran: Chevreux. 2,6. Coimbra: Gadow. 7. Giglio: Florence Mus. 8. Luras, Sardinia: Camerano. 8. Algiers: H4ron-Royer. 132 DISCOGLOSSIDZ. SKELETON.—Ethmoid short, not extending poste- riorly beyond the anterior third or two-fifths of the parasphenoid, embracing above a large fontanelle which extends to the line of the anterior borders of the orbits, and which remains exposed between the diverging anterior borders of the fronto-parietals ex- cept in very old specimens ; its upper lamina obtusely pointed and partly covered by the nasals, which are large and form a long suture in the median line. Fronto-parietals large, in contact along their posterior two-thirds or more. Zygomatic branch of the squa- mosal joining by suture an ascending process of the maxillary. Vomers large, their posterior toothed border nearly straight, and covering the feeble pala- tines, narrowly separated from each other in the median line; pterygoids regularly trifurcate, with a small round outer wing-like expansion at the meeting- point of the three branches, joining a corresponding process of the angular bone of the mandible; the inner branch of the pterygoid in contact with the parasphenoid, which is L-shaped, obtuse or truncated, or indentated anteriorly, and not quite reaching the vomers. Mento-Meckelians quite indistinct. Hyoid a large, broad cartilaginous plate with rounded lateral wings, small postero-lateral processes, and proximally slender, mesially broad and lamellar ceratohyal cornua without forward processes ; in addi- tion to the rather large thyrohyals, which meet with their inner angles, there is a V-shaped, slender, ventral ossification, which is in contact with the latter at its apex, and reaches forward to the anterior notch of the cartilaginous plate ; this ossification often paired, the two branches being disconnected at the apex. Vertebral column twice to twice and one-third as long as the skull. Vertebre imbricate and completely covering the spinal cord above, with a long postero- median neural process. Second, third, and fourth vertebre with short diapophyses, +0 which short ribs are attached. The first rib is the shortest, horizontal 133 DISCOGLOSSUS. Whole skeleton of female and fore limb of male. 184 DISCOGLOSSIDA. or directed a little forwards ; the second is the longest, nearly horizontal or directed a little backwards, and sends off from the middle of its upper border a short, slender process, directed outwards and backwards ; the third rib, which may be somewhat curved, is directed slightly backwards, and in length is intermediate be- tween the first and second. The following vertebrae have no distinct ribs, but the diapophyses are longer and more slender, the last two or three being turned more or less distinctly forwards. The diapophyses of the sacral vertebra, which cover the anterior extre- mities of the ilia, on which they may shift forwards and backwards to some extent, are moderately dilated, their diameter at the free extremity being somewhat less than their length. Two condyles for articulation with the urostyle. The latter is a little longer than the rest of the vertebral column, and bears on each side at its base a short, slender, transverse process directed obliquely backwards. Preecoracoids slender, curved, entering the glenoid cavity; coracoids stronger, feebly curved; supra- scapula ossified ; omosternum cartilaginous; sternum produced into two long, slender, diverging processes. Humerus once and a half to once and two-thirds as long as radius-ulna. Carpus with eight bones, two of which are in contact with radius-ulna ; a single bone in the pollex, very large in the male ; metacarpal of inner digit enlarged in the male, with strong inner crest. Pelvis nearly two-thirds the length of the vertebral column ; pubis cartilaginous ; acetabulum open. Femur and tibia with the epiphyses cartilaginous, the former strongly sinuous and shorter than the latter; astra- galus a little longer than calcaneum, from which it is pertectly distinct throughout, owing to the absence of calcified epiphyses, and one-half or rather more than half the length of tibia-fibula; three tarsalia in the distal row, one or two of which may ossify ; and two bones to the prehallux. Distal phalanges obtuse, slightly expanded at the apex. DISCOGLOSSUS. 1385 Meascrements or SkeLeron (in millimetres), Length of skull. 5s § ns is Width of skull ; ‘ : 24 sen 17 Least interorbital width 4 te 3 Dorsal vertebral column 18 Pe 17 Urostyle . : 19 wi 19 Humerus . 17 be 12 Radius-ulna 11 8 8 Manus : a7: 2 11 Pelvis a 7 a 23 Femur 26 mde SL Tibia 31 aes 24. Tarsus 17 he 13 Pes ; 25 21 Hasirs.—Discoglossus pictus resembles the true frogs im the quickness of its movements. It is active by day as well as by night, and usually found in or about water, objecting neither to brackish pools nor to cold running mountain streams. Like its relative Bombinator, and unlike all other European Hcaudates, it is able to seize its prey under water, in the same manner as newts do, and is therefore easily fed in confinement with earthworms dropped into the tank with which the terrarium must be provided. Although particularly shy when handled, it does well in captivity, and will usually take food immediately on arriving after a long journey in a-small box. When seized it becomes covered with a slimy secretion, which renders it very difficult to handle, but which has no special odour nor any irritating action on our mucous membranes, so far as I can judge from experi- ments upon myself. Like Lataste, who was the first to study the habits in captivity, and who later had the good fortune to observe this Batrachian in a wild state in Algeria, I have never heard the rutting male produce more than a feeble note, sounding to my ear as ha-a, ha-a-a, or wa, wa, wa-wa-wa, issued in rapid succession, although specimens have repeatedly paired and bred in my terrarium. But according to Héron- Royer, specimens under his observation uttered at night a constantly repeated.ra-a, ra-a, loud enough to disturb him when produced in a yard outside his 186 DISCOGLOSSID. window. Discoglossws cannot be described as mute, but its voice is certainly the feeblest among all our members of the tailless tribe. The pairing season extends from January, in Algeria, to September and October, and the female is able to spawn three times in the year. The embrace, which never lasts long, is lumbar, the Fie. 52. Male and female pairing. male’s hands joining on the pubic region. The re- markably small eges are produced slowly, one at a time, and feebly adhere to the bottom of the pool or tank, where they form a single layer, more or less closely set. Only exceptionally have I found eggs attached to the weeds with which I had furnished the small tank in the terrarium, the pair keeping to the bottom during oviposition. The spermatozoa, which from their large size can be detected with the naked eye, are ejaculated in several bundles. The larve, as a rule, live only from one to two months before transforming, and keep small, being in this respect comparable to those of the common toad ; but tadpoles born late in the summer may remain for nine or ten months in that condition. Specimens which I received from Algeria in January and February, 1897, although in apparently excellent health and readily feeding on worms and insects, did DISCOGLOSSUS. 137 not begin to pair, so far as I could observe, until the beginning of June. The first eggs, about 400 in number, I found on the bottom of the tank on the morning of June 7th, the embryos emerging on the 10th; whilst on the 14th, or exactly a week later, the young had lost their external gills and entered the tadpole stage, measuring 11 mm. total length, buds of the hind limbs being discernible on the 17th. On the 23rd the largest specimens measured 21 mm., and on the 3rd July some had reached their full length, viz. 25 to 30 mm. On the 5th July some had put on the striped livery. The fore limbs first appeared on the 6th, and the metamorphosis was completed by the 8th, when the first specimens left the water with a reduced caudal appendage, and measuring 8 to 10 mm. from snout to vent, the last metamorphosing as late as August 6th. Striped speci- . mens were in the proportion of one to four spotted or immaculate. On the morning of June 14th the tank again con- tained eggs, 895 in number, which, from their con- dition, must have been deposited in the night of the 12th—13th (I was absent on Sunday, 13th), and the embryos of which were liberated in the afternoon of the 14th, thus less than forty-eight hours after the deposition of the eggs—a process of development more rapid than has been observed in any other species of Batrachians. The young from this brood began to leave the water on July 20th, whilst others retained the larval condition until October 5th. None were of the striped form. A third lot of 936 eggs were deposited during the night of July 6th—7th; they hatched in the night of 8th—9th, and the young completed their metamorphosis between August 16th and October, a few being still in the tadpole condi- tion whilst these lines are passing through the press. About two-thirds of them belonged to the striped form. In all these cases I was struck by the ex- tremely small proportion of eggs that did not develop, 138 DISCOGLOSSIDE. and the very slight amount of mortality that took place among the embryos; whilst of the tadpoles all except such as I killed for study went through the metamorphosis. In fact, the success in rearing these broods of Discoglossus has been greater than I have experienced in dealing with any other kind of tailless Batrachians under similar conditions. I had no diff- culty in feeding the tiny young on aphides. On the 17th July eggs had again been produced during the night, but they numbered only 306. They hatched on the 18th; most of them were destroyed shortly after by a fungus. » endofsnout 6 ...5 2.6 ..6 0.5 2.6 ..6 0.5 Fore limb : OS Gp a2” al: wel Oy GO yea igenk 9 Hind limh : 51 ...60 2.57 2.52 46 2.51 ...58 246 Tibia. 216 15 17 16 TA 15 7 Foot ~16 2.15 2.17 16 Tk 15 17 1, 5. Mondorf, Luxemburg: Boulenger. 2,6. Marcellise, Prov. Verona: De Betta. > 3. 8. San Bruno, Calabria: Giglioli. 4,8. Parnassos: Kriiper. 7. Arsoli, Rome: Vinciguerra. SxeLeron.—The osteological differences between this and the preceding species are very shglt indeed. The most constant is found in the length of the tibia, which cquals that of the femur. The sacral diapo- physes are usually not quite so strongly dilated, and the bones are nearly white instead of yellow as usual in the allied species. Ag has been shown by Goette, whose great work ‘ Entwickelungsgeschichte der Unke’ deals with this species, by Camerano (‘ Atti Acc. Torin.,’ xv, 1880, p. 445) and by Sasserno (op. cit., xxiv, 1889, p. 703), there often occur curious individual anomalies in the posterior vertebra, which may result in asymmetry of BOMBINATOR. 157 the sacrum or in the presence of two sacral diapo- physes, the second of which is produced by enlarge- ment of the process at the base of the urostyle, and connected distally with the one in front of it by a common cartilaginous epiphysis. Both Camerano (loc. cit.) and Howes (‘ Proc. Anat. Soc. Gr. Brit.,’ 1890, p. xvi) have recorded a curious anomaly similar to that observed by Lataste in an Alytes obstetricans : the sacral process on the right side is normal, whilst that on the left side is formed by the tenth vertebra, or base of the urostyle. The latter bone is sometimes provided with two processes on each side instead of one. I have examined a specimen from Amboise, preserved in M. Lataste’s collection, which is re- markable for having nine presacral vertebra, and perfectly normal sacral vertebra and urostyle. MEASUREMENTS OF SKELETON (in millimetres). 3 g Length of skull. ‘ ‘ 12 i Il Width of skull , ‘ . 14 hag 12 Least interorbital width . : 25 ae. 2 Dorsal vertebral column . % 15 4 14 Urostyle . ~ . . : IL ws 12 Humerus . : ‘ , 10 = g Radius-ulna ‘ s 65... 6 Manus i é F ; 8 a 7 Pelvis A ‘ , : 17 a 17 Femur ; : ‘ 7 14 ae 13 Tibia y : ‘ A 14 a 13 Tarsus Z 5 ‘ 9 i 8 Pes . . ‘ j A 14 =e 13 Hasits.—B. pachypus has much the same habits as its congener, but is less particular in the choice of the localities. Any sort of pools, warm or cold, clear or dirty, with or without vegetation, even the merest puddles, appear a Satisfactory resort to this lively little Batrachian, which may be heard in spring and summer uttering its low and mournful note, hoo, hoo. This sound is often produced by large numbers in a small pool exposed to the fullrays of thesun. Pairing takes place two or three times in the year, and spawn is mostly found from the end of May to the middle 158 DISCOGLOSSIDZ. of July, sometimes as late as September, as observed by Pfliiger near Bonn. Specimens from Belgium kept by me in confinement were still pairing on August 18th, and Mr. A. Pam informs me that some brought home by him this summer from Switzerland were ob- served in embrace as late as October 13th. Méhely found spawn as early as the 2nd of May in Transylvania. Spallanzani, in the last century, ob- served it pairing in May in the mountains near Modena, and he tells us that during his journey to Switzerland in 4779 the same species was pairing in July and August. I have myself observed it in embrace in the end of May near Salzburg and near Bordeaux, and spawning in Belgium in June and July. ‘Two years ago, around Freiburg in Baden, the breeding seemed to be over in the beginning of August. On the whole, as observed by Werner, this species appears to breed a little later than its con- gener; near Vienna, where the two occur, the season has been observed by that authority to begin in the first half of May for B. igneus, in the second half of the same month for B. pachypus. The eggs are usually attached to weeds as in the other species ; but as they are sometimes laid in puddles without any trace of vegetation, they then simply drop to the bottom. The lack of vegetation is no hindrance to the development of the larve, which are mainly car- nivorous. Recently transformed young were found by Schrank in the last century in the salt water of the saline caves of Berchtesgaden, Austria. These Batrachians hibernate on land in holes or under stones. In early spring, in Belgium, I have found several specimens together in their winter quarters—deep recesses between stones, which they shared with Salamandra maculosa. They do not appear to emerge before the end of April, when they travel considerable distances in search of suitable breeding localities, breaking the journey in any small temporary puddle they come across. I much doubt BOMBINATOR. 159 their ever spending the winter in the mud at the bottom of pools, as they are believed by Fatio occa- sionally to do. Eeas.— Do not differ materially from those of the pre- ceding species. Upper hemisphere pale brown, lower yellowish-white. The figure here given is a copy of Héron-Royer’s. Fia. 59. Taprote (PI. I, fig. 3).—One very striking character distinguishes this tadpole from the preceding, viz. the shape of the mouth, which is elliptical as in other genera of Discoglosside. The tail is, as a rule, rather shorter, and the muciferous crypts are hardly distin- guishable in the tadpoles obtained by me; however, it is very probable that the latter difference would not prove constant if tested on more extensive material. Body 17 mm.; width of body 14; tail 20; depth of tail 10. Hasitat.—Bombinator pachypus has a very exten- sive range in Hurope. It is found nearly all over France with the exception of the north coast of 160 DISCOGLOSSIDA!. Brittany, in the plains as well as in the hills, but very local in many parts, and does not ascend the Pyrenees ; South-eastern Holland; Belgium south of the Sambre and the Meuse, and in the calcareous localities near Tournay ; Germany all along the Rhine, but not higher up than 2000 feet in Baden; South Germany, Switzerland, in the valleys as well as on the mountains up to 3900 feet. Hast of the Weser in Germany this species enters into competition with B. igneus, and, as first pomted out by Wolterstorff, becomes restricted to the hills; this being the case wherever the two species co-exist, as in Austria- Hungary and Moldavia. Its highest recorded oc- currence in Transylvania is at 3900 feet, as in Switzerland, whilst in the Tyrol it reaches to 4850 feet, and in Bosnia to 5500 feet. In Italy the species is found in Lombardy and Venetia,* in the Apuan Alps up to 4500 feet, and along the chain of the Apennines from Emilia to Calabria, between 600 and 3600 feet, as I am informed by Prof. Gigloli. It is on record from the Etna, but not from other parts of Sicily. East of the Adriatic it occurs in Dalmatia, Bosnia, Herzegovina, Montenegro, Turkey (Adria- nople), and Greece (Parnassos, 3000 feet). Méhely has recently expressed the opinion that B. pachypus is to be regarded as a typical hill or mountain form, which only descends to the plain in the cooler northern regions, like Rana temporaria. This is certainly not the case. In the south-west of France the species is quite abundant at sea level; I have myself collected it on the banks of the Garonne, in the immediate vicinity of Bordeaux, where it is plentiful in small ditches bordering the vineyards ; and H. de Betta found it numerous in the plains of Venetia. * The species is absent from Piedmont. Some hundred specimens from Venetia were turned loose in the neighbourbood of Turin by Count Peracca about ten years ago, BOMBINATOR. 161 Male (ventral view) and female (dorsal view) specimens from near Dinant, Belgium, are figured on the right side of Pl. V. Ventral views of specimens from the Serra San Bruno, Calabria (fig. 3), Basle, Switzerland (fig. 4), and Schemnitz, Hungary (fig. 5), are given on Pl. VI. Hysrivs.—Méhely has pointed out that in the few places in Hungary where the two species occur together, intermediate specimens are found such as are not to be procured from other localities. These specimens are no doubt hybrids. Héron-Royer suc- ceeded in crossing the two species in both directions, and four nearly adult hybrids are described and figured by him in ‘Mém. Soc. Zool. France,’ 1891, p- 81. These hybrids, which proved fertile, and pro- duced a second generation, were nearly intermediate between the parent forms, but somewhat nearer B. igneus, whether the latter was the father or the mother: the ventral spots were lemon-yellow on a black ground; the digits were not tipped with bright yellow ; the inner finger and the inner toe were yellow in one specimen, in the others the inner finger was black, but the inner toe yellow. 162 DISCOGLOSSID&. 5. ALYTES. Wagler, Syst. Amph., p. 206 (1830). Pupil vertical. Vomerine teeth in transverse or slightly oblique series behind the choane. Tongue circular, entire, slightly free behind. Tympanum distinct. Fingers free, toes webbed; outer meta- tarsals separated by web. Diapophyses of sacral vertebra strongly dilated. Urostyle articulated to two condyles. Two species, both of which inhabit Europe. The presence of three palmar tubercles in A. obstetricans, aud of two in 4A. cisternastt, and the short swollen outer finger in the latter easily distinguishes them, in addition to several other characters, external and osteological. The characters in which the latter species differs from the former are adaptations to more burrowing habits, and of too slight importance, in my opinion, to justify the genus Auemoryctis, proposed by Lataste in 1879 for the then newly discovered Alytes cisternasti, which has since proved to be endowed with the same extraordinary nursing habits as its Jong known and famous congener. It may be added, as a further argument against generic distinction, that the tad- poles of the two species resemble each other so closely as to be almost indistinguishable before the limbs have made their appearance; this being the only instance that I know among Batrachians of well- marked species not being differentiated in their larval condition. The contrary would, however, in my opinion, be no valid objection to generic association, for reasons explained in the Introduction, p. 110. COPA D 1427. Riga? ciptiyps 2 ALYTES. 163 4. ALYTES OBSTBTRICANS. (Plate VIT.) Demours, Hist. Acad. Se., 1741, p. 28, and Mém. Ac. Sc., 1778, p. 13. Bufo obstetricans, Laurenti, Syn. Rept., pp. 28 and 128 (1768) ; Brongniart, Bull. Soc. Philom., ii, 1801, p. 91, pl. vi, fig. 4; Daudin, Hist. Rain. Gren. Crap., p. 87, pl. xxxii, fig. 1 (1803), and Hist. Rept., viii, p. 176 (1803); Zawadski, Faun. Gal.-Bukow. Wirbelth, p. 156 (1840). Rana campanisona, Laurenti, 1. c., p. 30. oe Wolf, in Sturm, Deutschl. Faun., iii, Heft 4 (1805). Bombinator obstetricans, Merrem, Tent. Syst. Amph., p. 179 (1820); Gravenhorst, Delic. Mus. Vratisl., p. 68 (1829). Bufo campanisonus, Goldfuss, Handb. a. Zool., p. 484 (1820). Alytes obstetricans, Wagler, Icon. Amph., pl. xxii, figs. 3—5 (1830) ; Schinz, Faun. Helv., p. 145 (1837); Tschudi, Isis, 1837, p. 702; Bonaparte, Icon. Faun. Ital., Rett. Anf. (1838); Duméril & Bibron, Erp. Gén., viii, p. 467 (1841); Giinther, Cat. Batr. Sal, p. 38 (1858); Bruch, Wiirzb. Nat. Zeitschr., iv, 1863, p. 92, and Ber. Offenb. Ver., v, 1864, p. 51, fig.; Fatio, Vert. Suisse, iii, p. 358 (1872); Koch, Ber. Senck. Ges., 1872, p. 155; Schreiber, Herp. Eur., p. 102 (1875); De V’Isle, Ann. Sc. Nat. (3), xx, 1876, No. 7; Lataste, Herp. Gir., p. 249 (1876); Leydig, An. Batr., p. 64 (1877) ; Lataste, Bull. Soc. Zool. France, 1877, p. 281; F. Miiller, Verh. Nat. Ges. Basel, vi, 1877, p. 420; Boulenger, Cat. Batr. Ecaud., p. 448 (1882); Héron-Royer, Bull. Soc. Zool. France, 1883, p. 417, pl. xiii, and 1886, p. 671; Nehring, Sitzb. Ges. Nat. Fr. Berlin, 1887, p. 48; Heller, Zool. Gart., xxix, 1888, p. 180; Héron-Royer & Van Bambeke, Arch. Biol., ix, 1889, p. 285; Bedriaga, Bull. Soc. Nat. Mose., 1889, p. 596; Fatio, Vert. Suisse, v, App., p. 7 (1890); Wolter- storff, Zool. Anz., 1891, p.65; Erwin Schulze, Schr. Nat. Ver. Harz, vi, 1891, p. 42; Boulenger, Proc. Zool. Soc., 1891, p. 622, pl. xlvii, figs. 6 & 7; Wolterstorff, Zool. Anz., 1893, p. 151; Oudemans, Tijdschr. Dierk. Ver. (2), iv, 1893, Versl., p. xlii; Martin & Rollimat, Vert. Dép. Indre, p. 356 (1894); Werner, Rept. Amph. Oesterr.-Ung., p. 111 (1897); Dirigen, Deutschl. Amph., p. 558, pl. i, fig. 4, and pl. il, fig. 6 (1897). Obstetricans vulgaris, Dugeés, Rech. Ostéol. Batr., p. 7 (1834). Alytes obstetricans delislet, Lataste, Rev. Int. Sc., iv, 1879, p. 543. Alytes obstetricans boscai, Lataste, l. c., and Act. Soc. Linn. Bord., xxxiv, 1880, p. 181, pl. xi; Bedriaga, Amph. Rept. Portug., p. 25 (1889), and Larves Batr. Portug., p. 12 (1891). Alytes bescai, Hévon-Royer & Van Bambeke, 1. c., p. 289. Vomerine teeth in a transverse series, interrupted in the middle, behind the choane; the series extends 164 DISCOGLOSSIDA. outwards to the vertical of the inner or of the outer border of the choane. Tongue large, thick, circular, shehtly free behind. Fig. 60. Head large, moderately depressed, broader than long; snout rounded, slightly projecting, as long as the diameter of the orbit; canthus ros- tralis rounded ; loreal region grooved, nostril a little nearer the eye than the tip of the snout; eye large, very prominent; interorbital space as broad as the upper eyelid, and as broad as or a little broader than the distance between the nostrils; tympanum circular, three-fifths to four-fifths the diameter of the eye. Fingers rather short, depressed, obtuse, first shorter than second, as long as- fourth or slightly shorter, third longest; no subarticular tubercles ; Fre. 61. three prominent palmar tubercles, corre- sponding in position to the base of the first, My third, and fourth fingers, inner oval, outer SA 9 Open mouth. round and largest, median round and smallest. Lind limb short; the tibio-tarsal articula- Lover tion reaches the shoulder in the female, the view of tympanum in the male; the tarso-metatarsal jefihand, articulation reaches the eye in the female, the tip of the snout or between the eye and the tip of the snout in the male; tibia as long as the femur, the heels meeting when the legs are folded at right angles to the rhachis. Foot as long as or a little shorter than the tibia; toes short, depressed, obtusely pointed, one- third to one-half webbed, and bordered; no subarticular tubercles ; a small, rounded, inner metatarsal tubercle. Upper parts covered with small smooth warts, limbs nearly smooth, with a glandular thickening on the forearm and another on the calf; a small, more or less distinct parotoid gland behind the eye above the tym- panum, usually followed by a series of smaller glands ALY'TES, 165 along each side of the body; a more or less distinct round gland behind the angle of the mouth. ‘Throat, belly, and lower surface of thighs, or belly and pubic region granulate; a strong gular fold separating the throat from the breast. . Greyish or pale brown above, speckled with darker or with small greyish-olive or greenish spots, some- times with red or reddish centres; an_ ill-defined dark canthal streak sometimes present; a light band across the anterior half of the interocular space, and a triangular, cordiform, or A-shaped light marking between the shoulders are often dis- tinguishable, and between these two light markings a dark triangle or X; the glands of the lateral series often orange or red. Lower parts dirty white or greyish, with the granules of a pure white; limbs carneous ; throat and breast often speckled with grey, especially on the sides. Iris pale golden, rarely silvery, veined with black. Male difficult to distinguish from the female. The body is, however, somewhat shorter, and the fore limbs are a little stronger. GuocraPHicaL Vanriations.—'lhis species varies very little in France, Germany, and Switzerland. But the specimens from the Spanish Peninsula show less uniformity ; the more divergent may be distinguished at a glance, and have been named var. bosce, Lataste. The skin is usually smoother, the parotoid glands are smaller, and the dorsal spots are generally larger and better defined, sometimes forming mar- blings ; these characters combine to produce a physi- ognomy somewhat suggestive of a young Pelobates cultvipes. Iam, however, unable to confirm the abso- lute constancy of the various structural differences by which this variety is to be distinguished from the typical form; measurements of the skull and vertebral column of some Rhenish and Spanish specimens show me that the proportions may, exceptionally, be the same. 166 DISCOGLOSSID A. Measurements (in millimetres). g ; aS 1 2 3. 4, 5. 6. From snout to vent J 3b8 0... 460... 41 . 43°... 51 . 43 Length of head 5 Tb ase AG: nee WD oe AS ae TW: es DS Width of head 160... 18 2... 17 6 AP eee TO ns TT Diameter of eye 45... 6 45.0. 45... 5 45 Interorbital width 3 4 oo... Sb... 4°... 4 From eye to nostril 4... 45... 4 0, 4 45... 4 i . end of snout . 65. 8 7 7 8 7 Diameter of tympanum OD ge 3's 3 ap 3) age A 3 Fore limb ef ug BUN a Bb On D5 23 Hind limb . i BO an DY ce Hk Be 2 DS wea, “OY Tibia . ; 21S. VW. 16, 16. W168 Foot ‘i ‘ eh cee LO cc AD cee PD: cc WO: aan DA 1. Paris: Lataste. 4,6. Valencia, Spain: Bosca 2. French Jura: Ling Roth. (var. bosee). 3. Hameln, Weser; Wolterstorff. 5. Paris: Héron-Royer. According to Fatio the female reaches a length of 54 mm. Sxeteron.—Ethmoid short, embracing above a large fontanelle which extends nearly to the line of the anterior borders of the orbits; fronto-parietals very narrow, in contact with each other only quite poste- riorly, with an inward process in the middle which gives the fontanelle the shape of the sole of a shoe; ethmoid rounded or somewhat produced anteriorly, exposed between the posteriorly diverging inner borders of the nasals, which are large and in contact with each other in front or along the greater part of their length; a cartilaginous supra-orbital plate is articulated to the nasal and the anterior third of the fronto-parietal. Zygomatic branch of the squamosal very short. Vomers large, narrowly separated from each other on the median line, their posterior toothed borders partly covering the feeble palatines ; pterygoid regularly triradiate, the anterior branch joining the maxillary and the slender palatine ; parasphenoid i-shaped, obtuse or truncated in front, not quite reaching the vomers, and separated from the ptery- goids. Mento-Meckelians very indistinct. Hyoid a large, broad, cartilaginous plate with rounded lateral wings, small postero-lateral processes, and long cornua; a pair of diverging, ossified stylo- ALYTEs, 167 hyals, apart at the base, in front of which a slender V- or Y-shaped ossicle is developed on the ventral side. Vertebral column twice or a little more or a little less than twice as long as the skull. Spine closed Skeleton of male. above, the neural arch without or with a very short postero-median process. Second, third, and fourth vertebre with short diapophyses, with short auto- genous ribs. The first rib the shortest and directed 168 DISCOGLOSSID&. a little forwards ; the second the longest, nearly hori- zontal; the third intermediate in length between the other two, and horizontal or directed slightly back- wards; the fourth or fourth and fifth diapophyses nearly horizontal, the sixth and seventh, sometimes also the fifth, directed forwards. Diapophyses of the sacral vertebra triangular, their distal diameter equal- ling or slightly exceeding their length ; two condyles for articulation with the urostyle. Latter nearly as long as or a little shorter than the eight anterior vertebrae taken together, with a rather stro one, curved or posteriorly hooked transverse process at the base. Preecoracoids slender, curved, meeting at an acute angle, entering the glenoid cavity ; coracoids stronger, nearly straight; supra-scapula ossified ; omosternum absent; cartilaginous sternum with two feebly calci- fied posteriorly diverging styles. Carpus with eight elements, two of which are in contact with radius- ulna; a single bone in the pollex. Pelvis three-fifths to two-thirds the length of the vertebral column; pubis cartilaginous; acetabulum open. Femur and tibia equal in length, the former strongly curved; astragalus nearly two-thirds the length of the tibia ; three small cartilaginous tarsalia ; one cartilaginous element to the prehallux. Distal phalanges obtuse, slightly expanded at the apex. MEASUREMENTS OF SKELETON (in millimetres). F. typica. Var. bose. a, 3 g 3 2 Length of skull. 13... 14 ui 12)... 183 Width of skull 1... 17 ae 14... 16 Least interorbital width 4... 4 she 35... 35 Dorsal vertebral column iB... 15 oe Ve son 22 Urostyle LO ein. 1S ae TI gee 13 Humerus. : 10. ... 10 a 9 2... 10 Radius-ulna Lo sien |G Sie CO on 9 Manus : 2... #1 Be 10)... «1 Pelvis WW .., 18 on 16... «18 Femur ». 1 1. 45 ws 13) a. 15 Tibia ; . 15... «615 sis 13... 15 Tarsus : 10... 10 ei 8... 10 Pes ” ; I ce WO ce, Tee ae ALYTES, 169 Hastrs.—The life-history of this Batrachian is one of great interest, but difficult to observe owing to its nocturnal habits and shy disposition. First dis- covered in the act of parturition by Demours, in the middle of the last century, on the border of a small pond in the Jardin des Plantes, we have had to wait until the year 1876 for a truly scientific account, based on exact observations, of the mode of oviposition of the so-called ‘‘ crapaud accoucheur de sa femelle.” This we owe to that most excellent and patient naturalist, Arthur de l’Isle du Dréneuf, who, residing near Nantes, in a locality where the species is excep- tionally abundant, spent in three consecutive years more than fifty nights in the open in order to unravel the mystery, and whose notes taken on the spot cover 250 foolscap pages of small handwriting. ‘The account given hereafter of the pairing and oviposition is com- piled from that source. Although I have stayed at places where Alytes is not uncommon, and have made nocturnal excursions with a lantern in order to witness the operation, I have, in common with most herpetologists, hitherto failed in my object. Alytes is nocturnal and slow in its movements ; it progresses mostly crawling, but sometimes by short leaps, even when embarrassed with the eggs. It is able to burrow chiefly by means of the fore limbs, but usually selects for its retreat holes made by small mammals, or interstices between stones. Turning over large stones in the vicinity of the water where its tadpole attracts notice, is the surest means of securing specimens in the daytime. Towards evening it reveals its presence by aclear whistling note, which has often been compared to the sound of a little bell, or to achime when produced by numerous individuals. The breeding season lasts throughout spring and summer, and the female is able to spawn two, three, or even four times in the year. The season seems, however, at its height in May and June in the plain; later, of course, in the mountains. 170 DISCOGLOSSIDA. Pairing and oviposition take place on land. The male seizes the female round the waist; but during impregnation, or when he proceeds to lade himself with the ova, he clasps her round the head in front of the fore limbs. Males are more numerous, and often dispute about the females. When he has secured un- disturbed possession the male strongly bends the body so as to bring his heels close to the cloacal region of the female, which he proceeds to lubrify by rapid move- ments of the inner toes, at times introduced into the cloaca, the two feet working alternately. This sort of raking consists of 1100 to 1300 strokes for the two feet, divided into 15 to 21 periods of 45 to 103 strokes, with intervals of two or three minutes’ rest, and lasts about twenty-five minutes. Then the movements sud- denly stop, the female extends her hind limbs, tightly embracing those of the male, which are bent at angles at the knees, the tarsi erect and pressed close together ; the eggs are suddenly expelled, with noise, as if by ex- plosion, and fall into the rhomboidal receptacle formed between the pelvic limbs and bounded behind and beneath by the tarsi and feet of the male. The yellow egos, as if threaded together by elastic filaments, form a large mass, two to four layers of about ten, in this receptacle. The very moment the eggs are emitted the male unclasps the waist of the female and shifts his hold to the base of the head; the body then stretched out, but the legs remaining in the same position as before, fecundation commences; it takes place in two or three emissions at short intervals. he seminal fluid is much diluted with the contents of the urinary bladder, which soaks the envelops of the eggs. An interval of ten to fifteen minutes’ rest follows before the male proceeds to attach the eggs to his legs. This is done im the followmg manner :—Still holding his mate round the head, he draws out the legs go as to stretch the elastic connections between the eges sticking to his tarsi; then, folding one of the limbs, bringing the ALYTES. 171 knee to the level of the sacrum, and stretching it out again, he passes it, toes first, into the egg-mass; the other limb follows in like manner. These movements are repeated several times, and accompanied by one or two seminal ejaculations. Thus the strings of eggs come to be fastened round the legs, where they will remain until eclosion. All the time this operation has lasted, about ten minutes, the female has remained motionless, still connected with the eges by the last elastic filaments from the oviducts. ‘hese two threads stretch to a length of 4 to 12 inches before breaking when the pair separates. Thus laden, and yet so little impeded in _ his movements as to occasionally resort again to hymen during the nursing period, and successfully add on a second burden, the male retires to his usual retreat, going about at night in order to feed him- self and to keep up the moisture of the eggs, even resorting to a short immersion in the water during exceptionally dry nights. The development within the ege takes about three weeks, sometimes a little less, often a little more. ‘he male enters the water with his burden; the larve, in the full tadpole con- dition, measuring 14 to 17 mm., bite their way through the tough envelop, which is not abandoned by the father until all the young are liberated. ‘The tadpole is found in spring and summer, at all stages of development, in small reservoirs, cow-ponds, flooded quarries, pits in brick-fields, &c. The tadpoles of the late broods hibernate under the ice, concealed in recesses but not torpid. Some specimens, at least in confinement, remain nearly two years before trans- forming, but, as a rule, larvee born early in the spring accomplish their metamorphosis within three to five months. The duration of the larval life varies, how- ever, in individuals of the same brood; and as such as have not sufficiently developed their limbs to take to the land in the autumn have necessarily to post- pone the metamorphosis until the following spring, it 172 DISUOGLOSSIDA. follows that whereas some may have spent only three or four months in the water, their brothers will sojourn for a whole year or more in that element. Young reared by me measured, immediately after meta- morphosis, from 18 to 22 mm. from snout to vent. Eecs.—Large, the vitellus measuring 35 to 5mm. in diameter, varying in colour from pale straw-yellow to bright yellow. They are coated with Fre. 63. two transparent gelatinous envelops, ge the outer of which is tough and elastic, Q and forms the threads by which the eggs ‘are connected in two rosary-like series as they issue from the cloaca. These a connecting threads measure, without @ ra great tension, from 4 to 7 mm., and Tia according to the number of eggs, which varies between 18 and 54, the whole rosary has alength of 70 cm. to 2m. Males observed by Geisenheyner and Melsheimer with 126 and 150 egos were no doubt nursing double or treble broods. When fresh laid the eggs are nearly spherical, but they soon acquire a more transversely oval shape. Through the transparent capsules the whole development can be easily followed. An enormously large vitelline sac is present, and the em- bryo develops uncommonly long, unpigmented gills, one only on each side, with a large number of slender branches along the ventral side of the trunk. These gills are absorbed and replaced by internal ones, and the transformations which accompany the passage from the first or embryonic to the second or tadpole period are all effected within the egg-capsules. The embryos are at first uniform yellowish-white ; the pigment, when it appears, forms two brown stripes, but before hatching the little tadpole has put on his grey, more or less spotted coloration. Taprotz (PI. I, figs. 4 and 5).—Length of body once and one-third to once and a half its width, two-fifths to one-half the length of the tail. Nostrils nearly @OoeE@ ALYTES. 173 halfway between the end of the snout and the eyes. Hyes on the upper surface of the body, the distance between them about twice as great as that between the nostrils, and equal to or slightly greater than the width of the mouth. Spiraculum in the mid-ventral line, a little nearer the anterior than the posterior extremity of the body. Anal opening median, very much larger than the spiraculum. ‘Tail twice and two-thirds to thrice as long as deep, ending in an obtuse point ; the upper crest convex, usually a little deeper than the lower, and extending but. very slightly upon the back; the depth of the muscular portion, at its base, about half the total depth. Beak white, with a broad black margin. Lip entirely surrounded by a series of papille. Labial teeth in 3 series, occupying nearly the whole width of the inner surface of the lip, all continuous, or the third lower narrowly broken up in the middle; the first upper and the first lower series composed of one or two rows of teeth, the others of two or three. Lines of crypts usually very indistinct; all that can be distinguished being the usual lines from the end of the snout between the nostrils, bordering the eyes above, behind and below, and forming a loop on each side of the upper lip, a line beginning at a considerable distance behind the eye along each side of the back to the upper border of the muscular part of the tail, and another very short line close to and parallel with the anterior extremity of the latter. But im a fine specimen from Bellaigues, Switzerland, I find the lines much more distinct and blackish ; in addition to the series described above, it shows the second dorsal line prolonged to the base of the tail, which also bears two lines, the upper being on the supra-caudal crest ; a short series descends vertically from below the centre of the eye, another, curved, from below the anterior extremity of the dorsal lines, a third on each side of the mouth, and a fourth extends on each side of the belly, from the level of the spiraculum nearly 174 DISCOGLOSSID.®. to the origin of the hind limbs, its anterior extremity bent downwards and forwards. This specimen is figured above (p. 104, Fig. 46, ¢). Lead-grey to blackish above, uniform or with round blackish spots; sides with large silvery or pale golden spots; tail with numerous dark brown dots or round black spots, which are very apparent on the greyish- white crests. Nearly black tadpoles as well as albinos are on record. Total length 80 mm.; body, 28; width of body, 21; tail, 52; depth of tail, 19. Exceptionally grows to 90 mm. Hasirar.—The midwife toad is common in France nearly everywhere ; it is also found in Belgium (pro- vinces Namur, Liége, and Luxemburg), South-eastern Holland (Limburg), Luxemburg, Switzerland, Vorarl- berg, in Germany along the Rhine, and locally in hilly districts as far east as Brunswick and Thuringia (Hamicln on the Weser being the northernmost, Nord- hausen and Hisenach the easternmost points of its ascertained distribution), and all over Spain and Portugal. It ascends to 5000 feet in the Alps of Switzerland, and to 6500 feet in the Pyrenees (Lac Bleu, Hautes-Pyrénées), where the snow is not absent for more than three months. Specimens have been introduced by Knauthe near Schlaupitz, in Silesia, some years ago, and appear to have established themselves there. ‘The species 1s stated to occur in Bukowina, whence it was described by Zawadski in 1840, It would be highly interesting to ascertain whether it does really extend so far to the east; for the present the statement can only be accepted with caution. In France the midwife toad is found in the plains as well as in the mountains ; common at Biarritz, close to the sea, it extends high up the Pyrenees. I have also found it on the coast of Normandy, and, as Prof. Bavay informs me, it occurs on the coast of Finistére. East of the Rhine, however, its habitat, like that of ALYTES. 175 Bombinator pachypus, becomes restricted to the hills. It by no means avoids the dwellings of man, and may be found in great numbers in old walls of villages as well as in gardens and waste grounds of towns. It is still to be met with in the Jardin des Plantes, where it was first discovered in the middle of the last century. Four specimens are figured on PI. VII, viz. a male from Paris, laden with eggs; a female from Liége, on the left hand; a young albino from Génevilliers near Paris, reared by M. Lataste ; and on the right hand a specimen from Valencia, Spain (var. bosce). 5. ALYTES CISTERNASII. (Plate VIII.) Alytes cisternast, Bosca, An. Soc. Esp., viii, 1879, p. 217; Bou- lenger, Cat. Batr. Ecaud., p. 449 (1882); Héron-Royer & Van Bambeke, Arch. Biol., ix, 1889, p. 289; Bedriaga, Bull. Soc. Nat. Mosc., 1889, p. 617, Amph. Rept. Portug., p. 27 (1889), and Larves Batr. Portug., p. 141 (1891); Boulenger, Proc. Zool. Soc., 1891, p. 624, pl. xlvii, fig. 8. Ammoryctis cisternasi, Lataste, C. R. Ac. Se., Ixxxviii, 1879, p. 983; Bosca, Bull, Soc. Zool. France, 1880, p. 252, and An. Soc. Esp., x, 1881, pl. ii, figs. 1—6. Vomerine teeth in two transverse series, widely sepa- rated from each other in the middle, behind tlhe choane ; these series often slightly oblique, forming together an angle pointing forwards. Tongue large, circular, slightly free behind. Head large, moderately depressed, much broader than long ; snout round- ed, projecting, shorter than the dia- meter of the orbit; canthus rostralis rounded, loreal region grooved ; nos- tril nearer the tip of the snout than the eye; eye large, very prominent; interorbital space broader than the upper eyelid, as Fria. 64. Open mouth. 176 DISCOGLOSSIDA. broad as the distance between the nostrils; tympanum three-fifths to two-thirds the diameter of the eye. Fingers rather short, depressed, obtuse, first shorter than second, third a little longer than second, fourth shortest; fourth finger very thick, stump- like, brown and horny at the end; no sub- articular tubercles; two prominent palmar tubercles, inner small and oval, outer very large, oval, oblique. Brachium nearly en- tirely embedded in the skin of the body. Hind limb short ; the tibio-tarsal articula- tion reaches the axil or the shoulder; the ee a view tarso-metatarsal articulation reaches the tym- py panum or the eye; tibiaas long as the femur, the heels meeting when the limbs are folded. Foot as long as or a little shorter than the tibia; toes short, depressed, obtusely pointed, one-third webbed ; no subarticular tubercles; a small, oval inner meta- tarsal tubercle. Upper parts covered with very small smooth warts ; snout smooth; a series of enlarged warts along the upper eyelid; a very small parotoid gland. Lower parts coarsely granulate; a strong gular fold. Greyish or brownish above, with small dark spots ; a light cross-bar between the eyes, and three light spots ona line between the shoulders; the enlarged warts on the eyelids whitish or orange ; some of the warts on the sides also whitish or orange. Some specimens have the markings better defined, and a light triangular or cordiform spot between the shoulders is accompanied by an oblique, curved light band on each side of the back, beginning from the supra-temporal fold. These markings, which may also be observed, though less defined, in the preceding species, are highly suggestive of those of certain specimens of Discoglossus pictus, and afford further evidence of the close relationship, so long overlooked, which exists between the two genera. Lower parts whitish. Iris golden. Fig. 65. ALYTES. 177 Measurements (in millimetres). 1. 2. 3. 3 g 2 From snout to vent . . 38 w. 85 ase 40 Length of head : z 12 vi JL ee LD Width of head é 16 .. Lt ten Oe Diameter of eye ‘ 4 4 45 Interorbital width : 4 4 45 From eye to nostril. : 35 3 a5 es <3 end of snout ‘ 6 we = «6S 6 Diameter of tympanum : 25... 25 3 Fore limb i : 20 19 19 Hind limb ‘ ‘ ‘i 45 w. 42 w. 46 Tibia . , : 18 sue oko vw 14 Foot . ; 13 .. 1235 125 1, 2, Merida, Spain: Bosca. 8. Badajos, Spain: Bosca. SKeLteron.—Apart from the stouter and shorter limb-bones, the skeleton of this species differs from that of the preceding in the larger nasals, which over- lap the ethmoid and the inner borders of which meet along their entire length, in the constricted fronto- parietals, and in the shape of the fronto-parietal fontanelle, which is divided into two portions between which the fronto-parietals form a bridge; the anterior fontanelle is large and kite-shaped, the posterior very small, a mere foramen between the fronto-parietals. Fria. 66. Upper view of skull and lower view of left forearm and hand. The coracoids and precoracoids are more flattened and more robust, and the metacarpal and phalanges of the outer finger are shorter and more massive ; the terminal one in particular being extremely short, nearly hemispherical. The femur is slightly longer than the tibia. The more strongly ossified cranium is correlated’ M 178 DISCOGLOSSIDA. with the burrowing habits of this species, and the differences noticed between this species and 4. obstetricans are merely of degree. MEASUREMENTS OF SKELETON (in millimetres). 2 Length of skull . é : : : 12 Width of skull . : 16 Least interorbital width . $ 3 Dorsal vertebral column . 2 1g Urostyle . ; : 3 10 Humerus . ‘ 9 Radius-ulna 2 65 Manus ‘ 5 é 85 Pelvis aj : ; 16 Femur . : 12 Tibia 3 11 Tarsus ‘ i : 75 Pes 3 3 : : ‘ 12 Hasrts.—This species is a burrower in sandy locali- ties, digging by means of the outer edge of the hands. In accordance with these habits its lungs are more developed than in the allied species. Boscd having found the male carrying the eggs, in April, there is no doubt the breeding habits are much the same as in A. obstetricans, although the pairing has not yet been actually observed. The tadpole is to be found all the year round. Tappotez (Pl. I, fig. 6).—I am not able to detect any character by which this tadpole may be surely distin- guished from that of A. obstetricans. The spots on the tail are, however, smaller, more crowded, and forming vermiculations on the muscular portion, the space oc- cupied by the lateral groove being usually free from spots. Penal of body, 20 mm.; width of body, 15 mm.; tail, 42 mm.; depth of tail, 14. Hapitat.—Spain (Aragon, New Castille, Estrema- dura) and Portugal (Douro, Beira, and Alemtejo). The specimen figured on Pl. VIII is a female, one of the types from Spain, preserved in M. Lataste’s col- lection. [At page 179. 70 20, IN — PELODYTES PUNCTATUS. [| Pevopates Fruscus. NNWG CULTRIPES. 40 50 DISTRIBUTION OF EUROPEAN PELOBATID. PELODYTES. 179 Family 2.—PELOBATID A. Vertebree proccelous (in.the European genera), with- out autogenous ribs; diapophyses of sacral vertebra very strongly dilated. Upper jaw toothed. The Pelobatide occupy an intermediate position between the Discoglosside and the Bufonide. Seven genera are known, inhabiting Europe and South- western Asia, South-eastern Asia and New Guinea, and North America and Mexico.* The two European genera are easily distinguished :— Pelodytes, slender and frog-like, with deeply cleft toes, and a more or less distinct tympanum ; Pelobates, stout and toad-lke, with broadly webbed toes, no tympanum, the skin of the head adherent to the skuli, and a large, sharp-edged, shovel-shaped tarsal tubercle. Both genera must be looked upon as highly special- ised forms of a group which has, in all probability, sprung up from the Dvscoglossidxe, with which some of the exotic genera are nearer connected through a character to which great importance has been attached, viz. the opisthoccelous vertebree. A map is here given showing the range of the three Huropean species of this family. 4. PrELopytss. Fitzinger, in Bonaparte, Icon. Faun. Ital., Rett. Anf. (1838). Pupil vertical. Vomerine teeth in two small groups between the choanz. Tongue subcircular, entire or slightly nicked, and free behind. Tympanum present, * Exotic genera: Scaphiopus, Holby. EH. and 8. United States, Mexico. Batrachopsis, Blgy. New Guinea. Leptobrachium, Tsch. §S. China, Further India and Malay Archipelago. Megalophrys, Kuhl. Malay Peninsula and Archipelago, Asterophys, Tsch. New Guinea. The last two have opisthocelous vertebra. 180 PELOBATIDA. more or less distinct or hidden under the skin. Fingers free, toes webbed at the base and more or less distinctly bordered ; outer metatarsals separated by web. Proximal tarsal bones fused. Vertebree pro- ceelous ; diapophyses of sacral vertebra very strongly dilated; two condyles for articulation with urostyle. Omosternum cartilaginous; sternum with a bony style. This genus includes two species:—P. punctatus, Fitz., of Western Continental Europe, and P. caucasi- cus, Bler., recently discovered on the Asiatic side of the Caucasus at an altitude of 7000 feet. The latter has been made the type of a distinct genus, Pelody- topsis, by Nikolski, on grounds too trivial to deserve recognition. Perhaps this Pelodiytes caucasicus may yet be found in Cis-Caucasia, and yield a further addition to the European fauna. It is distinguished from its con- gener in the longer hind limbs, the tibio-tarsal articu- lation reaching the tip of the snout, and the somewhat more posterior position of the groups of vomerine teeth; the toes are not so strongly fringed in the male, the fore limbs are more robust, and the black horny nuptial excrescences are much more strongly developed on the warts and ridges of the upper sur- faces. 6. PBLODYLES PUNOTATUS. (Plate VIIT.) Rana punctata, Daudin, Hist. Rain. Gren. Crap., p. 51, pl. xvi, fig. 1 (1803), and Hist. Rept., viii. p. 100 (183). Ranu plicata, Daudin, ll. ec., pp. 53, 102. Rana dandinii, Merrem, Tent. Syst. Amph., p. 177 (1820). Bombinator plicatus, Fitzinger, N. Class. Rept., p. 65 (1826). Obstetricans punctatus, Dugés, Rech. Ostéol. Batr.. p. 7 (1834). Alytes punctatus, Tschudi, Class. Batr., p. 84 (1838). Pelodytes punctatus (Fitzinger), Bonaparte, Faun. Ital.. Rett. Anf, (1838); Duméril & Bibron, Erp. Gén., vili, p. 463 (1841) ; Thomas, Ann. Se. Nat. (4), i, 1854, p. 290; Ginther, Cat. Batr. Sal., p. 35 (1858); Fatio, Vert. Suisse, iii, p. 353 (1872); De Betta, Faun. Ital., Rett. Anf.. p. 68 (1874); & -eiber, Herp. Eur., p. 99 (1875); Lataste, Herp. Gironé a8 Po vm. oe c Vile y liv fecen elit tits, & PELODYTES. 181 (1878) ; Héron-Royer, Bull. Soe. Zool. France, 1878, pp. 128 & 299, pl. iii, and 1879, p. 229, pls. x and xi; Boulenger, Bull. Soc. Zool, France, 1880, p. 225, and Cat. Batr. Ecaud., p. 438 (1882); Peracca, Boll. Mus. Torin., i, 1886, No. 1; Héron-Royer, Bull. Soc. Et. Se. Angers, xv, 1886, p. 91; Héron-Royer & Van Bambeke, Arch. Biol., ix, 1889, p. 277, pl. xx, figs. 5—12; Bedriaga, Bull. Soc. Nat. Mosc., 1889, p. 533, and Amph. Rept. Portug., p. 20 (1890); Boulenger, Proc. Zool. Soc., 1891, p. 617, pl. xlvii, figs. 1 & 2; Bedriaga, Amph. Rept. Portug., Suppl, p. 11 (1893); Martin & Rollinat, Vert. Dép. Indre, p. 349 (1894). Pelodytes daudini, Bosca, Bull. Soc. Zool. France, 1880, p. 255. Vomerine teeth in two short transverse or slightly oblique series on a line with the anterior borders of the choanz; these series either close together or nearer the choane than each other. ‘Tongue large, circular, entire or feebly nicked behind. Head much depressed, as long as broad or slightly broader than long ; snout rounded, projecting beyond the mouth, as long as the diameter of the orbit; canthus rostralis rounded; loreal region grooved; nostril midway between — Open mouth. the eye and the tip of the snout or a little nearer the latter; eye large, prominent; interorbital space as broad as or a little narrower than the upper eyelid, and equal to the distance between the nostrils ; tympanum usually moderately distinct, sometimes hidden, one-half to three-fifths the diameter of the eye. Fingers rather elongate, somewhat swollen at the end, third much the longest, first a little shorter than second, which is a little shorter than fourth; basal subarticular tubercles distinct; three palmar tubercles, median smallest and circular. Hind limb rather slender; the tibio-tarsal articu- lation reaches the eye or between the eye and the nostril; tibia a little longer than femur, the heels overlapping when the legs are folded at right angles to the rhachis. Foot as jong as or a little longer than the tibia; toes slender, webbed at the base, the web extending as a fringe along each side to the Fia. 67. 182 PELOBATIDA. tips, which are slightly swollen; subarticular tubercles feeble or indistinct ; no tarsal fold; a small, rounded inner metatarsal tubercle. Upper surfaces covered with more or less prominent porous warts of unequal size; the larger warts on the back sometimes disposed in wavy longitudinal series ; a strong glandular fold trom the eye to above the shoulder, passing above the tympanum, and sometimes followed along each side of the body by a chain of large warts. “Lower surfaces smooth except the pos- terior part of the belly and the thighs, which are coarsely granulate. Greyish or pale greyish-olive above, with small irre- gular dark olive, bottle-green, or bright green spots (whence the name “ Persillé”’ by which this Batrachian is known in some parts of France), which may be confluent into more or less regular cross-bars on the limbs; between the eyes two oblique streaks of the dark colour, converging backwards, are usually dis- tinct; a dark streak extends from the tip of the snout to the eye, involving the nostril, and often reappears behind the eye along the supra-temporal fold; two or three dark vertical bars, sometimes broken up into spots, on the upper lip; the larger warts on the sides sometimes orange or rusty red ; a light X-shaped marking on the ground colour is usually more or less distinct on the anterior part of the body, the anterior branches extending to the upper eyelids, the posterior to the sacral region. Lower parts white, often yellowish on the limbs and rosy about the inguinal region. Iris golden, more or less pigmented with brown on the lower half, or bronzy-brown with a fine pale golden border to the pupil. Male distinguished from the female by a shorter body, longer and more robust fore limbs, a greater development of the dermal borders of the toes, and the presence of an internal vocal sac which communi- cates with the mouth through a long slit on each side of the tongue; owing to the pigmentation of the vocal PELODYTES. 188 sac the throat appears pale purplish-grey. During the breeding season horny excrescences are much de- veloped on the lower sur- Fra. 68 faces, and similar pro- ductions of small size are sometimes also scattered on the upper surfaces, the largest crowning the prin- cipal warts; no trace of them being found nor- mally in the females. These excrescences, which are of a dark purplish-brown or black colour, are disposed as follows on the lower sur- faces :—A rounded group on each side of the breast, at the base of the arm; another on the arm; a band along the inner side of \ the fore-arm ; and a band ee ee ee WW along the inner side of the wee two inner fingers; there are besides, very often, a band on the chin, isolated points on the abdominal granules, and two linear series along the toes, interrupted at the articulations. \ Moasvrements (in millimetres). I. 2. 3. 4. 5. 6. From snout to vent . .d8 ... 41 2. 40 2. 40... 43. 44 Length of head ; .12 2.18 2. 14 2. 18... 15... 14 Width of head : .12 2.15 2. 14 2... 18... 16 2... 14 Diameter of eye : » Bb... 4 0. 4 2. 4 4... 4 Interorbital width . 2 Db, Bh ee BD Shane Blae 28 From eye to nostril . » Bday Bbc Bas Bay Bban BO 3 oy end ofsnout . é os Z ade Bae , nig oe 2 Tympanum . : : les saa ND aes wen BOs, 2 ore lane a 4 324 ain 29) ae BO™ ges DB cee BY es 120 Hindlimb . : .59 2... 68... 68... 60... 71. 61 Tibia : ‘ AS nee 20 TO ae AB ce ZL. ds Foot : : + 20 exe PO igs BO ges AB guy 28 2 LP 1, 4. Paris: Héron-Royer. 8, 6. Valencia: Lilford 2,5. Nice: Bedriaga. 184: PELOBATIDA. SKELETON.—The skull, so far as the shape and the extent of ossification are concerned, is intermediate between that of Discoglossus and that of Bombinator, agreeing with the latter in the absence of palatine bones. Ethmoid short, not extending posteriorly beyond the anterior two fifths of the basisphenoid ; the lower anterior lamina rounded or truncate, and extend- ing nearly to the posterior border of the vomers; the upper obtusely pointed and extending to between the nasals. The large chondrocranial fontanelle is only partially covered by the fronto-parietals, these bones being in contact with each other to but a small extent either behind or in the iniddle. Nasals mode- rately large, tear-shaped, not meeting on the median ne. Zygomatic branch of the squamosal short. Vomers moderately large, narrowly separated from each other on the median line; pterygoids rather slender, the anterior branch longest and joining the maxillary; parasphenoid dagger-shaped, sharply pointed in front, and extending forwards to the line of the anterior borders of the orbits, separated from the pterygoids. Mento-Meckelian bones distinguish- able only from the inner side of the mandible. Hyoid a large cartilaginous plate with slender postero-lateral processes ; anterior processes much expanded, turned inwards and closely approximating in front, and confluent with the lateral wings, enclos- ing a small fenestra; thyrohyals slender, diverging, narrowly separated from each other at the base; a pair of slender ossifications in front of the thyroidals on the ventral side, distinct and L-shaped or united and H-shaped. Ceratohyal cornua with the posterior portion detached from the hyoid plate, the anterior portion being included in the hyoid plate. Vertebral column twice or a little less than twice as Jong as the skull. Spine closed above; neural arch without or with a very short postero-median pro- cess. First diapophysis obliquely directed forwards ; second, third, and fourth horizontal; second longest, PELODYTES. 185 fourth small; fifth, sixth, and seventh small, slender, and directed forwards. Skeleton of male. Sacral vertebra with very strongly dilated diapo- physis, the transverse diameter of which is nearly 186 PELOBATIDS. three times in the longitudinal, but little shorter than the urostyle, which articulates by two sockets, and measures three-fifths to two-thirds the length of the remainder of the vertebral column. Preecoracoids slender, strongly curved; coracoids nearly straight, dilated at both ends; a cartilaginous omosternum ; sternum with a bony style dilated proxi- mally, and a large discoid cartilage. Humerus con- siderably longer than radius-ulna. Carpus with seven elements, three of which are in contact with radius- ulna; a single bone in the pollex. Pelvis three-quarters the length of the vertebral column; ilia with an inner groove, embracing the sacral processes; the pubis does not ossify. Femur and tibia with the epiphysis cartilaginous; tibia a little longer than the femur, nearly as long as the pelvis; calceaneum and astragalus completely fused into a single slender bone with a minute foramen in the middle, resembling the tibia, of which it measures three-fifths the length; the elements of the distal row of the tarsus are not ossified, and the prehallux contains two cartilages. Distal phalanx of all digits slightly expanded at the apex. MeasvrEMeENTS OF SKELETON (in millimetres). 3 2 Length of skull , 11 ste 14. Width of skull : 11 bx 15 Least interorbital width 2 ie 3 Dorsal vertebral column . 12 17 Urostyle : 5 7 23 ll Humerus . : B10 aun 11 Radius-ulna A 6 ss 8 Manus ‘ 9 sis 12 Pelvis 15 ay 20 Femur : 14 an 17 Tibia ‘ 16 a 18 Tarsus . s 9 a 11 Pes ; ‘ ’ 4 17 sibs 20 Hasits.—This small and graceful Batrachian is essen- tially nocturnal; only during the breeding season is it met with in the daytime. Its long limbs enable it PELODY'ES. 187 to jump in the manner of the true frogs, covering 14 or 15 inches at one leap, and it is besides a good climber, ascending smooth vertical surfaces by adhesion of the abdominal surface. Nor, in spite of its nearly free toes, is it a bad swimmer, often frequenting deep ponds during the pairing time. Its dermal secretion bears a strong smell of garlic As often heard in the evening, its voice is feeble, a sort of creck-creck- ereck, which cannot be better compared than to the creaking of a leather shoe. But during the pairing the male produces, under water, a more sonorous and very different note—co-aci, to which the female re- sponds by a feeble coo. As a general rule, Pelodytes punctatus breeds, in France, between the end of February and the begin- ning of April; but instances of its breeding in May, July, August, September, October, and November have been placed on record by A. Thomas, Lataste, Bedriaga, and myself. Thomas, who first observed the breeding habits of this Batrachian, near Nantes, stated that it spawns twice in the year, in the spring and in the autumn. According to Héron- Royer, another French observer who devoted special attention to the habits of the same animal, Pelo- dytes breeds only once a year, in the spring. Bedriaga has observed the spawning to take place in the south of France in the spring and again in the autumn. On the 30th of October, 1896, he was so kind as to send me from Nice a large number of breeding specimens,—among them, however, only two females, one of which spawned on the way and the other reached me with eges in the oviducts. Having, besides, myself found Pelodytes pairing on the north coast of Brittany at the end of August, [ have no doubt Héron-Royer was wrong, and that the autumnal hymen will also be found to take place near Paris, where the species is so plentiful. The mode of embrace, which falls in the lumbar type, is peculiar. Owing to the thin waist of the 188 PELOBATIDA. female the elbows of the male join on the pubic region, and the forearms are carried forwards, pressed against the middle line of the belly, the hands meeting without intercrossing the fingers. Sometimes, however, the embrace is less close, the elbows fitting into the groins, and the forearms forming an angle on the belly. The pairing is usually of short duration, a few hours only, Fic. 70. Lower view of male and female pairing. and the male will, as a rule, let go if handled. Speci- mens kept in confinement by me last winter paired in January, and remained together for nearly three weeks. Stagnant water of all sorts is resorted to for the purpose of spawning, provided it be furnished with vegetation, preferably Graminacea, round the stems of which the bands of ova are twined. The larve of the spring broods transform in July and August; but late offspring of course hibernate in the larval condition, and tadpoles may therefore be found throughout the year. Males are more frequent than temales; out of 150 specimens collected by Lataste in the Gironde during the spring of 1877, only 20 were female. PELODYTES. 189 EKecs.—Small, measuring 14 to 2 mm. in diameter ; blackish, with the lower pole white. Laid in two strings which coalesce in the cloaca, thus forming a single lengthened band, often broken into several pieces, twined round the stalks of small reeds or grass below the surface of the water. The muci- laginous capsule surrounding each egg measures 23 to 3 mm.; the outer envelop, which in the toads forms a sort of tube protecting the egos, is here absent; and when, as sometimes happens, the band is deposited in fragments, the egg- @: masses embracing a stalk may “ed have more the appearance of little balls, resembling, but for the darker colour, those of Hyla arborea. Héron-Royer has ascertained the number of ova in a brood to vary between 1000 and 1600, and the embryo to leave the mucilaginous envelop on the fifth day, at a very low stage of development, tailless, and having not yet developed the gills. Tappotge (Pl. I, figs. 7 and 8).—Length of body rather more than once and a half its width, and not quite two-thirds the length of the tail. Nostrils half- way between the end of the snout and the eyes, or a little nearer the latter. Hyes on the upper surface of the body, equidistant from the end of the snout and the spiraculum, the distance between them about twice as great as that between the nostrils, and equal to the width of the mouth. Spiraculum on the left side, directed upwards and backwards, nearly equidistant from either extremity of the body, visible from above and from below. Anal opening median, much larger than the opening of the spiraculum. ‘ail twice and a half to three times as long as deep, ending in an obtuse point; the upper crest very convex, deeper Fia. 71. 190 PELOBATIDA. than the lower, and rarely extending forwards as far as the level of the spiraculum ; the depth of the mus- cular portion at its base one-third to two-fifths of the greatest total depth. Beak white, with a black margin. An inverted fold at the side of the lip; this is furnished with a single row of papille except on the upper border, which is toothed. Labial teeth in + or 2 series, the second and third, both above and below, the longest ; the first and second series in both divisions of the lip uninterrupted, or the second upper with very slight median interruption, the others separated in the middle and gradually decreasing in length to the fifth, which if present is short. According to Bedriaga, there may be as many as six series of teeth on the lower lip, the first three of which are uninterrupted. Lines of crypts usually very apparent, but some- tines very indistinct. On the head they approach each other between the nostrils and completely border the eye posteriorly, the anterior extremities of this naso-orbital hoop approaching each other above the upper lip. Of the two dorsal lines, which diverge posteriorly, the upper, extending to the dorsal edge of the muscular portion of the tail, is mterrupted at a short distance behind the eye; its anterior portion may even descend to join the lower line, which thus appears bifurcated in front; the lower line extends, usually uninterrupted, from behind the eye to the middle of the muscular portion of the tail, where it is lost; both lines, however, may stop short of the tail. A sinuous line, on the flanks, curved above the spira- culum, not bent upwards posteriorly, sometimes ex- tending nearly to the origin of the hind limbs. In addition to these lnes a short horizontal branch originates above the upper lip, and bifurcating below the vertical of the anterior border of the eye, forms a hoop which descends to the sides of the throat. Coloration usually varying from pale grey to olive- brown above, the sides with pale metallic spots; the PELODY'TES. 191 lines of crypts whitish; caudal crests greyish, with blackish spots and white dots, and pale metallic spots. Some specimens, however, have the tail almost spot- less; in others, on the contrary, it is very closely spotted, but always less abundantly on the lower crest than onthe upper. Lower parts pale grey with silvery spots. Tail and the greater part of the body with fine black decussating lines. I have observed at St. Eno- gat, in Brittany, a colony of abnormal tadpoles, not exceeding 40 mm. in length when fully developed, which were nearly black on the back, the tail dark brown with the black decussating lines very crowded, the belly of a beautiful steel-blue, and the lines of crypts quite indistinct (see Pl. I, fig. 8). The largest tadpole obtained by me in Brittany measures 57 mm., body 21, width of body 15, tail 36, depth of tail 14. One from Nice measures 65 mm. Hasrrat.—Pelodytes punctatus inhabits nearly the whole of France, from the coasts of Brittany, Nor- mandy, and the Pas-de-Calais to the foot of the Pyrenees and the Mediterranean littoral. It has not been found in the North-eastern Departments Nord, Ardennes, and Lorraine, which seem to be outside the eastern limit of its distribution. The only locaiities east of France whence this species has as yet been recorded are Castino in Southern Piedmont, and Loano in Hastern Liguria. In the Pyrenean peninsula it is on record from the Portuguese provinces Minho, Douro, Alemtejo, and Algarve, and the Spanish pro- vinces New Castille, Valencia, and Andalusia. In no part of its habitat does this species extend to any considerable altitude; in France it avoids the central plateau, and has not been found on the Pyrenees, although inhabiting Spain. The highest recorded altitude is 1450 feet at Castino, Piedmont. The specimens, male and female, figured on Pl. VIII, are from Nice, received from Dr. J. de Bedriaga. 192 PELOBATIDA. 5. PELOBATES, Wagler, Syst. Amph., p. 206 (1830). Pupil vertical. Vomerine teeth in short transverse series between the choane. Tongue circular, eutire or slightly nicked, and free behind. Tympanum absent. Fingers free, toes webbed ; outer metatarsals separated by web. Inner metatarsal tubercle shovel- shaped. Vertebre proccelous ; diapophyses of sacral vertebra very strongly dilated; urostyle usually fused with sacrum. Omosternum cartilaginous; sternum with a bony style. Three species of this essentially fossorial genus are known: two from Hurope, and one, P. syriacus, Boettger, from Asia Minor and Syria. The European species are easily distinguished from each other. In P. fuscus the occiput is humped, the orbit is not completely closed by bone behind, and the metatarsal spur or shovel is yellowish or pale brown. In P. cultripes the occiput is plane or gently arched, the cranial roof completely surrounds the orbit, and the metatarsal spur is black. P. syriacus agrees with P. cultripes in the form of the head and the structure of the skull, with P. fuscis in the colour of the spur. The genus Didocus, Cope (type Rana calearata, Michahelles), is founded on a young Pelobates cultiipes, and is therefore a synonym of Pelobates, as pointed out further on in a foot-note. The adult Pelobates cultripes has also been raised to generic rank by the same author, but quite unnecessarily in my opinion, the only distinctive feature being the greater develop- ment of the cranial roof—a mere difference of degree, the importance of which is outweighed by the close resemblance which this species bears in all other re- spects to its ally P. fuscus. ‘ponoens Ce uy a a PELOBATES. 1938 7. PeLoBares ¥uscus. (Plate IX.) Rosel, Hist. Ran., p. 69, pls. xvii—xix (1758). Bufo fuscus, Laurenti, Syn. Rept., pp. 28 & 122 (1768); Schneider, Hist. Amph., i, p. 196 (1799); Daudin, Hist. Rain. Gren. Crap., p. 81, pl. lxxx, fig. 1 (1803), and Hist. Rept., viii, p. 161 Ge3e)° Duvernoy, Régne Anim., Rept., pl. xxxviii, fig. 1 ). Rana vespertina, Pallas, Reise Russ. R., i, p. 458 (1771). Rana fusca, Meyer, Syn. Rept., p. 10 (1795); Sturm, Deutschl. Faun., i (1797); Gravenhorst, Delic. Mus. Vratisl., p. 32 (1829). Bufo vespertinus, Schneider, Hist. Amph., i, p. 225 (1799). Rana alliacea, Shaw, Gen. Zool., iii, p. 146, pls. xli & xlii (1802). Bombina fusea, Koch, in Sturm, Deutschl. Faun., ili (1828). Bombina marmorata, Koch, 1. c.; Hahn, Faun. Boic., pl. xxi, fig. b (1832). Bombinator fuscus, Fitzinger, N. Class. Rept., p. 65 (1826). Pelobates fuscus, Wagler, Syst. Amph., p. 342 (1830); Tschudi, Class. Batr., p. 83 (1838); Duméril & Bibron, Erp. Gén., viii, p. 477 (1841); Steenstrup, Isis, 1841, p. 900; Nilsson, Skand. Faun., Amf., p. 96 (1842); Ginther, Cat. Batr. Sal., p. 40 (1858); Bruch, Wirzb. Naturh. Zeitschr., ii, 1861, p. 178, and iii, 1862, p. 181; Collin, Naturh. Tidsskr. (3), vi, 1869, p. 316; Fatio, Vert. Suisse, iii, p. 376 (1872); C. Koch, Ber. Senck. Ges., 1872, p. 151; Cornalia, Atti Soc. Ital., xvi, 1873, p. 96, pls. ii & iii; De Betta, Faun. Ital., Rett. Anf., p. 71 (1874); Schreiber, Herp. Eur., p. 90, fig. (1875); Ley- dig, An. Batr., p. 77 (1877); Lessona, Atti Ac. Lincei, Mem. Sc. Fis., i, 1877, p. 1077, pl. iti; Boulenger, Cat. Batr. Ecaud., p. 437 (1882); Fatio, Vert. Suisse, iv, App., p. v (1882); Camerano, Mem. Acc. Torin. (2), xxxv, 1883, p. 217; Héron-Royer, Bull. Soc. Et. Sc. Angers, xv, 1886, p. 61; Camerano, Boll. Mus. Torin., i, 1886, No. 9; Boulenger, Bull. Soc. Zool. France, 1888, pp. 115 & 162; Peracca, Boll. Mus. Torin., iii, 1888, No. 46; Wolterstorff, Zool. Anz., 1888, p.672; Héron-Royer & Van Bambeke, Arch. Biol., ix, 1889, p. 269; Bedriaga, Bull. Soc. Nat. Mosc., 1889, p. 493; Boulenger, Proc. Zool. Soc., 1891, p. 614, pl. xlvi, fig. 7; Zander, Corr.-Bl. Nat. Ver. Riga, No. 38, 1895, p. 62; Werner, Rept. Amph. Oesterr.-Ung., p. 104 (1897) ; Diirigen, Deutschl. Amph., p. 519, pl. ii, fig. 7, and pl. iii, fig. 1 (1897). Cultripes minor, Miller, Isis, 1832, p. 588, and Zeitschr. f. Phy- siol., iv, 1832, p. 212. Pelobates insubricus, Cornalia, Atti Soc. Ven.-Trent., ii, 1873, . 44, Petobates latifrons, Héron-Royer, Bull. Soc. Zool. France, 1888, pp. 85, 108, 117, figs.; Héron-Royer & Van Bambeke, 1. c., p. 272. N 194 PELOBATID. Vomerine teeth in two strong, transverse, slightly oblique or slightly curved series between the choane, the inner borders of which they touch, Fre. 72. and narrowly separated from each other in the middle. Hustachian tubes very small. Tongue large, thick, cir- cular, sometimes entire, usually feebly nicked behind. Head very convex, broader than long, the occiput gibbose; the skin adherent to the skull; snout rounded, Open mouth. projecting beyond the mouth, as long as or a little longer than the diameter of the orbit; no canthus rostralis; nostril midway between the eye and the tip of the snout; eye large, prominent ; interorbital space convex, much broader than the upper eyelid, and a little greater than the distance between the nostrils; a sort of low knob behind the eye, formed by the squamosal bone. Fingers moderately elongate, pointed, third much the longest, first and fourth equal and slightly longer than second; subarticular tubercles very indistinct ; two rather indistinct carpal tubercles. Hind limbs robust and short, with swollen calves ; the tibio-tarsal articulation reaches the shoulder or the commissure of the mouth, and the tarso-metatarsal the eye, or between the eye and the nostril; tibia shorter than the femur, the heels being widely sepa- rated from each other when the legs are folded. Foot much longer than the tibia ; toes short, pointed, broadly webbed, at least two- thirds webbed in summer, the web reaching the tips of all the : toes in the spring; subarticular \) tubercles indistinct ; a very large, compressed, sharp-edged inner metatarsal tubercle, placed ob- liquely to the axis of the foot like the inner toe, which it equals or slightly exceeds in length. Fig. 73. a ey ea Lower view of foot. PELOBATES. 195 Skin smooth on the head and limbs; sometimes smooth on the back, usually covered with small flat warts of unequal size ; lower parts smooth, with a few eranules on the pubic region. Coloration very variable. Ground colour olive-grey, pale brown, yellow, yellowish-white, or brownish- white, with dark-edged chestnut-brown, olive-brown, or reddish-brown spots, which often simulate larger and smaller islands with numerous indentations, or may be confluent into a pair of longitudinal bands, bordering a pale area of the ground colour along the spine; sometimes the spots are very small and nume- rous, giving the animal a speckled appearance; the spots often form irregular bands across the hind limbs; a dark streak usually extends on each side of the snout to the eye, involving the nostril, and a dark, continuous or interrupted band sometimes extends from one eyelid to the other across the crown; a light streak of the ground colour is often present on the coccygeal region. In addition to these markings there are often brick-red or vermilion spots or dots, which may be confined to the sides, or so profusely scattered onthe whole of the upper surfaces as to almost obscure the ground colour. Lower parts dirty white, uniform, or spotted or speckled with greyish- Fra. 74. brown, the soles often greyish- brown; metatarsal spur yellowish, pale horn-colour, or reddish-brown. Iris golden or copper-colour, some- times fire-colour. Male distinguished from the fe- male by a large, oval, smooth gland on the upper surface of the bra- chium. Vocal sacs and copulatory excrescences are absent; during Upper view of fore limb the breeding season, however, small op Eacotino Stale. pearl-like, uncoloured excrescences are scattered on the upper surface of the antebrachium and the fingers. 196 PLOBATIDE. Measurements (in millimetres). é P a SS s 1. 2. 3. 4. 5. 6. From snout to vent BU sos DD ice 89: 34 OF vee C2ome GL Length of head sd oe LD ace QP ae RO ew BU ge AD Width of head DO sp OBE i Die WO. ang De as, BS Diameter of eye Ona Ol aae oF 6 7 6 Interorbital width GW veeite: (Ol Ace “Fo, ee ab 7 From eye to nostril J to 4 450 450. «5 5 a ss end of snout Si ce Or sek ig OD ey 10 10 Fore hmb : ‘ . 33... 383..., 38 1... 88 388... 37 Hind limb ; TU os FL -. BL, 81 85... 80 Tibia. NO. ee TD? sa, BE 93 ax, Bde 2. BE Metatarsal tubercle. Sas Ol ae Do un Stee Boe BD Inner toe (from tubercle) Aw. bon 8 wn 4b... 8... 45 1. Drancy (Seine-et-Oise): Paritre.! 4. Paris: Héron-Royer. 2,6, Turin: Peracca. | 5. Halle/S.. Wolterstorff. 8. Szamos-Ujvar (Transylvania): Méhely. SxeLveton.—Skull very strongly ossified, the maxil- laries, squamosals, nasals, ethmoid, and fronto-parietal rugose, granular, or pitted. Ethmoid entirely ossified, confluent with the likewise ossified nasal capsule, and produced forwards beyond the nasals to between the ascending processes of the pramaxillaries; only a small triangular portion of it is exposed between the nasals and the fronto-parietal. Nasals large, in con- tact with each other between the nares. Fronto- parietal single, widening behind into a more or less developed angular process on each side; this process is more or less widely separated from a corresponding one of the squamosal, with which it 1s connected by ligament only, the prootic being entirely exposed above. Transverse branch of squamosal expanded, plate-like, suturally united in front with the maxillary. The degree of roughness of the fronto-parietals and the development of the post-orbital processes vary according to individuals, the bone being generally less developed in this respect in specimens from France than in those from Germany, Austria, and Italy. In quite young specimens the fronto-parietal is nearly PELOBATES, 197 Skeleton of male from Transylvania. 198 PELOBATIDE. smooth, and the dermo-ossification, as it develops, produces granular asperities which may become con- fluent into alveolar ridges, producing a pitted appear- ance. In very old specimens the sutures between the nasals, the ethmoid, and the fronto-parietal may become obliterated. Vomers rather large, variable in shape, extending or not to the palatines, which are strongly ossified. On the palatal side the ethmoid extends posteriorly to about the middle of the length of the parasphenoid. Latter large, L-shaped, its anterior pointed extremity produced to between the palatines. Pterygoids not reaching the palatines, extensively in contact with the parasphenoid. Stapes absent. Mento-Meckelian or symphysial bones distinct on the inner side only. Hyoid alarge broad cartilaginous plate, with slender postero-lateral processes; anterior processes much expanded and confluent with the lateral wings, en- closing a small fenestra; ceratohyal cornua with short posterior portion detached from the body of the hyoid; thyrohyals large, massive, in contact at the base, diverging and more or less expanded poste- riorly. Vertebral column once and a half to twice as long ag the skull. Spine closed above, the neural arch produced posteriorly into a strong median process between the zygapophyses. Three anterior diapo- physes strong and long, especially the second, which bears a more or less distinct dorsal knob or process as on the corresponding rib of Discoglossus,—the first directed obliquely forwards, the second and third nearly horizontal; the following short and slender, directed forwards. Sacral vertebra with very strongly dilated diapophyses, the transverse diameter of which is twice and a half to nearly three times in the axial ; urostyle short, not longer than the sacral wings, and fused with the sacrum, to the diapophyses of which its anterior portion contributes if these processes be PELOBATES. 199 not entirely furnished by it, as is sometimes the case.* Precoracoids slender, strongly curved, not entering the glenoid cavity ; coracoids nearly straight, strongly expanded distally ; a cartilaginous omosternum; ster- num with a bony style dilated proximally and _a large discoid cartilage. Humerus considerably longer than radius-ulna. Hight bones in carpus, three of which are in contact with radius-ulna ; a single bone in the pollex. Pelvis five-sevenths to five-sixths the length of the vertebral column; ilia with an inner upper groove into which the sacral processes slide; pubis absent or reduced to a small bony nodule, not entering the acetabulum. Femur and tibia with cartilaginous epi- physes ; tibia shorter than the femur, which is shorter than the pelvis. Calcaneum slightly shorter than astragalus, half as long as tibia; three distal tarsal bones ; prehallux with a single very large, compressed, curved phalanx. Terminal phalanges pointed. MeAsvREMENTS OF SKELETON (in millimetres). 3 g Length of skull. : ; 14 ony 17 Width of skull Ps 3 18 Pas 24 Least interorbital width 5 Ns 6 Dorsal vertebral column . j li aa 18 Urostyle (with sacral vertebra) 105... 13 Humerus . : Z : 12 ae 15 Radius-ulna 8 - 10 Manus 12 te 16 Pelvis : 20 i 22 Femur 1? sey 22 Tibia 4 : 15 “ah 19) Tarsus y it wd 9 Pes é A : 23 ice 28 * I have long ago pointed out that the vertebral column figured by Gené in 1839 as of Bombinator belongs to a Pelobates. I should not allude to this again were it not for the fact that Gené’s error has crept into Bateson’s ‘Study of Variation,’ p. 127, in a paragraph marked with an asterisk as a sign of special importance to the example quoted, and with the addition, by the conrpiler, of a further error, viz. that the specimen came from Sardinia. The sacrum in the Gené specimen, as well as in others that have since been described and figured by Adolphi (‘Morphol. Jahrb.,’ xx, 1895, p. 449, pl. xix), is formed entirely by the processes at the base of the urostyle, and there are thus nine instead of eight presacral vertebre. 200 PELOBATID. Hasrrs.— Pelobates fuscus is a thoroughly burrowing Batrachian, spending the greater part of its existence several feet below the surface of the ground, the capacious lungs filled with a good provision of air ; it buries itself in a nearly erect squatting posture by means of the large sharp-edged horny shovels with which its feet are armed, the digging being effected by alternate lateral movements of the heels, inter- rupted now and then for a short rest; on sand or a very loose soil less than a minute may suffice for the animal to disappear from sight, the soil falling over and around it so as not to leave a trace of a burrow. The Englsh name “ spade-foot,” which is applied to its North American representative, Scaphiopus, would be a most appropriate one. Owing to these habits it is only found in localities where the soil is hght or sandy ; in cultivated districts the extensive growth of asparagus is an almost infallible indication of its pre- sence. In the summer months it emerges an hour or two after sunset, and hopping along frog-like, sets forth in search of its food, consisting of worms, slugs, and all kinds of insects, especially beetles. When suddenly seized it produces a dermal exudation, smell- ing like garlic, and usually utters a startling shrill cry much like that of a kitten, at the same time opening its mouth in a defiant attitude. Some specimens when repeatedly teased—pinches in the leg and light pokes in the sides being the best method to employ to witness the sight—can be roused to what appears to be a fit of anger lasting several minutes, the animal screaming in a most extraordinary manner, and jump- ing madly with open mouth as if to snap at its perse- cutor. The prolonged screams produced on those occa- sions can only be compared, in a diminutive way, to those of an infant. At the same time the lungs are inflated to their utmost, and the body, swollen like a ball, is raised above ground. A specimen is figured in this attitude on Pl. IX, right hand. These antics are very similar to those performed by Ceratophrys PELOBATES. 201 ornata, a South American Cystignathoid, of likewise nocturnal and burrowing habits, specimens of which have often been kept in confinement in this country ; an important difference being, however, that a large Ceratophrys is able to inflict a very painful bite on the finger of the unwary. This Batrachian appears in the daytime only during the breeding season, which takes place in March and even as late as May* in Italy, between the end of March and the beginning of May in Central and Northern Europe, stray pairs occurring exceptionally as late as the 21st of July, according to an observation made by Prof. Van Bambeke at Ghent, Belgium, in 1875. At the time of pairing, at which pools or deep ditches are resorted to, the males, much more nume- rous than the females, utter under water a mono- tonous, constantly repeated note—clock-cloch, clock- clock-clock,—produced by alternately shifting the air backwards and forwards from the capacious lungs into the buccal cavity. The female, which answers by a sort of grunt or a deep tock-tock-tock, is seized round the waist, and the eggs are expelled, more or less rapidly, either immediately or within a few days. The length of the larval life varies considerably ; as a rule, the final transformation takes place from the beginning to the end of summer, but several cases of larval hibernation have been observed by Pfliiger and Kollmann. Eaas.—The contents of the two oviducts fuse in the cloaca, and are expelled in a thick band 15 to 20 mm. in diameter when swollen up, in which the ova are irregularly disposed at small interspaces. These bands, which have a strong smell of fish, are twisted round weeds by the female as they are laid. The ova * According to Spallanzani, who described the oviposition in a pair brought to him in May, 1780, by a fisherman at Pavia, and who received afterwards other pairs of the same species (uvres de Spallanzani, III, ‘Expériences pour servir 4 l'histoire de la Généra- tion,’ Pavie, 1787, p. 187). As Crivelli and Camerano have shown, Spallanzani was the first after Résel to observe this curious Batra- chian, which was not rediscovered in Italy until ninety years later. 202 PELOBATIDS. are small, 2 to 24 mm. in diameter, grey or brown with white lower pole, and develop rapidly; the em- bryo, gill-less and tailless, breaks through about the fifth or sixth day, and attaches’ itself, through its adhesive subcephalic apparatus, to the outer surface of the mucilage. Tappote (PI. IT, fig. 1).—Length of body once and a half to twice its width, one-half to two-thirds the length of the tail. Nostrils a little nearer the eyes than the end of the snout. Eyes on the upper sur- face, equidistant from the end of the snout and the spiraculum, the distance between them at least twice, sometimes nearly three times as great as that between the nostrils, and greater than the width of the mouth. Spiraculum on the left side, directed upwards and backwards, equidistant from either extremity of the body or a little nearer the anterior extremity, visible from above and from below. Anal opening median, a little larger than the spiraculum, and close to the body. ‘Tail twice and a half to thrice and one-fifth as long as deep, acutely pointed; upper crest convex, slightly deeper than the lower, not extending far upon the back; the depth of the muscular portion at its base about half the greatest total depth. Beak black. Lip bordered with papille, which form two or more rows on the sides; the papillose border interrupted mesially by a narrow toothed descending lobe, which appears at first sight as continuous with the second upper series of teeth. This anterior series is followed by three or four other series of teeth, which are all widely interrupted in the middlé and PELOBATES. 203 gradually decrease in length,—the fourth, if at all present, being extremely short. On the lower lip we see likewise a short outer series, followed by three or four much longer ones, all of which, with the occa- sional exception of the first, are interrupted in the middle, and may be more or less broken up on the sides. The series of labial teeth may therefore be formulated as + or 8. Small isolated teeth may also be scattered on the papille at the angles of the mouth. I have not been able to distinguish lines of crypts beyond the two series which run from the end of the snout to the upper border of the eyes, passing be- tween the nostrils. J am therefore unable to judge whether the dorsal lines have been correctly figured by Lessona, who rightly regards the figure given by Cornalia as fanciful. The advanced tadpole is brown or olive-brown above, with or without small darker spots, greyish-white or bluish beneath; sides with roundish whitish or pale golden spots; tail pale brown, with small grey and whitish spots. The body usually reaches at least the size of a pigeon’s egg, but not unfrequently exceeds that size. The largest specimen in the British Museum, from Prague, measures 125 mm., body 38, width of body 25, tail 87, depth of tail 27. The largest specimen on record is one preserved in the Berlin Museum, obtained in December, 1867, on the Jungfernheide, near Berlin, and which measures 175 mm. Havitat.—The range of this species is an extensive but broken one, its occurrence being obviously depen- dent on the nature of the soil, and restricted to the plain, the highest elevation at which it has yet been recorded being 1400 feet (Val di Susa, Piedmont). P. fuscus also appears to avoid competition with its con- gener P. cultripes; the limit of the range of the two closely meet without anywhere overlapping. It in- habits the north-east and east of France, from Flanders 20-4 PELOBATIDE. to the Jura, westwards to Paris, the Dep. Sarthe, and along the Loire; Belgium (Prov. Limburg, Antwerp, and Kast Flanders), South Holland, Luxemburg, Swit- zerland near Basle, Germany (except Wiirtemberg), Austria-Hungary, Carniola, North Italy (Piedmont, Lombardy, Venetia, Emilia), Denmark, Southern Sweden and Gotland, and Russia, northwards to the Gulf of Riga and eastwards to the Ural River and the Kirghiz Steppes, the Emba River being its eastern- most limit. The species has quite recently been recorded from Lenkoran, south-west coast of the Caspian Sea, by Zander.* The reported occurrence of Pelobates fuscus in the Cyclades is probably to be ascribed to a confusion with Bufo viridis, some specimens of which closely resemble this species in their markings. The specimens represented on Pl. IX are from the environs of Turin, presented by my friend Count Peracca, to whom I am indebted for so much material utilised in the preparation of this work. The left- hand figure is that of a female; the others represent males, one in the act of disappearing under the sand, another in the irritated attitude described abeve. * Dr. Zander kindly informs me that tbe specimens have been pre- sented by him to the Senckenberg Museum, Frankfort (M.), and that his determination has been confirmed by Prof. Boettger. eoPiiggns fs ? Ce PELOBATES. 205 8. PkLOBATES CULTRIPES. (Plate X.) Rana cultripes, Cuvier, Régne Anim., 2nd ed., ii, p. 105 (1829). Rana calcarata, Michahelles, Isis, 1830, p. 807, pl. —. Cultripes provincialis, Miller, Isis, 1832, p. 5388, and Zeitschr. f. Physiol., iv, 1832, p. 212. Bufo calearatus, Schinz, Nat. Rept., p. 233, pl. xevi, fig. 2 (1833). Bombinator fuscus, Dugés, Rech. Ostéol. Batr., p. 7, pl. 11 (1834). Pelobates cultripes, l'schudi, Class. Batr., p. 83 (1838); Duméril & Bibron, Erp. Gén., viii, p. 483 (1841); Gunther, Cat. Batr. Sal., p. 41 (1858); Schreiber, Herp. Hur., p. 92 (1875) ; Lataste, Herp. Gir., p. 263 (1876); Boulenger, Cat. Batr. Eeaud., p. 4388 (1882); Héron-Royer, Bull. Soc. Et. Se. Angers, xv, 1886, p. 77; Héron-Royer & Van Bambeke, Arch. Biol., ix, 1839, p. 275; Bedriaga, Bull. Soc. Nat. Mosc., 1889, p. 519, and Amph. Rept. Portug., p. 19 (1889) ; Boulenger, Proc. Zool. Soc., 1890, p. 664, and 1891, p. 616, pl. xlvi, fig. 8. Pelobates fuscus, part., Bonaparte, Icon. Faun. Ital., Rett. Anf., 1888). ee calcaratus, Cope, Journ. Ac. Philad. (2), vi, 1866, p. 81. In the tongue and vomerine teeth this species entirely agrees with its congener, but it differs in the occasional presence of small, grain-like teeth on the pterygoids and parasphenoid; these teeth are quite rudimentary, and the mucous membrane hus to be removed to ascertain their presence. I have found them in three specimens from France (Nantes, Bor- deaux, south of France). In one specimen there are about ten teeth on the parasphenoid, at the base of the longitudinal branch of the L-shaped bone, and two pterygoid teeth close together on the left side; another has the same number on the parasphenoid and a series of four on the pterygoids; and in a third there are eight teeth on the right pterygoid, none being present on the parasphenoid nor on the right pterygoid. * The genus Didocus was founded upon a young specimen, on the erroneous assumption that in Pelobates cultripes the temporal roof is developed before the tail of the larva has disappeared. I have examined the skeleton of a young but fully transformed specimen from Béziers, Hérault, which shows both a large fronto-parietal fontanelle, bordered by smooth fronto-parietals, and un uncovered temple, thus answering in every respect the definition of the genus so rashly established by Cope more than twenty years ago, and still maintained by him in his latest writings. 206 PELOBATIDA. Head as in P. fuscus, but rather larger in propor- tion, and not humped on the occiput nor behind the eyes, the skull forming a complete, evenly curved rugose casque entirely surrounding the orbits; imter- orbital space nearly flat, usually much broader than the distance between the nostrils. Shape and proportions of the hmbs much as in P. fuscus, but digits more obtuse, and metatarsal shovel sharper and larger, always exceeding the length of the inner toe. Skin smooth or densely covered above with very small round warts. Yellowish, whitish, or greenish-yellow above, speckled, spotted or marbled with brown, olive, or greyish-olive, the spots small or large, and but rarely confluent into longitudinal bands. A small male specimen, collected by M. Lataste at Greilhan, Gironde, has four perfectly regular dark dorsal stripes, the median pair extending forwards to the interorbital region, where they meet a dark cross-bar. White beneath, uniform or speckled with greyish- olive; metatarsal spur black; toes often tipped with a black horny layer. Inis silvery or greenish-grey, vermiculate with black. External male sexual characters as in P. fiseus. In describing Pelobates syrweus, Boettger (‘ Zool. Anz.,’ 1889, No. 302) mentions breeding specimens of Pelobates cultripes to be possessed of nuptial asperities on the inner side of the three inner fingers and on the tips of the toes. This extraordinary statement is based on a confusion with the American Scaphiopus ham- mondit, Baird (dugesit, Brocchi), breeding males of which were erroneously labelled ‘ Pelobates cultripes, Spain,’ in the Senckenberg Museum. The so-called nuptial asperities of the tips of the toes are nothing but the blackish horny sheaths which cap them, as well as the tarsal spur, at all seasons. I am indebted to the kindness of Prof. Boettger for an opportunity of examining these specimens. PELOBATES. 207 Measurements (in millimetres). g g 1, 2. 3. 4. 5. 6, From snout to vent . 66. 77... 57... SL... 88... 62 Length of head 22 26 aie 20) wee TP ey 27 5. 20 Width of head . 26 29 26 25 4, QL. 86 au. 25 Diameter of eye Oe Ts Dee BR vee Decal! OOF aye Interorbital width OO “Bisa Fee CS. Pan 6 From eye to nostril Dae Pe DO On Blan Bi es Bs endofsnout .10 ... 13...10 ... 9 ... IL... 10 Fore limb 5 ; 400... 45... 85 0. 27 1. 47. BA Hind limb ; 88... 100... 80 ... 67... 118 ... 79 Tibia . ; F Ble cme SOB) pie WO de OK ane 32s OD Metatarsal tubercle . 2 Bove Boise 6 ue BS. ae Bia 6 Inner toe (from tubercle) Ol stecee (AG. Se Oe Say “OD ace Do enya 1. Nantes: Paris Museum. 5. Badajos: Bosca. 2. Seville: Calderon. 6. Abrantes: Gadow. 8, 4. Merida: Bosca. SKELEvoN.—The trunk and limbs agree with those of its congener, but the skull is very different, owing to the Fig. 77. Upper view of skull. more extensive dermo-ossification of the post-orbital region, which forms a complete roof over the prootic, only the posterior portion of the latter being visible from above. This roof, which completely closes the orbit behind, is formed by the fronto-parietal and the squamosal; the latter bone has a convex posterior border, separated from the lhkewise curved posterior border of the fronto-parietal plate by a notch, the angle of which corresponds to the suture between the two bones. 208 PELOBATIDS. All the bones of the upper surface are closely studded with granular or subconical asperities. A further important difference in the skull of this species compared to P. fuscus is the absence of the extra- ordinary forward prolongation of the ethmoid; this bone does not extend beyond the nasals. Dugés, basing his observations on this species, has denied the fusion of the sacral vertebra with the coccyx described by Mertens in Pelobates fuscus, with which P. cultripes was then confounded ; he observes, however, that the articulation, by means of one con- dyle, is an almost immoveable one. Ina specimen from Bordeaux, from which I prepared the first skeleton, I found matters as stated by Dugés, whilst in two other skeletons, from Bordeaux and Avignon, the two bones are as completely fused as in P. fuscus. As the ankylosis of the sacrum and coccyx has often been given as a generic character of Pelo- bates, it is important to note the imconstancy of the character in this species at least. MeasvRemEnts of SKELELON (in millimetres). 3 2 Length of skull. 23 sie 25 Width of skull i 31 Least interorbital width . 7 5: TS Dorsal vertebral column . of af 26 Urostyle (with sacral vertebra) iV sit is Humerus. wat 20 Radius-ulna : : 14 sh 18 Manus 20 se 19 Pelvis 34 dass 38 Femur 5 ‘ 28 Le 30 Tibia ‘ P Os ate 26 Tarsus 13 a 18 Pes ‘4 : : ; 34 i 36 Hasrrs.—Much the same as those of its congener. P. cultripes seems, however, more partial to the coast, being found in abundance, in France, on the sandy dunes of the Atlantic littoral. It breeds in the end of March or April in France, in March in Spain. Its note, which I have only heard in specimens captured PELOBATES, 209 in summer, may be well rendered by a guttural co, CO, CO, Co, co ; its nuptial call is said to be the same. Kacs.—According to Héron-Royer, they differ but slightly from those of P. fuscus; the mucilaginous band is a little narrower and flatter, thus much re- sembling that of Pelodytes punctatus. Tappote (Pl. II, fig. 2).—The differences from P. fuscus are but slight, and most of those that have hitherto been relied upon prove not to be absolutely constant. Thus the nostrils, as I now find from Portuguese specimens received from Dr. de Bedriaga since the publication of my ‘ Synopsis of the Tadpoles,’ may be as wide apart as in P. fuscus. The series of labial teeth are more broken up, and their arrange- ment is therefore less easily expressed by a formula. The tail is shorter, not more than once and a half the length of the body. The lines of crypts are usually more distinct, owing to the black colour of the tubules and the lighter colour of the body. heir disposition is well shown in the example figured. The following description is taken from five specimens obtained by M. Lataste near Bordeaux : Length of body once and two-thirds to once and three-fourths its width, once and one-fourth to once and two-fifths in the length of the tail. Nostrils half- way between the eyes and the end of the snout, or a little nearer the latter. Eyes on the upper surface, a little nearer the spiraculum than the end of the snout, the distance between them nearly twice as great as that between the nostrils, which equals the width of the mouth. Spiraculum equidistant from either ex- tremity of the body. Tail once and two-thirds to once and five-sixths as long as deep, pointed, the muscular portion not half the total depth. Specimens preserved in spirit, of which I have a good number before me at the present moment, are pale greyish or brownish above and on the sides, the belly dirty white with round white spots. Muscular part of tail pale reddish-brown, with some darker 0 210 PRLOBATIDA. brown spots; caudal crests greyish, more or less distinctly spotted or freckled with brown. The coloration in life is described by Lataste as reddish-yellow above, greyish or bluish-white beneath ; tail with small brown spots. Some twenty years ago I saw a number of large specimens in a small tank jn the Botanic Gardens at Bordeaux. My recollection is that they were of a very pale brown, without distinct markings. Total length 100 mm.; body 39; tail 61; depth of tail 25. According to Dugés the size of the body may equal a hen’s ege. Hacrrat.—France, Spain, and Portugal. In France this species has been found along the west - coast to a little north of the mouth of the Loire, and in the southern departments Haute-Garonne, Pyrénées- Orientales, Aude, Hérault, Gard, Bouches-du-Rhoéne, and Vaucluse; in Spain in the provinces Galicia, Estremadura, New Castille, Valencia, Andalusia; and in Portugal in the provinces Douro and Algarve. It probably occurs, as stated by Bosca, all round the coasts of the Pyrenean Peninsula. The record of the occurrence of Pelobates cultripes in Syria is due to a confusion with an allied species, Pelobates syriacus, to which allusion is made above, p. 192. ‘The two specimens figured on P]. X were obtained near Bordcaux by M. F. Lataste. PRINTED BY ADLARD AND SON, BARTHOLOMEW CLOSE, ¥.C., AND 20 HANOVER SQUARE, W. bus fay Basie tes ate ees ie gcetat era ey aetna Sj baie Es pars pee prea) ys rie boca bee et aa (ee f i Lae eR a ee are eC trem it sere CUR, galalasacP j Pal rasa : eran Mh hed 4 ily Hy Heaaatrtyireni ty niyo it BY Pt 5 vo hyphen iE ake Ph in coun Sans vatuntine | Cord ue at ‘ iy may Shai i aaah katate ci a wet Ee seinen mR it en ten) naa PT ae Sat