Taxonomy and Biology of the Genus CYCLORHAMPHUS b: DR. hea LUTZ (With Plates 1—5.) NAME AND CHARACTERISTICS OF THE GENUS. Though there has been a good deal of confusion in the literature, the genus Cyclorhamphus _(Cycloramphus TSCHUDI 1838) has undoubted priority and may be clearly distinguished from the other genera of the family Cystigna- thidae or Leptodactylidae, It precedes the names Pithecopsis BIBRON and Gry- piscus COPE, associated with the only species umbrinus. The name Iiodiscus MIR-RIB., though more characteristic, can not be maintained for the same rea- son. Cyclorhamphus receives also two species erroneouly included in Telmato. dius. The best characteristic of the genus consists in a raised dise with flat surface, found in the inguinal region of the males and becoming more distinct in propor- tion to the.age. (It is not found in fe- | males or yery young males.) The disc has a gelatinous aspect like the wheal raised by an intradermic inje diametre does not exceed 10 elevation is only a few mm. The disk must be looked for, as it is not always very evident, A similar formation is shown only by the males of Craspedoglossus Sanctae Catharinae LORENZ MUELLER. [The species ought to be called bolitoglossus, as it was named so by WERNER (who in cluded it in Borborocoetes), white it cer- tainly deserves a new generic name.] Some Paludicolinae show a somewhat similar organ, but in both sexes and ge- nerally distinguished by its colour, A confusion with Craspedoglossus can be excluded by examining the tongue. Another important character has been pointed out by MIRANDA-RIBEI- RO under the designation meniscus. It is a discoid lobule of the iris, insertedures ee in the middle of the superior pupillary margin, When it stands out over the black area of the pupil, as in a figure of M-R,, it is very striking, though it is always quite small, On the other hand, being connected by a narrow and flexible stalk, it is easily turned back which ex- plains why it may often fail to appear, even in living adults, and still more in: preserved material. Though probably al- ways existent, it is not equally distinct in every species. The tympanum and the rostral can- thus are wanting in all the species, but the teeth exist in the usual place and ar- rangement, i In the skeleton we find the sacral apophyses somewhat flattened and di- lated towards the apex, while the ster- num consists of a very thin hyaline car- tilage. The terminal phalanx is gene- rally T-shaped, but there are no real disks at the rounded tip of the finger, though it may be somewhat flattened and dilat- ed. On the hind feet an interdigital mem- brane is either well developed or alto- gether absent, The general form of the head and trunk is somewhat flattened and the snout large and rounded without a pro- jecting canthus. All this, together with the rather dull and often granular skin, the large trunk and the rather stout limbs of these frogs gives them a so- mewhat toadlike appearance, These pe- cularities are explained by the habits, as all the species like to hide in daytime in the clefts and interstices of stones and rocks, exposed by the action of the mountain brooks, full of larger and smal- Jer falls. The skin of these batrachians is ra- rely quite smooth or only finely granu- lar. Often it shows glandular warts of light colour and shaped like the seg- ments of a sphere or an egg, In other cases the back is densely covered with granulations, either opaque or translu- cent (like grains of boiled sago) which may show a corneous point, either black- ish or ochraceous, Sometimes they are grouped in rows forming short ridges. The ventral face may be light or vermi« culated with brown and this pigmenta- tion, which increases with age, may in some species reach a point where the ground becomes altogether blackish show- ing only a sprinklimg of white points. esh_ specimens may be distinguished by this pigmentation which however is liable to change in preserved material The dorsal side may be altogether sooty or show larger spots of lighter hue, principally in younger specimens. In as- pera WERNER the back is light brown with or without darker spots, The skin is generally very loose, as shown principally in little hardened al- cohol-specimens. (I must mention here a fact I have not found cited in the litterature on bra- zilian frogs. This is the frequent oceur- rence in their skin of red acarine larvae. I have knodwn them for more than 40 years and recognised them as closely al- lied to Lepfus and belonging perhaps to more than one species of fhrombidi- idae. Sometimes they are just below the lermis but in Cyclorhamphus where they are common, they may also invade the mucous glands, During live they are distinguished by the red colour, but when this has faded in alcohol-speci- mens, they may be confounded with di- lated mucous glands. Perhaps even the hypertrophic state of these glands may be due to the previous attack by lar- vae which have already escaped. I have not yet obtained adult speci- mens of the mites but have verified the transformation of the larvae in free-liv~ ing nymphae with four pairs of legs. All the species, I observed, have the same habits, In daytime they are hid- den in clefts with rocky or stony walls, in contact with swiftly running water, but their yoice is often heard comingSp } i . out of the clefts as a clucking or a short and rough croak. If one tries to catch them, these frogs retire to still more impenetrable places and when surprised by the lift: ing of a stone or a log under which they were hidden, they give quick and strong jumps. This explains the diffi- culty of catching them, even in places where they are numerous. When it be- comes darker, they seem to come out of their hiding places, to hunt, like the toads; but they alway stay near lo the water. In the period of propagation the males have big testicles while the fema- Jes contain numerous large eggs in which the cream color hides all the dark pig- ment which exists in the small eggs of recent formation. SPAWNING, FORMATION OF TADPO. LES AND METAMORPHOSIS. MIRANDA-RIBEIRO found in the Museu Paulista some eggs containing little fourlegged frogs which he attributed to a Cyclorhamphus. So he stated that the development in Cyclorhamphus res- sembled that of Craspedoglossus boli glossus the eggs of which contain small fourlegged frogs; they where described on the same occasion and shown in a dra. wing. As at that time I already possessed observations and drawings of the tad- pole of C. fuliginosus, 1 naturally con- cludes that the described eggs were not those of a Cyclorhamphus and more likely to belong to Craspedoglossus. The various species of Cyclorhamphus hav- ing quite similar habits, an absolutely different form of evolution was most un- likely. Moreover the young frogs of M- R. showed no interdigital membrane, * known to exist on the hindfeet of the two species described from S. Catharina, while my transitional stages between tad- poles and frogs showed them’ already most distinctly. LAs gee However, it seemed desiderable to find out about the development of those tadpoles which were always found cling- ing to the slanting and even perpendi- cular surface of rocks, irrigated by a thin film of running water. Near to those places and probably outside the water, the spawn might be looked for. ‘The researches made in appropriate places in December of 1927 and in the beginning of 1928 actually gave the solu- tion of the problem, masses of spawn. being found four times in three different and distant places. In October 1928 four more egg-masses were discovered in three localities, two of them quite new. They belonged partly to C. fuliginosus, partly to C. pinderi and were found in the pla- ces inhabited by the adults, sometimes exposed, sometimes under leaves or pro- jecting stones, generally near to, but ne- ver in the water. They contained seventy to more than a hundred eggs glued toge- ther in diskshaped clusters With a diameter of 10 and a maximal height of 2-3 cen- timeters, Each egg forms a gelatinous sphere bearing in its centre a vitelline globule of 4 mm. thickness and yellow- ish cream-color. Taken home and spread on moss or damp stones, they developed rapidly with exception of two which remained sterile and became mouldy, due either to lack of fecundation or improper conditions during the first days, Some spawnelusters were already in evolution and showed the stages seen on the pla- tes. There were also tadpoles caught on their way to the water. The first stage of development con: sists in the formation of a very crudely shaped embryo, bent around the yolk- mass. It shows external branchia with short ramifications in which the blood circulates very rapidly. Afterwards the tail becomes free and stands out in a rather flat curve. Already in this stage quite lively movements may be seen, Dut they cease soon and are followed by$90 Ss Jong periods of absolute rest, Next some pigment-spots appear as an indication of the eyes and jaws; the dorsal region may also show some pigmentation, In this stage the embryo shows head and trunk enclosing a subglobular vitelline mass, around which blood circulates acti- vely in some vessels, seen with low po- wer, There is also a long tail ending in a elongated and vertical, oarshaped blade. When the masses are sprinkled, the tad- poles may now be seen to come out of the egg membrane. They stay for some time in the surrounding jelly but finally wriggle of towards the water. The mouth, turning downwards, shows now the citcumoral combs of small roads. The exspiratory tube, located in the middle line shortly behind the mouth, is not easily seen and appears rudimen- tary. The nostrils are quite distinct. The anus seems to occupy the median line, On the upper part of the ventral side a red zone shows early, apparently cor- responding lo the heart and the Iungs which are well provided with vessels and may begin to function. The tadpoles are now quite strong swimmers but can not stay in the water unless they emerge at least with the an- terior part of the body. They adhere to vertical walls without using the tail which does not serve for fixation but only for locomotion. While absorbing the _yolkmasses which for a long period nourrish the lar- vae, these take a more familiar aspect helped by the increase of dark pig- mentation. Their elongated form becomes quite characteristical. The oval or obcor- date trunk looses in breadth but remains short, while the tail becomes four or five times longer showing awlshape with a narrow vertical oarlike blade at its end. The development of the limbs proceeds quickly, as the tadpoles are now less de- licate than in the beginning, when they die easily for want of appropriate con- ditions, ‘The young frogs are small, but quite lively and show in both the species dis. tinetly palmated hindfeet. They climb easily on inclined flags and on. vertical glasswalls, After the resorption of the tail which ensues quickly, they move in leaps, Their design and colouring is seen on plate 5. Failing direct observations of the evolution, all this stages may be collect- ed on inclined flags and slabs of stones in the small torrential streams where the adults are heard calling. The only other frogs found under similar condi- tions are Hylodes miliaris and petropoli- tanus but those are easily distinguished. TAXONOMY OF THE GENUS CYCLO- RHAMPHUS. The characteristics of the genus ha- ving been given above, we may dis- tinguish the different species of Cy- clorhamphus. Of such I found in my ma- terial one new and five already known ones. Our knowledge is still rudimentary, as the brasilian mountains (where this genus may be found) have not been suf ficiently explored and only by a stroke of luck or by special ability all the spe- cies existing in a given locality may be obtained. Thus we may expect that in time the distribution of each species may be ascertained and some new ones may also appear. All the species show a good deal of individual variation in skinstructure, co- lor and marking. Each specimen may change, not in a few hours as many spe- cies of Hyla, but during evolution, young ones being more variegated and oid ones more uniformly colored. The horny tips of the warts are not constant and the glandu- lar tubercules may vary in number and de- velopment. There may be in this some relation with the time of propagation, as in other batrachians, As a rule the de-—— a —— termination should be made from adults | not represented. He gave many drawings which unfortunately do not always re- | and photographs of the preserved mate- Present the largest number. However | rial, some specimens of which I haye half of the species are easily recognised | heen able to examine. In my own ma- and only those with complete interdi- | terial there are six species, five of which gital membranes may offer difficulties. | I saw in life. The aquarels and other MIRANDA-RIBEIRO made two pu- | ilustrations of the present paper were blications on the material collected in~| taken from living or recently preserved the museum of Sdo Paulo by its former | SPecimens. director and therefore already preserved | I give here first a key for the deter- for quite a time, The specimens, though | mination of the specics and the localities numerous, came from few localities and inhabited and afterwards a discussion the surroundings of Rio de Janciro where | of each of them. KEY FOR THE DETERMINATION OF FULLGROWN SPECIMENS OF CYCLORHAMPHUS. 1. Hind feet palmate. 22 Hind feet not palmate 5 2. Interdigital membrane shortened . asper WERNER. The same well developed... 5ae 3. Tooth in the midst of mandibula single 14 The same with two or three points. f . faliginosus 4. Back densely covered with small round warts... . Boulengeri Back with scattered grandular warts ore only with short and small wrinkles. . . Pinderi 5. Back densely covered with larger granules. .... granulosus Skin smooth .. Pa ee . eleutherodactylus HABITAT AND SYNONYMS OF THE (Serra de Cubatéo (Museu Paulista; ADULTS. LUTZ); Santa Catharina (WERNER, | L. MUELLER); Parané (Ypiranga DU- \-ANDERSSON). Cyclorhamphus pinderi_ M-R. 1. Cyclorhkamphus fulig nosus (TSCHUDI) 1838. Telmatobius | SE! brasiliensis STEIND, 1864, Grypiscus um- brinus COPE 1867. Pithecopsis BIBRON | 1920. 1858.—Brazil, leg. DELALANDE; Rio de Mha de S. Sebastiio (Type), Serra Janeiro leg. AGASSIZ; mountains of Rio | de Macahé (Cotype); Serra de Petro- de Janeiro and lower part of the moun- | polis at 800 m, Serra de Macahé near tains of Petropolis (LUTZ). to Nova Friburgo (LUTZ). 2. Cyclorhamphus Boulengeri LUTZ= 5. Cyclorhamphus eleutherodactylus asper BOULENGER 1907). MIR-RIB, tate Santa tharina (BOULEN- S, 5 oe Pane penta ee = » Seta) les» Cubiathoy GRUss Maule as e LUTZ); mountains near Rio and Serra da 3. Cyclorhamphus asper_ WERNER . 1899. Telmatobius Duseni ANDERSSON | Bocaina (LUTZ). 1914, Miodocus dubius et semipalmatus 6. Cyclorhamphus granulosus LUTZ, MIR-RIB, Bonito in the Serra da Bocaina (LUTZ).—— 22 —— DISCUSSION OF THE SPECIES. 1, Cyclorhamphus fuliginosus PLA, figs. 1&2 pl. 2, figs. 1 & 5; pl. 5 figs. 45. L use this name for the only species with palmate fect found in the mour tains near Rio de J iro. It is rather abundant and could quite eas'ly fall in the hands of DELALANDE, ASSIZ and NATTERER while making the clas: sic excursions to the Tijuea and the Cor- covado. DELALANDE brought the lwo specimens mentioned in DUMERIL and BIBRON, as being the types of TSCHU- DI. The description is not sufficient for determination, but the term fuliginosu applies very well and there is a draw- ing in the atlas showing a double tooth in the mandible. Grypiscus is charac- tevised by locality and by the double mandibular tooth, Also the name um- brinus and the photograph of the type RO reproduced in MIRANDA - RIB! (1926) agree well. (Telmatobius) brasi- liensis STEIND, was represented by a very young individual of only 28 mm. in length; however the drawing shows the palmated feet very plainly. The deserip- tion of brasiliens’s given by M. R not fit fuligincsus, bul refers partly to G. asper and partly to boulengeri, as shown by the features and the habitat given. He docs not seem to haye known the true fuliginosus, found only near Rio, Young specimens of juliginosus are much marked with ochraceous, as sveu in the figures on plates 1 and 5. Laler on, the upper side may become altoge- ther blackish with the exception of a constant lighter spot between the eyes and some larger whitish glandular warts, The under side may also become strongly pigmented, principally the gula, though this moslly shows very characte- rislical whitish dots. The color is either sooty or dark chocolate, The skin is rarely quite smooth; docs | in older specimens, preserved for some time, it mostly looks like shagreen, due to very short and aggregated wrinkles which might be mistaken for the round gra- nules existing in’ other species, ‘They hever show corneous tips. The ventral side is smooth or finely granulated. The larger specimens always show in the middle of the mandible a gsoup of two or three approximated and point ed teeth. They are typical for the spe- cies but appear rather late The palma- tion of the hind legs is well developed and the meniscus very distinct, 2. Cyclorhamphus pinderi MIR.-RIB. Pl. 1, fig. 35 pl. 2} fig. 25 pl. 4, figs. 1 & 2; ph. 5, figs. 1=3,'7 & Sl. The name pinderi was given to a specimen of the Museu Paulista collected by PINDER on the mointainous isle of Sebastifio which may be considered of the coast-range of the state lo, The channel between it and the mainland is neither deep nor large. ‘The description of the species is not sufficient but there is a figure. I have not compared the type but know a co type belonging to the same museum and collected in the Serra de Macahé. I have reasons to include in the same species numerous adults and spawnmasses, col- lected in the Serra de Petropolis at an 2 ation of 800 m. Pinderi comes near to fuliginosus but shows. less ochraceous pigment in va- rious stages of evolution, It may altain the length of nearly 7 cm. The skin is rarely smooth, generally it is roughened by groups of short and thin wrinkles, appearing like warts, principally on the upper eyelid, This structure may be more developed than in fuliginosus. There are light glandular warts with round or ellip- tical section, scattered over the whole surface and appearing distinctly in the inguinal disks. Between the eyes there is a light spot of varying shape. There also may appear lighter oblique bars onthe hindlegs. On the ventral side the black pigmentation appears generally lighter and less intense as in /uliginosus, but some larger specimens may show a sooty coloring on the whole of the ventral. side and principally on the throat. The meniscus is smaller and the dark black rim of the upper eyelid larger. The web of the toes is also at least as well developed as in the first and third species. There is never more than one mandibular tooth, even in the largest specimens, The aquarelle shows a specimen from the Serra de Macahé of 68 mm. in length with a smoother and lighter skin than usually observed, but the cotype of the. museum in io Paulo (which came from the same region) has a puck- red and almost completely black skin, However the corneous points seen in C. asper W R and boulengeri LUTZ never occur, 8. Cyclorhamphus boulengeri n. n. (Telmatobius asper BOULENGER). Comp.: Pl. 3, figs. 1 & 2 and copy of original deserip- tion in the appendix. alveady stated by BOULENGER, his species is different from Telmato- bius Duseni which corresponds to Cycto- rhamphus asper WERNER. It is much darker and has longer membranes be- tween the toes. m Pinderi it differs by larger warts which are rounded and crowded on the back, sometimes with forneous tips, like in Cyclorhamphus asper WERNER. His name has priority and I shall use the name C. boulengeri for the newer species. I have two quite good preserved spe- cimens, one with 30, the other with 37 mm. length from the tip of the snout to the vent. Plate 3 shows photographs ta- ken from an aquarelle of the second which has smail inguinal disks and a few horny tips. Lately I received also a living specimen, 38 mm. in length, alike to the others but with many spots of —— Be light chrome-yellow scattered over the body and more crowded on the sides and the posterior aspect of the thighs. They cor ond to glands and the lar- ger ones are elevated. On the venter and the gula they are smaller and flat- ter. The iris, metallic grey in color, supports a large meniscus at its upper pupillary margin which may be tilted backwards. It is strewn with glittering points while its ground is hardly percep- tible on the black pupillary field. The under pupillary margin has a very short median incision. Between the eyes there is an angular bar; some other narcow and yertical ones are seen on the upper lip and some others are broad and ath- wart to the thighs and legs. They are all of a light brownish ochraceous colour. The wellmarked disks and the ventral skin, as well as the lower side of the thighs, are translucent and look gelatinous. The back is sooty black, while the under sidé shows. a diffuse dark pigmentation. Abo- ve the skin is is covered with glandular warts, partly with white centers; below the glands are flat and quite small. There are no horny tips to the warts, probably because they are nupcial cha- racters. In this specimen, examined soon afterwards, the testicles where white and only moderately developed. The length in all my specimens is much below the measure given by BOULENGER but 1 believe that the males are smaller than the females. All my specimens came from the mountains of So Bento, the locality being different but situated in the same state S. Catharina as the habitat, given by BOULENGER. 4, Cyclorhamphus asper WERNER. Comp. PI 1, fig. 4, also the copies of WERNER’s and ANDERSON’s descriptions in the appet Of this species I have more than 50 specimens which may be recognised at first sight, They were all caught near the station Alto da Serra of the so cal-Seba fed Serra de Cubatio, The specimens in the museum of Sao Paulo came from the same reg’on while those of WERNER. and ANDERSSON are from the stctes of Parana and Santa Catharina In this species the interdigital mem- brane is somewhat variable but alwa distinclly shorter than in the other sp cies, (Pl. 2, fig. 3). The length from mouth to vent does not exceed 47 to 48 | mm. in my largest specimens; the others are considerably smaller. The prevalent pigment is ochraceous which forms the ground color. Black, when present on the back of older individuals, lies on top of it Generally the back has the co lor of mixtures of coffee with rather Iitle milk or diluted brown sepia, with or without darker spots. Fig. 4 on Pl. 1 shows a male of regular size but rather darker color, as seen in most specimens Dhere appears also the constant inter- ocular light spot and another on the back, qu’te frequently seen but occurring also in other species, The lighter oblique bands on the hind limbs are well shown The ochraceous skin is covered with tubercules occupying more space as the interstices. They may form some- what twisted rows. Their centre is either whitish hyaline or capped w'th brownish horny points, like those common in the genus Bufo. The ventral side is distinguished by a Y'ght color, altogether whitish or slight- ly vermiculated with light brown. 5. Gyclorhamphus eleutherodactylus (MIR.- RIB) Pl. 1, figs. 5-65 pl. 2, fig. 4; pl. 3, fig. 5, This species, represented by 41 spe- cimens from the Serra de Gubatio, was described under the generic name Ilio. discus, and differs from other species by having a’l the tocs free, The skin is either quite smooth or shows only a few granulations, widely scattered over the dorsum. The under side is strongly pigmented like in fuliginosus. This species appears in two rather different forms, ‘The first which might be ca'led forma variegata shows various, rather complicated markings. I have two adult and two young specimens from the Tijuca and the Serra da Bocaina. The f. 6, pl. 1 gives a very variegated speci- men while the one of f. 5 is little diffe- rent from the next form, Both are from: the Serra da Bocai Of the other form 1 have two specimens ‘Pl. 3, f. 5), one from Allo da Serra, the other, given by Dr. VELLARD, from the same region but from another place near the new auto: mobile road to Santos. The dorsum. is uniformly dark—brown like cinnamon- bark or the pod of Hymenaea courbaril, but marked wh'th small whitish or yel- lowisk streaks. There are round spots on the smaller articulations, the sides of the trunk and the under side of the tarsus, also in some other places. Gene- rally there length is not more than 5 mm. The ventral side is slightly infus- cated with oval light spots. This ra- ther si ing form might be called stri- gillata. M-R. gives pictures of a similar though much darker specimen. He in- dicates for this species a length of 50 mm., but my specimens hardly exceed 40. The species seems smaller and more delicate than the others. I dont think that the two forms have species value, as they may occur in the same place and the above mentioned species all show variations, sometimes of a simi- lar type. 6, Cyclorhampkus granulosus n. sp. Pi. 3, figs. 3 & 4. In my material from the Serra da Bocaina there are two more specimens, 44 and 27 mm. in length, which cannot enter in the species cleutherodaclylus M-R., though their toes are also quite Tree, Here however the skin of the back is altogether covered with submiliary,_ granulations, as in C. asper WERNER, but they are neither conical, nor capped with horny points, Seen from above their calibre is nearly equal, but the outlines of their bases are irregular. They look rather like shagrcen leather or the EXPLANATION PLATE I. | Figs. 1 & 2: Cyclorhamphus fuliginosus, not quite full sized, 1/1. Fig. 3: C. pinderi. Full sized male, 3/4. Fig. 4: C. asper WERNER. Fw! sized, rather dark male, 1/1. Figs. 5 & 6: C. eleutherodactylus show- ing varietions in markings, 1/1. PLATE II Fig. 1: Hind foot of C. fuliginosus, und- er s'de; 2: the same of C. pinderi; ‘J 3: the same of C. aspcr WERNER; 4: the same of C, eleutherodacty. lus. All the figures in nat. size. 5: Mouth of C. fuligirosus showing mandibular teeth. 1/1. PLATE IID Figs. 1 & 2: GC. boulengeri, small speci- men. 1/1. Figs. 3 & 4: C. granulosus n. sp. 1/1. Fig. 5: C. eleutherodactylus, forma stri- gillata. 1/1 pala oY shell of the pods of Hymenaea courba- ril. The color is dark chocolate and there is no sign of any larger markings. The ventral side is somewhat lighter and less rough OF PLATES PLATE IV Fig. 1: Spawn of . pinderi with un- developed eggs. 1/1. Fig. 2: The same after formation of the embryo, PLATE V Fig. 1: Egg with embryo of C. pinderi, upper side. 2/1, Del. PUGAS. Figs. 2 & 5: Metamorphosis of C. fuli- ginosus. 1/1, Del. SANDIG, Fig. 6: C. fuliginosus, quite small. 1/1. Del. SANDIG. Fig. 7: Tadpoles of C. pinderi, bred trom spawn. 1/1. Phot. PINTO. Figs. 1 & 11: Larger tadpole and meta- morphosis. 1/1. Del. PUGAS. REFERENCES A list of litterature is given at the end of the portuguese text, also a copy of some original descriptions.