Bijdragen tot de Dierkunde, 49 (2): 261-312 1979

Biosystematics of the Malagasy frogs.

II. The genus Boophis (Rhacophoridae)

by

R.M.A. Blommers-Schlösser

Institute of Taxonomic Zoology (Zodlogisch Museum), University of Amsterdam, The Netherlands

Abstract were always found in water), so characteristic of

Asiatic Rhacophorus, supports the opinion of the

This the 1938 = paper concerns genus Boophis Tschudi, ( latter authors. Consequently, I prefer also to use Malagasy Rhacophorus Kuhl & Van Hasselt, 1822). Apart the for the name Boophis Malagasy species. from a reconsidering of adult morphology, certain new colouration in the In earlier of the taxonomic characters, such as the life, an paper on the biosystematics morphology of the tadpole and some mating calls, are Malagasy Mantellinae (Blommers-Schlosser, 1979) presented. Karyotypes and nuclear DNA amount were re- it has been that is not elsewhere. argued already Boophis so ported upon

" ~ definition of endemic This study produces a sharper the closely related to Mantidactylus'' Boulenger, 1895, its clear from the Manti- genus, permitting separation genus as was presumed by previous workers. The main dactylus Boulenger, 1895. was the difference in reproductive Various sibling species are reported; as a result different argument great synonymies are revised and new species presented. Four new behaviour. Earlier taxonomic studies concerning B. B. are described, viz. Boophis mandraka, viridis, species the based genus Boophis Tschudi, 1838, were for reticulatus and B. hillenii. A lectotype is designated entirely on material (Ahl, 1931, and Rhacophorus majori Boulenger, 1896. preserved

The bonae and new following species, new synonyms Guibe, 1978). combinations are proposed: Boophis erythrodactylus (Guibe, The author and her husband collected 21 species comb. 1953), nov. (from Hyperolius); Boophis difficilis of this (about of those in (Boettger, 1892), bona species; Boophis miniatus (Mocquard, genus 80% known) 1902), bona species; Boophis goudoti Tschudi, 1938 (syn. Madagascar between November 1970 and May Rhacophorus fasciolatus Ahl, 1929 and Rhacophorus kanbergi 1973. The natural history, the larval morphology, Ahl, 1929); Boophis untersteini (Ahl, 1928), bona species; the and the in comb. mating call colouration life were Boophis pauliani (Guibe, 1953), nov. (from Hypero-

comb. nov. in lius) Boophis microtympanum (Boettger, 1881), investigated many species. A combination of (from Rhacophorus boettgeri Boulenger, 1882: substitute these new "field" taxonomic characters and the 1881). name for Hylambates microtympanum Boettger, morphology of the species are sometimes essential

for identification, because clear morphological

INTRODUCTION characters separating sibling species are often

missing.

Tschudi for A careful of the (1838) created the genus Boophis a study Malagasy Boophis single Malagasy species, Boophis goudoti. The Tschudi, 1838, has demonstrated the impractica-

has been considered of of a dichotomous to all genus Boophis a synonym bility attempting key Rhacophorus Kuhl & Van Hasselt, 1822, by most species. The earlier keys of Ahl, 1931, and Guibe, authors not useful for identification of all (among others: Boulenger, 1882, Ahl, 1978, are very

characters used 1931, and Guibe, 1978). species, since the might include

Laurent (1943) noted differences in the osteo- some features of several closely allied species as logical characters between the Asiatic and the well.

I Malagasy Rhacophorus species; so he proposed to Therefore have attempted to assemble the

the into several natural based reserve the name Boophis for latter. Liem species groups, on affin-

(1970), studying mainly osteological and muscular ity in structural characters, often together with

with Laurent's I then characters, agreed opinion. colour similarities. have presented a diag-

Our failure to observe foamy nests of eggs (eggs nosis for the species within each group. 262 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

The B. Summarizing: Measurements and characters of both tad- following groups are recognized: rap- poles and adults and the terminology concerning the sona- luteus rhodoscelis piodes group, group, group, in accordance with Duellman's In de- grams are (1970). goudoti group, tephraeomystax group, opisthodon scribing the larvae, I have defined the stages according to

A Gosner formula: the numbers 1, 2i, 2e group and pauliani group. remaining group (1960). Webbing refer to the first toe (or finger), the inner side of the which their in consists of species have occurrence second, the outer side of the second and the numbers in high mountains common. refer number of free web. in parentheses to the phalanges of The following species were neither studied nor Tooth formula: the tooth rows are numbered from top to both and The bottom in the upper lower lip. separation in of the B. leucomaculatus included any groups: between both lips is indicated by a double slant line. A B. microtis B. (Guibe, 1975), (Guibe, 1974), marked a median gap between the lateral tooth rows is by brygooi (Guibe, 1974), and B. laurenti Guibe, plus sign. The tooth formula is given from stage 28 onwards of (complete). The bioclimatic areas correspond to those 1947. Koechlin (1972). Lotic means living in brook or river. Lentic,

in living swamp, pool or pond.

Since this after the of paper comes out shortly publication

Guibe "Les Batraciens de in which a ACKNOWLEDGEMENTS (1978) Madagascar", description is given of all frog species occurring in Mada-

I detailed all gascar, have not given a very description of The Foundation for investigations were supported by the Boophis species. The study in the field revealed that there Fundamental Research which is sub- Biological (B.I.O.N.), Guibe estimated are more species than (1978) on the base of sidized by the Netherlands Organization for the Advance- alcohol material only; it also appeared that the juvenile of ment of Pure Research (Z.W.O.). the one resembles much the adult of the other, resulting in

Material was made available Mrs. Dr. R. Roux-Esteve by of Therefore I have a changing synonymy. given a very and Prof. Dr. J. Guibe, Museum National d'Histoire Natu- detailed description of juveniles of several species, of sibling relle, Paris (MNHP), Prof. Dr. G. Peters, Zoologisches removed and of species, species from previous synonymy Museum, Berlin (ZMB), Mr. A. F. Stimson and Mrs. A. G. course of the new species. C. Grandison, British Museum (Natural History), London The description of other species is concise, and placed (BMNH), Prof. Dr. K. Klemmer, Naturmuseum und For- identification mainly within the group description, making schungsinstitut Senckenberg, Frankfurt (SMF), and Dr. F. possible. Tiedemann, Naturhistorisches Museum, Wien The (WM). four included in the The species not present paper (see author is much indebted for these loans. very known from their introduction) are not common and only I Dr. M. S. for am very grateful to Hoogmoed (Leiden) type localities (see Guibe, 1978). examining the holotype of Rhacophorus femoralis Bcmlenger,

1882, in London.

I wish to thank in particular Prof. Dr. J. Guibe, who Genus to before Boophis Tschudi, 1838 allowed me read his manuscript it was published.

I am very grateful to Mr. Andriamampianina and his staff

d'Eaux their Guibe, (Dept. et Foret, Repoblika Malagasy) for per- Boophis Tschudi, 1838: 36, 76; Laurent, 1943: 13; mission and in the to stay work some of most beautiful 1947: 438; Guibe, 1978: 59 (in synonymy of Rhaco- forests of Madagascar. phorus); Bachmann & Blommers-Schlosser, 1975: 17

I express my gratitude to Dr. D. Hillenius (Amsterdam), (nuclear DNA amount); Blommers-Schlosser, 1978: 30-37 for encouragement during this study, to Prof. Dr. J. H. Stock (chromosomes). (Amsterdam) and Prof. Dr. J. T. Wiebes (Leiden) for Rhacophorus Kuhl & Van Hasselt, 1822: 104 (part.).

to critically reading my manuscript, Mr. W. J. van der Wey- Type species: Boophis goudoti Tschudi, 1838: 77, by mono- den for the (Free University, Amsterdam) making sonagrams typy. of the tape recordings, to Mr. L. A. van der Laan (Amster- dam) for making the photographic prints and to Mr. A. Definition. — Vertebral column procoel. Ameling (Amsterdam) for typing a part of the manuscript. Second tarsal not free, but connected with third

and fourth tarsal. Omosternum with bony style, MATERIAL AND METHODS with not or only slightly forked at base. Sternum

long bony style, not forked. Length of style of The material is deposited in the Zoologisch Museum Am- sternum than that of omosternum. Maxil- sterdam (ZMA). The ZMA registration numbers are given greater under number each species with the of specimens in paren- lary teeth present. Vomerine teeth mostly present. theses. Tongue distinctly bifid and free behind. Pupil The definitions of measurements and characters of both horizontal. Intercalary cartilage between tadpoles and adults and the terminology concerning the sona- present

and the bioclimatic areas in are the same disc and grams Madagascar adjacent phalanx. Digital pads completely as used previously (Blommers-Schlosser, 1979). For certain surrounded by ventro-marginal groove. Fingers localities the distance to Tananarive (in km) alongthe various with small trace of web to three-fourths webbed. highroads is provided. a 49 - BIJDRAGEN TOT DE DIERKUNDE, (2) 1979 263

TABLE I

the Key to groups.

123 4 56789 10

B. + — — — a rappiodes group a a a 1/3-1/2 a ± B. luteus baa 1/2 + b + — a group

B. rhodoscelis + + ± — a group a a+b a 1/3-1/2 a ± ± B. b a+b + a + a goudoti group a 2/5-3/4

— — b B. tephraeomystax group a+b b b 1/3-2/3 + a +

— B. b b b + + — b? opisthodon group a

— b? b — + + B. pauliani group a a <1/3 a

a — — a Montane species a+b b b <1/3 —

= b = 1. Snout-vent length of adults: a mm; JS35 mm.

= the of the snout b = not far 2. Tibiotarsal articulation: a reaching tip or beyond; reaching as

as the tip of the snout.

= webbed, web at least the base 3. Webbing of the fingers: a one-third or more the reaching of the penultimate phalanx of the fourth and the middle of the antepenultimate phalanx of the

third finger; b = trace of web, less than one-third webbed.

4. Size of in the ratio of the horizontal diameter of the that tympanum, expressed tympanum to

of the eye.

= = teeth: + — absent. 5. Vomerine present;

= b = 6. External subgular vocal sac: a single, median; paired, lateral.

= = 7. Transverse bars on the limbs: + present; — absent.

= — = absent. 8. Dermal fringe along the lateral edge of lower arm and tarsus: + present;

= heel: + = — absent. 9. Dermal flap on present;

10. Breeding site: a = streaming water; b = standing water.

Toes half to fully webbed. Outer metatarsal tuber- Vomerine teeth are present. The horizontal diam-

cle absent. Lateral metatarsalia separated. Males eter of the tympanum is one-third to the half of

than females. in horizontal diameter of the The vocal smaller Nuptial pads present the eye. sac

males. is single, median, subgular. The snout is short and

Amplexus axillary, sometimes lumbal. Eggs rounded. The commissure of the mouth is slightly

small, laid in water. Tadpoles in streams (lotic) curved upwards ((rappiodes, erythrodactylus, man-

viridis and in stagnant water (lentic). Chromosomes: 2n draka) or straight ( ). The tibiotarsal artic-

= 26; N.F. (number of chromosome arms) - ulation extends beyond the tip of the snout. The

7.0-9.9 discs well half 52. Nuclear DNA amount: pg (diploid). are developed. The fingers are

A the is in table between the third and fourth the key to species groups presented webbed; finger

I. The colouration is also characteristic in most webbing extends to the bases of the penultimate

included in the since it is three-fourths almost groups, but not key, phalanges. The toes are to

difficult to in the the discs of express symbols. entirely webbed, webbing reaching

the first and fifth toe.

Contents: B. rappiodes (Ahl, 1928), B. man-

B. Boophis rappiodes group draka sp. nov., erythrodactylus (Guibe, 1953)

and B. viridis sp. nov.

— this Diagnosis. The members of group

dor- Differential — The colour are small-sized, stream-breeding species. The diagnosis.

is in in life is the best character to distinguish the sum bright green (yellowish preservative) and distinctive IV but much of it dis- the bones are green. There are no species (pi. figs. 1-3, 5),

the in alcohol. Therefore I have tried to find markings on the posterior surface of thighs appears

the limbs. Dermal other characters for each of the and no transverse bars on diagnostic species.

The B. viridis is than the other the fringes and flaps are lacking on the limbs. larger species, and the of skin on the dorsum is smooth or finely granular. nostrils are strongly protruding length 264 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

TABLE II

characters in the Diagnostic Boophis rappiodes group.

B. erythro-

B. viridis B. rappiodes dactylus B. mandraka

Snout-vent length of adult $ (mm) 29-31 24 19.5-25.0 21-26

Ratio hand length/body length (mean) 0.33 0.26 0.33 0.33

Ratio hor. diameter tympanum/eye (mean) 0.35 0.33 0.47 0.32

Length of snout compared with distance

between anterior of the shorter shorter shorter corners eye equal

of of Position nostrils in respect tip of nearer tip nearer tip nearer tip nearer eye

snout and anterior corner of eye of snout of snout of snout

Character of nostrils strongly slightly slightly slightly

protruding protruding protruding protruding

Colouration cf. text description species and pi. IV figs. 1-3, 5

I the snout is longest of all. B. rappiodes has much believe that the present couple belongs to this

smaller hands than the other B. It with the of species. erythro- species. agrees original description

dactylus has the largest tympanum. B. mandraka R. rappiodes. differs from the other three in the position of the which the than the table for nostrils, are nearer to eye to Description (see IV measure-

in — the tip of the snout. These characters are shown ments). The ratio of hand length to snout-

table II. vent length is 0.25-0.28. The ratio of the tibia

length to snout-vent length is 0.58-0.59. The ratio

Boophis rappiodes (Ahl, 1928) of the foot length to snout-vent length is 0.42-

Text-figs. 1-2; pi. IV fig. 1. 0.44. The ratio of the horizontal diameter of the

that of the is Snout- tympanum to eye 0.33-0.34. Rhacophorus rappiodes Ahl, 1928: 315; Guibe, 1978: 78 vent of male 24 of female 32 mm. (as dubious species, because of bad condition of the type). length mm,

The body is in the region equal to Material. — ZMA 7164 (1 9 and 1 $), 14.XI.72; postaxillary

7167 (eggs deposited by 7164), collected near Perinet (high- the head width. The head is wider than long. The

road R.N. 2 at km 142) alt. 900 m; ZMA 7165 (tadpoles), head width is about one-third of the body length. 24.IX.72 at the same locality, but alt. 1100 m. — ZMA 7168 is rounded in lateral reared from collected The snout bluntly dorsal and (tadpoles and young them), IV-VIII.72 in the Mandraka valley (highroad R.N. 2 at km 69); ZMA profile. The snout is equal to the horizontal di- but km 7161 24.IX.72 at the same locality, at 70, (tadpoles), of the ameter eye, and shorter than the distance

1100 — collected alt. m. ZMA 7166 (tadpoles), 29.VIII.72

between the anterior corners of the The along the road from Moramanga to Anosibe at km 27. eye.

nostrils are much nearer to the tip of the snout, than the anterior of The — the canthus Natural history. The couple was to corner eye.

is found in axillary amplexus, in the shrubbery along distinct and concave. The loreal region is slightly

interorbital is a stream in woodland, during a rainy night. One concave. The region slightly convex.

the female 260 The internarial distance is smaller than day later, deposited glutinous eggs slightly

in (2 mm diameter) of a light green colour in the the interorbital distance. The interorbital distance

The is about times the width of vivarium. eggs dried up quickly on land, 11/2 an upper eyelid.

the is directed which might indicate that normal oviposition oc- The nostrils are protruding, opening

in the commissure the mouth is curs in water. The tadpoles were collected laterally. The of curved

still where is parts of streams, abundant vegetation upwards. The pupil horizontally elliptical.

the is on side shaded the water. There a straight weak dermal fold over the

tympanum, reaching a point behind the insertion

Taxonomy. — Although the holotype of R. of the arm. The fold covers the dorsal margin of rappiodes is in a bad condition (Guibe, 1978), the tympanum. The tympanum is rather indistinct, BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 265

its hori2ontal diameter is one-third of the hori- The dorsum is covered with minute dark spots.

the diameter of the The distance from the A white line runs from the nostril over zontal eye. eyelid

the is about three-fourths of the and the and ends behind the insertion eye to tympanum tympanum

The horizontal diameterof the tympanum. of the arm. The upper eyelids are creamish.

diameter the disc the surrounded The largest of of third iris is silvery grey with brown pigment,

is the diameter of the black The rim of the finger equal to horizontal by a ring. upper palpebral

and is The lower is tympanum, the discs of the fourth second membrane slightly pigmented. rim

and first and in the finger are about equal of the smaller. pigmented anteriorly posteriorly same

The relative lengths of the fingers from shortest colour as the skin.

colour is with red to longest are 1-2-4-3. The subarticular tubercles The in life yellow-green

round. is IV The are small, distinct and There an inner pigment (pi. fig. 1). white lines (in flat oval metacarpal tubercle. A small unpig- alcohol) are yellow. In both male and female much

and mented, glandular nuptial pad is present dorso- red pigment is present on the eyelids around

of male. The the nostrils. The red is in on the first finger the pigment present irregular

of the hand: 2i and the formula webbing of the 1 ( 1), ( 1), spots markings on whole dorsal surface,

the 2e (1), 3i (2), 3e (1), 4 (1). The web is glan- more extensively in the female than in male.

is The dular. The belly uniformly coloured. bones are

The and slender. limbs arms legs are When the green.

are laid along the side, the knee and elbow overlap slightly. The adpressed heels meet. The tibiotarsal Tadpoles. — Thirteen tadpoles in stage 25 articulation extends just beyond the tip of the have body lengths of 5-9 mm and total lengths snout. The inner metatarsal tubercle is small, of 15-27 mm. Twenty tadpoles in stages 26-30

and elliptical. A small web connects the tip of the inner have body lengths of 8.5-13.0 mm total

first middle Eleven in metatarsal tubercle with the toe to the lengths of 24-37 mm. tadpoles stages of the antepenultimate phalanx. There is no outer 31-41 have body lengths of 9-13 mm and total

of The ratio of the metatarsal tubercle. The discs on the toes are lengths 28-36 mm. average

in 44 equal to those on the fingers. The subarticular tail length to the body length the tadpoles tubercles small and round. are The fifth toe is listed above is 1.9 (range 1.75-2.10). Six meta-

the third. of the from slightly longer than The formula morphosing young measure 11.5-14.0 mm webbing of the foot: 1 (0), 2i (0), 2e (0), 3i (0), snout to vent (mean 12 mm).

3e (0), 4i (1), 4e (1), 5 (0). The ventral surface The mouth and lateral view of a tadpole in

in of the web is glandular. The lateral metatarsalia stage 31 (ZMA 7165) are shown text-figs. 1-2. are separated. The body is depressed. The nostrils are dorsal, The anal opening is directed posteriorly at the the opening is anterolaterally, situated slightly

the anterior of the than to upper level of the thighs. The anal sheath is short. nearer to corners eyes

The is and bifid the of the snout. The small and tongue heart-shaped distinctly tip eyes are

The situated directed The is sinis- posteriorly. vomerine teeth are me- laterally. spiracular opening dially, behind the choanae in two small straight tral, situated at about 5/8 from the tip of the

The vocal slits situated in the snout and the end of the The cloacal tube is groups. are small, body.

the wide The caudal floor of mouth, near the angles of the jaws. and dextral to the caudal fin.

The vocal sac is single medially, subgular and musculature is robust, especially in the first half moderately distensible. of the tail. The dorsal caudal fin is reduced

skin dorsum is The on the smooth. The skin on anteriorly. At midlength of the tail, the height of

the caudal musculature of the the throat is finely granular. The skin on the belly, represents about 2/5 and proximal posteroventral surfaces of the thighs total tail height. The mouth is small and directed and the anal is elsewhere has wide below opening granular; ventrally. The papillary border a median

the the skin is smooth. There on venter gap anteriorly. are 1-2 rows of papillae.

The colour in preservative is light yellowish. The oral disc is indented laterally and medio- 266 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

ZMA Figs. 1-7. 1-2, Boophis rappiodes: 1, mouth of tadpole, ZMA 7165.X.25; 2, tadpole, 7165.X.3; 3-4, B. mandraka:

3, mouth of tadpole, ZMA 7121.X.12.5; 4, tadpole, ZMA 7121.X.3; 5-6, B. erythrodactylus: 5, mouth of tadpole, ZMA

7208.X.12.5; 6, tadpole, ZMA 7208.X.3; 7, B. luteus: mouth of tadpole, form A, ZMA 7005.X.12.5. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 267 posteriorly. The tooth formula isl/3 + 3//l + l/2 length is 0.30-0.36 (mean 0.33). The ratio of tibia

is well is 0.54-0.59 or 1/2-f 2//1 + 1/2. The horny beak length to snout-vent length (mean developed. 0.57). The ratio of foot length to snout-vent The colour in life of the tadpole is blackish, length is 0.43-0.55 (mean 0.47). The ratio of the

with the to that of covered small bright green spots. horizontal diameter of tympanum

the is Snout-vent Metamorphosis occurred in November (begin- eye 0.29-0.34 (mean 0.32). ning of hot season). length in males 21-26 mm.

is 6.8 head width is Holotype: head length mm;

— colour of is Young. The the living young internarial distance is inter- 8.1 mm; 2.8 mm; is nearly identical to that of the adults. The ground orbital distance is 3 mm; snout-vent length is colour is and thebones are The green green. yellow 23.5 mm. line at each side of the head is already present. The head is wider than the body, and somewhat is the and Red pigment on eyelids between present wider than long. The head width is one-third of the The back is covered with blackish eyes. the body length. The snout is rounded in dorsal nostrils the speckles. The are nearer to the tip of snout is and lateral profile. The slightly longer snout than to the anterior of the The corners eye. than the horizontal diameter of the and eye, are assigned readily to B. rappiodes. young distinctly shorter than the distance between the

nostrils anterior corners of the eye. The are nearer

the anterior of the than to the mandraka to corners eye tip Boophis sp. nov. of the snout. The distance from the nostril to the Text-figs. 3, 4, 8, 41; pi. IV fig. 2. anterior corner of the eye is distinctly shorter than

Boophis aff. ; horizontal diameter of the The canthus is erythrodactylus Blommers-Schlosser, 1978: 35 the eye. (chromosomes). distinct and rounded. The loreal region is slightly

Material. — ZMA 7119A, male holotype and ZMA concave. The interorbital region is flat. The inter-

7119B, six male paratypes 4.III.73; ZMA 7118 six male narial distance is slightly shorter than the inter- collected Mandraka paratypes, 27.111.73; all in the valley orbital distance. The interorbital distance is about (highroad R.N. 2 at km 67), alt. 1200 m.

ZMA 7120 (tadpoles and reared from them); ZMA times the width of The young ll/ 2 an upper eyelid. 7121 (one tadpole); all collected from June till October 1972 nostrils are slightly protruding, the opening is in the rivulet Vokanatetezandava, in the nursery garden of laterodorsally. The commissure of the mouth is the city of Tananarive in the Mandraka valley (highroad

R.N. 2 at km 69), alt. 1200 m. curved upwards. The pupil is horizontally elliptical.

is There a dermal fold from the posterior edge The present species is closiest allied to the sibling of the eye, over the tympanum, curving down- species B. erythrodactylus (Guibe, 1953). It was wards to a point just behind the shoulder. The discovered in the field that the mating call and the fold covers the dorsal margin of the tympanum. larva are clearly different in both. B. mandraka The tympanum is distinct and round. The hori- can be distinguished from erythrodactylus by its of zontal diameter the tympanum is about one- difference in colouration (see text description and third of the horizontal diameter of The the eye. pi. IV figs. 2-3), by the smaller size of the tym- distance from the eye to the tympanum is equal to and the which is panum, the position of nostrils, the horizontal diameter of the tympanum. nearer the than the of the to eye to tip snout. The diameter of the disc of the third finger is

the larger than the diameter of the tympanum, Natural history. — The males are calling discs of the fourth and second finger are slightly from shrubbery alongside rapids, on rainy nights. smaller, and the disc of the first is much The in finger tadpoles are clearly lotic and found rapids. smaller. The relative of the from those lengths fingers The adults and tadpoles occur mingled with shortest to longest are 1-2-4-3. The subarticular of B. erythodactylus (see B. erythrodactylus). tubercles are small, round and flat. There is an

IV for flat There Description (see table measure- inner, oval, metacarpal tubercle. is a

small ments). — The ratio of hand length to snout-vent unpigmented, glandular nuptial pad, dorso- 268 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

first is laterally on the finger. Supernumerary tu- green. The bones are green. The iris silvery

bercles are present on the base of the fingers. The white with a brown reticulation.

webbed. fingers are about half The formula of the The colour in preservative is yellowish. The

the hand: 2i and the in webbing of 1 (1), (1), 2e (1), brown spots are still visible yellow line

3e 4 The ventral surface of life has become whitish. The rim of the 3i (2), (1), (1-1 y2 ). upper

the web is glandular. palpebral membrane is pigmented. The lower rim

slender. limbs in colour the skin The arms and legs are When the is broadly pigmented the same as

knee and the dorsum. The iris is with are laid along the side, the elbow overlap. of silvery grey brown

The adpressed heels meet. The tibiotarsal articula- reticulation, surrounded by a black ring.

tion extends beyond the tip of the snout. The inner

is and with — The metatarsal tubercle distinct, small, elliptical Comparison paratypes.

There is holo- protruding. no outer metatarsal tubercle. paratypes agree with the description of the

A small web connects the tip of the inner meta- type. The distal tubercles of the fingers are often

first those third and tarsal tubercle with the toe till the base of the bifid, especially on the fourth

In penultimate phalanx. The terminal discs are some- finger. some specimens there are a few large

than dorsum what smaller those on the fingers. The fifth yellow spots on the (in life). toe is slightly longer than the third. The sub-

tubercles Voice. — The articular are small, flat and round. The males were heard on an evening formula of the of the foot: 1 2i in March. The made of male webbing (0), (0), sonagram (fig. 8) a

2e 3i 3e 4i 4e 5 4 March at 12 shows that (0), (1), (l/ 2 -0), (1), (1), (0). (ZMA 7119) on p.m.

The lateral metatarsalia are separated. The ventral the duration of one call is about 0.75 sec. One call surface of the web is glandular. The ventral surface consists of three loud melodious clicks followed of the foot is soft click. The dominant is granular. by a very frequency The anal opening is directed posteriorly at about 3500 Hz. upper level of the thighs. The dermal flap above

in — the anal opening continues a fold on each side Etymology. B. mandraka derives its name of in the anal opening, together forming a A-shaped from the river flowing the Mandraka valley. figure.

The is tongue heart-shaped and distinctly bifid Tadpoles. -— Fourteen tadpoles in stage 25 posteriorly. The vomerine teeth are situated me- have body lengths of 8.5-11.0 mm and total

in dially, far behind the choanae, two small oblique lengths of 26.2-34.5 mm. Four tadpoles in stages

The vocal slits in the groups. are small, situated 27-28 have body lengths of 10.5-12.0 mm and floor of the mouth, near the angles of the jaws. total lengths of 31.5-35.0 mm. Four tadpoles in

vocal is The sac median, subgular and moderately stages 33-37 have body lengths of 11.0-12.5 mm

and in distensible. total lengths of 33-35 mm. Four tadpoles

The skin 40-42 on the dorsum is shagreened, on the stages have body lengths of 12.5-14.0 mm throat and chin and the and total of 37-40 The ratio finely granular on belly lengths mm. average granular. The ventral surface of the limbs is of tail length to body length in the twenty-six smooth, except the ventral surface of the hands, tadpoles listed above is 2 (range 1.8-2.2). One feet and of the which is has of posterior part thighs, metamorphosing young a body length granular. The skin below the anus is coarsely 13 mm.

and in granular. The mouth lateral view of a tadpole stage

The colour in is shown in The life of the dorsum bright green, 34 (ZMA 7121) are text-figs. 3-4. with scattered brown spots (pi. IV fig. 2). The body is depressed. The nostrils are dorsal, the

brown. A upper eyelids are yellow with moderately opening is anterolaterally, situated at about 2/3

the wide yellow line running from the tip of the snout from the tip of snout and the anterior corner over the nostrils the anterior corner of the of the The are situated The to eye eye. eyes laterally. is bordered with situated brown spots. The venter is pale spiracular opening is sinistral, at about BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 269

Fig. 8. Sonagram, filter wide, Boophis mandraka.

2/3 from the tip of the snout and the end of the Boophis erythrodactylus (Guibé, 1953) body. The cloacal tube is long and dextral to the Text-figs. 5, 6, 9; pi. I figs. 1-2, pi. IV fig. 3. caudal fin. The caudal musculature is robust. The Hyperolius erythrodactylus Guibe, 1953: 102; Guibe, 1978: caudal fin is shallow and at anteriorly deepest 90. about 2/3 of its length. At midlength of the tail, Boophis erythrodactylus;; Blommers-Schlosser, 1978: 35 (chro- mosomes). the height of the caudal musculature represents

1/2 of the total tail height. The mouth is broad Material. — MNHP 53-171, four syntypes of Hypero- lius forest of erythrodactylus. Type locality: Mahajeby, near and directed ventrally. The papillary border has Morafenobe, West Madagascar. Coll. Paulian, 1952. a wide median There are 2-4 rows of gap. papillae ZMA 6924 (1 9 and 1 $), 18.11.73; 6925 (6 $$), laterally and 5-6 posteriorly. The tooth formula is 27.111.73; 6926 (2 $ $), 27.111.73; 6927 (1 S ), 26.XII.72; 6928 (4 3 3), 4.III.73; 6930 (1 $), 6.1.73; 7207 (tadpoles 5/2H-2//3. The upper row of the upper lip and and reared from young them), VI-X.72; 7208 (one tadpole), the lower row of the under consist of lip very VII.72; all collected in the Mandraka valley (highroad R.N.

2 at km alt. 1200 — small teeth. The beak is and 67-69), m. ZMA 7205 (tadpoles), upper horny narrow VIII.72; 7206 (one tadpole), 21.III.73; collected at Manja- moon-shaped. The lower horny beak is Y-shaped kotompo, forest station, Ankaratra mts. (near Ambatolampy), with heavy serrations. alt. 1800-2000 m. — ZMA 6929 (2 $ $ ), 13.XI.72 collected

in 2 The colour of the tadpole life is brown on near Perinet (highroad R.N. at km 142), alt 1100 m. the back; the belly is black in stage 25 and silvery Natural history. — The species was never in later The caudal musculature has grey stages. the noticed during day. The males were calling brown with large well-defined dark patterns a from leaves of shrubs and trees alongside rapids, creamish background. The caudal fin is transparent in the evening. The female (gravid) was also with a few dark brown markings. in captured such a place. In the Mandraka valley at The in tadpoles were collected rapids from June km 67 the species occurs together with B. man- till with the of B. October, together tadpoles ery- draka. They are collected in the same thicket, throdactylus. The rearing was only successful with while sitting at short distance from each other. far developed tadpoles. The calls mating of both are much alike, but could

— The is Brown Young. young frog green. be distinguished. When touched, both secrete a

is the and around the pigment present on eyelids strong smelling fluid, which serves probably also nostrils. There white are spots on the black. The for species recognition. The two species resemble bones The hindlimbs other but in are green. are relatively each in morphology, differ coloura- shorter than in the adult frog. tion. The tadpoles of both are clearly lotic, and 270 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

occur in the same torrents with stony bottom where largest diameter of the disc of the third finger

in the rivulet Vokana- the diameter of the Hydrostachys spp. grows: equals tympanum. of skin and chin is in the tetezandava, in the nursery garden of the city The on the throat smooth

tad- Tananarive, at km 69, Mandraka valley. The female, finely granular in the male (as in B.

in poles collected at Manjakotompo were found mandraka).

with The colour in life of the dorsum is rapids gravelly bottom. bright green,

covered with numerous red spots, which are sur-

thin rounded by a yellow circle (pi. IV fig. 3). with — Our Taxonomy. specimens agree There is a yellow line from the nostril to the the of type material Hyperolius erythrodactylus. anterior of the Another corner eye. straight yellow The is but description of Guibe, 1953, very brief, dorsolateral line extends from the posterior corner the typical red punctation is described and still of the over the where it deviates eye tympanum, Vomerine teeth in visible in the type. are present from the tympanic fold, which is curved, and and our material. Both sternum and the syntypes fades behind the shoulder. Two other yellow the of the omosternum have a bony style; style where parallel lines run from the level the former sternum is longer than the style of the omoster- ends, but nearer to the midline, over the entire num. Thus the species belongs to the genus with back. The upper eyelids are yellow, brown Boophis. spots. In some specimens there are also brown

shanks and around spots on the the nostrils. The

Description (see table IV for measure- discs of the two lateral fingers and fourth toe are

— in discs The ments). The ratio of hand length to snout- red; some specimens all are red. belly

The ratio of is The bones The iris vent length is 0.30-0.38 (mean 0.33). pale blue-green. are green.

is 0.54-0.67 is with blue tibia length to snout-vent length silvery grey a periphery.

(mean 0.59). The ratio of foot length to snout- In preservative, the general colouration is ratio of with vent length is 0.45-0.56 (mean 0.49). The yellowish. The dorsal surface is covered

the horizontal diameter of the tympanum to that silver-white spots. There is a silver-white line from of the is 0.41-0.59 Snout-vent the nostril to the The are white eye (mean 0.47). eye. upper eyelids

of males of female mm. with brown. The other lines invisible. The red length 19-5-25.0 mm, 32 are

Of the ratios mentioned, only the last is different colour has disappeared completely in the ZMA

from that in B. mandraka, in which species the collection, probably because the collection was the the for time. tympanum is smaller and eye larger. exposed to light some

and B. Since B. erythrodactylus mandraka are

have the the — Novem- much alike, I restricted description of Voice. The males were heard from

B. adult features of erythrodactylus mainly to those ber till April in the evening. The sonagram (fig.

in which it differs from B. mandraka. All re- 9) made of a male (ZMA 6928) on 4 March 1973

features identical those mentioned 11 shows that the duration of call maining are to at p.m. one above for B. mandraka. varies from 0.5-0.7 sec. One call consists of 4 to

The snout is bluntly rounded in lateral profile 5 rather melodious clicks. The dominant frequency

in the male and rounded in the female. The is about 2500 Hz.

nostrils are slightly nearer to the tip of the snout

than the anterior corners of the The distance E — In the dissected female the to eye. g g s . eggs are

from the nostril the anterior of the cream-white and in their number to corner eye 2 mm diameter;

is subequal to the horizontal diameter of the eye. is about 200.

The canthus is distinct and slightly angular.

— ZMA The horizontal diameter of the tympanum is Tadpoles. 7207, 7208. Twenty-

of the horizontal diameter of the four in 25 have of 2/5 to 2/3 eye. tadpoles stage body lengths

The the and distance from eye to the tympanum is 5.0-10.2 mm total lengths of 15.0-30.5 mm.

1/2 to 3/4 of the diameter of the tympanum. The Five tadpoles in stages 27 and 28 have body BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 271

Fig. 9. Sonagram, filter wide, Boophis erythrodactylus.

lengths of 10.5-11.8 mm and total lengths of large papillae laterally, and 5-6 rows of large

in 31-36 have The oral disc is round. The 32.0-35.5 mm. Five tadpoles stages papillae posteriorly.

total body lengths of 10.5-11.0 mm and lengths tooth formula is 4/4 + 4//3. The upper row of

One in 41 has lower of of 29.5-34.0 mm. tadpole stage a the upper lip and the row the under lip total of consist of small teeth. The beak is well body length of 31-5 mm and a length very horny

All beak with 41.0 mm. these tadpoles were caught in the developed. The lower is U-shaped

Mandraka valley, at 1200 m alt. Those collected in robust serrations.

in the Ankaratra mts. are larger; two tadpoles stages The colour of the tadpole in life is brown on

and 14 in 31 and 34 have body lengths of 12 mm the back, the belly is black stage 25 and becomes

The caudal musculature and total lengths of 36-39 mm (ZMA 7205). silver-grey in later stages. The

ratio of tail to in the has dark brown and the caudal fin is average length body length patterns thirty-seven tadpoles listed above is 2 (range 1.7- transparent.

Five have The were collected in from 2.1). metamorphosing young body tadpoles rapids June

lengths of 13-15 mm (mean 14 mm). till October, together with those of B. mandraka.

in occurred in The mouth and lateral view of a tadpole stage Metamorphosis October (at the begin-

The the 28 (ZMA 7208) are shown in text-figs. 5-6. ning of hot season). Only fully developed tad-

body is depressed. The nostrils are dorsal, the poles could be brought through metamorphosis in opening is anterolaterally, situated at about 3/4 the vivarium.

from the tip of the snout and the anterior corner

the The The — It of eye. eyes are situated laterally. Young. was rather difficult to assign the

is sinistral, situated at about reared from the different spiracular opening young frogs tadpoles 3/4 from the tip of the snout and the end of the of the two sibling species B. erythrodactylus and

is and the B. I have body. The cloacal tube long dextral to mandraka. assigned the young from the caudal fin. The caudal musculature is robust. The tadpole described above to B. erythrodactylus,

caudal fin is shallow and at because the in life shows the anteriorly deepest young frog typical about three-fourths of its length. At midlength of yellow line from the nostril to the anterior corner

the the of the caudal musculature of the and another tail, height eye straight yellow line from

of the total The mouth is the corner of the to behind represents 1/2 height. posterior eye the broad and The shoulder. This directed ventrally. papillary border green froglet has yellow spots all

has wide median There 3-4 of the dorsal and hands and a gap. are rows over surface, yellow feet. 272 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

The bones The canthus is The width of The nostrils are are green. angular. an upper eyelid. strongly

in the hindlimbs are relatively shorter than adult, protruding, the opening is laterally. The commis-

in of the mouth is The is hori- which is a common phenomenon young frogs. sure straight. pupil

zontally elliptical.

There is a weak dermal fold from the posterior viridis Boophis sp. nov. of edge the eye, over the tympanum, curving PI. IV fig. 5. of downwards to a point just behind the insertion

the arm. The fold covers the dorsal of the Material. — ZMA 7100A, male holotype, ZMA 7100B, margin

two male paratypes, collected 14.XI.72 near Perinet (high- tympanum. The tympanum is rather indistinct, its 2 km road R.N. at 142), alt. 900 m. horizontal diameter is one-third of the horizontal

The resembles the other of diameter of the The distance from the present species species eye. eye

the B. the is about rappiodes group. viridis can be distinguished to tympanum 3/4 of the horizontal

from rappiodes, mandraka and erythrodactylus by diameter of the tympanum.

its larger body size (snout-vent length 29-31 mm The largest diameter of the disc of the third

adult the the of males), by strongly protruding finger equals the horizontal diameter of tym-

and its the discs of nostrils, by the relatively longer snout, by panum, the fourth and second finger

difference in colouration (see text description). are about equal and of the first much smaller. The relative lengths of the fingers from shortest

— The males Habitat. were found on a rainy to longest are 1-2-4-3. The subarticular tubercles

shrubs and and round. distal evening, calling on leaves of trees, are small, distinct The tubercles

along running water, in forest. on the fourth and third finger are bifid. There is

A an inner flat, oval metacarpal tubercle. small

IV is Description (see table for measure- unpigmented, glandular nuptial pad present

ments). — The ratio of hand length to snout-vent dorsolaterally on the first finger. There are some

is The ratio of tibia tubercles the of the length 0.32-0.34 (mean 0.33). supernumerary on base fingers.

length to snout-vent length is 0.58-0.62 (mean The formula of the webbing of the hand: 1 (1),

0.60). The ratio of foot length to snout-vent 2i ( 1 ), 2e ( 1 ), 3i (2), 3e ( 1 ), 4 ( 1 ). The web is

length is 0.45-0.48 (mean 0.46). The ratio of the glandular.

horizontal diameter of the tympanum to that of The arms and legs are slender. When the limbs

the eye is 0.32-0.36 (mean 0.35). Snout-vent are laid along the side, the knee and elbow overlap

males length of 29-31 mm. noticeably. The adpressed heels meet. The tibio-

head is head width is tarsal articulation extends the of the Holotype: length 9 mm; beyond tip

11 mm; internarial and interorbital distance is snout. The inner metatarsal tubercle is small,

3 mm; snout-vent length is 31 mm. distinct, elliptical. A small web connects the tip of

The body in the postaxillary region is little the inner metatarsal tubercle with the first toe to

than the width the The the narrower greatest of head. middle of the antepenultimate phalanx. There

is head width is discs the head wider than long. The one- is no outer metatarsal tubercle. The of

in third of the body length. The snout is truncate toes are equal to those of the fingers. The sub-

and dorsal lateral profile. The snout is longer than articular tubercles are small, distinct and round.

the horizontal diameter of the and to The fifth is than the third. The formula eye, equal toe longer

the distance between the anterior corners of the of the of the foot: 1 2i 2e webbing (0), (l/ 2 ), (0),

nostrils much The are nearer to the of 3i 3e 4i 4e 5 The eyes. tip (l/ 2 ), (0), (1), (1), (0).

the snout than to the anterior of the ventral surface of the web is The corners eyes. glandular. lateral

The canthus is and rounded. A tarsal is distinct The loreal metatarsalia are separated. weak fold region is slightly concave. The interorbital region present.

is slightly convex. The ratio between the inter- The anal opening is directed posteriorly at the

narial and interorbital distance is 1. The inter- level of the The anal sheath is upper thighs. short. orbital distance is about and is one a half times the The tongue heart-shaped and distinctly bifid BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 273

posteriorly. The vomerine teeth are situated me- Differential diagnosis

dially, far behind the choanae, in two small 1. Large species (75-95 mm), toes entirely webbed, the webbing the discs between the third and fourth The vocal slits situated reaching straight groups. are small, finger B. albilabris (Boulenger) in the floor of the near the of the mouth, angles 2. Medium-sized species (40-55 mm), one phalanx of and jaws. The vocal sac is single, median subgular fourth toe free of web, the webbing reaching the bases of the penultimate phalanges between the third and noticeably distensible. fourth finger B. luteus (Boulenger) dorsum and throat is The skin on the finely

granular. The skin on the belly and posteroventral

— have Remarks. Although I not seen B. surfaces of the thighs is granular. The skin beneath albilabris in life, I consider the species closely the anal opening is coarsely granular. Elsewhere related to B. luteus. Boulenger (1888) in the the the on venter, skin is smooth. original description, mentioned the paired, lateral, colouration in life is The general light green, vocal the white lateral subgular, sac, fringes along- the is and the dorsum is upper eyelid yellow the lower and side arm tarsus, and supposed a covered with red speckles (pi. IV fig. 5). The iris colour in life. green is orange-brown with a blue periphery, surrounded

The bones by a black ring. are green. The colour in preservative is entirely light Boophis luteus (Boulenger, 1882) is yellowish, the upper eyelid whitish, especially Text-figs. 7, 10-12; pi. I fig. 3, pi. IV fig. 4. the border. The iris is beige-brown, surrounded

The rim by a black ring. upper of the palpebral luteus Rhacophorus Boulenger, 1882: 468; Guibe, 1978: 73, membrane is The lower rim is slightly pigmented. figs. 172-177 (equals isabellinus). in the colour the skin broadly pigmented same as Rhacophorus isabellinus Boettger, 1913: 277, pi. XXIII fig. 6. luteus Bachmann & of the Boophis ; Blommers-Schlosser, 1975: 17 upper eyelid. (DNA); Blommers-Schlosser, 1978: 30 (chromosomes).

Material. — SMF 6821, holotype of Rhacophorus Etymology. — The specific name refers to isabellinus. Type locality: Moramanga. Coll. Voeltzkow. the colour of the green species. ZMA 6840 (3 3 3), 27.111.73; 6838 (1 3,1 9), 4.III. of 73; 7094 (eggs 6838); collected in the Mandraka valley

R.N. 2 at km (highroad 67); 7005 (tadpoles and young from reared them), 1.IV.-1.X.72 same locality at km 69;

1100 7092 (tadpoles), 24.IX.72 same locality at km 70, alt. Boophis luteus group m. — ZMA 6839 (5 3 3), 13.XI.72; 7090 (tadpoles and

reared from young them), 24.IX.72; collected near Perinet

The members of this (highroad R.N. 2 at km 142), alt. 1100 m. — ZMA 7091 Diagnosis. — group and reared from 12.IX.71 collected (tadpoles young them), are moderate- to large-sized, stream-breeding spe- near Tampoketsa dAnkazobe forest station (highroad R.N. cies. The dorsum is dark the bones are km — green, 4 at 125), alt. 1600 m. ZMA 7093 (tadpoles), 19.IX.

the 71 collected at forest station, Ankaratra There are distinctive on Manjakotompo, mts., green. no markings alt. 1800 m. — ZMA 7146 (tadpoles), 29.VIII.72 collected white posterior surface of the thighs. There are along the road from Moramanga to Anosibe at km 27, alt.

lateral the lateral of the lower 800 fringes along edge m.

arm and tarsus. The skin on the dorsum is smooth.

vocal is — The Vomerine teeth are present. The sac paired, Natural history. males are calling

lateral, subgular. The horizontal diameter of the from the leaves of shrubs and trees along rapids in

The is the The female tympanum is half that of the eye. snout forest, during rainy nights. was

the The tibiotarsal artic- found with a male in The tad- rounded, eyes are large. axillary amplexus.

in ulation extends to the tip of the snout. The discs poles occur swiftly flowing waters.

are well developed. The fingers are half or more

almost Measurements — webbed. The toes are entirely or entirely (see table IV). The

webbed. ratio of hand length to snout-vent length is 0.28-

Contents: B. luteus (Boulenger, 1882) and B. 0.32 (mean 0.29). The ratio of tibia length to albilabris (Boulenger, 1888). snout-vent length is 0.51-0.57 (mean 0.54). The

i 8 274 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

ratio of foot snout-vent is 0.44- E s -— The female about 200 black, length to length g g . deposited

0.51 The ratio of the horizontal in The (mean 0.48). very sticky, eggs a collecting bag. eggs diameter of the to that of the is in diameter. of the tympanum eye measure 2 mm Development

0.44-0.49 Snout-vent of did not occur. (mean 0.48). length eggs

female males 37-40 mm, of 54 mm.

Tadpoles. -— Tadpoles, which developed into

Sexual — The male has of B. luteus in the structure dimorphism. young are dimorphous

the first of the I discover in a white nuptial pad latero-dorsally on mouth. could not differences

slits situated in the from the different finger. The vocal are small, the young developing tadpoles,

is that B. floor of the mouth, near the angles of the jaws. but it possible one belongs to albilabris.

is is Form B The vocal sac paired subgular. The male A (ZMA 7005, 7092, 7146) and form smaller than the female. (ZMA 7090, 7091, 7093). A: Fifty-seven tad-

in have poles stage 25 body lengths of 4.5-11.0

Colour — colouration of 14-28 in life. The general mm and total lengths mm. The average

IV tail is is dark, slightly bluish, green (pi. fig. 4). The ratio of length to body length 1.7 (range bones The iris is whitish in in are green. orange, or 1.5-1.9). Twenty-nine tadpoles stages 26-30

red in the middle surrounded with dark red, or have body lengths of 9-0-14.5 mm and total

middle and whitish around. Ten the lengths of 24-30 mm. tadpoles in stages 31-40

have body lengths of 14-20 mm and total lengths

of of — call B. is of The tail Voice. The luteus conspicuous 38-59 mm. average ratio length to and characteristic. It is melodious in the 26-40 is 1.7 very a sound, body length stages (range

and for Thirteen have repeated quickly constantly as long as 30 1.5-1.9). metamorphosing young

It New B: seconds. reminds of a York police car. body lengths of 18-22 mm (mean 20.5 mm).

At first, we did not suspect a frog to produce it. Fourteen tadpoles in stage 25 have body lengths

The males could be heard the of 7-13 and total of Three every evening during mm lengths 19-38 mm.

made in 14- rainy season. The sonagram (fig. 10) of a tadpoles stages 26-30 have body lengths of

4 and total Five tad- male (ZMA 6838) on March 1973, at 11.30 15 mm lengths of 39-41 mm.

shows of the The in p.m. only a part very long note. poles stages 35-38 have body lengths of 18-22 dominant here also fundamental fre- and total of The frequency, mm lengths 50-65 mm. average

is about 3000 Hz. ratio of tail in the quency, length to body length twenty- two tadpoles listed above is 1.7 (range 1.5-1.9).

Three metamorphosing young measure 21.0-24.5

mm from snout to vent.

The mouth (form A) of a tadpole in stage 36

and mouth (ZMA 7005) the (form B) of a tad-

pole in stage 27 (ZMA 7090) are shown in text-

figs. 7 and 11.

A: The mouth is small and directed ventrally.

The border has wide median ante- papillary a gap

and riorly. There are two rows of papillae laterally

The there are numerous supernumerary papillae.

oral disc is indented very slightly medioposteriorly.

The is tooth formula 1/5 + 5//3 or 1/4 + 4//3,

the lower is the first row of lip sometimes slightly

interrupted in the middle. The horny beak is well developed.

B: The mouth is small and directed ventrally.

The border has wide median Fig. 10. Sonagram, filter wide, Boophis luteus. papillary a gap BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 275

Figs. 11-18. 11-12, Boophis luteus: IX, mouth of tadpole, form B, ZMA 7090.X.12.5; 12, tadpole, ZMA 7005.X.3;

13-14, B. sp.: 13, mouth of tadpole, ZMA 7154.X.12.5; 14, tadpole, ZMA 7154.X.3; 15-16, B. majori: 15, mouth of

tadpole, ZMA 7096.X.12.5; 16, tadpole, ZMA 7096.X.3; 17-18, B. goudoti: 17, mouth of tadpole, ZMA 7209.X.12.5;

18, tadpole, ZMA 7209.X.1.5. 276 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

of anteriorly. There are 1-2 rows fringing papillae Boophis albilabris (Boulenger, 1888)

inside and a supernumerary row, more laterally

with The oral disc albilabris 1888: VI, a few supernumerary papillae. Rhacophorus Boulenger, 105, pi. fig. 1; Guibe, 1978: 71, figs. 164-165 is indented medioposteriorly. The tooth formula is

first of the This collected 1/2 + 2//3 or 1/3 + 3//3, the row species was not by the author.

lower lip is sometimes slightly interrupted in the

middle. The horny beak is well developed. Since the morphology of both forms (A, B) is The tadpole of the following unknown species similar in the other features, the description is resembles much the tadpole of B. luteus, therefore

given once. The lateral view of a tadpole in stage its description follows here.

36 (ZMA 7005) is shown in fig. 12. The body

is depressed. The nostrils are dorsal, the opening Boophis sp. is large and directed to the front, situated slightly Text-figs. 13-14; pi. I fig. 4. of the than to nearer to the anterior corners eyes

of the The and directed the tip snout. eyes are large

Material. — ZMA 7154 and reared (tadpoles young The is sinistral, laterally. spiracular opening from them), 1.IV-I.X.72; 7147 (tadpoles), 24.IX.72; col- situated at about 5/8 from the tip of the snout lected in the Mandraka valley (in the Mandraka and Vokana- tetezandava river), (highroad R.N. 2 at km 69-70), alt. and the end of the body. The cloacal tube is wide 1200 m. and dextral to the caudal fin. The caudal muscula-

is in the first half — in ture robust, especially of the Habitat. The tadpoles were collected

tail. The dorsal caudal fin is reduced anteriorly. rather large streams with stony bottom. An adult

At midlength of the tail, the height of the caudal specimen was never noticed.

musculature represents about 2/5-1/2 of the total

— in tail height. Tadpoles. Thirty-two tadpoles stage 25

The colour of the tadpole is brownish or have body lengths of 5.5-9.5 mm and total lengths

in it of blackish life; becomes greenish in the late 16.0-26.5 mm. The average ratio of tail length

is in metamorphic stages. The belly transparent to body length in this stage is 1.85 (range 1.65-

and in the in have the early stages silvery later stages of 2.00). Twelve tadpoles stages 26-30 body

caudal is and development. The musculature densely lengths of 9-11 mm total lengths of 24-29

with and the in pigmented brown or black spots mm. Seventeen tadpoles stages 31-40 have body

few brown of caudal fin has a dispersed spots. lengths of 11.0-13.5 mm and total lengths

in October and the 29.0-36.5 The ratio of tail Metamorphosis occurs begin- mm. average length to

The in 26-40 is 1.60- ning of November. tadpoles were collected body length the stages 1.7 (range

from streams with or bottom. Those Three measure 13- stony muddy 1.85). metamorphosing young

collected from the streams with a stony bottom are 15 mm from snout to vent.

blackish; theothers are brownish. The mouth and lateral view of a tadpole in stage

in 31 (ZMA 7154) are shown figs. 13-14. The

— In Young. life the colour of the back is body is depressed. The nostrils are dorsal, the

with red between the is green a spot eyes; sometimes opening large and directed to the front, situated

the of the red pigment covers more back. There slightly nearer to the anterior corners of the eyes lateral lines from of the than of the The are two yellow the tip to the tip snout. eyes are large

snout the the insertion of the hind- and directed The is over eye to laterally. spiracular opening

limbs. The is The bones are situated at about from the of the belly greenish. green. sinistral, 5/8 tip

iris is The white tube The orange. lateral folds from the snout and the end of the body. The cloacal elbow to the disc of the fourth finger and from the is wide and dextral to the caudal fin. The caudal

the disc of the fifth is in heel to toe are already visible musculature robust, especially the first half

The in some specimens. webbing of hand and feet of the tail. The dorsal caudal fin is reduced ante-

is identical to that of adults of B. luteus. riorly. At midlength of the tail, the height of the BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 277

caudal musculature represents about 2/5-1/2 of posterior surface of the thighs. There are white

the total tail height. The mouth is rather broad lateral fringes along the lateral edge of the lower

The border The and directed ventrally. papillary has a arm and tarsus ( difficilis, majori, miniatus).

wide median gap anteriorly. There are three rows ventral surface of the hindlimbs, the web between

of papillae and laterally there are three rows of the toes, the posterior and anterior surface of the The oral disc is violet-red. The skin the supernumerary papillae. slightly thighs are usually on

indented medioposteriorly. The tooth formula is dorsum is smooth or slightly granular. Vomerine

teeth diameter the 2/4-f4//3 or 2/4 + 4//1 + 1/2. The two upper are present. The horizontal of

is half that of the rows of the upper lip are situated distinctly above tympanum one-third to eye.

The vocal is The the horny beak. The teeth are large. The horny sac single, median, subgular.

beak is well developed. snout is short and rounded. The commissure of

The colour of the tadpole is brownish in life. the mouth is slightly curved upwards except in The caudal musculature is densely pigmented with rhodoscelis, in which the commissure of the mouth

and the has brown spots caudal fin a few dis- is straight. The tibiotarsal articulation does not

persed brown spots. extend beyond the tip of the snout. The discs are

in well The one-third Metamorphosis occurs October (beginning developed. fingers are webbed, of the hot season). The tadpoles resemble those the web connects the middle of the antepenulti- of B. luteus (Boulenger, 1882), collected from the mate phalanx of the third finger with the base of

same streams. The diagnostic feature is the mouth the penultimate phalanx of the fourth. The toes and size the tad- One the smaller of metamorphosing are two-thirds to three-fourths webbed. to

It is not to the with two of the fourth toe are free of web. poles. possible identify young phalanges

certainty, since we collected no adults at the same Contents: B. rhodoscelis (Boulenger, 1882), place. B. difficilis (Boettger, 1892), B. majori (Boulen-

and B. miniatus ger, 1896), (Mocquard, 1902).

Young. — In life the colour of the back is

beige with brown spots. The venter is equally — Remarks. B. difficilis, B. majori and B. creamish. There are two lateral light beige lines miniatus are hard to distinguish morphologically. from behind the to the The nostrils eyes groin. They were collected, when calling, during rainy are slightly nearer to the tip of the snout than to nights. It was heard that the mating call is clearly the anterior corners of the The tibiotarsal eye. different in each of them. The nuclear DNA articulation extends to the centre of the The eye. amount 7.8 in B. and 9-1 (diploid) was pg majori

heels meet. The formula of the ~ ~ adpressed webbing in B. Pg difficilis (Bachmann & Blommers- of the foot: 1 (0), 2i (1), 2e (0), 3i (1), 3e (0), Schlosser, 1975). Once ascertained that there are 4i 5 The formula of the of (1), (0). webbing made three species, I have an attempt to find the hand: 3e (2), 4 (1). A trace of web between diagnostic features for each of them. the others. The discs are moderately developed.

The skin of the back and chin and throat is finely Differential diagnosis granular. The skin of the belly and the ventral

Tibiotarsal articulation the lateral surface of the thigh is coarsely granular. The 1. reaching eye, fringes alongside the lower arm and tarsus absent, commissure of remaining ventral surface is smooth. mouth straight, canthal and temporal region dark brown.

B. rhodoscelis (Boulenger)

2. Tibiotarsal articulation reaching beyond the eye, lateral

fringes alongside the lower arm and tarsus present, com- rhodoscelis Boophis group missure of mouth curved upwards, canthal and temporal

region same colour as dorsum . B. difficilis (Boett-

B. miniatus — The this ger), majori (Boulenger), B. (Mocquard) Diagnosis. members of group

are small-sized, stream-breeding species. The dor-

is red. There bars Between the latter three small sum beige or are transverse on are very differences,

the which in the limbs, but no distinctive markings on are shown table III. - 278 R. M. A. BLOMMERS-SCHLOSSER BIOSYSTEMATICS OF BOOPHIS

TABLE III

Diagnostic characters between Boophis difficilis, B. majori and B. miniatus.

B. difficilis B. majori B. miniatus

Snout-vent length of adult $ (mm) 27-31 22.0-26.5 31-34

Ratio hor. diameter tympanum/eye (mean) 0.36 0.35 0.50

weak Fringes alongside lower arm and tarsus well developed well developed developed

Position of nostrils of in respect of tip of much nearer tip midway tip slightly nearer

and anterior and of snout corner of eye of snout snout eye tip snout

Character of nostrils strongly slightly slightly protruding protruding protruding

Skin on the back, chin and throat smooth finely granular smooth

Colouration cf. text description species

Boophis rhodoscelis (Boulenger, 1882) Natural history. — The species was never

heard or noticed during daytime. The males were Rhacophorus rhodoscelis Boulenger, 1882: 466; Guibe, 1978: found on rainy evenings, calling in the vegetation 74, figs. 181-182 (equals andrangoloaka and brevirostris). in along a brook, forest. They were seen moving Rhacophorus andrangoloaka Ahl, 1928: 311; Guibe, 1978: 74

of (in synonymy rhodoscelis). down from the trees.

Rhacophorus brevirostris Ahl, 1928: 313; Guibe, 1978: 74 ZMA IV (in synonymy of rhodoscelis). Description of material (see table

for — The ratio of hand to — measurements). length Material. BMNH 1947.2.7.66-67, two syntypes of Rhacophorus rhodoscelis. Type locality: East Betsileo. Coll. snout-vent length is 0.29-0.32 (mean 0.305). The Cowan. ratio of the tibia length to snout-vent length is ZMB 30510, two syntypes of Rhacophorus andrangoloaka. 0.52-0.59 The ratio of the foot Type locality: Central Madagascar. Coll. Hildebrandt. (mean 0.56).

ZMB 30518, holotype of Rhacophorus brevirostris. Type length to snout-vent length is 0.43-0.50 (mean

locality: N.W. Madagascar. Coll. Hildebrandt. 0.46). The ratio of the horizontal diameter of the ZMA reared from 12.IX.71 7194 (one young tadpole), to that of is forest the eye 0.34-0.38 (mean collected near Tampoketsa d'Ankazobe, station (high- tympanum Snout-vent road R.N. 4 at km 125), alt. 1600 m, in a stream. 0.36). length of males 27-31 mm.

The head is wider than the body, and somewhat

Yo u n — The hatched measures 19 mm g . young wider than long. The head length is one-third of

from snout to vent of this about (adult species the body length. The snout is rounded in dorsal

35 mm). It has already the crimson colouration on and profile, sloping to truncate in lateral profile. the ventral surface of the hindlimbs. The snout is slightly longer than the horizontal

diameter of the somewhat shorter than eye, and

the distance between the anterior corners of the

difficilis bona Boophis (Boettger, 1892) species The nostrils much eye. are nearer to the tip of the Pl. IV fig. 6. than the anterior of The snout to corners the eye.

distance from the nostril to the anterior corner

Rhacophorus difficilis 1892: 1913: Boettger, 14; Boettger, of is the eye shorter than the horizontal diameter XXIX 1947: 439 276, pi. fig. 1; Guibe, (in synonymy of of the The canthus is rather distinct and Guibe, 1763 of eye. depressiceps); 1975a: (elimination synon- ymy of depressiceps); Guibe, 1978: 65 (in synonymy of rounded. The loreal region is slightly concave. tephraeomystax; in part). The interorbital region is flat. The ratio of the Boophis sp. 2; Bachmann & Blommers-Schlosser, 1975: 17 internarial distance to the interorbital distance is (DNA). 4/5. The interorbital distance is somewhat wider Material. — SMF 6762, holotype of Rhacophorus than the width of an The nostrils difficilis. Type locality: Foizana, E. Madagascar. Bought from upper eyelid.

Schneider, Basel, 1880. are strongly protruding, the opening is laterally.

ZMA 7098 (1 $), 14.XI.72; 7099 (6 $ $), 24.XII.72; The commissure of the mouth is slightly curved collected near Perinet (highroad R.N. 2 at km 142), alt. The is horizontally 900 m. upwards. pupil elliptical. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 279

TABLE IV

Dimensions in mm of Boophis rappiodes, B. mandraka, B. erythrodactylus, B. viridis, B. luteus, B. difficilis, B. majori

A = = and B. miniatus. snout to vent; B length of lower arm; C = hand length; D = length of hind limb; E = tibia length;

= F length of foot and G = foot = I = of tarsus; length; H horizontal diameter of tympanum; horizontal diameter eye.

ABCD EFGH I

B. rappiodes

1 $ ZMA 24 11 6 42 14 16 10 1 3

1 9 ZMA 32 16 9 55 19 22 14 1.2 3.5

B. mandraka

3 holotype ZMA 23.5 12.5 7.5 41 13 16.5 10.5 1 3

12 3 3 paratypes ZMA 21-26 12.0-13.5 7-8 39-42 12.5-14.0 16-18 10.5-11.5 0.8-1.0 2.8-3.0 B. erythrodactylus

25 3 3 ZMA 19.5-25.0 11-13 7-8 40-44 12.5-14.5 15.5-18.5 10.5-12.0 1.0-1.3 2.2-2.9

1 9 ZMA 32 17 9.5 55.5 17 22.5 15 1.8 3.2

B. viridis

3 holotype ZMA 31 15.5 10 57 18 23 14 1.2 4

2 3 3 paratypes ZMA 29 15 10 52-53 17-18 22 13-14 1.2-1.3 3.6

B. luteus

9 3 3 ZMA 37-40 18-20 10.5-12.0 61-68 19-22 27-30 17-20 2.0-2.2 4.1-4.5

1 9 ZMA 54 25 15 90 29 39 25 2.2 5

B. difficilis

holotype SMF 28 13.5 8.5 44 14.5 18 12 1.2 3.2

7 $ 3 ZMA 27-31 13.5-15.0 8.5-9.5 46-52 14.5-16.5 19.0-21.5 12-14 1.2 3.2-3.5

B. majori

lectotype BMNH 3 24 12 7 41 13 18 11 1.1 3

32 3 3 ZMA 22.0-26.5 12-13 7-8 40-45 13-15 16.5-19.0 10-12 1.1-1.2 2.8-3.2

1 9 ZMA 32 16 10 57 18 25 15 1.2 4

B. miniatus

holotype MNHP 27 13.5 8 45 15 20 11.5 1.8 3.5

3 3 3 ZMA 31-34 15-16 9-11 49-56 16.5-18.0 22-24 13.5-15.0 1.9-2.2 3.8-4.1

weak dermal fold from the of web the the fourth There is a posterior between fingers. Between

of the the down- and third the web extends from the middle edge eye, over tympanum, curving finger

wards behind insertion of the to a point just the of the of the antepenultimate phalanx third to the

of the arm. The fold covers the dorsal margin base of the penultimate phalanx of the fourth.

is distinct and tympanum. The tympanum round, The web is glandular.

its diameter is about one-third to two-fifths of the The arms and legs are slender. When the limbs

from horizontal diameter of the eye. The distance are laid along the side the knee and elbow overlap

the the is about half the diameter The heels eye to tympanum strongly. adpressed overlap slightly.

the articulation of tympanum. The tibiotarsal extends to a point

The largest diameter of the disc of the third between the nostril and the tip of the snout, or to

finger equals the diameter of the tympanum; the the tip of the snout. The inner metatarsal tubercle disc of the fourth finger is subequal, of the second is distinct, small, elliptical and protruding. There

finger slightly smaller and of the first much is no outer metatarsal tubercle. A small web

smaller. The relative lengths of the fingers from connects the middle of the inner metatarsal

shortest to longest are 1-2-4-3. The subarticular tubercle with the first toe to the base of the

and terminal discs tubercles are distinct round. The distal tu- penultimate phalanx. The are some-

what bercles of the fourth and third finger are some- smaller than those on the hand. The fifth toe

is inner is times bifid. There an oval, flat metacarpal somewhat longer than the third. The subarticular

tubercle and no palmar tubercle. There is a small tubercles are distinct and round. The formula of

unpigmented, glandular nuptial pad dorsolaterally the webbing of the foot: 1 (0), 2i (1), 2e (0),

on the first tubercles are 3i 3e 4i 4e 5 The finger. Supernumerary (1), (0), (2), (U/2 ), (l/ 2 ).

on the base of the There is lateral metatarsalia The present fingers. a trace are separated. ventral 280 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

is surface of the web glandular. The ventral surface toes being about half the size of the discs on the

of the foot is A white Examination of the showed that slightly granular. fringe goes fingers. holotype

from the the the elbow along side of lower arm the discs on the toes are shrunken. This specimen,

and fourth finger to the disc. Another white collected in 1880, probably already had shrunken tuberculous from the heel when fringe goes along the discs Boettger saw it. The hindlimbs are side of the tarsus and fifth toe to the disc. relatively slightly shorter in the holotype than in

The anal opening is directed posteriorly at the our specimens.

the upper level of thighs. The dermal flap above the anal opening is short.

The tongue is heart-shaped and distinctly bifid Boophis majori (Boulenger, 1896)

vomerine teeth situated I posteriorly. The are me- Text-figs. 15-16; pi. fig. 5.

dially just behind the large choanae in two small

slits situated Rhacophorus majori 1896: 402; Guibe, 77 oblique groups. The vocal are small, Boulenger, 1978: (equals miniatus; in part). in the floor of the mouth near the angles of the Bachmann Boophis sp. 3; & Blommers-Schlosser, 1975: 17 The vocal sac is median, and jaws. subgular (DNA). moderately distensible. Boophis aff. majori; Blommers-Schlosser, 1978: 35 (chro- mosomes) . The skin on the dorsum, chin and throat is

— of smooth. The skin on the belly, below the anal Material. BMNH 1947.2.7.67, male lectotype Rhacophorus majori by designation. Type locality: and is present opening on the ventral surface of the heel Ambohimitombo forest (near Perinet). Coll. F. Major. —

and white. The skin on the very coarsely granular BMNH 1947.2.7.66, paralectotype, same data as lectotype;

posteroventral surface of the thigh is moderately other two original syntypes not examined. ZMA 7095 (2 $ $), 6.1.73; 7096 (one tadpole), VI.72; granular. Elsewhere on the limbs the skin is and reared from 7097 (tadpoles young them), VI-X.72; 7102 smooth. (1 g), 15.XI.72; 7103 (7 $ $ and 1 9), 26.XII.72; 7104 The colour in preservative is yellowish. There is (15 $ $), 18.11.73; 7105 (7 3 3), 27.111.73; 7143 (one Mandraka tadpole), 24.IX.72; all collected near valley brown line from the of a transverse edge one eye (highroad R.N. 2 at km 69), rivulet Vokanatetezandava, in to two brown lines from the other, and the of the of alt. oblique nursery garden city Tananarive, 1200 m.

behind the eyes form a V-shaped figure, ending

behind the forelimbs. From this point the back Natural history. — The species was never

is less heard the The males more or brown spotted, the border of this or noticed during day. were

is in in the spotted region A-shaped anteriorly. There are found the evening, calling vegetation

Brown the rivulet. The female found with brown crossbands on the limbs. pigment is along was a

The present around the nostrils, around the anal male in axillary amplexus. female was gravid.

in The opening and on the ventral surface of the tarsus, The tadpoles were found the rivulet. and sometimes the canthus. The rim call resembles of birds. along upper mating high chirping of the palpebral membrane is entirely pigmented, and the lower rim only anteriorly and posteriorly. Description of ZMA material (see table IV

The iris is with for — The ratio of hand silvery grey some dark pigment, measurements). length

is surrounded by a black ring. to snout-vent length 0.29-0.33 (mean 0.31).

In is with is life, the back orange-red numerous The ratio of tibia length to snout-vent length

red The ratio of foot dark spots, the markings (brown in preserva- 0.52-0.59 (mean 0.55). length

is tive) are blackish (pi. IV fig. 6). The fringes to snout-vent length 0.40-0.50 (mean 0.44).

the along the lower arm and tarsus are white. The The ratio of horizontal diameter of the tym-

is white. The iris is surrounded of is belly silvery, by a panum to that the eye 0.31-0.40 (mean 0.35). black ring. Snout-vent length of males 22.0-26.5 mm, of

female 32.0 mm.

Remarks. — The holotype of Rhacophorus The head is wider than the body, and somewhat

than width is difficilis agrees largely with the description above. wider long. The head one-third of

described in Boettger, 1892, the terminal discs on the the body length. The snout is rounded dorsal BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 281

profile and bluntly rounded in lateral profile slightly. The adpressed heels overlap slightly. The

is in the male, rounded in the female. The snout tibiotarsal articulation extends to a point between

slightly longer than the horizontal diameter of the the nostril and the tip of the snout in males and

and somewhat shorter than the distance between the and the nostril in the female. eye, eye

between the of The inner metatarsal tubercle is small anterior corners the eye. The distinct,

the and There is nostrils are at equal distances from the tip of elliptical protruding. no outer meta-

and the anterior of the The tarsal tubercle. A small web connects the middle snout corners eye.

distance from the nostril to the anterior corner of of the inner metatarsal tubercle with the first toe

of the base of the eye is shorter than the horizontal diameter to the penultimate phalanx. The discs

the The canthus is distinct and rounded. The the almost those the eye. on toes are as large as on

loreal region is slightly concave. The interorbital fingers. The fifth toe is somewhat longer than the

region is flat. The ratio of the internarial to the third. The subarticular tubercles are distinct and

interorbital distance is 7/10. The interorbital round. The formula of the webbing of the foot:

distance is and half times the width of an 1 2i 2e 3e 4i - one a (V2)' (1/2). (0), 3i (1), (0), (H/ 2

The nostrils are 4e 5 The lateral metatarsalia upper eyelid. slightly protruding, 2), (11/ 2 ), (l/2"0)-

The the opening is dorsolaterally. The commissure of are separated. ventral surface of the web is the mouth is slightly curved upwards. The pupil glandular. The ventral surface of the foot is

is A white the horizontally elliptical. granular. tuberculous fringe goes from

There is a weak dermal fold from the posterior elbow along the side of the lower arm and fourth

of the the down- the disc and another from the heel edge eye over tympanum, curving finger to along

and wards to a point just above the insertion of the the side of the tarsus fifth toe to the disc.

arm. The fold covers the dorsal margin of the The anal opening is directed posteriorly at the

The is rather indistinct and level of the The dermal above tympanum. tympanum upper thighs. flap

round; its diameter is about one-third to two-fifths the anal opening is short.

of the horizontal of and diameter the eye. The distance The tongue is heart-shaped distinctly bifid.

from the is half three- The vomerine teeth situated eye to the tympanum a to are medially just

the in small fourths of the horizontal diameter of tym- behind the large choanae, two oblique

vocal slits in the panum. groups. The are small, situated

The largest diameter of the disc of the third floor of the mouth near the angles of the jaw.

the the The vocal is finger equals diameter of the tympanum; sac median, subgular and moderately disc of the fourth finger is subequal, of the second distensible.

finger slightly smaller, and of the first much The skin on the back, chin and throat is finely

The smaller. The relative lengths of the fingers from granular. skin on the ventral surface of the

shortest The shank skin the to longest are 1-2-4-3. subarticular and upper arm is smooth. The on

tubercles are distinct and round. The distal tuber- posteroventral surface of the thigh, lower arm and

cles on the fourth and third finger are often bifid. tarsus is moderately granular. The skin below the

is heel There an inner oval flat metacarpal tubercle anal opening, on the elbow and and on the

is and and no palmar tubercle. There is a small un- belly white coarsely granular.

pigmented glandular nuptial pad, dorsolaterally The colour in preservative is beige with dark

and dark on the first finger. Supernumerary tubercles are beige brown. The back is beige with some

the base the There brown present on of fingers. is a trace spots. There is a dark beige transverse line of web between the Between the third and from the of other. the fingers. edge one eye to the Behind

fourth web extends is finger the from the middle of eyes a more or less X-shaped figure with a broad

the in antepenultimate phalanx of the third to the transverse line behind it the same dark beige base of the penultimate phalanx of the fourth. colour. A dirty white vertebral line is in general

The web is the back. There dark glandular. present on are beige cross- The arms and legs are slender. When the limbs bands on the limbs. Dark brown pigment is usually

laid the and around and are along side, the knee elbow overlap present the anal opening on the ventral 282 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

surface of the tarsus and feet. The venter is uni- at about three-fifths of its length. At midlength of

creamish. The lower rim of the the of the caudal musculature form the palpebral tail, height rep-

membrane is pigmented. The iris is silvery beige resents two-fifths to the half of the total tail

surrounded is and with brown pigment, by a narrow height. The mouth broad directed ventrally.

dark ring. The whole contour of the lip has papillae: about

3-4 5-6 and In life, the colour is about the same, except the rows anteriorly, rows laterally poste-

ventral surface of the hindlimb and the web riorly, except in the median part of the lower lip,

between the toes, being violet-red, the venter light which has about 4 rows of papillae of larger size.

and the The The which is when blue-green bones light green. iris is oral disc, contracted preserved

brown with surrounded in shows lateral notches and orange, by a narrow brown alcohol, a strong ring. median notch in the lower lip. The tooth formula

is 4/3 + 3//3. The upper row of the upper lip

Remarks. — of under of The lectotype of Boophis majori and the lower row the lip consists very

is conform to the description above. It has retained small teeth. The horny beak is well developed,

the skeleton and the is brown the beak is less and green body light upper more or moon-shaped with dark brown markings. It is questionable the lower beak is U-shaped with robust serrations.

The of in is with whether the paralectotype belongs to the same colour the tadpole life beige

species. It differs from the lectotype in colour, dark brown markings; the belly is golden-coloured.

reddish with dark in being brown markings. The The tadpoles were collected a fast-running

throat October. The chin and are not granular, but smooth. current from June till metamorphosis

and Tuberculous folds along lower arm tarsus occurred only in October (beginning of the hot

are not discernible. It is neither entirely similar to season). The rearing was only successful when

B. with difficilis nor to B. miniatus. starting the late metamorphosis stages.

Eggs. — In dissected female, 2 mm in diameter, Young. — The metamorphosed froglet is creamish; number about 120. coloured as the adult, except that the hands and

— is Tadpoles. Twenty-seven tadpoles in stage feet are yellow, and that the red-violet colour

4.1-8.0 ventral hind- 25 have body lengths of mm and total not yet present on the surface of the

lengths of 13.5-24.5 mm. Ten tadpoles in stages limbs. The fringes along the lower arm and tarsus

26-31 have body lengths of 8.0-10.0 mm and total are already present. The legs are relatively shorter

lengths of 23.2-30.0 mm. Two tadpoles in stages than in the adults.

40 39 and have body lengths of 10.5-11.0mm and

total of The ratio lengths 32.0-32.5 mm. average Boophis miniatus (Mocquard, 1902) bona of tail length to body length in the thirty-nine species PI. I fig. 6. tadpoles listed above is 2 (range 1.8-2.2). Five

have of metamorphosing young body lengths 10.5- Rhacophorus miniatus Mocquard, 1902: 19, pi. II fig. 1; 12.0 mm (mean 11.5 mm). Guibe, 1978: 77 (equals majori; in part) The and in mouth lateral view of a tadpole stage Material. — MNHP 1901-386, holotype of Rhacophorus 31 are shown in 15-16. The (ZMA 7096) figs. miniatus. Type locality: forest between Isaka and valley of

body is ovoid. The nostrils are dorsal, the opening Ambobo, near Fort-Dauphin, S. Madagascar. ZMA 7162 (2 3 3), 4.IV.72; 7163 (1 3), 14.XI.72; is dorsolaterally, situated at about two-thirds from collected near Perinet (highroad R.N. 2 at km 142), alt.

the of the snout and the anterior corner of tip 900 m.

the The are situated The eye. eyes laterally. spirac- ular is three- — opening sinistral, situated at about Natural history. The species was never

fourths from the of the and the heard tip snout end of nor noticed during the day. The males were

the body. The cloacal tube is long and dextral to found in the evening, calling in vegetation along

the caudal fin. The caudal musculature is strong. a brook in the forest. They were observed moving is shallow The caudal fin anteriorly and is deepest down from the trees. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 283

Description of ZMA material (see table IV tubercle. There is a small unpigmented, glandular

— in first for measurements). The ratio of hand length nuptial pad, a dorsolateral position on the

to snout-vent length is 0.29-0.32 (mean 0.30). finger. Supernumerary tubercles are present on

The the base web ratio of tibia length to snout-vent length is of the fingers. There is a trace of

0.53. The ratio of foot length to snout-vent length between the fingers. Between the fourth and third

is 0.43-0.44. The ratio of the horizontal diameter finger the web extends from the middle of the

of the that of the is 0.48-0.54 of the third to the tympanum to eye antepenultimate phalanx finger

(mean 0.50). Snout-vent length of males is 31-34 base of the penultimate phalanx of the fourth one.

The is mm. web glandular.

head and limbs The head is wider than the body, and the The arms legs are slender. When the

width is equal to the head length. The head length are laid along the side the knee and elbow overlap

is one-third of the body length. The snout is slightly. The adpressed heels overlap slightly. The

rounded acuminate in articulation extends to slightly dorsal profile, tibiotarsal to a point between

in and and sloping to truncate lateral profile. The the eye the nostril. The inner metatarsal

snout is as long as the horizontal diameter of the tubercle is distinct, small, elliptical and protruding.

There is metatarsal tubercle. A small web eye, and somewhat shorter than the distance no outer

between the anterior of the The connects the middle of the inner metatarsal corners eye.

nostrils are somewhat nearer to the tip of the snout tubercle with the first toe to the base of the penul- than anterior of timate The discs the somewhat to the corner the eye. The distance phalanx. on toes are

from the nostril the anterior of the smaller than those the The fifth is to corner eye on fingers. toe

is shorter than the horizontal diameter of the than the third. The subarticular eye. slightly longer

The canthus is distinct and slightly angular. The tubercles are distinct and round. The formula of

is the of 2i loreal region slightly concave. The interorbital webbing the foot: 1 (V2), (V2), 2e (0),

is flat. The internarial distance is 3i 3e 4i 4e 5 The region equal (1), (l/ 2 ), (1), (1), (0).

to the interorbital distance. The interorbital dis- ventral surface of the web is glandular. The lateral

tance is somewhat wider than the width of an metatarsalia are separated. The ventral surface of

The nostrils the foot is A weak white fold upper eyelid. are slightly protruding, slightly granular. and opening laterally. The commissure of the goes from the elbow along the side of the lower

mouth is slightly curved upwards. The pupil is arm and fourth finger to the disc, and another

horizontally elliptical. from the heel along the side of the tarsus and

fifth There is a dermal fold from the posterior edge toe to the disc. of the The anal eye, over the tympanum, curving downwards opening is directed posteriorly at the

behind the insertion of the level of the The dermal above to a point just arm. upper thighs. flap

the anal is The fold covers the dorsal margin of tym- the opening short.

The is distinct and its The is panum. tympanum round; tongue heart-shaped and distinctly bifid

is teeth diameter about half the horizontal diameter of posteriorly. The vomerine are situated me- the The distance from the the behind the in small eye. eye to tym- dially just large choanae, two

is about diameter the The slits panum one-fifth of the of oblique groups. vocal are small, situated

in the tympanum. floor of the mouth, near the angles of the

The largest diameter of the disc of the third jaws. The vocal sac is median subgular, moderately

is of finger about 4/5 the diameter of the tym- distensible.

The skin on the back, chin panum; the discs of the fourth and second finger and throat is smooth. are subequal, and of the first much smaller. The The skin on the belly and below the anal opening

the is and skin relative lengths of fingers from shortest to coarsely granular white. The on the longest are 1-2-4-3. The subarticular tubercles are posteroventral surfaces of the thigh, heel and distinct and The distal elbow is round. tubercle of the moderately granular and the skin on the

is is of is fourth finger sometimes bifid. There an inner ventral surface the limbs smooth. oval flat and The colour is metacarpal tubercle no palmar in preservative yellowish. There 284 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

are dark brown spots on the back and dark brown articulation extends to the nostril or beyond. The

is crossbands on the limbs. Dark brown pigment fingers are one-third to half webbed, the web

also present on the ventral surface of the tarsus and reaching the base of the penultimate phalanx of

around the anal opening. It was conspicuous that the fourth finger in all species. The toes are two-

beige specimens preserved in formalin became red thirds to three-fourths webbed. One to two pha-

the back with blackish life the of the fourth free of on spots (in species langes toe are web.

was also able to change its colour. Sometimes the Contents: B. goudoti Tschudi, 1838, B. mada-

back was red at night and beige during the day). gascariensis (Peters, 1874), B. untersteini (Ahl,

" " " The rim of the membrane is B. B. callichromus upper palpebral pig- 1928), hyloides (Ahl, 1929),

mented and the lower rim and B. reticulatus entirely only anteriorly (Ahl, 1928) sp. nov.

iris is and posteriorly. The silvery grey with some

dark surrounded a black pigment, by ring. Differential diagnosis

In life, the back is usually beige to brownish Large species (80-100 mm) in the adult stage. with darker brown spots and markings, sometimes 1. White of and fringes along the lateral edges lower arm red with blackish at The ventral sur- spots night. tarsus and distinct triangular dermal flaps on elbow and

faces of the feet heel B. hindlimbs, and hands, and the present .... madagascariensis (Peters) 2. Fringes and dermal flaps absent . B. Tschudi and anterior goudoti posterior surfaces of the thighs are

violet-red. The iris is creamish and brownish in the However, the half-growns of B. goudoti show low

middle, surrounded by a blackish ring. fringes along the lateral edge of lower arm and

tarsus and small dermal flaps on the elbow and

Remarks. — The holotype of Rhacophorus heel. The half-growns of B. madagascariensis

miniatus with show and than agrees the description above. Only less developed fringes flaps the

its adults. the the relative length of hindlimbs is slightly Therefore, four remaining species are

larger. difficult to distinguish from the half-growns of

goudoti and madagascariensis. In the following

key to the medium-sized species those half-growns

are included. Boophis goudoti group

Nostril from and 1. equidistant tip of snout eye.

Diagnosis. — The members of this group a. Dorsum covered with tubercles and ridges, forming an network, are medium- to large-sized species, with a pref- irregular interorbital distance equals the

width of the eyelid, arm robust, small The upper upper erence for more quiet parts of streams. triangular dermal flaps on heel and elbow . differ from the adults. The dorsum young greatly B. reticulatus sp. nov.

b. is in the Dorsum smooth with a few tubercles, inter- usually tan or grey and often green young. dispersed orbital distance 1V2 X the width of the eyelid, distinct upper There are transverse bars on the limbs. tympanic fold curved, upper arm slender, tubercles on The surface of the is dark posterior thighs usually heel and elbow B. untersteini (Ahl)

Dorsum interorbital distance X the width with light spots. The temporal region and the c. smooth, 11/2 of the fold upper eyelid, tympanic straight, upper arm streak along the canthus is usually dark brown. dermal elbow and robust, small triangular flaps on One or more white subocular blotches triangular heel B. madagascariensis (Peters), half-grown

2. in the and still in of the Nostril nearer to than to tip of snout, are present young some eye a. Dorsum granular B. hyloides (Ahl) adults. There are white fringes along the lateral b. Dorsum smooth, blue-grey with round red-brown spots of the lower arm and tarsus and a small edge (in alcohol) B. callichromus (Ahl)

the Dorsum brown with dark dermal flap on heel and elbow in the half- c. smooth, grey or irregular

and in brown markings (in alcohol) growns some of the adults. Vomerine teeth B. goudoti Tschudi, half-grown are present. The horizontal diameter of the tym- two-fifths is — In and panum to three-fourths of that of the Remarks. the field in the aquarium

The skin the dorsum is I eye. on smooth, granular noticed slight differences among robust tadpoles

with or covered irregular reticulate ridges. The with a moderately developed tail musculature and vocal sac is single median subgular. The tibiotarsal a medium-sized mouth, occurring in quieter parts BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 285

ZMB of callichro- of streams and occasionally in adjacent pools. 30508 A, B, two syntypes Rhacophorus

mus. Type locality: N.W. Madagascar. Coll. Hildebrandt. These differences, mainly in size and relative tail ZMA 6802 (1), 4.IV.72; 6804 (1), 6.II.72; 6805 (2), to distinguish three forms, which length, permitted 19.1.73; 6792 (1 young), 6.XII.70; 7209 (tadpoles and

reared from all collected Perinet were often collected simultaneously. Young frogs young them), 22.X.72; near

(highroad R.N. 2 at km 142), alt. 900 m. — ZMA 6807 were reared from several batches of each form (1), 13.X.72; 6809 (2), 15.XI.72; 6852 (1), 6.1.73; 7211 from different localities, mixed batches being (one young reared from tadpole), VIII.72; 7212 (one

after collection. The juvenile), 15.XI.72; collected in the Mandraka valley (high- segregated soon young emerg-

road R.N. 2 at km 69), alt. 1200 m. — ZMA 7210 (tadpoles ing from each type of tadpole were also clearly and reared from 29.VIII.72 collected young them), along batches. distinct, and so, in the various constantly the road from Moramanga to Anosibe at km 27, alt. 900 m.

Thus it doubt that these ZMA 6801 6.IX.71 collected in Tananarive, alt. appears beyond tadpoles —• (5),

— ZMA 7004 1300 m. (1), 19.IX.71; 6799 (1), 21.111.73; represent three different species. However, the collected at Manjakotompo forest station, Ankaratra mts. reared were difficult to relate with the young 1800-2200 — ZMA 6808 (near Ambatolampy), alt. m. (4),

adults. They showed a typical juvenile colour pat- 14.VIII.71; 6806 (7), 14.VIII.71; collected in Tsinioarivo

km S.E. of Ambatolampy) alt. 1600 m. older. (46 tern, gradually disappearing as they grew

of Since the juveniles these species are of a rather

Natural — The large size, I have compared the whole series from history. species was

adult with found and in or near stagnant or young to the type material of many during day night

water in both forests and ricefields. species. In that manner I have noted that some slowly running

in described in of other Being touched, they inflated themselves. Pairs species are, fact, juveniles

were found in Both nominal species. I found the tadpoles belonging axillary amplexus August. and males and females are calling during day to B. goudoti, B. madagascariensis and B. under- found in steini. night. The tadpoles were slowly running

waters and adjacent pools. Arnoult & Razarihelisoa

(1966) have observed that B. goudoti feeds on

like moth and beetle Boophis goudoti Tschudi, 1838 large prey, grasshoppers,

larvae. found B. also in al- Text-figs. 17-19; pi. I fig. 7, pi. II fig. 1. They goudoti trees, this is climber. though species a poor Reproduction

started at the end of November. observed Boophis goudoti Tschudi, 1838: 77; Guibe, 1947: 439 They

(equals flavoguttatus); Bachmann & Blommers-Schlosser, lumbal amplexus, which I have never noticed. 1975: 17 (DNA); Blommers-Schlosser, 1978: 36 (chromo- The in of attached eggs were deposited lumps 30, somes) .

to rocks and roots in water. Rhacophorus goudoti; Boulenger, 1882: 76; Arnoult & Razari-

helisoa, 1966: 114 (tadpole ecology); Razarihelisoa, 1969:

953 (tadpole); Guibe, 1978: 71-72, figs. 166-171 (equals

obscurus and flavoguttatus). Synonymy. — Rhacophorus kanbergi and

Rhacophorus obscurus Boettger, 1913: 275, XXIII 3; with pi. fig. Rh. fasciolatus were synonymized Rh. calli-

Guibe, 1978: 72 (in synonymy of goudoti). chromus by Guibe, 1978, but are identified with Rhacophorus fasciolatus Ahl, 1929: 478 (new synonymy); B. in the the entire Guibe, 1978: 76 (in synonymy of callichromus). goudoti present paper. Studying Rhacophorus flavoguttatus Ah!, 1929: 479; Guibe, 1947: 439 ZMA series of specimens of Boophis goudoti, I (in of goudoti): Guibe, 1978: 71-72 (in synonymy g synon- have noticed transitional forms from the juvenile ymy of ,goudoti).

to the adult. So it was not difficult to relate the Rhacophorus kanbergi Ahl, 1929: 482 (new synonymy);

Guibe, 1978: 76 (in synonymy ofcallichromus). the Transitional char- young to present species.

exist till the size acters specimens have reached a Material.Boophis— MNHP 4578, female holotype of of about 50 mm. The A and C of Rhaco- goudoti. Type locality: Madagascar. syntypes SMF 6763, male of lectotype Rhacophorus obscurus, phorus fasciolatus are conform to the adults of designed by Mertens, 1967. Type locality: Fianarantsoa, S.E. B. and B still goudoti syntype shows some colour Madagascar. Coll. Voetzkow. characters of the The of Rhaco- ZMB 30527 A, B, C, three syntypes of Rhacophorus juveniles. holotype

fasciolatus. Type locality: N.W. Madagascar. Coll. Hilde- with the B. phorus kanbergi agrees young of brandt. below. goudoti,, of which a description follows ZMB 30517, juvenile holotype of Rhacophorus kanbergi with the material of Rhaco- Type locality: N.W. Madagascar. Coll. Hildebrandt. Comparison type 286 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

TABLE V

callichromus, B. A = Dimensions in mm of Boophis goudoti, B. hyloides, B. untersteini and B. reticulatus. snout to vent;

= of lower C = hand D = hind E = tibia = of foot and B length arm; length; length of limb; length; F length tarsus;

= H = = G foot length; horizontal diameter of tympanum; I horizontal diameter of eye.

ABCD EFGH I

B. goudoti

9 holotype MNHP 80 39 25 132 42 57 38 5 8

$ lectotype obscurus SMF 50 25 15 86 27 38 23.5 3.5 6.5

syntypes fasciolatus' ZMB A 56 31 20 106 36 46 30 3.8 8

B 52 26 17 86 29 35 24 3 6.5

C 49 28 16 91 32 40 26 4 7

15 adults ZMA 56-87 30-43 18-29 105-175 37-55 46-68 31-50 4-6 7-11

holotype kanbergi ZMB 32 15.5 10 52 17 21 15 1.5 3.5

7 juveniles ZMA 30-32 15-16 9.5-10.5 50-54 16-18 21-22 13-15 1.5-1.8 3.5-4.0

2 halfgrown ZMA 40-41 20-21 12.5-14.0 71-75 23 31 18.5-21.0 2.0-2.5 5

B. callichromus

ZMB A 14 69 syntype 38 20 22 31 19.5 2 5 B. hyloides

holotype ZMB 42 20 13 68 23 29 19 2.5 5

B. untersteini

$ holotype ZMB 26 13.5 8.5 45 15 19 12.5 2 4

3 halfgrown $ $ ZMA 27.5-36.0 14.0-19.5 9-12 45-62 15-21 19-25 12.5-17.0 2-3 4.0-5.5

B. reticulatus

s holotype ZMA 35 18.5 11 61 20 26 16 2 4

ZMA 30-36 15-19 9-12 54-61 23-26 14-16 1.8-2.2 3.8-4.2 8 $ $ paratypes 18-20

phorus callichromus shows no morphological dif- The body in the post-axillary region is equal to

ferences from R. kanbergi, but the colour (blue- the head width. The head is somewhat wider than

with red-brown is as The head is one-third of the grey spots) quite different, long. length body

Ahl In fact the The is in lateral was already described by (1928). length. snout round dorsal and

colour resembles that of preserved Boophis micro- profile. The snout is longer than the horizontal

which is with diameter of the and the distance tympanum (Boettger, 1881), green eye, equals

but differs in Mor- between the anterior corners of the The red spots in life, morphology. eye.

phological features to distinguish between species nostrils are nearer to the anterior corner of the

but since the than of the The distance from are scanty in the genus Boophis, eye to the tip snout.

is useful I the nostril to the anterior of the is colour a very character, regard Boophis corner eye

shorter than diameter callichromus (Ahl, 1928) a distinct species. slightly the horizontal of the

is eye. The canthus distinct, concave. The loreal

Description of the juvenile of Boo- region is oblique and slightly concave. The inter-

phis goudoti (see table V for measurements). — orbital region is slightly convex. The ratio between

the of R. kan- the internarial and interorbital distance is about Based on ZMB 30517, holotype 1.

ZMA The is and times bergi, seven metamorphosing young, 7209, interorbital distance one a half

and width The 7210 and 7211, two half-grown specimens the of an upper eyelid. nostrils are

ZMA 7212 and 6792. The ratio of hand length to protruding with lateral openings. The commissure

snout-vent length is 0.31-0.34 (mean 0.33). The of the mouth is straight. The pupil is horizontally ratio of tibia length to snout-vent length is 0.53- elliptical.

is 0.58 (mean 0.55). The ratio of foot length to There a distinct dermal fold from the

is The of the the snout-vent length 0.43-0.51 (mean 0.47). posterior edge eye, over tympanum,

the horizontal the in- ratio of diameter of the tympanum curving downwards to a point just above

the is to that of eye 0.40-0.50 (mean 0.45). Snout- sertion of the arm. The fold covers the dorsal vent length is 30-41 mm. margin of the distinct and round tympanum, its BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 287

diameter is two-fifths to half that of the horizontal The skin on the back, chin, throat and on the

of the is ventral surfaces of the limbs the diameter eye (the tympanum not yet (except thighs)

distance from the is smooth. The skin the and visible in all young). The eye to on belly postero-

diameter ventral the is the tympanum is about half the of the surface of thighs coarsely granular.

The skin anal is tubercular. tympanum. below the opening The largest diameter of the disc of the third The dorsal colour of preserved specimens is

is the diameter of the brown with dark brown finger equal to tympanum; or grey irregular markings.

is in is live the disc of the fourth finger about equal size, The venter uniformly yellowish (in spec-

that of the second finger slightly smaller and of imens the ventral surface of the hindlimbs is

The relative the first finger much smaller. lengths orange). There are distinct, dark brown transverse

of the fingers from shortest to longest are 1-2-4-3. bars on the limbs. Dark brown streaks are present

lateral The subarticular tubercles are small, distinct and ventrally along the white fringes of lower round. is flat tubercle and The surface of the There an inner metacarpal arm tarsus. posterior thighs

distinct is with also and no palmar tubercle. Many supernumer- brown light spots. There are light spots

tubercles the base of the on the flanks. The streaks the canthus and ary are present on fingers. along

dark A The formula of webbing of the hand: between 1 the tympanic region are brown. triangular

2i trace of web, 2e 3i 3e white is both and and a (1), (2), (1 ]/2 ), spot present anteriorly posteriorly

4(1). The web is glandular. of the tympanum. A white band often runs from

The is robust. underneath the anterior of the upper arm When the limbs are part eye obliquely

the dark brown band laid along the side, the knee and elbow overlap. to front, separated by a from

behind. The The adpressed heels overlap slightly. The tibio- the triangular white spot upper and

rim tarsal articulation extends to a point between the lower of the palpebral membrane are pig-

and the of the The inner mented. The iris is with dark brown nostril tip snout. meta- silvery grey a tarsal tubercle is small, elliptical and protruding. reticulation.

There is no outer metatarsal tubercle. A small web The colour in life is similar, but more intense.

connects the tip of the inner metatarsal tubercle

B. V with the first toe to the base of the penultimate Adults of goudoti (see table for measure-

— Ratios of phalanx. The discs on the toes are nearly as large as ments). 20 specimens (holotype

those on the fingers. The fifth toe is slightly goudoti, lectotype obscurus, syntypes of fasciolatus,

longer than the third. The subarticular tubercles and 15 specimens ZMA). The ratio of hand length

The formula is are small, round and protruding. of to snout-vent length 0.30-0.38 (mean 0.33).

the webbing of the foot: 1 (0), 2i (1), 2e (0), The ratio of tibia length to snout-vent length is

3i 3e 4i 4e 5 The 0.53-0.66 The ratio of foot (1), (1), (ll/ 2 ), (H/ 2 ), (l/2 ). (mean 0.60). length lateral metatarsalia are separated. The ventral sur- to snout-vent length is 0.46-0.57 (mean 0.52).

the is A white The ratio of the horizontal face of web glandular. fringe goes diameter of the tym- from the side of the lower and to that of the is 0.46-0.66 the elbow along arm panum eye (mean 0.60).

and another the fourth finger to the disc goes from Snout-vent length is 49-87 mm. the heel along the side of the tarsus and the fifth The adults differ from the juveniles in the

A the toe to the disc. small white tubercle is present on following features: the white fringes along the heel and the elbow. side of the lower arm and fourth finger and along

The anal opening is directed posteriorly at the the tarsus and fifth toe and the white tubercles

level of the The dermal above the elbow and heel upper thighs. flap on disappear during growth.

in In adults heel the anal opening continues a fold on each side some there are still tubercles on the of the anal opening, which results into a A-shaped and elbow. The large white spots underneath the

and the figure. eye near tympanum disappear also. The

The tongue is heart-shaped and distinctly bifid. horizontal diameter of the tympanum is three-

The vomerine teeth are situated behind fifths of the horizontal diameterof the in the medially eye the choanae in two in the diameter of small, oblique groups. adults; consequence largest the 288 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

the is disc of the third finger is two-thirds of di- about 7 sec. The duration of one call about

of the The hindlimb is 0.5 The to about 1500 Hz. ameter tympanum. longer sec. frequency goes up

than in the the young, tibiotarsal articulation Eggs. — In dissected females about 1000, 2 extends to the tip of the snout or beyond. mm in diameter, dark brown with cream.

Sexual dimorphism. — The males have Tadpoles. — Four tadpoles in stage 25 have

well-developed white nuptial pads laterodorsally body lengths of 12-23 mm and total lengths of

in in 26 have on the first and dorsally on the second finger 31-61 mm. Two tadpoles stage body

The and the breeding season. back, chin and throat are lengths of 26-27 mm a total length of 68 mm.

covered with small sharp tubercles in that period. Two tadpoles in stages 31 and 37 have a body

of The males are much smaller than the females. length of 27 mm and a total length 73 mm.

One in 41 tadpole stage has a body length of 27

in — The is and Colour life. ground colour mm a total length of 73 mm. The average ratio

highly variable, from nearly entirely blackish to of tail length to body length in the eight tadpoles

yellowish with blackish spots. The venter is uni- listed above is 1.65 (range 1.5-1.7). Seven meta-

with or without blackish 30-32 30 formly yellow or orange morphosing young measure mm (mean

spots. mm) from snout to vent.

The mouth and lateral view of a tadpole in

— in in Voice. The species was heard August stage 37 (ZMA 7209) are shown figs. 17-18. and and The is ovoid. September during the day evening. The body The nostrils are dorsal, the

made of in are situated at distan- sonagram (fig. 19) a couple axillary openings dorsolaterally equal

in 1971 at 9 from of the and the anterior amplexus (ZMA 6808) August p.m. ces the tip snout

shows the call of the male and the female. The of the The situated dorso- corner eye. eyes are

duration is about It is is of one male call 0.3 sec. a laterally. The spiracular opening sinistral,

pulsed low-pitched note with about 10 pulses in situated at about from the tip of the snout

one note. These notes are repeated about one and the end of the body. The cloacal tube is long

caudal The minute and their frequency is about one note per and dextral to the fin. caudal muscu-

sec. The indistinct dominant frequency is from lature is moderately developed. The caudal dorsal

500-1000 Hz. The call of the female is a low fin is shallow anteriorly. At midlength of the tail,

buzzing sound. It is repeated with intervals of the heigth of the caudal musculature accounts for

1/4 to 3/10 of the total tail height. The mouth is

small and directed ventrally. The margin of the

oral disc is indented laterally. The papillary border

has wide median and a gap anteriorly a narrow

median There of gap posteriorly. are two rows

and papillae laterally, one posteriorly. The horny

beak is heavy. The tooth formula is 1/3 + 3//1 +

1/2 or 1/4 + 4//1 + 1/2.

in Metamorphosis was observed October and the

beginning of November.

The colour of the tadpole is brownish. The tad-

poles were found in gently running and in stagnant

water.

Arnoult & Razarihelisoa (1966) described the

B. well. tadpole of goudoti as The whole develop-

ment took three months in captivity. Their descrip-

with the tion agrees largely more detailed descrip-

tion above. Fig. 19. Sonagram, filter wide, Boophis goudoti. given BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 289

Boophis callichromus (Ahl, 1928) R.N. 2 at km 142), alt. 900 m; and 6836 (3), 13.XI.72; 7002 (tadpoles), 24.IX.72; 7003 (young reared from 7002),

at the same locality, but alt. 1100 m. — ZMA 6848 (1), Rhacophorus callichromus Ahl, 1928: 314; Guibe, 1978: 76, 20.IX.71 collected near Anjozorobe, alt. 1300 m. — ZMA figs. 188-191 (equals untersteini, fasciolatus, kanbergi 7064 and reared from (tadpoles young them), 13.X.71 in part). collected near Foulpointe, sea-level. -— ZMA 7066 (tadpoles

and reared from collected Tam- young them), 29.111.72 near Material. — ZMB 30508 A, B, two syntypes of Rhaco-

poketsa dAnkazobe, alt. 1600 m. — ZMA 7069 (tadpoles and phorus callichromus. Type locality: N.W. Madagascar. Coll.

reared from 1.VII.71 collected near Ranoma- Hildebrandt. young them),

fana, alt. 800 m.

Remarks. — I unable to find mor- was any

phological difference between the half-grown of Natural history. — The males were

B. goudoti and B. callichromus (40 mm). How- noticed by their groaning call, in the evening. They

the colourationof the back is different: call in low less than m above shallow, ever, quite vegetation, 1

bluish with in B. small the the grey reddish brown spots calli- muddy streams. During day species

chromus (measurements table V). was occasionally found in the leaf-axils of large

and in the plants. The eggs tadpoles were found

same streams; occasionally the tadpoles were found

Boophis hyloides (Ahl, in It is 1929) also adjacent shallow pools. a real forest

species, difficult to keep alive in the vivarium

Rhacophorus hyloides Ahl, 1929: 481; Guibe, 1978: 75, where it incessantly attempted to move upwards. fig. 183. In the embrace the breeding season, males tried to

Material. — ZMB 30495, holotype of Rhacophorus any frog, including those of other species who hyloides. Type locality: central Madagascar. Coll. Hildebrandt. happened to be in the same bag. The amplexus

Remark. — The only difference between B. was always lumbal.

hyloides (40 mm) and the half-grown of B.

goudoti is the granular skin of the dorsum in Colour in life. — The ground colour of

B. hyloides (measurements table V). the back in the adults varies from beige to reddish

brown. The posterior part of the thigh is black

with light blue spots. On the limbs dark brown

crossbands Boophis madagascariensis (Peters, 1874) are present, and a transverse dark brown

II line the is The Text-figs. 20-23, 42; pi. figs. 2-3. behind eyes mostly present. venter

is creamish.

Rhacophorus madagascariensis Peters, 1874: 618, pi. I fig. 3;

Guibe, 1978: 73, figs. 178-180 (equals brachychir and

Voice. — The males were heard from herthae). Sep- tember and in in the Rhacophorus brachychir Boettger, 1882: 480; Guibe, 1947: till April July, evening,

439 of (in synonymy madagascariensis); Guibe, 1978: 73 the The occasionally during day. sonagram (fig. of (in synonymy madagascariensis). in 20) made of a male (collected Ranomafana Rhacophorus herthae Ahl, 1929: 480; Ahl, 1931: 157, fig.

104; 1978: sent to the Paris at 9 Guibe, 73 (in synonymy of madagascariensis). 4.VII.71, museum) p.m. on Boophis madagascariensis; Guibe, 1947: 439 (equals bra- the same day, shows that the duration of one note chychir); Bachmann & Blommers-Schlosser, 1975: 17 is about consists 1 sec. The note of a series of (DNA); Blommers-Schlosser, 1978: 35 (chromosomes). rather melodious clicks. The indistinct dominant

Material. — ZMA 6820 (3), 6834 (3), 15.XI.72; frequency is about 1000 Hz. 6.1.73; 6845 (1), 2.IV.72; 6847 (2), 18.11.73; 6849 (1),

26.VIII.72; 7065 (tadpoles), 14.111.72; 7068 (one tadpole),

7070 and reared from 22. VIII.72; (tadpoles them), — Numerous black of about young Eggs. single eggs all collected in the Mandraka X.71; valley, in the nursery 3 mm in diameter were collected in shallow (depth garden of the city of Tananarive (highroad R.N. 2 at km 69), 20-30 in the alt. 1200 — ZMA cm) slowly m. 6835 (1), 17.11.72; 6837 (7), 6.II.72; running water, shade,

6846 13.11.72; 7062 and reared the (1), (eggs, tadpoles young at same place where the males were calling from them), 11.72; 7063 (eggs and embryos), 6.II.72; 7067 the before. The hatched after evening eggs one (tadpoles and young reared from them), 20.X.71; 7071 week. The measures 9 (tadpoles), 6.XII.70; all collected near Perinet (highroad hatchling (fig. 21) mm;

19 290 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

Fig. 20. Sonagram, filter wide, Boophis madagascariensis.

it has ZMA 7070. in 25 have no external gills. The nostrils, the rudiment Two tadpoles stage body

of and of of two suckers, the beginning mouth, eyes lengths of 6.0-6.5 mm and total lengths 17.0-

After the Four in anus are present. fourteen days, tadpole 18.5 mm. tadpoles stages 26-29 have body

reached of 16 The tail is the and a length mm. twice lengths of 9-5-11.5 mm total lengths of 28.5-

tooth is in and have body length. The formula then 1/1 + 1// 33.5 mm. Two tadpoles stages 33 37

1/2 + 2. body lengths of 11.5-14.0 mm and total lengths

of 34.5-41.0 Two mm. metamorphosing young

Tadpoles. — Since the tadpoles and meta- have body lengths of 15 and 16 mm.

show rather variation ZMA in have morphosing young a large 7002. Eight tadpoles stage 25

in size in the different batches, they are treated body lengths of 5.5-15.0 mm and total lengths of

in 26-28 separately. 16.5-45.0 mm. Three tadpoles stages

ZMA 7062 (reared from eggs). Twenty-five have body lengths of 14-15 mm and total lengths

tadpoles in stage 25 have body lengths of 5-9 mm of 42-45 mm. Thirteen tadpoles in stages 34-40

and total lengths of 15-25 mm. Five tadpoles in have body lengths of 15-19 mm and total lengths

9-12 of Sixteen stages 26-28 have body lengths of mm and 45-57 mm. metamorphosing young

Four in reared total lengths of 27-36 mm. tadpoles stages (ZMA 7003 from 7002) have body lengths

and 17-24 34-40 have body lengths of 13-15 mm total of mm (mean 20 mm).

lengths of 39-46 mm. Three metamorphosing ZMA 7068. One tadpole in stage 25 has a body

of of young have body lengths of 15-16 mm. length 13 mm and a total length 39 mm.

in have ZMA in ZMA 7065. Three tadpoles stage 25 7064. One tadpole stage 25 has a body

of body lengths of 7-10 mm and total lengths length of 11 mm and a total length of 32 mm.

22-31 mm. Two tadpoles in stages 28-29 have Three tadpoles in stages 32-37 have body lengths

body lengths of 10-11 and total lengths of 33-35 of 14-15 mm and total lengths of 42-45 mm. Six

Two in and have have of 13- mm. tadpoles stages 30 38 a metamorphosing young body lengths

and of body length of 12 mm a total length 17 mm.

in 37-40 mm. ZMA 7067. One tadpole stage 25 has a body

ZMA 7069. Two tadpoles in stage 26 have body length of 9 mm and a total length of 27 mm. One

and in lengths of 8.5-9-0 mm total lengths of 26-27 tadpole stage 37 has a body length of 18 mm

mm. Two have and total of 54 Three metamor- metamorphosing young body a length mm.

17-18 have lengths of mm. phosing young a body length of 18 mm. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 291

Figs. 21-28. 21-23, Boophis madagascariensis: 21, hatchling, ZMA 7063.X.6; 22, mouth of tadpole, ZMA 7002.X.25; 23, tadpole, ZMA 7066.X.1.5; 24-25, B. untersteini: 24, mouth of tadpole, ZMA 7090.X.12.5; 25, tadpole, ZMA 7090.X.1.5;

26-27, B. tephraeomystax: 26, mouth of tadpole, ZMA 7011.X.25; 27, tadpole, ZMA 7011.X.1.5; 28, B. hillenii: mouth of tadpole, ZMA 7122.X.25. 292 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

ZMA in 26-28 have behind the The fold and the can- 7066. Five tadpoles stages eyes. tympanic

thus bordered with broad brown body lengths of 13-15 mm and total lengths of are a dark band;

Seven in the underneath this band is white 38-46 mm. tadpoles stages 33-37 have region milky

under the body lengths of 14-20 mm and total lengths of with a dark brown longitudinal band

have The hands and feet and the ventral surface of 42-60 mm. Two metamorphosis young body eye.

of 19-21 The ratio of tail the hindlimbs or The lengths mm. average are yellow light orange.

listed is There weak tubercles length to body length in the tadpoles above belly very pale green. are

heel the character- is 2 (range 1.8-2.3). on and elbow, developing into

in the istic The mouth (ZMA 7002) stage 28 and triangular dermal flaps of Boophis madagas-

lateral view in cariensis At the (ZMA 7066) stage 36 are shown during growth. an early stage

in is nostrils colour from into the figs. 22-23. The body depressed. The young's ground changes green are situated dorsally with lateral openings slightly adult's beige or brown. Just after metamorphosis

than the of the The the lateral folds lower arm and are nearer to the eye to tip snout. along tarsus spiracular opening is sinistral, situated at equal not yet present, but during growth they soon be-

distances from the snout and the end of the body. come apparent.

The cloacal tube is dextral to the caudal fin. The caudal musculature is slender. The caudal dorsal Boophis untersteini (Ahl, 1928) bona species fin is shallow anteriorly. At midlength of the tail, Text-figs. 24-25; pi. II figs. 4-6. the height of the caudal musculature accounts for

2/7 of the total tail height. The mouth is small Rhacophorus untersteini Ahl, 1928: 317; Guibe, 1978: 76 of the and directed ventrally. The margin oral of (in synonymy callichromus) . disc is indented laterally. The papillary border has Material. — ZMB 30497, half-grown holotype of Rha- wide median Two of a gap anteriorly. rows pa- cophorus untersteini. Type locality: central Madagascar. Coll. The Hildebrandt. pillae are present. The homy beak is heavy. ZMB 30508 A, B, syntypes of Rhacophorus callichromus. tooth formula is in general 1/4 + 4//1/2 + 2; it Type locality: N.W. Madagascar. Coll. Hildebrandt.

2 - + varies from 1/3 + 3//1/2 + 1/6 + 6//1/2 2, ZMA 7190 and reared from (tadpoles young them),

26 ob- August collected near Mandraka R.N. from stage onwards. Metamorphosis was 1972 valley (highroad 2 at km 69), rivulet Vokanatetezandava, in the nursery served in May and in October and the beginning

of 1200 — ZMA garden the city of Tananarive, alt. m. 7191 of November. The whole development took about (tadpoles and young reared from them), 24.IX.72 collected

2 km 1100 — three months in the vivarium in the hot season. near Perinet (highroad R.N. at 142), alt. m.

ZMA and 7189 (tadpoles one young reared), 29.VIII.72 The development is retarded in the cool dry season collected along the road from Moramanga to Anosibe at km

in which we did not (June-September), period alt. 800 — ZMA and one 25, m. 7193 (tadpoles young observe The of the reared), 12.IX.71 collected near the forest station of Tam- any metamorphosis. tadpoles

poketsa d'Ankazobe R.N. 4), alt. 1600 m. in of the (highroad last eggs (April) hatched the beginning hot season. collected in Habitat. — The tadpoles were The colour of the is brownish. tadpole all slowly running water near forest, at localities.

No adult specimens were caught.

— The hatched Young. young frogs are quite

distinct from the adults. with the ■— The reared from the They agree Taxonomy. young

have description of Rhacophorus herthae Ahl, 1929. tadpoles a typical juvenile colour pattern,

Some hatchlings were maintained in the vivarium which changed drastically during subsequent for another 2-3 which reared 2-3 months, after period they growth. Some young were during had reached of 41 and a body length mm, were months. I have compared these half-grown speci-

B. with the resembling the adults of madagascariensis. mens type material. They appeared to be The ground colour of the back of just meta- similar with the holotype of Rhacophorus unter-

is dark with B. morphosed young light green. There are steini (not R. callichromus, see goudoti).

the brown spots on back, dark brown crossbands The largest half-grown specimen in our collection

line on the limbs and a transverse dark brown was already calling and made attempts to mate. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 293

table V for The Description (see measure- to longest are 1-2-4-3. subarticular tubercles

of tubercle ments). — Based on the holotype R. untersteini, are small, distinct and round. The distal

the two largest specimens of ZMA 7190, and the of the fourth finger is sometimes bifid. There is

inner oval and largest of ZMA 7191. All are half-grown males, an flat metacarpal tubercle no pal-

vocal slits. tuber- having The ratio of hand length to mar tubercle. Many distinct supernumerary

The ratio snout-vent length is 0.33. of tibia length cles are present on the base of the fingers. The

to snout-vent length is 0.55-0.58 (mean 0.58). formula of the webbing of the hand: between 1

The ratio of foot length to snout-vent length is and 2i a trace of web, 2e (1), 3i (2), 3e (1), 4

0.45-0.48 (mean 0.47). Theratio of the horizontal (1). The web is glandular.

diameter of the to that of the is The is slender. When the limbs tympanum eye upper arm are 0.49-0.55 (mean 0.51). laid along the side, the knee and elbow overlap.

The head is much wider than the body, and The adpressed heels overlap slightly. The tibio-

wider than extends the somewhat long. The head length is tarsal articulation to a point between one-third of the body length. The snout is broadly nostril and the tip of the snout. The inner meta- rounded in and in lateral tubercle is and dorsal profile sloping tarsal distinct, small, elliptical pro-

than is profile. The snout is slightly longer the truding. There no outer metatarsal tubercle. A

horizontal diameter of the and somewhat small web the of the inner metatarsal eye, connects tip

shorter than the distance between the anterior tubercle with the first toe to the base of the penul-

nostrils distan- corners of the eye. The are at equal timate phalanx. The discs on the toes are nearly as

from the anterior corner of the and the those the The third and fifth ces eye tip large as on fingers. of the snout. The distance from the nostril to the toe are about equal. The subarticular tubercles are

of the is shorter than the round and anterior corner eye small, protruding. Many distinct super-

horizontal diameter of the The canthus is tubercles the base of the eye. numerary are present on

distinct, angular, somewhat concave. The loreal foot. The formula of the webbing of the foot:

The is concave and inclined inter- 1 2i 2e 3e 4i region laterally. (l/ 2 ), (1), (0), 3i (1), (0), (U/ 2 ),

is between inter- orbital flat. The ratio the 4e The lateral metatarsalia are region (11/ 2 ), 5 (l/ 2 )-

narial and interorbital distance is about 1. The separated. The ventral surface of the web is glan-

interorbital distance is one and a half times the dular. A white fold runs from the elbow along the

width of The nostrils side of the lower the fourth an upper eyelid. are strongly arm and finger to the

the protruding, their openings are lateral. The com- disc, and another white fold runs from heel

missure of the mouth is straight. The pupil is along the side of the tarsus and the fifth toe to

disc. horizontally elliptical. the A small white tubercle is present on the

There is distinct dermal fold from the heel and a poste- elbow.

rior edge of the eye, over the tympanum, curving The anal opening is directed posteriorly at the downwards above the insertion of the level of the The dermal above to a point upper thighs. flap

the the in each arm. The fold covers the dorsal margin of anal opening continues a fold on side

tympanum. The tympanum is distinct and round; of the anal opening, thus forming together a its diameter is half the horizontal diameter of the A-shaped figure.

The distance from the to the The is and bifid. eye. eye tympanum tongue heart-shaped distinctly

is about one-fourth of the diameter of the tym- The vomerine teeth are situated medially behind

the in two small panum. choanae, straight groups. The

The largest diameter of the disc of the third vocal slits are small, situated in the floor of the

about the The vocal is finger is two-thirds of the diameter of the mouth near the angles of jaw. sac

tympanum; the disc of the fourth finger is about median, subgular and moderately distensible.

the of that the The the smooth with equal to diameter the tympanum, of skin on back is dispersed second is smaller and that the skin and finger slightly of small tubercles. The on the chin, throat

first the finger much smaller; the discs are truncate. on ventral surface of the limbs (except the

shortest is smooth. skin the The relative lengths of the fingers from thighs) The on belly and 294 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

of the is the of the caudal fin. The caudal muscula- posteroventral part thighs coarsely gran- on right ular. The skin below the anal opening is covered ture is moderately developed. The dorsal caudal fin

with small white tubercles. There are white tuber- is shallow anteriorly. At midlength of the tail,

heel and the of caudal for cles on the elbow. height the musculature accounts

of total tail is The colour in preservative is beige or brown. 1/3 the height. The mouth small

is distinct and directed The venter uniformly cream. There are ventrally. The margin of the oral

brown dark transverse bars on the limbs. Dark disc is indented laterally, and somewhat medio-

brown streaks are present ventrally along the white posteriorly. The papillary border has a wide

lateral folds lower and The median and on arm tarsus. posterior gap anteriorly a very narrow gap

the is brown with surface of thigh or without posteriorly. There is one row of papillae with

The The cream-coloured spots. region below the can- a few dispersed supernumerary papillae. horny

A is thus and the tympanic region are dark brown. beak heavy. The tooth formula is 1/4 + 4//1 +

from underneath the broad white band runs poste- 1/2 or rarely 1/5 + 5//1 + 1/2. Metamorphosis

rior of the backwards and some- was observed in October and the of part eye obliquely beginning

times another white band runs from underneath November.

of the the front. The colour of the is brownish. the anterior part eye obliquely to tadpole

each These white bands are separated from other

— The hatched rim the Young. young are quite by a dark brown patch. The upper of distinct from the half-grown, especially in coloura- palpebral membrane is pigmented. The iris is tion. resemble the of B. with dark brown reticulation and They young madagas- silvery grey a cariensis in small tubercles the heel and dark brown having on surrounded by a ring. elbow, whereas the lateral folds the lower The colour in life is similar, but more intense. along

arm and tarsus are not yet present. The amount of

the and is the webbing of hands feet also same Tadpoles. — (ZMA 7189-7191, 7193). in B. Seventeen as madagascariensis. tadpoles in stage 25 have body lengths

The colour on the back of just of 12-17 mm and total lengths of 35-47 mm. The metamorphosed

is with a broad chocolate-brown ratio in young olive-green average of tail length to body length stage in marking the middle, extending over the 25 is 1.8 (range 1.7-1.9). Sixteen tadpoles in upper The of this is variable have and eyelids. shape marking (see stages 26-29 body lengths of 18-23 mm II The sides are chocolate-brown total of 47-61 The ratio of pi. figs. 4-5). lengths mm. average interrupted by two light anteriorly and tail length to body length is 1.7 (range 1.6-1.9). green spots

in posteriorly underneath the eye. The dorsal surfaces Nine tadpoles stages 32-39 have body lengths of of the limbs with dark brown The are olive-green cross- 18-26 mm and total lengths of 48-80 mm. bands. The hands and feet The ratio of tail to is 1.8 are orange. belly average length body length is The ventral surface of the hindlimbs Seven light green. (range 1.65-1.85). metamorphosing young is measure 21-25 mm (mean 24 mm) from snout orange.

to vent (ZMA 7189, 7190, 7193). Two metamor-

18 and 19 phosing young (ZMA 7191) measure reticulatus Boophis sp. nov. from snout to vent. mm PI. Ill fig. 1. The mouth and lateral view of a tadpole in

stage 30 (ZMA 7090) are shown in figs. 24-25. Boophis sp. 1; Bachmann & Blommers-Schlosser, 1975: 17 (DNA) nostrils The body is ovoid. The are dorsal, the

openings are situated dorsolaterally at equal distan- Material. — ZMA 7101A, male holotype and ZMA male 13.XI.72 collected Perinet from the of the 7101B, eight paratypes, near ces tip snout and the anterior (highroad R.N. 2 at km 142), alt. 1100 m. corner of the eye. The eyes are situated laterally.

The spiracular opening is sinistral, situated at The present species resembles the other species of

from the of B. is about 1/2 tip the snout and the end the goudoti group. reticulatus readily distin- of the body. The cloacal tube is long and situated guished from untersteini, madagascariensis, hy- BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 295

loides, callichromus and goudoti by the structure finger is equal to the horizontal diameter of the

with of the skin of the dorsum, which is covered tympanum; the disc of the fourth finger is about

and network. tubercles ridges, forming an irregular equal, of the second finger slightly smaller, and of

the first much smaller. The relative lengths of

Habitat. ■— The males were found on a rainy the fingers from shortest to longest are 1-2-4-3.

The evening, calling on leaves of shrubs and trees, subarticular tubercles are small, distinct and

in woodland. oval. The distal the and third along running water open tubercles of fourth

is fingers are bifid. There an inner flat, oval meta-

V Description (see table for measure- carpal tubercle. A white nuptial pad is situated

ments). — The ratio of hand length to snout-vent dorsolaterally on the first finger. There are two

is ratio of oval length 0.29-0.32 (mean 0.305). The flat, palmar tubercles. Many distinct super-

tibia to snout-vent is 0.53-0.60 tubercles of length length numerary are present on the base the

(mean 0.57). The ratio of foot length to snout- fingers. The formula of the webbing of the hand:

is ratio between 2i vent length 0.42-0.50 (mean 0.45). The 1 and a trace of web, 2e (1), 3i (2),

of the horizontal diameter of the tympanum to 3e ( 1), 4 ( 1). The web is glandular.

that of the is 0.48-0.52 Snout- The is robust. When the limbs eye (mean 0.50). upper arm are

vent length in males is 30-36 mm. laid along the side, the knee and elbow overlap.

11 The heels Holotype: head length is mm; head width adpressed overlap. The tibiotarsal artic-

is internarial and interorbital distance ulation extends to between the nostril and 12.5 mm; a point

is 4 mm; snout-vent length 35 mm. the tip of the snout. The inner metatarsal tubercle

The body in the postaxillary region is little is small, distinct and oval. A small web connects

than width the the the inner metatarsal narrower the greatest of head. The tip of tubercle with the

head is somewhat wider than long. The head first toe to the middle of the antepenultimate

is length one-third of the body length. The snout phalanx. There is no outer metatarsal tubercle.

is rounded in round in The discs the dorsal profile, to sloping on toes are slightly smaller than

is the hori- lateral profile. The snout longer than those on the fingers. The subarticular tubercles are

zontal diameter of the and the eye, equal to distinct, small and round. There are some super-

distance between the anterior of the tubercles the base of corners eyes. numerary on the toes. The

The third and fifth about in The nostrils are at equal distances from the tip of toe are equal size. the the anterior of formula of snout and corner the eye. The the webbing of the foot: 1 (0), 2i (1),

canthus is is 4i distinct, concave. The loreal region 2e (0), 3i (1), 3e (1), (1), 4e (1), 5 (0).

distinctly concave and oblique. The interorbital The ventral surface of the web is glandular. The region is slightly convex. The ratio between the lateral metatarsalia are separated.

internarial and interorbital distance is 1. The inter- The anal opening is directed posteriorly at the orbital distance is about the width of dermal of an upper upper level the thighs. The flap above

The nostrils with anal is continues in eyelid. are protruding, lateral the opening tubercular, and a

The the mouth is openings. commissure of straight. fold on each side of the anal opening, together

The is pupil horizontally elliptical. forming a U-shaped figure.

There is a distinct straight dermal fold from the The tongue is heart-shaped and distinctly bifid.

of the the The vomerine teeth situated posterior edge eye, over tympanum, are medially behind

to a behind the insertion of the choanae in two The reaching point just large oblique groups. vocal the The fold the dorsal of the arm. covers margin slits are small, situated in the floor of the mouth,

The is rather its tympanum. tympanum distinct, near the angles of the jaws. The vocal sac is single, horizontal diameter is about half the horizontal median, subgular, and noticeably distensible. diameter of the The distance from the eye. eye to The skin on the back is covered with elevated the is about half the tympanum horizontal diameter ridges and tubercles, forming an irregular net- of the work. The tubercles tympanum. are most numerous on the

The diameter of the disc of the third and largest upper eyelid the end of the back. There are 296 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

and also tubercles on the knee, a row of three or four Fringes dermal flaps are lacking on the limbs.

Vomerine teeth situated longitudinally on the middle of the shank, are present. The horizontal diam-

side of is half that of a row along the of the tarsus, a row along the eter the tympanum one-third to

lower and tubercles the The skin the dorsum is side of the arm, dispersed on eye. on smooth, finely

vocal is the dorsal middle of the lower arm. Large white granular or verrucose. The sac single,

the median and is triangular tubercles are present on heel and subgular. The snout rounded. The

elbow. There are fringes along the side of the fifth tibiotarsal articulation extends to the centre of

is the but extends far the toe and fourth finger. The skin on the belly eye or beyond, never as as

coarsely granular. The skin on the chin and postero- tip of the snout. The discs are well developed.

ventral part of the thigh is granular. There are The fingers either have a trace of web, or they are

The three- distinct, large white tubercles beneath the anal to one-fourth webbed. toes are about

the opening, and on the anal sheath. The remaining fourths webbed, one to two phalanges of fourth

ventral surface of the limbs is smooth. toe are free of web.

The colour in preservative, same as in life: the Contents: B. tephraeomystax (Dumeril, 1853),

B. B. back is dark brown, the flanks are light brown idae (Steindachner, 1867), granulosus

with whitish and the limbs brown B. hillenii spots are light (Guibe, 1975), sp. nov.

with indistinct dark brown crossbands. There is a

white line along the canthus and from the nostril Differential diagnosis

to the of the snout. There is a white band from of tip 1. Nostrils much nearer to tip snout than to eye . underneath the anterior of the B. tephraeomystax (Dumeril) part eye, obliquely 2. Nostrils from of and equidistant tip snout eye. to the front. A large white triangular patch extends a. Skin of dorsum verrucose . B. granulosus (Guibe) from underneath the of the posterior part eye, b. Skin of dorsum finely granular ($), tibiotarsal

underneath the the insertion of the articulation the centre of the . tympanum to reaching eye B. idae (Steindachner) arm. The ventral surface is cream with some c. Skin of dorsum smooth ( $ ), tibiotarsal articulation scattered brown The rim of the pigment. upper B. hillenii nov. reaching the nostril .... sp. palpebral membrane is pigmented. The iris is

with dark brown Remarks. — it observed that silvery grey a reticulation, sur- Initially, was

rounded black In three distinct from tad- by a ring (in preservative). life, young Boophis developed

the iris is silvery in the middle, with an orange poles, collected in temporary rainpools, in the

(mixed with brown) periphery. forest around Perinet. High numbers of just

The with of of these B. hille- paratypes agree the description above. emerged young one species ( nii), together with single specimens of the other

— B. and B. Etymology. The specific name is derived two ( granulosus idae)) were collected

the It from latin reticulum meaning net. refers to on the vegetation above one pool during daytime

the tubercles and ridges on the back, forming an on April 4, 1972. Three different tadpoles were

irregular network. collected at various occasions, and could be linked

to the B. young Boophis species by rearing. granu-

losus is readily distinguished from the other two

the skin of the which is Boophis tephraeomystax group by back, coarsely granular,

feature visible in the The other a already young.

— this Diagnosis. The members of group two are difficult to distinguish as adults, but the

are medium-sized which breed in tem- and that species, tadpoles young are so clearly different,

The dorsum is I have porary pools. yellow-brown (often described one of them as a new species,

in silver-grey preservative), mostly covered with B. hillenii. B. tephraeomystax is a coastal species,

dark speckles. There are indistinct transverse bars breeding also in temporary pools. Its tadpole

the on limbs. The flanks and the anterior part of closely resembles the tadpoles of the afore-men-

the thighs are mostly also dark with light spots. tioned species. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 297

rather but still in the Boophis tephraeomystax (Duméril, 1853) open country, shrubbery, Text-figs. 26-27, 29; pi. II fig. 7. often close to human habitation. Some specimens

while seated in the were caught sun during dry

Polypedates tephraeomystax Dumeril, 1853: 153; Boettger, in the The spells rainy season. species has appar- 1881: 505. ently the greatest tolerance for drought and heat Polypedates crossleyi Peters, 1874: 618. of all This with the low Polypedates dispar Boettger, 1879: 86; Boettger, 1881: 507, Boophis species. agrees

V fig. 18. DNA of this pi. amount (7 pg diploid) species, Rhacophorus tephraeomstax; 1882: Boulenger, 78; Boettger, which could be an adaptation to fast development 1913: 276 (equals dispar); Ahl, 1931: 200, fig. 105

" - - - - in an area of limited rainfall. B. (equals dispar); Guibe, 1978: 65, figs. 140-147 (equals tephraeomystax

crossleyi, boulengeri, difficilis,hildebrandti,doulioti). has the smallest size of the genome Boophis species Rhacophorus hildebrandti Ahl, 1925: 18. examined (Bachmann & Blommers-Schlosser, Rhacophorus doulioti Angel, 1934a: 169.

Boophis tephraeomystax; Bachmann & Blommers-Schlosser, 1975).

1975: 17 (DNA); Blommers-Schlosser, 1978: 37 (chromo- The call of the males is heard in rainy nights, somes). it resembles the yelping of a small dog. Calling Boophis aff. tephraeomystax; Blommers-Schlosser, 1978: 37 males of this situated few (chromosomes). species were always a

each metres from other. They were observed

Material. — MNHP 1660 and 4584, syntypes of Poly- moving down trees in the forest edge towards the pedates tephraeomystax.Type locality: Nossi Be and Mada- ground, near pools. The amplexus is gascar. Coll. Boivin. axillary.

of in SMF 6766, 9 lectotype (designed by Mertens, 1967) The tadpoles were found sunlit temporary Polypedates dispar. Type locality: Nossi Be. Coll. Stumpff. rainpools with abundant vegetation. ZMB 10472, $ holotype of Rhacophorus hildebrandti.

Type locality: N.W. Madagascar. Coll. Hildebrandt. dou- MNHP 1891-356 and 357, syntypes of Rhacophorus

table VI for measure- lioti. Type locality: Morondava. Coll. Douliot. Description (see

ZMA 6766 (5), 1-15.IV.71; 6767 (1), 19.IV.71; 6758 ments). — The ratio of hand length to snout-vent (11), 26.IV.71; 6759 (2), 5.IV.71; 6765 (7), 22.IV.71; length is 0.26-0.31 (mean 0.28). The ratio of tibia 6775 (1), 15.1.71; collected near Tulear, S.W. coast. — to snout-vent is 0.49-0.57 ZMA 6785 (3), 23.IV.72; 6786 (10), 29.XI.72; 7010 (tad- length length (mean

reared from them), 2.V.72; 7011 The ratio of foot poles and young (tad- 0.52). length to snout-vent Ankarafantsika poles), all collected near Ampijoroa, 2.V.72; length is 0.37-0.47 (mean 0.42). The ratio of the forest (highroad R.N. 4 at km 465). — ZMA 6787 (1 9 ), horizontal diameter of the to that of the 10.11.72 collected near Tamatave, E. coast. — ZMA 6783 tympanum

km of (2 S $), 13.11.72 collected near Foulpointe, 60 N. eye is 0.45-0.60 (mean 0.50). Snout-vent length is

Tamatave. — ZMA 6784 (1), 1.VIII.72 collected near 35-50 mm. Ivoloina, 13 km N. of Tamatave. B. tephraeomystax resembles closely the species

of the both in Mantidactylus depressiceps group, Natural history. — B. tephraeomystax is colouration and external morphology. However, in a common species all coastal areas of Mada- the of the M. have both species depressiceps group visited It is the gascar by us. probably only species an inner and outer metatarsal tubercle, while B. of Boophis occurring in the S.W., almost the driest tephraeomystax has only an inner metatarsal tu- it is found in part of the island, where the irrigated like all of the bercle, just species genus Boophis area around Tulear. There, as well as in the de- (see Blommers-Schlosser, 1979). ciduous western coastal forest (Ankarafantsika), it

leads a rather hidden life, sheltering in leaf-axils

of plants, like bananas, during the dry season. Sexual dimorphism. — The male has

But when the weather turns rainy, the frogs are white nuptial pads, located laterodorsally on the

first and The slits found on the vegetation, also in daytime. Popu- second fingers. vocal are small,

this situated in floor the lations of species (perhaps subspecies: dif- the of mouth, near the angles

ferences in the karyotypes are found between of the jaws. The vocal sac is single, median sub-

Blommers- and The specimens of the S.W. and E. coast, see gular moderately distensible. female is

humid than skin Schlosser, 1978) from the eastern coastal larger the male, the on the back is

far from hidden. are found there in while that of the male is area, are They smooth, finely granular.

20 298 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

TABLE VI

B. idae and B. A = = of Dimensions in mm of Boophis tephraeomystax, B. hillenii, granulosus. snout to vent; B length

= = = lower arm; C = hand length; D length of hind limb; E tibia length; F length of foot and tarsus; G = foot

H = of = horizontal diameter of length; horizontal diameter tympanum; I eye.

ABCD EFGHI

B. tephraeomystax

2 9 9 syntypes MNHP 48-49 23 14 77 25-26 32 22

9 lectotype dispar■ SMF 49 23 14.5 81 28 36 23

$ holotype hildebrandti 40 18 11 64 21 25 17 2.5 5.5

syntype doulioti MNHP 38 17 11 55 20 24 16.5

7 9 9 ZMA (6758,

6786-87) 41-50 18-24 11-16 65-84 21-27 26-35 16-24 2.5-3.5 4.5-6.0

3 $ $ ZMA (6783,6786) 35-42 17-18 10-11 56-64 19-22 23-26 16-18 2.5-3.0 5.0-5.2

B. hillenii

8 holotype ZMA 30 15 10 50 15 21 14 1.4 3.8

3 paratype ZMA 30 15 10 50 16 20 14 1.5 4

B. idae

juvenile holotype WM 18 8 5 26 9 12

holotype femoralis BMNH 29 44 13.5 18.5 12.5

- - holotype catalai- ■ MNHP 30 13.5 42.5 15 20 12.5

15.5-16.0 4 5 $ $ ZMA 33-36 10.5-11.0 51-52 16-17 23-24 15.0-15.5 1.5-1.7

B. granulosus

7 $ $ ZMA 35-40 17-19 10.0-11.5 55-61 18-20 22.5-26.5 16-18 1.8-2.2 4-5

1 9 ZMA 36 17.5 10.5 56 18 25 16.5 1.9 4

in life. — The colour is ratio of tail is 2.1 Colour ground average length to body length

brown. The back is covered with beige or light (range 1.8-2.3). Twenty metamorphosing young

irregular brownish spots and markings. There are measure 15-20 mm from snout to vent (mean

The irregular brown crossbands on the limbs. 18 mm).

with A mouth and in flanks are covered light yellow spots. light The lateral view of a tadpole stage

yellow longitudinal lateral band is mostly present. 37 (ZMA 7011) are shown in figs. 26-27. The

is ovoid. Thebelly is cream coloured. body The nostrils are situated dorsally

with anterolateral openings at equal distances from

— The Voice. species was heard on rainy

evenings, in April in Ampijoroa, in January in

Tulear, in February and July in Foulpointe. The

of male sonagram (fig. 29) made a (ZMA 6775)

in 1971 10 shows that the call January at p.m. con-

sists with of a series of melodious notes, a duration

short clicks of 0.3-0.4 sec. each, interrupted by very

and of 0.06 sec. The major frequencies are 1500

Hz. A similar made of another 3000 sonagram was male (ZMA 6785).

Tadpoles. — Four tadpoles in stages 26-29 have of and total body lengths 8-10 mm lengths of The ratio tail 24-29 mm. average of length to body length is 1.9 (range 1.8-2.0). Eleven tadpoles in stages 38-42 have body lengths of 12-

16 and total mm lengths of 39-49 mm. The filter Fig. 29. Sonagram, wide, Boophis tephraeomystax. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 299

the tip of the snout and the anterior corners of the The present species is closiest allied to the sibling

The and directed B. It dis- eyes. eyes are large laterally. species, idae (Steindachner, 1867). was

The spiracular opening is sinistral, situated at covered in the field that the larva and juvenile are

the the in B. about 5/8-2/3 from tip of the snout and clearly different both. hillenii can be distin-

is the end of the body. The cloacal tube located on guished from B. idae by the relatively longer

right side of the caudal fin. The caudal muscula- hindlimbs.

ture is slender. The dorsal and ventral caudal fin

are thickened alongside the caudal musculature in

— The Natural history. males were anterior in the part of the tail (see dotted line calling in the shrubbery around a temporary rain- fig. 26). At midlength of the tail, the height of pool, in forest, during a rainy night. The tadpoles the caudal musculature represents about 2/5 of found in that and the the were pool young on the total tail height. The mouth is small and leaves of marshy plants, above that pool. The directed anteroventrally. The papillary border has juvenile (7125) was collected from the leaf-axil median and small a wide gap anteriorly a very one banana. is of a The species sympatric with B. medioposteriorly. There are two rows of papillae. granulosus (Guibe, 1975b) and B. idae (Stein- The oral disc is indented laterally. The tooth for- dachner, 1867). mula is 1/4 + 4//1 + 1/2, or 1/3 + 3//1 + 1/2.

beak is well The The horny developed. upper part

the of lower beak is black. Description (see table VI for measure-

in — The ratio Metamorphosis was observed the beginning ments). of the lower limb length

of May in Ampijoroa. I could not find tadpoles to snout-vent length is 0.50. The ratio of the hand

in Tulear between February and April; they prob- length to snout-vent length is 0.33. The ratio of

ably had already metamorphosed, the wet season the hindlimb length to snout-vent length is 1.67.

being shorter in the southern part of Madagascar. The ratio of the tibia length to snout-vent length

The colour in life of the tadpole is yellowish is 0.50. The ratio of the foot length to snout-vent

with beige spots and becomes mostly greenish in length is 0.47. The ratio of the horizontal diameter the late The is white, of the to that of the is stages. belly milky speckled tympanum eye 0.37-0.38. with in its minute dark spots anterior part. Snout-vent length of males 30 mm.

head head width 10 Holotype: length 9 mm,

internarial and interorbital distance is 3 — The dorsum of the is dark mm, mm; Young. young

snout-vent 30 mm. I never length green (a juvenile colour, saw a green The body in the postaxillary region is little adult), and sometimes beige (as the adult). A light narrower than the greatest width of the head. The yellow longitudinal lateral band is present as in head is somewhat wider than The snout is the adult. The limbs are beige with dark brown long.

rounded in dorsal and crossbands. acuminately profile, sloping

in lateral profile. The snout projects beyond the

leading edge of the lower jaw. The snout is much

than the horizontal diameter of the longer eye Boophis hillenii sp. nov. and equal to the distance between the anterior Text-figs. 28, 30; pi. Ill fig. 4. of the The corners eye. nostrils are at equal

distances from the of snout and the anterior Boophis aff. idae•;; Blommers-Schlosser, 1978: 37 (chromo- tip somes). corners of the The canthus is eye. distinct, straight

and slightly angular. The loreal region is barely

Material. — ZMA 7123A, $ holotype, ZMA 7123B, The is concave. interorbital region flat. The ratio $ paratype, 24.XII.72 collected near Perinet (highroad

2 between the internarial distance and interorbital R.N. at km 142), alt. 900 m.

ZMA 7122 (tadpoles and young reared from them), 20.1. distance is The interorbital distance is 1 times 1. y2

73; 7124 (young), 4.IV.72; collected at the locality. — type the width of an upper eyelid. The nostrils are ZMA 7125 (1 juvenile), 3.VII.71 collected near Ranomafana the is lateral. The (highroad R.N. 25), alt. 800 m. barely protruding, opening 300 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

is commissure of the mouth is straight. The pupil small, situated in the floor of the mouth, near the horizontally elliptical. angles of the jaws. The vocal sac is median,

There is the distensible. a dermal fold from posterior edge subgular and barely of the the downwards The skin on the dorsum and the ventral eye, over tympanum, curving on

the insertion The fold surfaces of the is smooth. to a point above of the arm. limbs (except thethighs)

The skin chin covers the dorsal margin of the tympanum. The on the and throat is finely granular.

its diameter is The skin and tympanum is distinct, horizontal on the belly on the posteroventral about 2/5 of the horizontal diameter of the eye. surface of the thighs is coarsely granular.

distance from the the is The colour in The eye to tympanum preservative is silvery grey with

dark about half the horizontal diameter of the tym- small round brown spots. The flanks are dark

brown with whitish The limbs panum. large spots. are

The largest diameter of the disc of the third light brown with irregular dark brown transverse

dark finger is equal to the horizontal diameter of the bars and brown spots. The anterior part of

the is with tympanum; the discs of the fourth and second thighs whitish large brown spots. The

fingers are about equal, but those of the first posterior part of the thighs is brownish. The fingers are much smaller. The relative lengths of venter is evenly cream. The iris is blackish and the

from shortest 1-2-4-3. rim of the is the fingers to longest are upper palpebral membrane pig-

The subarticular tubercles are small, round and mented.

this protruding. The distal tubercle on the fourth Unfortunately we made no diapositives of

is bifid. One distinct tubercle the colour in is the finger supernumerary species; pattern alcohol same as is the base of each There is in B. that the back is in present on finger. an idae, except silvery grey

and flat and in inner flat, oval metacarpal tubercle, two the present species brown or beige B. idae.

small oval palmar tubercles, a large and a one. is Two An oval unpigmented glandular nuptial pad Tadpoles. — tadpoles in stage 25 have

the first There of 3.5-4.0 present laterodorsally on finger. body lengths mm and total lengths of

the is a trace of web between the fingers. Between 9.5-10.0 mm. Eight tadpoles in stages 26-30 have fourth third from and total and finger the web extends the body lengths of 4.0-6.5 mm lengths of middle of the antepenultimate phalanx of the 10.0-18.5 mm. Eleven tadpoles in stages 31-41

the base have fourth to of the antepenultimate phalanx body lengths of 6-9 mm and total lengths of of the third. The web is 17-23 The ratio of tail glandular. mm. average length to body

When the laid knee in limbs are along the side, the length the twenty-one tadpoles listed above is and elbow overlap. The adpressed heels overlap 1.7 (range 1.5-1.9). Thirty metamorphosing noticeably. The tibiotarsal articulation extends to young have body lengths of 9-10 mm.

metatarsal tubercle is The mouth and the nostril. The inner small, lateral view of a tadpole in

There is in elliptical and protruding. no outer meta- stage 35 (ZMA 7122) are shown figs. 28 and 30.

The discs The is ovoid. The the tarsal tubercle. on the toes are subequal body nostrils are dorsal, to those on the fingers. The subarticular tubercles opening is directed to the front and situated at

fifth is are small, round and protruding. The toe equal distances from the tip of the snout and the

to the third in The formula of the anterior corners of the The equal length. eye. eyes are directed

of the foot: 1 2i 2e The is webbing (y 2 ), (1), (0), 3(1), laterally. spiracular opening sinistral,

3e 4i (1), 4e 5 ( 1). The ventral surface situated at about 2/3 from the of the snout (V2) > (1), tip of the web is glandular. The lateral metatarsalia and theend of the body. The cloacal tube is located are separated. on the right side of the caudal fin. The caudal

The anal opening is directed posteriorly at the musculature is slender. The caudal fin is well

level of the The anal sheath is short. and the The upper thighs. developed goes on to body. dorsal

is and ventral caudal The tongue distinctly bifid posteriorly. The fins are thickened alongside vomerine teeth are situated medially behind the the musculature in the first half of the tail (see

in The vocal slits dotted in choanae two oblique groups. are line fig. 30). At midlength of the tail, BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 301

and the height of the caudal musculature represents Natural history. — Only in October

is in about 1 /5 of the total tail height. The mouth November we heard the males calling the small and directed anteroventrally. The papillary evening, near stagnant, sunlit pools. One specimen border has median There in a wide gap anteriorly. are was found resting the leaf-axil of a banana plant

1-2 rows of papillae. The oral disc is round. The near the same pool, during the day. The tadpoles tooth is which the formula 1/4 + 4//1 4-1/2 or 1/5 + 5// were found in the same pool near adults

beak is well The found in Mandraka. In 1 + 1/2. The horny developed. were Perinet we found

the lower black. with B. upper part of beak is the species together granulosus (Guibe,

~ The colour in life of the is and B. hillenii tadpole yellowish, 1975) sp. nov. with a broad blackish middorsal line and blackish lateral lines. There is a white spot on the tip of

— I with Guibe Taxonomy. agree (1978) the snout. The belly is silvery. that R. femoralis (holotype examined by Dr. M. S. took in The entire development 3-5 weeks our catalai Hoogmoed, Leyden) and R. are synonyms rearings. and B. of B. idae. Since Boophis idae hillenii are

much I of the very alike, give only a description Young. — The ground colour in life is silvery. features of B. idae in which it differs from B. The back is covered with There light green spots. examined and hillenii. All remaining features are are brown crossbands on the limbs. A large brown be the in B. hillenii. proved to same as is between spot present the eyes and a brown line is the canthus and present along above the tym-

is VI panum. The webbing formula the same as in Description (see table for measure-

the adult. — The tibiotarsal articulation extends ments). The ratio of the lower arm to snout-

the is 0.44-0.47 The ratio beyond eye. vent length (mean 0.45).

of the hand length to snout-vent length is 0.28-

— The Etymology. present species is ded- 0.32 (mean 0.30). The ratio of the length of the

the Dr. is icated to my colleague, herpetologist D. hindlimb to snout-vent length 1.42-1.55 (mean

Hillenius. 1.50). The ratio of the tibia length to snout-vent

length is 0.44-0.50 (mean 0.47). The ratio of the

foot length to snout-vent length is 0.40-0.45

0.43). The ratio of the horizontal diameter Boophis idae (Steindachner, 1867) (mean of the tympanum to that of the is 0.33-0.42 Text-figs. 31-32, pi. Ill figs. 2, 5. eye (mean 0.38). Snout-vent length 29-36 mm.

The snout is rounded in dorsal and Hyperolius idae Steindachner, 1867: 52, pi. V fig. 10. profile

Rhacophorus idae; Guibe, 1978: 68 and (equals femoralis bluntly rounded in lateral profile. The snout is catalai) slightly longer than the horizontal diameter of the Rhacophorus femoralis Boulenger, 1882: 466. The canthus is indistinct, rounded and Rhacophorus albiventer.; Ahl, 1929: 478 (substitute name eye. slightly

for femoralis). concave. The interorbital distance is times the

Rhacophorus catalai Angel, 1934b: 247. width of of an upper eyelid. The commissure the

mouth is curved Material. — WM 14845, the holotype of Hyperolius upwards. idae. Type Coll. Pfeiffer. is its locality: Madagascar. The tympanum rather indistinct, horizontal BMNH 1947.2.27.60, the holotype of Rhacophorus femo- diameter is about 1/3-2/5 of the horizontal di- ralis. Type locality: East Betsileo. Coll. Cowan. ameter of the MNHP 1933-237, the holotype of Rhacophorus catalai. eye. Fianarantsoa. Coll. Catala. the the Type locality: When limbs are laid along side, the knee

ZMA 7014 (1), 13.X.72; 7015 (1), 7196 (tad- 15.XI.72; and elbow meet. The adpressed heels meet or pole), 13.X.72; 7198 (tadpoles), 13.X.72; 7199 (tadpoles tibiotarsal and reared from collected The articulation ex- young them), 13.X.72; in the slightly overlap.

Mandraka 2 km 1100 — valley R.N. at alt. m. the formula (highroad 69), tends to centre of the eye. The of the

ZMA 7197 (1), 13.XI.72 collected near Perinet (highroad of the foot: 1 2i 2e webbing (0-l/ 2 ), (1), (0), 2 km 1100 7200 R.N. at 142), alt. m; ZMA (one young), 3i 3e 4i 4e 5 4.IV.72 at same alt. (1), (0), (U/ -2), (U/ 2), (0-l/ ). collected the locality, but 900 m. 2 r 2 - 302 R. M. A. BLOMMERS-SCHLOSSER BIOSYSTEMATICS OF BOOPHIS

The skin is the of and there on the back finely granular (in row large papillae, are some super-

The oral disc male). numerary papillae lateroposteriorly.

The ground colour in life of the dorsal surface is indented laterally. The tooth formula is 1/3 +

is yellowish brown. The back is covered with 3//1 + 1/2 - 1/5 + 5//1 + 1/2. The horny beak is

minute surrounded with dark well the rim of the lower beak light green spots developed; upper

brown. The flanks, groin and anterior part of the is black.

The in is thigh are turquoise with large brown spots. The colour life of the tadpole greyish.

A white posterior part of the thighs is brownish. There are white spot on the tip of the snout and a

irregular brown crossbands on the limbs. The transversal bar on the middle of the back is most

discs The throat is and the distinct in the earlier of The are orange. yellow stages development. belly white. The iris is orange-brown. belly is silvery.

In preservative, the general colouration is beige The entire development took two months in our

or brown with irregular small brown spots. The rearings.

turquoise area is whitish, with dark brown spots.

The is The iris is with — The colour in life of the belly cream. silvery grey a Young. ground

rim of is covered dark brown ring. The upper the palpebral back greenish or brownish. The back is

and with with membrane is pigmented, sometimes the lower minute green spots, which are rimmed

rim black. On the heel and too. upper arm, thigh, tarsus

turquoise longitudinal bands are present. Irregular

— Seven in brown the Tadpoles. tadpoles stage 25 dark crossbands are present on limbs.

have body lengths of 3-6 mm and total lengths The hands and feet are yellow. The belly is uni-

ratio of tail creamish. The is in of 8.0-14.5 mm. The average length formly webbing the same as the

in this is adult. to body length stage 1.5 (range 1.4-1.6). The tibiotarsal articulation extends to the

Nine in 26-29 have of centre of the tadpoles stages body lengths eye.

6.5-9-0 mm and total lengths of 16.5-25.0 mm.

The ratio of in average tail length to body length

Thirteen tad- these stages is 1.7 (range 1.6-1.8). Boophis granulosus (Guibé, 1975)

poles in stages 31-41 have body lengths of 9-13 Text-figs. 33-34; pi. Ill fig. 3, pi. IV fig. 7.

and The mm total lengths of 25-38 mm. average

ratio of tail length to body length is 1.9 (range Rhacophorus granulosus Guibe, 1975b: 1083, fig. 3; Guibe, 1978: 66, figs. 148-149. 1.75-2.10). Eight metamorphosing young have

Material. — MNHP the of R. body lengths of 14.5-16.0 mm (mean 15 mm). 1953-150, holotype granulosus. Type locality: forest of Moramanga. and view in The mouth lateral of a tadpole ZMA 6853 (8), 13.XI.72; 7201 (eggs deposited by ZMA

28 are shown in 31-32. and reared 22.X. stage (ZMA 7196) figs. 6853); 7202 (tadpoles young from them),

2 km The body is ovoid. The nostrils are dorsal, the 72; collected near Perinet (highroad R.N. at 142), alt.

1100 m; 7203 (one young), 7204 (tadpoles and and situated 4.IV.72; openings are directed to the front at reared collected the young from them), 20.1.73; at same equal distances from the tip of the snout and the locality, but alt. 900 m.

anterior corners of the eyes. The eyes are directed

laterally. The spiracular opening is sinistral, Natural history. — The species was

in situated at about 1/2-5/8 from the tip of the found on a rainy evening November on leaves

and of snout the end the body. The cloacal tube of shrubs and trees around a pool, near forest.

is located the fin. on right side of the caudal The males were calling loudly. One day later,

The caudal musculature is slender. The caudal female In a deposited blackish, very sticky eggs. fin is well At developed. midlength of the tail, the same pool we had found jelly, containing very

the of the caudal musculature larvae weeks which height represents young (3 before), developed

1 of the total in about /5 height. The mouth is small our rearings to the present species. Close to the

and B. directed anteroventrally. The papillary border same pool one specimen of idae (Steindachner, has wide median is a gap anteriorly. There one 1867) was found on the leaves, and on the BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 303

Figs. 30-37. 30, Boophis hilleni i: tadpole, ZMA 7122.X.6; 31-32, B. idae: 31, mouth of tadpole, ZMA 7196.X.25; 32, tadpole, ZMA 7196.X.3; 33-34, B. granulosus: 33, mouth of tadpole, ZMA 7202.X.25; 34, tadpole, ZMA 7202.X.3; 35-37,

B. microtympanum: 35, hatchling, ZMA 7086.X.12.5; 36, mouth of tadpole, ZMA 7085.X.12.5; 37, tadpole, ZMA

7006.X.1.5. 304 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

ground Aglyptodactylus madagascariensis (Du- have body lengths of 7.5-9-0 mm and total lengths

of which also found larvae of 21.5-25.0 The ratio of tail meril, 1853), we young mm. average length in the pool. The tadpoles collected in January to body length is 1.85 (range 1.7-2.0). Eighteen

in forest. were found a pool as well, near tadpoles in stages 31-40 have body lengths of 9-

12 mm and total lengths of 26-36 mm. The av- table VI for Description (see measure- ratio of tail in these erage length to body length ments). — The ratio of hand length to snout-vent stages is 1.95 (range 1.8-2.0). Sixteen metamor- is The ratio of tibia length 0.28-0.30 (mean 0.29). 11-14 phosing young measure mm (mean 13 mm)

to snout-vent is 0.50-0.53 length length (mean from snout to vent.

The ratio of foot to snout-vent 0.51). length mouth in and in The stage 35 the lateral view length is 0.44-0.47 (mean 0.45). The ratio of the stage 31 of a tadpole (ZMA 7202) are shown in horizontal diameter of the to that of the tympanum figs. 33-34. The body is ovoid. The nostrils are is 0.43-0.49 Snout-vent length eye (mean 0.46). dorsal with anterolateral openings, situated at

35-40 mm. equal distances from the tip of the snout and the The is related B. idae species closely to (Stein- directed anterior corners of the eyes. The eyes are dachner, 1867) and B. hillenii nov.; therefore sp. laterally. The spiracular opening is sinistral, sit- only the differential characters are describedbelow. uated at about 2/3 from the tip of the snout and The skin of the dorsum is The verrucose. the end of the body. The cloacal tube is located is nostrils are strongly protruding. The foot en- on the right side of the caudal fin. The caudal the of tirely webbed, except penultimate phalanx musculature is slender. The caudal fin is well the fourth toe, which is free of web. The anal developed. At midlength of the tail, the height of is directed midlevel opening posteroventrally at the caudal the musculature represents about I/4 of of the thighs. total tail height. The mouth is small and directed The ground colour in life is yellowish brown anteroventrally. The papillary border has a wide in The back is with (silvery grey alcohol). covered median There of gap anteriorly. are two rows and irregular blackish markings, which are spots papillae. The oral disc is indented laterally. The bordered with There dark brown light are tooth green. formula is 1/4 + 4//1 + 1/2 or 1/5 + 5// crossbands over the limbs. The discs and the first 1 + 1/2. The horny beak is well developed. The

and second are The is fingers orange. pupil rim of lower beak is upper the black. There the flanks orange-brown. are light on spots The colour in life of the tadpole is yellowish, (pi. IV fig. 7). covered with brown spots. The belly is silvery.

white the and There is a spot on tip of the snout

Sexual dimorphism. — The male has there are white lines from the nostrils to the white nuptial pads laterodorsally on the first and of anterior comers the eye. situated second fingers. The vocal slits are small, took The entire development two and a half to in floor the the of the mouth, near angles of the three months in captivity. jaws. The vocal sac is median subgular and mod- erately distensible. The arms are robust in the male •— in is Young. The colour life beige or silvery and slender in the female. The skin on the throat with grey covered irregular black spots and is in the male and smooth in the female. granular which are bordered with The markings, green.

is skin of discs are yellow. The pupil orange. The

E — Diameter in . 2 and about 200 g g s mm the dorsum is warted as in the adult. number. The nostrils are strongly protruding and the

webbing is the same as in the adult. Tadpoles. — Twenty-six tadpoles in stage

25 have body lengths of 3.0-7.5 mm and total Boophis 8.0-20.5 opisthodon group lengths of mm. The average ratio of tail

in is — This length to body length this stage 1.7 (range Diagnosis. group comprises only

which 1.5-1.9). Twenty-one tadpoles in stages 26-30 one large-sized species (90 mm), breeds BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 305

The dorsum is probably in temporary pools. Boophis pauliani group

the brown. There are transverse bars on limbs.

— This is created for the Diagnosis. group only Fringes and dermal flaps are lacking on the limbs. of without vomerine teeth. It is is white the The species Boophis a There a margin on upper jaw. small-sized Vomerine species (20 mm), breeding probably snout is long and acuminate. teeth are in The colour in life is The is its horizontal temporary pools. greenish, present. tympanum very large, and the bones Indistinct transverse bars The are green. diameter is three-fourths of that of the eye.

the limbs. There is row of white The are present on a vocal sac is single, median and subgular. discs granules along the lateral edge of the tarsus and are well developed. The fingers have only a trace lower The is small. The snout the arm. tympanum very of web, toes are three-fourths webbed, reaching short and is is rounded. The vocal sac single, the discs of the first and fifth toe. The tibiotarsal

median and subgular. The fingers are one-third articulation extends to the nostril. Guibe (1978)

webbed and the toes are three-fourths webbed. described that the tibiotarsal articulation extends

The tibiotarsal articulation does not extend the does beyond beyond the tip of snout. This not agree the of the snout. with observation and the tip my original description Contents: B. pauliani (Guibe, 1953) by Boulenger (1888).

Contents: B. opisthodon (Boulenger, 1888).

Boophis pauliani (Guibé, 1953) comb. nov. Boophis opisthodon (Boulenger, 1888) Text-fig. 43. PI. II fig. 8.

Rhacophorus opisthodon Boulenger, 1888: 105; Guibe, 1978 Hyperolius pauliani Guibe, 1953: 101; Guibe, 1978: 89. 63, figs. 134-135.

Boophis opisthodon; Bachmann & Blommers-Schlosser, 1975 Material. — MNHP 1953-169-170, two syntypes ($

17 (DNA); Blommers-Schlosser, 1978: 36 (chromosomes) and 9) of Hyperolius pauliani. Type localities: forests of

Moramanga and Perinet. Coll. Guibe.

Material. — ZMA 6932 (1 $), 2.VIII.72; 6851 ZMA 6915 (2 S3), 24.XII.72 collected near Perinet, alt km of (4 $ £), 13.11.72; collected near Foulpointe, 60 N. 900 m. Tamatave, E. coast of Madagascar.

Natural — The males were col- — history. Natural history. The males were call- lected in shrubs in in shallow calling near a pool forest, ing in the shrubbery 30-50 cm above the evening. Guibe (1953) found the male also pools in February (these pools had disappeared in a shrub, near a pool in forest in January 1951. by July), in forest, during a rainy night. The sound

and resembled the blow of was very loud a trum-

Colour in life. — The colour is males rather ground pet. Calling were situated at large greenish, the bones are distances (about 10 m) from each other. Once green.

captured they clasped each other in axillary am-

— Our with the Taxonomy. material agrees plexus, even other species were clasped.

type material. The species has no vomerine teeth,

Colour in life. — The colouration is but the style of the omosternum is entire and brown. dark in- The temporal and loreal regions are shorter than the sternal style (see fig. 43),

A white line borders the There that it and brown. upper jaw. dicating belongs to Boophis not to

dark brown crossbands the limbs. The in- has are on Hyperolius. The male type specimen a well-

and the ventral hind- guinal spots surface of the developed nuptial pad, just as in the males col-

limb iris is The are orange. The yellow-orange. lected by us.

is belly cream.

Male. — The back is covered with small tuber- Montane species group cles (smooth in the female according its descrip-

There is small in Five tion). a blackish nuptial pad medium-sized Boophis species are known only

dorsolateral position on the first finger. from the High Mountains Area (cf. Koechlin, 306 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

1972). We have foundfound B. microtympanum (Boett- Colour in life. — The ground colour of

B. in the is dark the ger, 1881) and williamsi (Guibe, 1974) the male green; of female yellow-

Ankaratra mountains, above 2000 m, in wooded green. The irregular spots on the back, on the

bushes of ericoid flanks the limbs and under the fold country, characterized by appear- over tympanic

is also known from the surrounded line. ance. B. microtympanum are blackish, by a narrow orange

Between the Andringitra mountains. the blackish spots on the flanks skin

B. and B. is The iris is brygooi (Guibe, 1974) laurenti orange. discs are orange too. The

Guibe, 1947 are only known from the Andringitra greenish.

is mts.; B. microtis (Guibe, 1974) apparently The back of the Sexual dimorphism. — restricted to the Chaines Anosyennes. These three male is covered with small tubercles, the back of species are not further considered here. the female is smooth. An unpigmented glandular B. microtympanum and B. williamsi are stream- is the nuptial pad present dorsally on first finger breeding species. They can be distinguished readily in the male. The male is much smaller than the by their completely different colouration (also in female. alcohol, see under the species description).

Voice. •— The species was heard in April

the The made during evening. sonagram (fig. 38) microtympanum Boophis (Boettger, 1881) male the 13th of 1972 of a (ZMA 6850) on April

comb. nov. 10 shows that the call consists of number at p.m. a Ill 6. Text-figs. 35-38; pi. fig. One of short notes. call group consists of 4-9 notes.

from 2-4.5 The duration of one call group varies Hylambates microtympanum Boettger, 1881: 47. Hz. sec. The dominant frequency is about 3000 Rhacophorus boettgeri; Boulenger, 1882: 467 (substitute

name); Guibe, 1978: 69, figs. 156-158 (equals arboreus).

— The measure 3 mm in diameter, Boophis boettgeri; Blommers-Schlosser, 1978: 36 (chromo- Eggs. eggs

somes). 5 mm with the capsule; their number amounts to Rhacophorus arboreus Ahl, 1928: 312 colour is black with white. The ± 100. The eggs

hatch within 7-10 days. The hatchling (fig. 35) Material. — ZMB 30498, holotype of Rhacophorus has external mouth two small arboreus. Type locality: central Madagascar. Coll. Hilde- no gills; a opening,

brandt. The suckers and an anal tube are present. eyes ZMA 6850 (1 $ and 1 9), 13.IV.72; 6931 (1), 21.111. are visible through the skin and the external nares 73; 7006 (tadpoles and young) from rearing during nine are about to months, coll. 13.IV.72-3.VI.72; 7084 (eggs), 13.IV.72; 7085 open.

7086 and 21.111. (tadpole), 21.111.73; (eggs young larvae),

and 7088 in 73; 7087 (eggs young larvae), 13.IV.72; (tad- Tadpoles. — Thirty-two tadpoles stage 25 poles), 13.IV.72; 7089 (tadpole and juvenile), from rearing have body lengths of 5-14 mm and total lengths during several months, coll. 21.III.73; all collected at Manja- The of 13.5-41.0 mm. ratio of tail length kotompo forest station, Ankaratra mts. (near Ambatolampy), average

alt. 2200 m. to body length in this stage is 1.7 (range 1.5-1.9).

Eight tadpoles in stages 26-28 have body lengths

Natural history. — The species was col- of 12-20 mm and total lengths of 34-57 mm. Eight

lected in clear mountain brooks, with stony bottom, tadpoles in stages 30-40 have body lengths of

in wooded The male The country with ericoid bushes. 19-24 mm and total lengths of 53-71 mm. and in tail in the female collected were axillary amplexus average ratio of length to body length

in the water. The males call in the evening. This stages 26-40 is 1.9 (range 1.8-2.0). One meta-

is and climbs The 24 from snout species a poor jumper badly. morphosing young measures mm

and the found in the tadpoles eggs were same to vent. brooks the the attached around The mouth and lateral view of a as adults, eggs were (ZMA 7085) in a twig in a clutch of about 100 in a fast flowing tadpole (ZMA 7006) in stage 37 are shown

the with is part of current. The clutch was covered figs. 36-37. The body ovoid. The nostrils are

small particles. dorsal, their dorsolateral openings are situated at BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 307

Fig. 38. Sonagram, filter wide, Boophis microtympanum.

equal distances from the tip of the snout and the Boophis williamsi (Guibé, 1974)

of the The situated Ill 7. anterior corners eyes. eyes are Text-figs. 39-40; pi. fig. dorsolaterally. The spiracular opening is sinistral, Rhacophorus williamsi Guibe, 1974: 1178, fig. 6; Guibe, situated at about 5/8 to 2/3 from the tip of the 1978: 68, figs. 152-155. and the end of the The cloacal tube is snout body. Boophis williamsi; Blommers-Schlosser, 1978: 36 (chromo-

the side of the somes). long and situated on right caudal

fin. The caudal musculature is in robust, especially Material. — MNHP 1973-1091, the holotype of Rhaco-

fin is phorus williamsi.Ambohimirandana, Ankara- the anterior part of the tail. The caudal Type locality: tra mts. Coll. Ch. P. Blanc. reduced anteriorly. At midlength of the tail, the ZMA 6933 (one young reared from tadpoles ZMA 7012); caudal musculature height of the accounts for 7012 (tadpoles), 21.111.73; 7195 (one tadpole), 21.111.73;

collected at Manjako- about 1/2 of the total tail height. The mouth is 7013 (tadpoles), 13.IV.72-3.VI.72; forest Ankaratra tompo station, mts. (near Ambatolampy), small and directed ventrally. The oral disc is alt. 2200 m. slightly indented medioposteriorly. The papillary

Natural — The border has wide median and history. tadpoles were a gap anteriorly a very collected in clear mountain brooks with There 1-7 of narrow gap posteriorly. are rows stony in wooded with bushes of ericoid papillae laterally and 1-2 posteriorly. The homy bottom, country

The were found together beak is well developed. The tooth formula is appearance. tadpoles

with those of B. 1/5 + 5//1 + 1/2 generally; but also 1/4 + 4// microtympanum (Boettger,

We failed to adult 1 + 1/2 or 1/6 + 6//1 + 1/2 occurs. 1881). capture frogs.

The colour of the tadpole in life is greyish with Tadpoles. — Fifteen tadpoles in stage 25 dark grey spots. have body lengths of 7-15 mm and total lengths The the development proceeds very slowly; of 17.5-40.0 The ratio of tail mm. average length estimated duration is about one year, as we could to body length in this stage is 1.5 (range 1.4-1.7). in in observe our rearings and the field. This is Eighteen tadpoles in stages 26-28 have body also indicated by the large amount of growth in lengths of 14-30 mm and total lengths of 38-85 each stage of development. The mm. average ratio of tail length to body length

in these stages is 1.7 (range 1.6-1.8). One tadpole

— The reared resembles the in 42 has Young. young stage a body length of 32 mm and a adult that it could be so much, easily assigned to total length of 82 mm. One metamorphosing this species. young measures 30 mm from snout to vent. 308 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

Figs. 39-43. 39-40, Boophis williamsi: 39, mouth of tadpole, ZMA 7012.X.12.5; 40, tadpole, ZMA 7012.X.1.5; 41-43,

ventral views of bony elements of the shoulder girdle. From top to bottom: bony style of omosternum, procoracoids,

coracoids, bony style of sternum. 41, B. mandraka, ZMA 7119.X.12.5; 42, B. madagascariensis, ZMA 6849.X.6; 43, B. pauliani, ZMA 6915.X.12.5.

mouth and view of in dian There The lateral a tadpole gap anteriorly. are 3-4 rows of papillae.

is stage 27 (ZMA 7012) are shown in figs. 39-40. The horny beak is heavy. The tooth formula

The body is depressed. The nostrils are dorsal, 2/5 + 5//3 or 2/6 + 6//3.

their openings dorsolateral, situated at equal dis- The colour of the tadpole in life is black with

tances from the tip of the snout and the anterior some bright yellow patches, in the early stages.

the situated dorso- These colours fade with The corner of eye. The eyes are gradually age.

is situat- of the laterally. The spiracular opening sinistral, development tadpole proceeds very slowly, ed the of in at about 1/2-5/8 from tip the snout and and considerable growth occurs each stage of

the end of the body. The cloacal tube is long and development. The estimated duration is at least

could observe situated on the right side of the caudal fin. The one year, perhaps even longer, as we

in the and in caudal musculature is robust, especially in the field our rearings.

anterior The caudal fin part of the tail. dorsal

the At of the extends on to body. midlength tail,

caudal — Our is the height of the musculature accounts for Young. metamorphosed young very

the total about 1/2 of height. The mouth is broad large (30 mm), while the known adults reach a and directed The of the oral disc size of The colour in life of the ventrally. margin 38 mm. young

is if with It round; folded inside it has a notch medio- was olive-green brown spots. could be

border has B. posteriorly. The papillary a wide me- easily identified as williamsi. BIJDRAGEN TOT DE DIERKUNDE, 49 (2) - 1979 309

DISCUSSION Cytogenetic studies showed that the karyotypes

within the genus Boophis (12 species) are very In earlier the an paper on biosystematics of similar (2n = 26), and that acrocentric chromo- the Malagasy Mantellinae (Blommers-Schlosser, somes never occur (Blommers-Schlosser, 1978). 1979), it has been argued already that Boophis The quantitative nuclear DNA amount shows little Tschudi, 1838, is not so closely related to the variation; 7-9.9 in eight species (Bach- Mantella diploid genera Boulenger, 1882, Mantidactylus mann & Blommers-Schlosser, 1975). The char- Boulenger, 1895 ( = Gephyromantis Methuen,

~ mentioned is acters above indicate that Boophis a 1919), and Trachymantis Methuen, 1919, as has natural unit at generic level. been presumed by most previous workers but The distribution of the of the genus Boopbis appears Laurent, 1946. Mainly on the basis peculiar to be confined to the rain forests of the largely (East way mating, genera Mantella, Mantidactylus Area, Centre Area and Sambirano Area) and the and Trachymantis were considered to form the High Mountains Area of Madagascar. Only two subfamily Mantellinae of the Ranidae, while also be found in farmland B. considered of the Rhaco- species may ( gou- Boophis was a genus in the doti) or savanna and deciduous woodland phoridae (Blommers-Schlosser, 1979). The pres- (B. tephraeomystax). The more arid biotopes are ent study confirms this opinion. of the Heter- A normal of is if not occupied primarily by species genus sort amplexus common, ixalus Laurent, 1944, which likewise belongs to the general, in Boophis. We watched axillary amplexus Rhacophoridae (Hyperolius Rapp, 1842, probably in B. rappiodes , luteus, majori, goudoti, tephraeo- does not in Blommers-Schlosser and occur Madagascar, mystax, opisthodon microtympanum. We in preparation). watched lumbal amplexus in B. madagascariensis ; Most in the species Boophis are Arnoult & Razarihelisoa, 1966, recorded lumbal genus genuine which active tree frogs are at night. B. amplexus in B. goudoti. Calling males of various Only gou- doti, B. untersteini and the montane species B. species showed a pronounced clasping reflex. microtympanum are found normally on the ground. The accessory sexual characters, generally con- Some diurnal B. nected with this of in the activity was observed in way mating, are present goudoti and B. madagascariensis only. Various brown species of the genus Boophis. Every male collected

~ species ( B. untersteini and te- in the breeding season has nuptial pads. The males madagascariensis, have also phraeomystax) a juvenile hue, which is are distinctly smaller than the females. Only

and serves for were collected. In the predominantly green probably submerged eggs contrast, camouflage. species of Mantellinae lack a real amplexus, the The differentiation within this is mani- males femoral but genus possess glands nuptial pads are fested of about size and most in the of absent, the sexes are equal eggs clearly variety aquatic

habitats, the While some were found outside the water. Nevertheless, mor- occupied by tadpoles.

differences Mantidac- of are confined in this to phological between some groups species respect

others are found in streams of and some are stagnant waters, tylus species groups Boophis species varying velocity. Because stagnant water is scanty. rarely in the rain in The best criterion I could find is the shoulder permanent hilly forest eastern

Madagasacar, the fauna connected with this sort girdle, which I examined in B. mandraka, erythro- of sites dactylus, luteus, majori, goudoti, madagascariensis, breeding seems dependent mainly on the of and The reticulatus,opis-tephraeomystax, granulosus, idae, availability temporary pools puddles. development in these species is rather the thodon, pauliani and microtympanum. The bony rapid, hatch before the of drier style of the sternum is always longer than the young onset the winter of this those in bony style of the omosternum, the latter being season. Species biotope are notably

while in the B. and also never really forked, Mantidactylus the tephraeomystax group probably B. and B. The bony style of the omosternum is broadly or mod- pauliani opisthodon. webbing of

than the is reduced in these erately forked; its length is equal to or greater fingers frogs. that of the sternal style. The larvae of both B. goudoti and B. madagas- 310 R. M. A. BLOMMERS-SCHLOSSER - BIOSYSTEMATICS OF BOOPHIS

found also in these larvaires cariensis are occasionally pools, de Rhacophorus goudoti (Tschudi). Bull. Mus.

natn. Hist. found in nat. Paris, (2) 38 (2): 111-117. although they are most often small, shal- K. R. BACHMANN, & BLOMMERS-SCHLOSSER, 1975. Nuclear streams and in the lentic of low relatively parts DNA amounts of the frogs of Madagascar. Zool. Anz., large streams together with the tadpoles of B. 194 (1/2): 13-21.

BLOMMERS-SCHLOSSER, R. M. A., 1978. of the rappiodes. Cytotaxonomy Ranidae, Rhacophoridae, Hyperoliidae (Anura) from Larval in these to development species appears with Madagascar a note on the karyotype of two am-

is of be slower, or the egg-laying period more ex- phibians the Seychelles. Genetica, 48 (1): 23-40.

1979. of the I. Mantel- , Biosystematics Malagasy frogs. tended than that of real pool dwellers. Hatching linae (Ranidae). Beaufortia, 29 (352): 1-77. shows two one at the onset of the winter, peaks, BOETTGER, O., Die 1879. Reptilien und Amphibien von the other at the start of the next hot season. During Madagascar, 2. Nachtrag. Abh. senckenb. naturforsch.

Ges., 11: 457-497, pi. I. the dry and cold winter (May-September), these 1881. Die Reptilien und , Amphibien von Madagascar, and takes tadpoles slowly no metamorphosis 3. Abh. grow Nachtrag. senckenb. naturforsch. Ges., 12: 435- place. 558, pis. I-V. 1882. Batrachiorum , Diagnoses et While the of B. B. and of Reptilium novorum tadpoles luteus, sp. insulae Nossi-Be Madagascariensis. Zool. Anz., 5 the montane B. and B. species microtympanum (120): 478-480.

williamsi live in currents and mountain brooks, 1892. der im Museum , Katalog Batrachier-Sammlung der Senckenbergischen Naturforschenden Gesellschaft in the larvae of B. erythrodactylus, B. mandraka and Frankfurt am Main: i-x, 1-73 (Gbr. Knauer, Frankfurt B. in majori are found real torrents. Here larval a. M.). takes for the development nearly a year, young , 1913. Reptilien und Amphibien von Madagascar, den Inseln und dem hatch in the of the hot while Festland Ostafrikas. In: Reise in Ost- beginning season, Afrika von Prof. A. Voeltzkow, 3 (4): 269-376, pis. laid new eggs are a couple of months later. XXIII-XXX (E. Schweizerbart, Stuttgart). in the Development montane species may even BOULENG.ER, G. A., 1882. Catalogue of the Batrachia Sa-

lientia s. Ecaudata in the collection of the British take two Mu- years (B. williamsi). seum (2nd ed.): i-xvi, 1-503, pis. I-XXX (Trustees A comparison of the habitats occupied by the British Museum, London).

1888. larvae of of the and of the Descriptions of new reptiles and batrachians from species genus Boophis ,

that Madagascar. Ann. Mag nat. Hist., (6) 1 (2): 101-107, genus Mantidactylus shows the tadpoles of pis. V-VI. in both concur largely relatively lentic streams and 1895. On a of peculiar to , genus frogs Madagascar. in In and temporary pools. fast running currents Ann. Mag. nat. Hist., (6) 15 (89): 450.

1896. of new batrachians in the British , Descriptions torrents in Madagascar, I only found tadpoles Museum. Ann. Mag. nat. Hist., (6) 17 (101): 401-406, belonging to Neither arboreal Boophis species. pi. XVII. direct tadpoles nor development, both known in DUELLMAN, W. E., 1970. The hylid frogs of Middle Amer-

ica. Monogr. Univ. Kan. 1 the in the Mus. nat. Hist., (1): i-xi, genus Mantidactylus, occur genus 1-427; (2): [i-iv], 429-753, pis. I-LXXII. Boophis. DUMERIL, A., 1853. Memoire sur les Batraciens anoures,

de la famille des Hylaeformes ou Rainettes, comprenant

la d'un de description genre nouveau et onze especes

nouvelles. Annls. Sci. nat., 19: REFERENCES (3, Zool.) 135-179, pi. VII.

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Madagascar. Zool. Anz., 75 311-318. 1947. d'un (11/12): GUIBE, J., Description Batracien nouveau de

1929. Beschreibung neuer Frosche aus Madagascar. laurenti , Madagascar (Boophis n. sp.), et synonymie de Mitt. zool. Mus. Berl., 14 (3/4): 469-484. plusieurs especes du genre Boophis. Bull. Mus. natn.

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ANGEL, F., 1934a. d'un du Naturaliste Description Viperide nouveau, gache (Batraciens). malgache, 5: 101-103. et de Congo Beige, deux Batraciens de 1974. Batraciens nouveaux de Madagascar. , Madagascar. Bull. Mus. Bull. Soc. zool. 59: Fr., 169-172. natn. Hist. nat. Paris, (3) 1973 (171) (Zool. 116):

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Hist. Paris, of snake from natn. nat. (3) 1975 (323) (Zool. 230) Methuen, P. A., 1919. Descriptions a new

1081-1089. the Transvaal, together with a new diagnosis and key

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Flora and of KOECHLIN, J., 1972. vegetation Madagascar. Mocquard, F., 1902. Sur une collection de Reptiles et de

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Frankfurt nebst a. M. einem Verzeichnis ihrer Typen. pis. I-VI. Senckenberg. biol., 48 (Sonderheft A): 1-106, figs. 1-12.

Received: 18 May 1979 LEGENDS TO THE PLATES

PLATE I

ZMA B. hatched ZMA hatched 1, Boophis erythrodactylus, tadpole, 7207; 2, erythrodactylus, young, 7207; 3, B. luteus, . . ,

~ ZMA B. hatched ZMA 5, B. ZMA B. ZMA young, 7005; 4, sp., young, 7154; majori,' male,' 7102; 6, miniatus, male,

' ~ and female ZMA 7162; 7, B. goudoti, male in amplexus, 6808.

PLATE II

1, Boophis goudoti, hatched young, ZMA 7210; 2, B. madagascariensis, male, ZMA 6837; 3, B. madagascariensis, hatched

~ B. ZMA young, ZMA 7069; 4, untersteini, hatched young, 7189; 5, B. untersteini, hatched young, ZMA 7190; 6, B. untersteini, male, three months old, ZMA 7190; 7, B. tephraeomystax, male, ZMA 6775; 8, B. opisthodon, male, ZMA 6851.

PLATE III

ZMA 7101; B. ZMA B. hatched ZMA 1, Boophis reticulatus, male, 2, idae, male, 7014; 3, granulosus,j young, 7204;

" ~ ~ B. ZMA B. hatched ZMA B. ZMA B. 4, hillenii, tadpole, 7122; 5, idae, young, 7199; 6," microtympanum, male, 6850; 7, williamsi,tadpole, ZMA 7013.

PLATE IV

1, Boophis rappiodes, male and female, ZMA 7164; 2, B. mandraka, male, ZMA 7119; 3, B. erythrodactylus, male, ZMA

' * ~ ~ ' " ~ ~ B. ZMA 6838; B. ZMA B. ZMA B. 6930; 4, luteus, female, 5, viridis, male, 7100; 6, difficilis, male, 7099; 7, granu- losus, male, ZMA 6853. PLATE I PLATE II PLATE III PLATE IV