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 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). 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 — — =

11111111111111 1 2-1 107 110 103 15.4 67 82 42 34 thieli height index = 2

= 1<1 2-1 101 101 1111 — 89 18 tuberculata of and scales present, head the 0 2-1 93 95 102 19.0 63 66 35 35 vadoni 32 D. HILLENIUS - RELATIONSHIPS OF CHAMAELEONIDAE

10. Interorbital crest: from one orbital crest to

the in front of the often other, eyes, accen-

tuated those with on crests pointed ex-

crescences.

11. Axillary pits: little pockets in the axillary

region.

12. Inguinal pits: little pockets in the inguinal

region.

of the 13. Flexible excrescence on the tip snout.

14. Guiar cones or 'tufts'; Loveridge, 1942:

"beard-like 'tuft' of scales forming a flexible

the chin". process on

15. Guiar cones in a double row.

16. Form of the scales. In most pygmy Fig. 2. Height of head measured in a vertical line at the chameleons I found scales with in- corner of the mouth between the underside of the lower deeply

surface of lateral dentedborders table I indicated jaw and the the skull (a); b, crest; c, tem- (in by 2, see

poral crest. (Drawing Miss N. Pruim.) figs. 4 to 7); sometimes the scales were

polygonal (indicated by 1).

7. Lateral the continuationof the orbital 17. Lateral series of crest: pointed excrescences or

scales the Parker crest backwards (see fig. 2). Werner (1911) larger on body. (1942) de-

calls this character in Brookesia "Oc- scribed stumpffi strong spines projecting laterally

cipitalkante". Comparison with the skull of from the prezygapophysis of the dorsal

Brookesia superciliaris in the Paris Museum vertebrae 3 to 9 and penetrating the skin. I

(see also Siebenrock, 1893) made it clear do not doubt that most or all of the above-

that the lateral coincides with the mentioned to be crest up- pointed excrescences are

border of the fused and Parker's transverse per postorbital regarded as processes,

squamosal bones. but indeed — as Klaver (1979) remarked —

8. Parietal crest: in the mid-line of the broad we do not know the skeleton of all Brookesia

I its parietal. species, so limit this character to exter-

9. Form of the broad parietal: trigonal or nal manifestations.

trapezoid (see fig. 3). 18. Lung diverticula (after Klaver, 1979).

19. Lung septation (after Klaver, 1979).

20. Bicuspid claws.

21. Single spines at the base of the claws.

' ' ' Werner (1911) called these second claws '.

22. Spiny cones on the soles of hands and feet.

the indices the In table II average and

absence of certain characters presence or are

compared in the African dwarf chameleons, the

Madagascan dwarf chameleons, African

Chamaeleo minus the representatives of the

minus nasutus-group, Madagascar! Chamaeleo the

representatives of the «aiutai- group and the Fig. 3. Parietals, left: trapezoid in Brookesia (schematized group around Ch. nasutus, including Ch. nasutus after Siebenrock, 1893), right: trigonal in Rhampholeon & Ch. Ch. (schematized after Frank, 1951). (Drawing Miss N. (Duméril Bibron), fallax (Mocquard), boett- Pruim.) gallus (Günther), Ch. linotus (Müller), Ch. 56 - 1986 33 BIJDRAGEN TOT DE DIERKUNDE, (1)

4 5

6 7

Fig. 4. Squamation of Rhampholeon marshalli.

Fig. 5. Squamation of Rhampholeon spectrum.

Fig. 6. Squamation of Rhampholeon kersteni.

Fig. 7. Squamation of Brookesia stumpffi. (Photographs by L. A. van der Laan, ZMA.)

geri (Boulenger), Ch. guibei Hillenius, Ch. DISCUSSION AND CONCLUSIONS

spinosus (Matschie) and Ch. tenuis (Matschie).

The latter taken unit 1. Did Brookesia s.l. from branch group was as a separate originate a or

because of Klaver's remark (1979) that Brookesia branches of Chamaeleo or was it the reverse?

s.l. has more characters in common with the Table II shows that Brookesia s.l. and

around Ch. than with other all of these group nasutus any Chamaeleo, though not species

of chameleons. The measurements of have a number of characters in com- group groups,

lateral the head + body and the tail indices of Chamaeleo are mon: temporal crest, crest (on

computed from literature (Werner, 1911; homology of these crests I hope to comment

De To later flexible Brygoo, 1971, 1978; Witte, 1965). get at on), a single parietal crest, a

of least an indication of the indices of the length rostral appendage, axillary pits, inguinal pits,

in mouth, the width of mouth, and the height of guiar cones double rows, lung diverticula.

head, I took Ch. chamaeleon (Linnaeus) O* ZMA These characters confirm the close relationship

but do in- 14719 as an example for African Chamaeleo, Ch. of Chamaeleo and Brookesia s.l., not

oustaleti ZMA 14302 for the dicate which is the of the other. (Mocquard) C group source

the char- Madagascan Chaemaleo, and Ch. fallax 9 ZMA More important in this context are

which mentioned in table 15339 as a representative of the group around acters are not II

well known the Ch. nasutus. because they are to belong to 34 D. HILLENIUS - RELATIONSHIPS OF CHAMAELEONIDAE

TABLE II

African the of the around Comparison of Rhampholeon, Brookesia, Chamaeleo (minus species group Ch. nasutus),

the of the around Ch. and Chamaeleo For of Madagascan Chamaeleo (minus species group nasutus) nasutus c.s. length

of head of the African chameleons Ch. chamaeleon is chosen of the mouth, width of mouth, and height as representative,

chameleons Ch. and of the around Ch. Ch. is chosen. 3 = Madagascan oustaleti, group nasutus, fallax Squamation: not

like = Chamaeleo. = indented, not polygonal, in Form parietal: 3 narrow, as in Ch. chamaeleon, 4 broad, as in Ch. pumilus.

Other terms see table I.

AFRICA MADAGASCAR AFRICA MADAGASCAR

(Rhampholeon) (Brookes ia ) ( Chamaeleo) (Chamaeleo ) Ch. nasutus c.s.

n av. n av. av. n av. SD SD SD n av. SD n SD

8 14.0 19 28 17 8 head + body O* 44 9.7 38 111 47 46 142 70 54

9 54 8.5 8 43 12.4 17 117 53 37 107 50 26 53 8.3 6

tail index cr 42 14.1 8 72 11.6 19 110 21.0 46 119 14.3 28 101 18.4 8

9 36 13.0 8 64 10.9 17 100 22.1 37 104 18 26 102 6.5 6

mouth length index 19.3 2.5 8 18.4 2.8 17 24.8 18.5 19.2

mouth width index 73.5 8.8 8 92.5 9.2 17 66.7 70.2 68.8

head height index 81 7.3 8 93.7 9.0 17 114 130 82.3

height : width 112 17.0 8 102 10.7 17 171 185 120

temporal crest 1 1-0 1-0 1-0 1-0

lateral crest 0 1-0 1-0 1-0 1

parietal crest 1 0 1-0 1-0 1-0

form parietal 2 1 3-4 4 ?

interorbital crest 110 0 0

axillary pit 1-0 0 0 1-0 1-0

inguinal pit 1-0 0 0 0 1-0

0 rostral excrescence 1-0 0 0 1-0

double guiar crest 1-0 1-0 1-0 1-0 0

squamation 2 1-2 3 3 3

vertebral processus 0 1(0) 0 0 0

lung diverticula 1-0 0 1-0 1-0 1-0

lung alveoles/septa 1-0 1-0 2 2 2

bicuspid claws 1-0 0(1) 0 0 0

second claws 1-0 0 0 0 0

spinose soles 1-0 1(0) 0 0 0

situated in the fron- common heritage of all Chamaeleonidae: the vertebrae, pineal foramen

pincer shaped hands and feet, the in- tal or absent, etc." (Klaver, 1981) and the low

movable the fusedbut number of ribs. In table III I list of dependently eyes, eyelids numbers

for small in front of mentioned in the literature a opening the pupil, the ribs, as or observed

projectile tongue, the prehensile tail (in most personally in skeletons, in whole animals that

Brookesia s.l. too short to be effective, but even have been opened or in X-ray photographs.

here observes one attempts to grip twigs or The latter two sources are less accurate as the

It that these number of shorter ribs in the lumbar is grass stems). seems probable are region arboreal retained in the difficult discern. It is clear the adaptations which are to that number

secondarily ground-dwelling Brookesia s.l., most of ribs in Brookesia and in Rhampholeon is lower

the notably highly modified hands and feet. than in any species of Chamaeleo.

The tail of Brookesia, more or less prehensile but Methuen & Hewitt (1914) remark on this:

short to be smaller too of much use, is clearly derived "The numbers most probably represent

the But do from the functional Chamaeleo tail. more primitive conditions". they

Other characters of which the derived status not give justification for this opinion. Most is clear are "the absence of Jacobson's organ, probably the smaller numbers are derived. In

of the dorsal I 19 ribs accessory transverse processes Amblyrhynchus (Iguanidae) counted on 56 - 1986 35 BIJDRAGEN TOT DE DIERKUNDE, (1)

TABLE III would then have arisen in Chamaeleo only after

number of ribs found in literature Brookesia s.l. had off. The as (authors) or split

skeleton in bodies by myself on (s), opened (b) or on In the Agamidae, one of the families most The latter numbers less X-ray photographs (X). two are find: closely related to the Chamaeleonidae, we accurate as the number of shorter ribs in the lumbal "The consists of parietal a flat, square to region is difficult to discern. trapezoidal, anterior portion which roofs the

cranial vault" (Moody, 1980). But in the Br. superciliaris 11 (s) Iguanidae, the other closely related family, both Br. stumpffi 9 (Werner, 1911), 11 (X) and narrow broad parietals are represented. Rh. spectrum 12 (Werner, 1911), 12 (s)

Romer writes: "The often a Rh. marshalli 12 (X) (1956) parietal,

Rh. 12 broad be narrowed varied temporalis (X) plate, may to a

Ch. lateralis 14 (b) degree, sometimes — as in the iguanid Con- Ch. oustaleti 18 (X) olophus — developing a median sagittal crest". Ch. verrucosus 17 (X) If there are no strong reasons for assuming Ch. chamaeleon 16, 17 (s) for the character Ch. jacksoni 15 (X) monophyly state narrow

Ch. bifidus 16 (X) parietal, we might as well assume the possibility

Ch. 16, 17 (b) of fischeri reverse development. More important seems Ch. goetzei 15 (X) the of the in to me state parietal Sphenodon punc- Ch. fuelleborni 14 (X) The tatus. Rhynchocephalia are generally Ch. melleri 17 (X)

as the sister of the Ch. cristatus 15 (X) regarded group Squamata. The in Ch. johnstoni 16 (b) parietal Sphenodon is quite narrow and

Ch. pardalis 16 (X) Romer (1956) states: "modern Sphenodon ap- Ch. brevicornis 14 (X) be of in pears to a persistently primitive type Ch. ellioti 15 (s) and be most regards considered as Ch. Hewitt, may pumilus 14 (Methuen & 1914) characteristic of the order". Ch. dilepis (quilensis) 16,17 (Methuen & Hewitt, 1914) As the I do Ch. gastrotaenia 13 (Methuen & Hewitt, 1914) to lung septation not understand

Ch. nasutus 13 & (Methuen Hewitt, 1914) — the why on this character alone neglecting all

other characters mentioned above — we have to

decide that Brookesia s.l. is the older from group

which recent Chamaeleo stems. In the smaller

and in of Chamaeleo observe decrease in the a skeleton Amphibolurus (Agamidae) 20. species we a

Romer number of In (1956) it is stated that the number of septa (see Ch. spinosus, Ch. guibei,

in 15 and It is the of ribs Squamata generally runs from to 20. Ch. fallax). striking that lungs

The number of ribs in Chamaeleo are closer to Ch. tenuis, the largest representative of the

the number in It show much elaborate general Squamata. seems nasutus- group, a more

therefore that the low numbers in Brookesia and septation than the smaller species.

be derived. In 1979 Klaver the Rhampholeon must was reminded by en-

Klaver (1981) mentions two character states larged alveolar walls found in the lungs of

in Brookesia which considers various Brookesia s.l. of the small s.l. he plesiomor- species septa

in the found in Ch. and He phic compared with the states Chamaeleo : e.g. fallax Ch. guibei. broadened flat lack of in writes: "The results of studies parietal and the septa my on

have that less this view the lungs. Consequently we to assume chameleon-lungs more or support

the ancestral chameleon looked like recent [Beddard's view "that the simplicity in lung-

Chamaeleo but for the lack of in the correlates with the small size of septa lungs structure the

in all the (which occur recent Chamaeleo) and animals"], as few or no diverticula and few or broad in different no found in small to intermediate parietal (which occurs, a septa are

form, only in Ch. pumilus s.l.). The septal species. Theoretically this might have been ex-

the of small-sized animals development and narrowing the parietal pected, as have a relatively 36 D. HILLENIUS - RELATIONSHIPS OF CHAMAELEONIDAE

and and The large lung-surface area probably can do Madagascan African species. broad

is less in without structures that increase the surface area parietal more or trigonal Africa,

further, such as septa." trapezoid in Madagascar. The transverse

Klaver does the (1981) not give reasons why his spines projecting laterally from dorsal

of 1979 valid. So and arguments are not longer my vertebrae penetrating the skin, or at least

conclusion is that Brookesia s.l. developed out of the indications of these spines on the outside of

and is there- the in the form one or more branches of Chamaeleo body of enlarged scales, can

fore than Chamaeleo. be found in The younger only Madagascan species. only

Madagascan species in which I could not

discern it is Brookesia nasus. 2. Is Brookesia s.l. monophyletic? in Some other characters occur only Africa or From table II we can see that a few characters only in Madagascar but not in all species: the in of Brookesia s.l. which occur all species are not lateral crest is absent in Africa, present in most known in Chamaeleo. These include the broad, Madagascan species, the single parietal crest is flat parietal, quite different in form from the found in and in- only Africa, as are axillary broad parietal of Ch. pumilus; the interorbital guinal pits, flexible rostral processes and lung in Rh. crest (absence or poor development diverticula. To these be added the dif- may brachyura is probably secondary); the indented ferences in indices (table II): the tails in scales (the polygonal scales in Br. decaryi and Br. Madagascan species are relatively longer than ebenaui are probably derived from these, as Br. in Africa, the heads of the Madagascan species thie Br. tuberculata and Br. vadoni both li, possess are relatively higher and broader than those of polygonal and indented scales). Africa. Partial exceptions to this division are A number of characters occurs in some the African Rh. kersteni and the Madagascan Br. species of dwarf chameleons, both African Rh. kersteni has the tail indices in nasus. largest (Rhampholeon) and Madagascar! (Brookesia s.s.), Africa and it is the only species with guiar cones in but are unknown Chamaeleo. Among these are in in a pattern that is quite common claws and scales the soles of bicuspid spinose on Madagascar. Moreover the body is much less and the in hands feet, and great difference flattened laterally than those of other African tail indices. It is Klaver average true, as (1979) species and more like those of Madagascar. On that few chameleons have shorter tails states, a the other hand it has a trigonal parietal. Br. than some Brookesia, but the overall difference is less flattened nasus has a more or body (less so clear. in the subspecies pauliani) , no transverse pro- Brygoo (1971) mentions a particular kind of the vertebrae far be cesses on (so as can seen ex- behaviour he observed in Brookesia superciliaris, ternally), a narrow head which is higher than Br. thieli and Br. vadoni: "une certaine vibration broad, a somewhat flexible snout that is less perceptible au toucher mais pratiquement in- developed but perhaps comparable with the audible. Cette vibration n'a jamais été con- rostral processus in Rh. temporalis. However the statée avec des Chamaeleo”. I observed this same parietal has a Madagascan form: trapezoid. vibration in Rhampholeon kersteni. So with some hesitation I conclude that the I conclude that the characters that differen- division in Brookesia and Rhampholeon is valid. tiate Brookesia s.l. from Chamaeleo are autapo- Even with a faint knowledge of the pygmy morphies and thus Brookesia s.l. has to be chameleons one might decide on appearance regarded as a monophyletic group. from whether a specimen comes Madagascar

from Africa (Brookesia) or (Rhampholeon).

3. Is the separation of Brookesia (Madagascar) Of course, to strict cladists all these lizards and Rhampholeon (Africa) valid? have to be called Chamaeleo until eventually this

In table II that there few we see are only a genus is divided into a number of other genera.

that characters consistently differentiate But, until much more is known about osteology 56 - 1986 37 BIJDRAGEN TOT DE DIERKUNDE, (1)

and other I do the than the morphological characters, not genus Chamaeleo, probably younger

island. feel inclined to subdivide it in this way. age of Madagascar as an

This of suggests an invasion Madagascar by

4. To the above last be of — for questions a one may a representative Rhampholeon, perhaps

added: what is the group of chameleons prob- reasons mentioned above — most closely

ably most closely related to Brookesia + Rham- resembling Rh. kersteni. pholeon?

I referred already to Klaver's remark that

Brookesia 8.1. has more characters in common ACKNOWLEDGEMENTS

Ch. than with with the group around nasutus any I Miss Alice C. Grandison am very grateful to (British other chameleon. In that Klaver same paper Museum (Natural History), London) and Dr. E. G. "demonstrate that due to hoped to ... similarity Brygoo (Muséum National d'Histoire Naturelle, Paris) the parallelism is more acceptable proposi- for the kind hospitality offered in their rich collections. I

thank Dr. G. Peters University, East Berlin), tion". However, I have not been able to find (Humboldt Dr. C. Crumly (Museum of Comparative Zoology, Har- this demonstration in the succeeding para- Instituut vard), Dr. J. P. Gosse (Koninklijk Belgisch voor graphs. Natuurwetenschappen, Brussel) and Dr. M. S.

As shown in table II, Brookesia and Rham- Historie, Hoogmoed (Rijksmuseum van Natuurlijke pholeon both have the following characters in Leiden) for lending indispensable material. Miss N.

Pruim and Mr. L. A. van der Laan common with Ch. nasutus c.s. whereas other (drawings)

(photographs) are thanked for their clear illustrations. I African or Madagascan chameleons do not: owe special thanks to Dr. Mills Tandy, who by sending in relative of in relation size, height head, height 1971 number of laid the seeds for this a living Rhampholeon in other to width of the and Dr. E. N. Arnold Museum head, axillary pits (also paper, to (British (Natural

and who kind read and Madagascan chameleons), inguinal pits History), London) was so to correct my English. flexible rostral protuberance. This leads me to

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Received: 28 June 1985