The Relationship of Brookesia, Rhampholeon and Chamaeleo
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Bijdragen tot de Dierkunde, 56 (1): 29-38 1986 The relationship of Brookesia, Rhampholeon and Chamaeleo (Chamaeleonidae, Reptilia) by D. Hillenius Institute of Taxonomic Zoology (Zoologisch Museum), University of Amsterdam, P.O. Box 20125, 1000 HC Amsterdam, The Netherlands of Abstract Klaver himself provides an example the changeable opinions inspired by these little Comparing the species of Brookesia and Rhampholeon with lizards. Chamaeleo it is concluded that Brookesia + Rhampholeon form from branch of In 1979 he considered Brookesia s.l. to be a monophyletic group, arising a Chamaeleo, probably most related to the around Chamaeleo group derived from "a fully arboreal Chamaeleo-like nasutus. The separation between Rhampholeon and Brookesia ancestor". In 1981 he still considers Brookesia is confirmed. s.l. descendant of "a fully arboreal form" which, because of the "true chameleon feet" Résumé andother details (eyes, tongue), might be called En les de Brookesia de least Chamaeleo-like. the of comparant espèces et Rhampholeon at Although lungs avec Chamaeleo, on arrive à la conclusion Brooke- que Brookesia s.l. are simpler than those of Chamaeleo sia + constituent un monophylétique, Rhampholeon groupe this first "This seems at sight no problem; descendant d’une branche de Chamaeleo, probablement reversed trend the return des affinités accentuées le de Ch. evolutionary (viz. ayant avec groupe autour from arboreal life to may also nasutus. La séparation de Rhampholeon et de Brookesia a été ground dwelling) furnish for confirmée. an explanation the simple lung structure of the Brookesia species". According to Klaver (1981) this secondarily simple lung INTRODUCTION structure includes only the reduction and, in The of the the loss of the diverticula. systematic position pygmy most species, even chàmeleons has been uncertain for than the absence of more a However, one single character, the Brookesia century. Although genus was de- lung septation, leads Klaver to the opposite scribed by Gray in 1864 and Rhampholeon in conclusion: "I do not think that Brookesia lungs 1874 by Günther, several species that belong lost their septation secondarily, because there the chameleons does be unmistakably to pygmy were not seem to a correlation between septa- later Chamaeleo. For tion described as belonging to and body form as in the case of diver- instance, in 1911 Werner regarded temporalis ticula". (Matschie) as a Chamaeleo. Even in Mertens' list Klaver's opinion of 1981 is best expressed in of 1966 of the chameleons (marshalli his of one pygmy diagram of a hypothesized phylogeny included in A based In Boulenger) was Chamaeleo. chameleons on lung septation (fig. 1). number of other genera will not be considered that scheme Brookesia s.l. is closer to the original in this Chamaeleonidae branches of paper for reasons given by Klaver than all the (1979). Chamaeleo. Klaver and excellent In this (1979 1981) gives an paper I want to discuss the following survey of the various opinions — sometimes of questions: the author in — the 1. Is Brookesia s.l. than same subsequent years on more primitive different that Klaver did genera were proposed and after Chamaeleo, as (1981) suggested, or time considered be Brookesia from branch or branches of some to synonyms. Indeed, originate a Downloaded from Brill.com09/28/2021 04:38:56PM via free access 30 D. HILLENIUS - RELATIONSHIPS OF CHAMAELEONIDAE RESULTS I took note of the following characters (see table I): 1. of head Length + body: from tip of snout to foremost border of the vent. 2. Length of tail: from the foremost border of the vent to tip of tail. Tail index expressed in of the percentage length of head + body. 3. Length of mouth cleft: from tip of snout to corner of mouth, index expressed in of the percentage length of head + body. 4. of measured the Fig. 1. Schema of hypothesized phylogeny of chameleons, Width mouth: at corner of based on lung septation. B, C, D, E and F the index in of represent mouth, expressed percentage of Chamaeleo, A is Brookesia (and Rhampholeon). groups the length of mouth. After Klaver (1981). 5. of Height head: measured in a vertical line at the corner of the mouth, from the under- side of the jaws to the surface of the skull In other is index Chamaeleo? words Brookesia older or (see fig. 2), expressed in percentage of than Chamaeleo? the of mouth. younger length 2. Is Brookesia s.l. 6. in with (including all pygmy Temporal crest: accordance Werner's in chameleons, Rhampholeon etc.) a monophyletic (1911) use of this term group? Chamaeleo I regard as temporal crest the one 3. How valid is the separation of Brookesia that "traverses the mid-lateral temporal (Madagascar) and Rhampholeon (Africa)? region from the middle of the posterior border of the orbit the horizontally to poste- rior border of the skull" (Raw, 1976). Klaver in his MATERIAL (1981) comment on the de- scription of Chamaeleo intermedius Hillenius, All African species and all but four of the Madagascan 1978, made objections to my use of this species were examined. As far as possible I analyzed the the of this character is four missing Madagascan species from literature. The term, as homology following species are considered: Rhampholeon brachyurus uncertain. I will return to this problem Günther, 1892, Rh. brevicaudatus (Matschie, 1892), Rh. but it be more extensively elsewhere, may kersteni (Peters, 1866), Rh. marshalli Boulenger, 1906, Rh. stated here that the temporal crest in Rham- nchisiensis (Loveridge, 1953), Rh. platyceps Günther, 1882, pholeon is most probably homologous with Rh. spectrum (Buchholz, 1874), Rh. temporalis (Matschie, the crest in at least Chamaeleo 1892), Brookesia antoetrae Brygoo & Domergue, 1971, Br. temporal betschi s.l. and in all Brygoo, Blanc & Domergue, 1974, Br. bonsi pumilus Ch. tigris since, these Ramanantsoa, 1979, Br. decaryi Angel, 1938, Br. dentata cases, the crest is based on the lateral ridges Mocquard, 1900, Br. ebenaui (Boettger, 1880), Br. griveaudi of the postorbital and the squamosal bones Brygoo, Blanc & Domergue, 1974, Br. karchei Brygoo, Blanc (see Engelbrecht, 1951; Frank, 1951; & Domergue, 1970, Br. lambertoni Brygoo & Siebenrock, Domergue, 1969, Br. legendrei Ramanantsoa, 1979, Br. 1893). minima Br. In Brookesia the Boettger, 1893, nasus Boulenger, 1887, Br. (the Madagascan species) Br. perarmata (Angel, 1933), peyrierasi & Brygoo Domergue, situation is more complicated. The fused 1975, Br. ramanantsoai Brygoo & Br. Domergue, 1975, and postorbital squamosal bones are rather stumpffi Boettger, 1894, Br. superciliaris (Kuhl, 1820), Br. broad Siebenrock, the therezieni (see 1893), probably Brygoo & Domergue, 1970, Br. thieli Brygoo & crest is based the lower of Domergue, 1960, Br. tuberculata Mocquard, 1894, and Br. temporal on ridge vadoni Brygoo & Domergue, 1968. these bones (see fig. 2 and the next section). Downloaded from Brill.com09/28/2021 04:38:56PM via free access 56 - 1986 31 BIJDRAGEN TOT DE DIERKUNDE, (1) tail lung lung form form head 0 head are spinose second double rostral axillary lateral height mouth mouth = bicuspid vertebral inguinal parietal temporal + index septa squamation guiar interorbital : index pit crest body indented. soles claw guiar pit parietal crest height width length absent, of claws diverticula crest width male Comparison crest female expressed the processus cones crest excrescence male index index index female Lung of in length the 1 1 1 1 1 113 89 79 16.0 20 22 45 32 brachy urus 01 of species 1 1 brevicaudatus 1 1 1 120 91 76 20.1 27 30 48 53 the septa/alveoles: percentages of 0 1 kersteni 1 1 1 1 137 85 62 16.2 48 63 54 41 developed, mouth 0 of = 1 1 1 0 93 68 52 53 64 30 marshalli 0 000<1 < 73 18.9 or the are Rhampholeon 0 1 nchisiensis 0 1 1 1 82 30 67 43 AFRICA 98 80 19.6 24 absent, only length with 1 1 1 1 85 30 36 46 55 0 91 77 19.8 platyceps in expressed of the 00000011 00000101 <1111111 00000000 111111<1 a 00 00000000 22222222 02100000 22222222 1 1 1 70 34 52 58 54 spectrum 113 79 19.8 in 10 1< 1<0 few species 1 1 61 0 130 79 23.8 52 46 48 47 temporalis mouth. of 1 1 106 108 102 16.6 76 36 antoetrae present, specimens. percentages <1 Height: 00 01 00 00 1< 2 betschi 1 97 97 68 65 34 34 Brookesia. 100 17.0 of =alveoles 1 bonsi Form the 1 22-1 101 1 56 76 61 67 septa width For 00100000<10001 is 0 93 79 85 21.7 51 47 53 43 decaryi length present. parietal: the detailed 11111— — 87 23 dentata of Table 1<1 1<1 1 01 I ebenaui = 1 97 91 94 19.2 55 69 50 48 height head 00 of description 1 1 1 and 2 102 93 91 17.6 67 76 55 59 griveaudi trapezoid, the of karchei 1 < 125 105 84 15.4 58 73 23 30 body, 2 head the lambertoni = 0 1 1 86 87 101 17.3 55 44 whereas 1 1 0 95 91 96 20.1 68 77 41 35 legendrei trigonal expressed characters the minima 1 0 96 86 90 16.1 60 20 MADAGASCAR (see in see 0 1<100010 90 72 32 58 37 31 nasus 125 25.1 indices 0<1 fig. text. 1 1 102 99 97 67 66 perarmata 3). of 20.6 percentages It the is < 1 1 111 92 83 17.5 72 77 25 22 peyrierasi of 1 1 width < 1011001?<11 71 78 26 23 ramanantsoai the stressed Squamation: of 1 1 1 1 — — — 0 —0 —001— 0 -——000000000 —0 85 81 89 46 47 77 —— stumpffi —— —— 1 ——105 — — =<feebly — 22.5 — width —— that 2222222222 — 1 = mouth 1 1 1 1 000000000000000000000111111111111111111111 100 103 103 15.6 70 79 50 53 superciliaris of the 00000000000<000000000 000000000000000000000 000000000000000000000 and 1 1 1 1 tail <11<111111<1001101 93 82 88 15.6 62 69 42 51 therezieni 0000000000000000 0000000000000000 1010 mouth.