A NEW SPECIES OF THE Agama agama GROUP (SQUAMATA

Russian Journal of Herpetology

Vol. 12, No. 2, 2005, pp. 143 – 150

A NEW SPECIES OF THE Agama agama GROUP (SQUAMATA: AGAMIDAE) FROM WESTERN KENYA, EAST AFRICA, WITH COMMENTS ON Agama lionotus BOULENGER, 1896 Wolfgang Böhme,1 Philipp Wagner,1 Patrick Malonza,2 Stefan Lötters,3 and Jörn Köhler4 Submitted June 9, 2005. We describe a new species of Agama from westernmost Kenya. It is a member of the Agama agama species group characterized by small size (males up to 89 mm snout-vent length and 245 mm total length) and a unique male breeding coloration: flame- to scarlet-red head, neck and forelimbs, jet- or velvetly-black body, hindlimbs and tail root, and again a scarlet-red tail the terminal third of which is again black. Morphologically, the new species is similar to A. planiceps from southwestern Africa but has a much less depressed body. It also resembles typical A. agama from West and Central Africa, but is much smaller, less stoutly built and differently colored. It is strikingly different from the two parapatric species of the Agama agama group, viz. A. caudospinosa and A. mwanzae, and differs also considerably from the sympatric representatives of the Agama agama complex itself: from the geographically neighboring A. a. elgonis and A. a. lionotus. We provide evidence that the latter taxon deserves full species rank and that the other East African subspecies of A. agama (i.e., elgonis, dodomae, usambarae, ufipae) should be subordinated under a full species Agama lionotus. Keywords: Reptilia: Squamata: Agamidae: Agama finchi sp.n.; A. agama species group; A. agama (subspecies) complex; East Africa: Kenya, Uganda, Tanzania; taxonomy.

INTRODUCTION In September 2001, some strikingly colored agamas caught the eye of Brian W. Finch at a rocky hillside near Malaba in western Kenya, when he was in the field for observing birds. He took photographs of the attractive lizards and subsequently tried to identify them with the East Africa field guide by Spawls et al. (2002) but failed. He correctly concluded that a new species unknown to science might be involved and published his photos in the popular natural history journal “Swara” (Finch, 2003a). Its editor requested the opinion of the senior author of the field guide, S. Spawls, who confirmed that the photos would show an “absolutely distinctive” new species. When we became aware of Finch’s (2003a) article (through the courtesy of H. Dintelmann, Bonn), we were able to secure some specimens from near Malaba which is situated only 40 km NW of Kakamega Forest where 1

2 3 4

Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany; National Museums of Kenya, P. O. Box 406 58, Nairobi, Kenya Institut für Zoologie, Saarstraße 21, D-55099 Mainz, Germany Hessisches Landesmuseum, Friedensplatz 1, D-64283 Darmstadt, Germany.

the BIOTA East Africa project, run by the Zoologisches Forschungsmuseum A. Koenig holds a field station. A first visit by one of us (S. Lötters, together with B. A. Bwong and M. Muchai) resulted in the capture of three juveniles, and during a second visit, the BIOTA staff members C. Bensch, B. A. Bwong, M. Hagen, V. Muchai, and M. Peters collected three adult males and three more juveniles. This material enables us to name and to describe the new species below. MATERIAL AND METHODS The type material of the new species is deposited in the National Museum of Kenya (NMK), Nairobi, and in the Zoologisches Forschungsmuseum A. Koenig (ZFMK), Bonn. It is compared with samples of related taxa of the ZFMK collection, a sample of the Naturhistorisches Museum Wien (NMW) and with literature data. Measurements were taken with a dial calliper to the nearest 0.1 mm. Measurements and scale counts were done according to Grandison (1968) and Moody and Böhme (1984). Color pattern in life is described using field notes and life photographs. Mid-dorsal scales for REM imaging (Hitachi S-2460N at ZFMK) were taken

1026-2296/2005/1202-00143 © 2005 Folium Publishing Company

— — — — — — 12 11 12 97 52 85 86 37 119 90 42 112 17-17 18-18 17-18 18-18 18-17 18-18 19-18 18-18 18- – ZFMK 2820931 ZFMK 82094 NMK L/2534/6 NMK L/2534/3 NMK L/2534/1 NMK L/2533/3 ZFMK 82091 ZFMK 82092 NMK L/2534/2

No.

TABLE 1

Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Holotype

43.97 44.63 52.44 38.61 46.03 59.38 78.8 88.64 87.71

— — — — — 103.91 — — —

112.29 83.92 129.58 95.01 76.44 163.29 201.83 158.41 242.31

5.93 4.93 6.84 6.3 6.55 7.14 10.0 9.47 9.82

10.38 11.87 11.47 9.28 9.46 13.59 16.25 17.72 15.68

14.17 12.56 16.68 11.2 13.55 17.46 23.53 21.49 23.19

81 81 80 78 83 83 84 86 82

77 — 78 78 80 80 — 79 79

78 76 71 70 76 74 – 79 79

9-9 9-9 8-9 9-9 10-10 9-9 9-9 10-10 9-9

9-9 10-10 9-9 9-10 8-9 9-9 9-8 9-9 8-8

4(2)-4(2) — 5(2)-5(2) — 5(2)-5(2) 5(2)-5(2) 5(2)-5(2) 4(2)-5(2) 5(2)-5(2)

20-19 21-21 21-20 21-21 20-20 19-21 20-20 22-22 21-22

Femo ral pores 4th finger SnoutTail vent length length

Total length

Head height

Head width

Head length

Ventrals longi tudi nally

Dorsals longi tudi nally

Mid body scales

Supra Sub Lobules labials labials around ear

4th toe

Sub caudals

juv. juv. juv. juv. juv. subadult female male male male

Wolfgang Böhme et al.

Sex

144

from a paravertebral scale row of the posterior half of dorsum of the following specimens: A. finchi (ZFMK 82092), A. cf. finchi (Uganda, NMW 17034:1), A. agama (Cameroon, ZFMK 15222; Ethiopia, ZFMK 8709; Sudan: ZFMK 2630), A. (agama) lionotus (Kenya: ZFMK 2651); A. mwanzae (Rwanda: ZFMK 55797); and A. planiceps (Namibia: ZFMK 21961). Terminology of scale description follows Grandison (1968). RESULTS AND DISCUSSION In this section, we shall name and describe the new species and discuss its taxonomic relationships. In a second section, we shall discuss the taxonomy of the East African nominal subspecies of A. agama. 1. Description of the New Species Agama finchi sp.n. Holotype. NMK L/2534/2, adult male, km 11 on road from Malaba to Busia, Western Province, Kenya (00°34¢ N, 34°11’E, 1500 m a.s.l.), collected by C. Bensch, B. A. Bwong, M. Hagen, V. Muchai and M. Peters, 12 October 2003. Paratypes. ZFMK 82091-092, two adult males, and NMK L/2534/1, 3, and 6, three juveniles, collected with the holotype; NMK L/2533/3 and ZFMK 82093-094, three juveniles, same locality as the above specimens, collected by B. A. Bwong, S. Lötters and V. Muchai, 27 September 2003. Diagnosis. A small species of Agama (total length of adult males below 25 cm) which is characterized by a bright nuptial coloration of breeding males, the females retaining a characteristic juvenile pattern. A. finchi sp.n. differs from A. agama sensu lato (see below) not only by the small size but mainly by the nuptial colors of breeding males: flame-to scarlet-red head and forelimbs (vs. yellow, orange or greenish), jet- or velvety-black body, and hindlimbs (vs. bluish) and a bicolored, viz. red and black tail (vs. three-colored: whitish, orange/yellow, and black in West and Central African forms or bluish with narrow light rings in East African forms). Moreover, the new species has less sharply keeled dorsal scales than the A. agama populations from Ethiopia, Sudan, Central and West Africa). A. finchi sp.n. differs from A. planiceps (Namibia, S. Angola) in having keeled (vs. smooth) dorsal scales and a different color pattern; from the geographically neighboring, parapatric species A. mwanzae by stronger mucrones at the dorsal scales and a different color pattern, from the likewise parapatric A. caudospinosa by a less spiny tail scalation and again a strikingly different

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color pattern. All three species attain a considerably larger size than A. finchi sp.n. (see Table 1). It should be noted here that also the taxon kaimosae Loveridge, 1935 (synonymized with A. caudospinosa by Loveridge 1936, but regarded as a full species by Moody 1980) the type locality of which (Kaimosi, Kakamega) is close to Malaba, is also much bigger than A. finchi sp.n. (35 vs. less than 25 cm maximum total length) and is also differently colored. Description of the holotype (Fig. 1). Habitus slender, snout-vent length (SVL) 87.71 mm, tail length (TL) 154.60 mm, head length (HL) 23.19 mm, head width (HW) 15.68 mm, head height (HH) Fig. 1. Male holotype of Agama finchi sp.n. (NMK L/2534/2) in life (photo M. Hagen). 9.82 mm. Nostril slightly above canthus rostralis, pierced in hindlimbs, and tail blackish above and below. Middle the posterior part of a big nasal scale, directed obliquely piece of tail reddish, last third black above and whitish upwards. Between nostrils, a longitudinal, sharply below. keeled shield, followed by slightly keeled, irregularly arVariation. For measurements and scale counts of ranged scales in the interorbital region and on the rethe type series see Table 1. The two other adult males maining upper side of head. Supraocular scales smooth. agree with the holotype in color pattern. ZFMK 82092 A pentagonal, well differentiated parietal shield. The has the tail mutilated, ending in a club-shaped regenerscales originating from both sides of the parietal midline ate and thus lacking a black terminal third of tail; morehave their imbrications anteriorly directed, the keels in over, the head color extends a bit more onto the neck as the upper temporal region have their mucrones at the anin the holotype and the other male paratype. Coloration terior margins. The free, anterior margins of these scales in life is documented by the brilliant photographs made bear several sensory pits. Tympanum large, superficial, by B. W. Finch which are partly reproduced here (Figs. 2 about the same size as the eye, its anterior margin being and 3). In one male (see Fig. 3) there is clearly some composed of spiny, mucronate scales. On each side five black spotting on the otherwise red forelimbs. spiny tufts, three behind the tympanum, and one each on The remaining paratypes are juveniles with a charthe nape and on the neck side. Nuchal crest low, consistacteristic pattern on the upper side of head and neck. ing of 14 weakly differentiated lanceolate scales. Gular There is a white supralabial line and a yellowish-white scales flat, juxtaposed and becoming smaller towards temporal band that may pass above the shoulder into a the gular fold. Dorsal scales weakly keeled, becoming broad longitudinal flank band extending to the rear of larger posteriorly, more pointed and with stronger keels hindlimbs. Upper side of head and neck with whitish in the sacral region and on the tail base. Scales on tail arflecks and spots. This color pattern is most distinct in the ranged in weakly but nonetheless distinctly differentismallest juveniles and becomes less expressed in the ated whorls that consist of three scale rings each, dorlarger ones; particularly the bright dorsolateral bands besally strongly, ventrally weakly keeled. Ventral scales coming broken and indistinct. Dorsum dark grayish with smooth, slightly imbricate at their posterior margins. four pairs of indistinct dorsolateral flecks. Hindlimbs, One row of 12 preanal pores. Scales on forelimb with feet and upper side of tail also with darker variegations. strongly keeled scales, on the upper arm twice as large as Lower parts grayish to whitish, with some irregular gray the dorsals, becoming smaller towards the underside and lateral vermiculation on the throat which become more the manus. 3rd and 4th finger longest, equal-sized, digital distinct longitudinal gray lines in the older juveniles length decreasing 2-5-1, subdigital lamellae keeled. (which seem to be subadult females, at least ZFMK Hindlimbs also covered by distinctly keeled scales, on 82094). An adult female in live coloration, with reddish the upper thighs as large as dorsals, becoming larger todorsolateral bands, is shown in Fig. 4). wards the lower thighs and feet. Distribution. Agama finchi sp.n. is known with cerColor (in alcohol, after 15 – 16 months of preservatainty only from the type locality near Malaba in westion). Head, nape, and forelimbs flesh-colored; body, ternmost Kenya, in parapatry to A. caudospinosa (con-

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(J. Köhler) and tentatively consider the new species to occur in the Murchison Falls NP, too (see Remarks). Habitat. B. W. Finch’s first photographic documentation and the type series originated from the rocky hillside outside Malaba, and the agamas exhibited a rupicolous lifestyle. They were associated with five-lined skinks, Trachylepis quinquetaeniata (not T. margaritifera; see Finch, 2003a; Spawls, 2003; if their taxonomic split on the species level is justified at all). The Murchison Falls NP population, however, was also found on buildings, on the ground and also on treetrunks. Fig. 2. Agama finchi sp.n., male (not collected) from Malaba, western Kenya, basking on rocky outcrops (photo from Finch 2003a). Etymology. The new species is dedicated to Brian W. Finch, Nairobi, who first to saw this remarkable agama in the stantly misnamed as “A. caudospina” by Spawls et al. field and recognized at the same time that it was a still 2002, following the erroneous citations by Loveridge undescribed. 1929, 1935, 1957) and A. mwanzae, and in sympatry to Remarks. According to our scarce data (see above), A. agama lionotus (Fig. 5). In addition, Brian Finch was the Murchison Falls NP agama is indeed similar to the able to take photographs of similar agamas in the MurMalaba population, but seems to be of somewhat stouter chison Falls NP, Uganda (Fig. 6) which he considered appearance. In contrast to the Malaba population, the conspecific with the Malaba population (Finch, 2003b). forelimbs have the same blackish color as the body and As there is no voucher specimen of this population the same red head coloration as in the latter. However, as available to us, we can rely here only on his photographs at least one male from Malaba (Fig. 3) has some black and on the unpublished observation of one of us speckling on its forelimbs, and also a male from Murchison Falls NP (Fig. 6) has some red speckling there, one might expect some geographic variation in this character. On the other hand, the Murchison Falls agamas have clearly bicolored tails (Fig. 6) (without a third light proximal zone as in Agama agama, see Fig. 10) so that Finch (2003b) may have justifiably regarded them

Fig. 3. Agama finchi sp.n., male (not collected) from Malaba, western Kenya, basking on rocky outcrops (photo from Finch

Fig. 4. Agama finchi sp.n., female (not collected) from Malaba, western Kenya, basking on rocky outcrops (photo M. Hagen).

A New Species of the Agama agama Group (Squamata: Agamidae) from Western Kenya conspecific with the Malaba population. This argues for a much wider distribution of A. finchi sp.n. But new voucher material from Uganda is essential to corroborate this assumption. The Naturhistorisches Museum of Vienna (NMW) holds three specimens of “Agama planiceps” from “Yale River, Uganda,” which are small-sized and have weakly keeled scales as in A. finchi sp.n. Also their scale counts fall within the range of the latter. Unfortunately, we were unable to trace the locality on our maps. It cannot be identified with the Kenyan Yala River which crosses Kakamega Forest because the latter comes from the east and flows westwards into lake Victoria. Although the nuptial coloration is not preserved in these specimens, we assign them here, with some reservation, also to A. finchi sp.n. Taxonomic relationships. At this stage, taxonomic relationships of A. finchi sp.n. can only be described on the basis of morphological similarity. Apart from its striking male breeding coloration, it can be well diagnosed by the weakly keeled scales (Fig. 7A, B) (strongly keeled in A. agama from Cameroons, Sudan and Ethio-

Fig. 5. Distribution map of Agama finchi sp.n. (encircled star), Agama mwanzae, and Agama caudospinosa. Stars indicate type localities of the nominal taxa described. Outlines of shaded areas after Spawls et al. (2002).

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pia (Fig. 8), smooth in A. planiceps (Fig. 7E, F) and nearly so in A. mwanzae (Fig. 7G, H). However, scale counts as well as scale morphology are subjected to ecological selection, and similar conditions of characters might be the result of convergence rather than of relationship. This may also be true for coloration characters, even if they are the result of sexual selection. Therefore, scalation and coloration characters are indeed reliable to delimit and diagnose taxa, but for determining phylogenetic relationships additional character sets, particularly from molecular genetics, are needed. The use of adaptive characters (including body size and proportions, scale morphology and color pattern) has led to an unstable taxonomic concept of the Agama agama species group, as can be seen from the synonymy-chresonymy list in Wermuth (1967). Rock-dwelling forms such as A. caudospinosa or A. mwanzae have both

Fig. 6. Agama cf. finchi, male (not collected) from Murchinson Falls NP, western Uganda, climbing on tree trunk (photo B. Finch).

148

Fig. 7. Dorsal (left column) and dorsolateral view (right column) of middorsal scales of Agama finchi sp.n. (Kenya, paratype, ZFMK 82092, A and B), Agama cf. finchi (Uganda, NMW 17034:1, C and D), Agama planiceps (Namibia, ZFMK 21961, E and F), and Agama mwanzae (Rwanda, ZFMK 55797, G and H).

been regarded earlier as subspecies of A. planiceps (see Loveridge, 1932), but their status as full species is today beyond any doubt (Moody, 1980; Broadley and Howell, 1991). The same is true for A. spinosa which was formerly regarded as a subspecies of A. agama (Wermuth 1967). The taxonomic situation of the latter complex remains chaotic and will be briefly discussed in the following section of this paper. 2. The Agama agama (Sub)Species Complex The East African taxa related to Agama agama are currently still regarded as subspecies of a single, polymorphic and wide-ranging species (e.g., Spawls and Ro-


Fig. 8. Dorsal (left column) and dorsolateral view (right column) of middorsal scales of Agama lionotus (Kenya, ZFMK 2651, A and B), Agama agama (Ethiopia, ZFMK 8709, C and D), Agama agama (Sudan, ZFMK 2630, E and F), and Agama agama (Cameroon, ZFMK 15222, G and H).

tich, 1997; Razzetti and Msuya, 2002; Spawls et al., 2002). However, a basic difference separates two distinct groups: Breeding males of the populations from Uganda, Kenya and Tanzania (see the distribution map in Spawls et al., 2002) differ from more northern (Ethiopia, Sudan), Central and West African populations in having tails of the same (mostly bluish) ground color as their bodies, but with numerous narrow, light bluish to whitish crossrings (Fig. 9). Males from the last-named regions have in contrast typically tricolored tails, i.e., the (also mostly bluish) body is followed by a light whitish proximal part of the tail, a yellowish or red (corresponding to the respective head coloration) median and a

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black terminal part of the tail (Fig. 10). Because this striking color pattern of either ringed or tricolored tails has certainly behavioral significance in these strongly territorial lizards and is thus certainly subjected to sexual selection by the females, we regard both groups as two distinct species. This view is supported by the fact that within each of these two species, there is geographically correlated variation which has led to the delimitation of several nominal subspecies in the past. These two groups of names (see Wermuth, 1967) can be hierarchically subordinated under Agama agama (e.g., africana, congica, mucoFig. 9. Male Agama lionotus from central Kenya (photo B. Finch). soensis, savattieri) or under Agama lionotus as characterized above (e.g., dodomae, elgonis, turuensis, ufipae, usambarae). For the latter, we have marked their type localities within the entire lionotus range (Fig. 11) as recently given by Spawls et al. (2002). The validity of these forms or even a full species status of some of them is still under debate. However, the chorological relationships between elgonis (misnamed “elgonensis” in Spawls et al., 2002) and turuensis [the latter described by Loveridge (1932) and again synonymized with the former by himself: Loveridge (1939, 1957)] in regard to dodomae elucidate again the urgent need of a systematic revision of these agamas which will be extremely interesting but can of course not be the aim of this paper. Fig. 10. Male Agama agama from Mokolo, northern Cameroon (photo W. Böhme). Agama finchi sp.n., however, can easily be separated from all East African repB. A. Bwong, M. Hagen, V. Muchai, and M. Peters, provided resentatives of the A. agama (sub)species complex as us with the necessary voucher material and made thus the dewell as from the A. agama species group. If inserted into scription possible. H. Grillitsch and F. Tiedemann of the Naturthe key of the genus by Spawls et al. (2002), it should alhistorisches Museum, Vienna, kindly loaned us some comparready key out at the first dichotomy where large, climbative specimens from Uganda. Karin Ulmen and Ursula Bott, ing forms of more than 30 cm with vividly colored heads ZFMK Bonn, helped with skillfully preparing the REM photoin the males are opposed to small, ground-dwelling graphs, the manuscript and the figures for print, and A. Burforms below 30 cm without vividly colored heads. The mann (ZFMK) gave useful information. small size and the nonetheless climbing life habits and extremely vivid colors in males make A. finchi sp.n. a unique, unmistakable species. REFERENCES Acknowledgments. We thank H. Dintelmann, Bonn, for bringing our attention to the first article by B. W. Finch on the remarkable new agama. Brian Finch himself helped us with additional information on the Malaba and the Murchison Falls populations and provided us with his brilliant photographs of the new species and other Kenyan agamas. The members of the BIOTA East project at Kakamega Forest, C. Bensch,

Broadley D. G. and Howell K. M. (1991), “A checklist of the reptiles of Tanzania, with synoptic keys,” Syntarsus, Bulawayo, 1, 1 – 70. Finch B. W. (2003a), “A dazzling display,” Swara, 26(1), 12 – 14. Finch B. W. (2003b), “Also in Uganda,” Swara, 26(3), 13.

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Wolfgang Böhme et al. Moody S. M. (1980), Phylogenetic and Historical Biogeographical Relationships of the Genera in the Family Agamidae (Reptilia: Lacetrilia). Unpublished Ph.D. Thesis, Univ. of Michigan. Moody S. M. and Böhme W. (1984), “Merkmalsvariation and taxonomische Stellung von Agama doriae Boulenger, 1885 and Agama benueensis Monard, 1951 (Reptilia: Agamidae) aus dem Sudangürtel Afrikas,” Bonn. Zool. Beitr., 35(1/3), 107 – 128. Razzetti E. and Msuya C. A. (2002), Field Guide to the Amphibians and Reptiles of Arusha National Park (Tanzania), Negri, Varese. Spawls S. (2003), “Absolutely distinctive,” Swara, 26(1), 14. Spawls S., Howell K., Drewes R., and Ashe J. (2002), A Field Guide to the Reptiles of East Africa, Acad. Press, San Diego – San Francisco – New York – Boston – Sydney – Tokyo. Spawls S. and Rotich D. (1997), “An annotated checklist of the lizards of Kenya,” J. East Afr. Nat. Hist., 86, 61 – 83. Wermuth H. (1967), “Liste der rezenten Amphibien and Reptilien: Agamidae,” Das Tierreich, 86, i – xiv, 1 – 127.

MATERIAL EXAMINED Agama cf. finchi: Uganda: NHMW 17034:1-3, Yale (?) River

Agama agama: Fig. 11. Distribution map of the Agama agama (sub)species complex in East Africa. Stars indicate type localities of available names. Outline of shaded area after Spawls et al. (2002).

Grandison A. G. C. (1968), “Nigerian lizards of the genus Agama (Sauria: Agamidae),” Bull. Br. Mus. Nat. Hist. Zool., 17(3), 67 – 90. Loveridge A. (1929), “East African reptiles and amphibians in the United States National Museum,” Smithsonian Inst. U.S. Natl. Mus. Bull., 151, 1 – 135. Loveridge A. (1932), “New reptiles and amphibians from Tanganyika territory and Kenya colony,” Bull. Mus. Comp. Zool. Cambridge, 72(10), 376 – 387. Loveridge A. (1935), “Scientific results of an expedition to rain forest regions in eastern Africa. I. New reptiles and amphibians from East Africa,” Bull. Mus. Comp. Zool. Cambridge, 79(1), 1 – 19. Loveridge A. (1936), “Scientific results of an expedition to rain forest regions in eastern Africa. V. Reptiles,” Bull. Mus. Comp. Zool. Cambridge, 79(5), 209 – 337. Loveridge A. (1957), “Checklist of the reptiles and amphibians of East Africa (Uganda; Kenya; Tanganyika; Zanzibar),” Bull. Mus. Comp. Zool. Cambridge, 117, 153 – 362.

Ghana: R07759-61, Quitta (Kitta) = Keta; Nigeria: R07778, Nigerdelta; Sudan: ZMH R07777, Mongalla; Tanzania: ZMH R04699, 04701, 04704, Kwa Mtoro: Usandaui;

Agama lionotus: Kenya: ZFMK 2650 – 52, Nairobi; ZFMK 82063, Cherangani Hills; NHMW 24024:1 – 2, Tsavo West; ZMB 11482, Tana; ZMB 55651, Kilimatinde; ZMB 22466, Kibwezi; ZMB 28825, Kibwezi; ZMB 55574, Kibwezi;

Agama caudospinosa: Kenya: ZFMK 9025; ZFMK 8701;

Agama mwanzae: Rwanda: ZFMK 55797 – 98, Kibungo: Rugarama Comm. Rukira; ZFMK 51195, Mpanga; ZFMK 61664, Prov. Kibungo: Ntaruka; Kenya: ZFMK 82075 – 77, Masai Mara area; Tanzania: ZMB 30771, Mwanga, Shandwa

Agama planiceps: Namibia: ZFMK 21961, Kaoko: Werda; ZFMK 55062, Windhoek; ZFMK 18395, Frauenstein; NHMW 35763:2, Windhoek: Tintenpalastpark; NHMW 35109:1, Erongo; NHMW 35800:2, Distr. Windhoek: Gobabisweg; NHMW 17035, Windhoek; ZMH R07765 – 69, Okahandja; ZMH R07779 – 89; ZMB 18269; ZMB 35490, 92, Okahandja; ZMB 54560 – 61, Walfish Bay; ZMB 18257 – 60; ZMB 18263 – 64; ZMB 55741; ZMB 55744 – 49; ZMB 4200, Neu Barmen; ZMB 44045, Distr. Windhoek: Lichtenstein; Angola: ZMH R07737 – 40, R07751 – 58, Otschinjau; ZMH R07764; ZMH R07770 – 6, Alto Cubal.