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Rhacophoridae 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.
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