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iVl iul s e tul iml oi L ounparatti we /^oology 


US ISSN 0006-9698 

Cambridge, Mass. 

4 November 2011 

Number 527 


Philipp Wagner 1 - 2 and Aaron Bauer 1 ' 3 

Abstract. A new dwarf lizard of the genus Agama Daudin, 1802 (Sauria: Agamidae), is described from central 
Ethiopia. This dwarf agama (< 59 mm snout-vent length [SVL]) was compared with other dwarf congeners (< 65 mm 
SVL) in West, Central, and East Africa and to medium-sized agamas (< 92 mm SVL) occurring in the Horn of 
Africa. The new species is characterized by a homogeneous body scalation; keeled but nonmucronate dorsal scales; 
smooth ventral, gular, and upper head scales; and having the nasal scale on the canthus rostralis. It is 
morphologically well differentiated from all congeners to which it was compared in having a nuchal crest, few tufts of 
short spinose scales, and the nasal scale on the canthus rostralis and can further be identified by its large occipital 
scale, smooth gular and ventral scales, and keeled, but not strongly mucronate or spinose, dorsal scales. The new 
species may live in colonies, like many of the larger rock Agama species, and based on the presence of multiple age 
classes at the same time of year, it is probably not an annual species, as are some other dwarf Agama. The affinities of 
the new species are unclear but, on zoogeographic grounds, may lie with congeners to the west of the Rift Valley. 

Key words: Agamidae; Agama; A. bottegi; A. hartmanni; A. persimilis; A. rueppelli; A. gracilimembris; A. 
weidholzi; Agama sp. nov.; Africa; Ethiopia; new species 

The Horn of Africa is a peninsula in East 
Africa jutting into the Arabian Sea and lying 
along the southern side of the Gulf of Aden. 
It is the easternmost projection of the 

1 Department of Biology, Villanova University, 800 
Lancaster Avenue, Villanova, Pennsylvania 19085, 
U.S.A.; e-mail: 

2 Zoologisches Forschungsmuseum A. Koenig, Ade- 
nauerallee 160, D-53113 Bonn, Germany; e-mail: 
philipp . wagner . zfmk@uni-bonn . de 

3 Research Associate, Department of Herpetology, 
Museum of Comparative Zoology, 26 Oxford Street, 
Cambridge, Massachusetts 02198, U.S.A. 

African continent and encompasses the 
countries of Eritrea, Djibouti, Ethiopia, 
and Somalia. Its lowlands are dominated 
by arid and semi-arid habitats in spite of 
their proximity to the Equator. Monsoon 
winds lose their moisture before reaching 
Djibouti and Somalia, with the result that 
most of the Horn receives little rainfall 
during the southeast monsoon season (June 
to end of August). 

The Horn is the greatest hotspot of 
African agamids and certain other lizards 

The President and Fellows of Harvard College 2011. 


No. 527 

(Lanza, 1983), e.g., the gekkonid genus 
Hemidactylus (Bauer et al., 2010). Apart 
from Laudakia Gray, 1845, all African 
agamid genera are present in the area (see 
Wagner, 2010), including the regionally 
endemic Xenagama Boulenger, 1885. The 
highest species diversity occurs in Agama 
Daudin, 1802 (seven species), and Acantho- 
cercus Fitzinger, 1843 (six species), whereas 
Trapelus Cuvier, 1816, and Pseudotrapelus 
Fitzinger, 1843, are each represented by only 
a single species (Largen and Spawls, 2010; 
Wagner, 2010). All agamid lizards inhabiting 
the Horn are adaptated to arid habitats, and 
most of them occur in rocky savannahs. 
Uromastyx Merrem, 1820, inhabits deserts 
and Xenagama dry woodland to semidesert, 
whereas Acanthocercus atricollis minutus 
(Klausewitz, 1957) and A. atricollis loveridgei 
(Klausewitz, 1957) are most likely tree 
dwellers. For most species, our knowledge 
of ecology is too meager to discuss ecological 
adaptations, but a high degree of variation in 
morphology is obvious. The size of the 
agamid lizards in the Horn ranges from the 
small Agama per similis Parker, 1942 (snout- 
vent length [SVL] max. 64 mm), to medium- 
sized lizards (e.g., Xenagama; SVL max. 
90 mm), to the large Uromastyx (SVL max. 
174 mm), the latter both characterized by 
thick, armored tails, but most of the other 
species are very conservative in body shape 
and similar to the widespread genus Agama 
Daudin, 1802. 

Because of the limited information avail- 
able about agamid lizards in Africa and 
especially those occurring in the Horn, 
museum vouchers from the area are of 
special interest. The Museum of Compara- 
tive Zoology (MCZ) holds a series of 
specimens initially identified as "Agama 
agama agama' from Ethiopia. According to 
Largen and Spawls (2006, 2010) Agama 
agama (Linnaeus, 1758) is present in Ethio- 
pia, but this is, in fact, an artifact of the use 

of an older taxonomic concept of this species 
(see Wagner, 2011). Wagner et al. (2011) 
have shown that Agama finchi Bohme et al., 
2005, is the only representative of the A. 
agama s. str. species group in the Horn. The 
MCZ series consists of quite small specimens 
(SVL 25.0-58.6) that might be mistaken for 
juveniles, but it includes one obviously 
gravid female. Therefore, this series is 
distinctly smaller than most other Agama 
species known from the Horn and compara- 
ble in size only to dwarf species of the genus. 

Only a few dwarf species of Agama are 
known. Agama weidholzi Wettstein, 1932 
(SVL max. 65 mm), is only known from a 
restricted area in Senegal, Gambia, western 
Mali, and Guinea-Bissau (for a detailed 
distribution, see Wagner et al., 2009). It is 
easy to identify by its large white-bordered 
dark patch on the shoulders and the thin 
vertebral stripe in males. Another dwarf 
Agama is A. gracilimembris Chabanaud, 
1918 (SVL max. 57 mm), described from 
Benin and now known from further east in 
Nigeria (Grandison, 1969; Gartshore, 1985), 
Cameroon (Bohme, 1975), and the Central 
African Republic (Joger, 1990; see also 
species account herein). Both species are 
among the rarest members of the genus in 
Africa, and information about their habitat 
preferences, behavior, and distribution is 
limited. Relationships between the two dwarf 
species have not yet been investigated. 
Grandison (1969) assumed a close relation- 
ship based on morphology, but Wagner et al. 
(2009) have shown that A. weidholzi is closely 
related to the Guinean Agama cristata 
Mocquard, 1905, and Agama insularis Cha- 
banaud, 1918 (however, A. gracilimembris 
was lacking from their analysis). 

Three other dwarf to medium-sized Agama 
occur in the Horn of Africa: A. persimilis 
(SVL max. 64 mm), Agama hartmanni Peters, 
1869 (SVL max. 92 mm), and Agama 
rueppelli Vaillant, 1882 (SVL max. 88 mm). 



Their relationships are also unknown but are 
under investigation by the authors. All three 
are solitary, only establishing territories 
during the breeding season (Wagner, unpub- 
lished data) and are, therefore, distinct in 
behavior relative to the mostly larger (SVL 
80-145 mm) rock Agamas. All are poorly 
known, but it is uncertain if they are truly 
rare or only underrepresented in museum 
collections. Agama rueppelli is the most 
common, but A. persimilis and A. hartmanni 
are only known from a small number of 
vouchers from a handful of localities. Two 
other species occurring in the Horn, Agama 
cornii Scortecci 1928 and Agama bottegi 
Boulenger, 1897, are only known from a 
few specimens each, and their taxonomic 
status is questionable. Because all these 
lizards are poorly known, a short review of 
each species including information from 
museum vouchers and literature is presented 
herein to facilitate the comparison of these 
taxa with the MCZ series of "Agama agama 
agama" from Ethiopia and to help establish 
the identity of the members of this series. 


Material. Specimens from the following 
institutes were examined or referenced for 
locality data (see Appendices 1 and 2): 
California Academy of Sciences (CAS), San 
Francisco, California, U.S.A.; Museo Civico 
di Storia Naturale di Genova, Italy 
(MSNG); Museum of Comparative Zoology 
(MCZ), Harvard University, Cambridge, 
Massachusetts, U.S.A.; Museum d'Histoire 
Naturelle (MHNG), Geneve, Switzerland; 
Museum Nationale d'Histoire Naturelle 
(MNHN), Paris, France; Naturhistorisches 
Museum Wien (NHMW), Vienna, Austria; 
Zoologisches Forschungsmuseum A. Koenig 
(ZFMK), Bonn, Germany. 

Name-bearing types of the following taxa 
were examined for comparison: A. bottegi, 

A. cornii, A. gracilimembris, A. hartmanni, 
A. persimilis, A. rueppelli rueppelli, Agama 
rueppelli septentrionalis and A. weidholzi (see 
Appendix 1). 

Moreover, data and descriptions presented 
by Parker (1942), Grandison (1968, 1969), 
Lanza (1978), Gartshore (1985), and Largen 
and Spawls (2010) and relevant original 
descriptions (Peters, 1869; Vaillant, 1882; 
Boulenger, 1897; Chabanaud, 1918; Scortecci, 
1929; Wettstein, 1932; Parker, 1942) were 
consulted. Distributional data were based on 
specimens examined (Appendix 1), reliably 
determined but unexamined specimens in 
museum collections (Appendix 2), and addi- 
tional literature sources (Appendix 3). 

Digital X-ray images of specimens were 
obtained using a Faxitron closed cabinet X- 
ray (LX-60, Faxitron Corp.) with a Varian 
flat-panel digital X-ray detector. 

Measurements. Scale counts and terminol- 
ogy follow Grandison (1968), and measure- 
ments were taken with callipers to the 
nearest 0.1 mm. The following values were 
used: SVL, from tip of snout to cloaca; head 
width (HW), measured at the point of 
greatest width; head height (HH), measured 
at the point of greatest height; head length 
(HL), measured from behind the tip of 
retroarticular process to tip of snout; tail 
length (TL), measured from posterior lip of 
cloaca to tip of tail; supralabials (SL), 
number of all supralabial scales; infralabials 
(IL), number of all sublabial scales; scales 
around midbody (SaM), number of scale 
rows around body midway between the 
limbs; precloacal pores (PP), number of 
rows and number of pores in total; sub- 
digital lamellae (SDL), number of lamellae 
under the fourth toe or finger, respectively; 
ventral scales (VS), longitudinal ventral 
scales along midbody from shoulders to 
cloaca; dorsal scales (DS), longitudinal 
dorsal scales along midbody from shoulders 
to cloaca. 


No. 527 


The specimens of the unknown dwarf 
Agama from Ethiopia were compared in 
detail with the other dwarf congeners (A. 
gracilimembris, A. persimilis, A. rueppelli, A. 
weidholzi, see also table 1 ) as well as with the 
medium-sized A. hartmanni, with which it 
shares similarities of head and body scala- 
tion. Two other rarely mentioned taxa, A. 
cornii and A. bottegi, were also included in 
the comparison. As noted by other authors 
(e.g., Largen and Spawls, 2010, see also 
species account below), A. cornii has been 
recognized as a synonym of A. hartmanni 
because the type and sole specimen of the 
former species is identical with respect to 
relevant characters with the types of the 
latter. The status of A. bottegi is still 
doubtful. The name-bearing type was exam- 
ined, and additionally the original descrip- 
tion and illustration shows that A. bottegi is 
clearly distinct from the other taxa compared 
herein (e.g., large nuchal and dorsal crests 
present; see species account below). A 
comparison of the head scalation between 
dwarf species mentioned above revealed 
differences in scale tufts, nuchal crest, and 
head scales (see Fig. 1) with respect to MCZ 
R- 146685-89 (see species accounts). Other 
regional congeners could be excluded on the 
basis of size and other features, and we thus 
recognize this Ethiopian dwarf Agama as a 
new species, which we describe below. 


Agama persimilis Parker, 1 942 

Holotype. BMNH 1937.12.5.64, adult fe- 
male, from "45°50'E X 8°N in the Haud 
[north-eastern part of Ogaden, Somali region 
Ethiopia]," collected by Capt. R.H.R Taylor 
on 25 XI 1934. 

Diagnosis (partly fide Parker, 1942). A 
dwarf Agama reaching a total length of 
about 160 mm, females (SVL 56-64) are 

larger than males (SVL 43-54). Body strong- 
ly depressed, hind limbs long. Gular pouch 
absent. Tail about 65% of the total length, 
very broad at its base, tapering sharply. 
Head convex with the upper surface of the 
snout flat. Head scales on the upper parts 
moderately large, with the occipital scale 
enlarged. Nostril, directed upward and back- 
ward, pierced in the posterior part of a nasal 
scale above the canthus rostralis. Usually 
with one scale between the nasal and the first 
supraciliary scale. Scales of the supraocular 
region longitudinally elongate, with a dull 
keel. Nuchal and dorsal crests absent. Ear as 
large as the eye opening, superficial and 
completely exposed. Lower and posterior 
borders of the ear as well as the sides of the 
neck with tufts of spines, longest spines 
about half the length of the eye opening. 
Tufts of spines not encroaching upon the ear 
opening. Body scales large, homogeneous, 
imbricate, keeled, and mucronate, with 52- 
57 scale rows around midbody. Thirty 
vertebral scales between the insertions of 
the limbs. Gular and ventral scales smooth, 
ventrals much smaller than the dorsals. 
Lamellae 16-18 under fourth toe, third and 
fourth toes nearly equal in length. Males 
with one row of 10-12 precloacal pores. 

Distribution. Eastern Ethiopia, Somalia, 
and E/NE Kenya (see Fig. 2). 

Agama hartmanni Peters, 1869 

Syntypes. ZMB 4355 (three specimens, 
holotype not designated), from "Dongola 
[Sudan]," collected by Dr. Hartmann and 
[Adalbert] von Bamim. 

Synonym. Agama cornii Scortecci, 1929, 
Holotype: MCM 1193, from "Urn Ager, 
[Om Ager, Eritrea]," collected .by the "Mis- 
sione Corni, Calciati." 

Diagnosis (partly fide Peters, 1869). A 
poorly known, medium-sized agamid lizard 
reaching a total length of 230 mm and a SVL 
up to 92 mm, with a depressed body and 



Figure 1 . Comparison of head scalation in dwarf and selected medium-sized agamas discussed in this paper 
(images are not the same scale): A, Agama sp. nov. (MCZ R- 146689); B, Agama hartmanni (BMNH 1913.9.16.9); C, 
Agama rueppelli (BMNH 1946.8.24.50, syntype); D, Agama per similis (BMNH 1937.12.5.64, holotype); E, Agama 
gracilimembris (MNHN 1904.0115, syntype); F, Agama weidholzi (MCZ R-44311, paralectotype). 


No. 527 




= A 






= A 



= A 



= A 



= A 

lucyae sp. n. 





26° 27 



3„v 30 


7 33 


?a 36 

34 t7 7 

AA 61 





7 65 
5U.67 59 58 

43^4^41 50 


K-T ■ 

5 V 



48 V 




Figure 2. Distribution of relevant dwarf or medium-sized Agama lizards. A, the entire distribution area of A. 
weidholzi; for locality details, see Wagner et al. (2009). 1, Wa; 2, Pendjari NP; 3, Igbetti; 4, Ado-Awaiye; 5, Kura; 6, 
Samaru; 7, Falgore; 8, Zaria; 9, Anara; 10, Kano; 11, Ringim; 12, Yankari; 13, Zonkwa; 14, Lafia; 15, Pandam; 16, 
Shendam; 17, Hung; 18, Wukari; 19, Boki; 20, Benue NP; 21, between Kotissako and Sakoumba; 22, Ndele; 23, 
Koumbala; 24, Bahr el Ghazal; 25, Boma Hills; 26, Bahr el Zeraf; 27, Taufikia; 28, Omhajer; 29, Dongola type 
locality; 30, Boorama; 31, Hargeysa Airport; 32, Buq Village; 33, Buran District; 34, Bahr-el-Gebel; 35, Lokichoggio; 
36, Lokomarinyang; 37, Lokitaung; 38, Lokori; 39, Lodwar; 40, Loyengalani; 41, Marsabit; 42, Loperot Kalabata; 
43, Kakuma; 44, Kanapoi; 45, Lake Baringo; 46, Kajiado; 47, Voi; 48, Mbunyi; 49, Ngomeni; 50, Wajir; 51, 
Mandera; 52, Haud, type locality; 53, Kodok; 54, Geriban; 55, Har Addei; 56, 55 km N-NE of Dusa Mareb; 57, El 
Bur [Ceel Buur]; 58, Bud Bud; 59, Iesdmma; 60, Hard; 61, Bohodle; 62, Sassabana; 63, Gumboworen; 64, Dabanac; 
65, confluence of the Web and the Ganana [Ganale]; 66, Dinsor; 67, Lugh [Luug], type locality; 68, 58 km NW of 
Soddo, type locality; 69, Singa; 70, Karin. 

moderately long limbs. Gular pouch absent. 
Head convex, not as flat as in A. rueppelli or 
A. persimilis. Tail about 60% of total length. 
Head scales moderately large, smooth, with 
the occipital scale usually enlarged. Nostril 
nearly as large as the nasal scale, round, 
directed dorsally and laterally, pierced in the 
middle of the nasal scale, situated on the 
canthus rostralis. Usually one to two scales 
between nasal and first supraciliary scales, 
supraciliary scales smooth. Nuchal and 
dorsal crest absent. Ear opening smaller than 
the eye, superficial and completely exposed. 
No tufts of spinose scales near the ear or on 
the neck, a few single erect scales scattered 
near the ear opening. Body scales homoge- 
neous, keeled, mucronate, and erect, with 
76-81 scale rows around midbody, ventral 
smaller than dorsal scales. Gular and ventral 
scales smooth. Males with 9-12 precloacal 

pores. Third and fourth toes nearly equal in 
length, 16-21 lamellae under fourth toe. 

Distribution. Sudan, South Sudan, Eritrea 
(see Fig. 2 for details). 

Remarks. Wermuth (1967) considered A. 
cornii a doubtful species and the status 
remained dubious until Moody (1980) syn- 
onymized it with A. hartmanni without 
explicit justification. Because Moody's 
(1980) doctoral dissertation was never pub- 
lished, this taxonomic act has been largely 
overlooked. Barts and Wilms (1997) men- 
tioned it again as a valid species and later 
Largen and Spawls (2010:239) mentioned A. 
cornii as an invalid name and treated it as a 
synonym of A. hartmanni. We reexamined 
the holotype of A. cornii and compared it 
with the description and types of A. hart- 
manni (Peters, 1869). Based on this exami- 
nation we follow Moody (1980) and Largen 



and Spawls (2010) in recognizing A. cornii as 
synonym of A. hartmanni. No differences 
were found between the single type specimen 
and specimens of the latter nominal species, 
and both share all characters mentioned 

Agama rueppelli YaiMsLnt, 1882 

Syntypes. MNHN 5897; BMNH 
1946.8.24.50 (syntypes, see remarks below), 
from "Bender-Meraya, Pays Comalis [= So- 

Synonym. Agama vaillanti Boulenger, 
1895. Lectotype: MSNG 28850, adult male, 
from "Ogaden," Ethiopia. 

Diagnosis. A medium-sized Agama with a 
maximum total length of 280 mm. Males 
(58-88 mm SVL) smaller than females (76- 
88 mm SVL). Body depressed, hind limbs 
long. Tail more than 65% of total length, 
very broad at its base, tapering sharply. 
Gular pouch absent. Head convex, with the 
upper surface of the snout flat. Head scales 
smooth, equal to ventral scales, with the 
occipital scales variable in size (equal to 
other head scales to slightly enlarged). 
Nostril directed posterodorsally, completely 
visible from above, pierced in the posterior 
part of a large nasal scale, situated above the 
canthus rostralis. Usually with one to two 
scales between nasal and first supraciliary 
scales. Supraciliary scales smooth. Nuchal 
and dorsal crest absent. Ear opening only 
half the size of the eye, surrounded by tufts 
of spinose scales hiding the tympanum. Body 
scales large, homogeneous, keeled, mucro- 
nate, and erect, in 54-64 scale rows around 
midbody. Vertebral scales 59-66. Gular and 
ventral scales smooth, ventrals smaller than 
dorsal scales. Lamellae 14-16 under the 
fourth toe, third and fifth toes nearly equal 
in length. Males with one row of 9-13 
precloacal pores. 

Distribution. Sudan, South Sudan, Ethio- 
pia, Somalia, and Kenya (see Fig. 2). 

Remarks. Three different subspecies are 
currently recognized (Largen and Spawls, 
2010), but their status is questionable. 
Therefore, we only refer to A. rueppelli, but 
a review of the species is needed. 

Guibe (1954), in his agamid type catalogue 
of the Paris museum, incorrectly regarded 
the single MNHN syntype as the holotype. 
Later, Brygoo (1988) indicated that the 
second syntype of A. rueppelli left the Paris 
collection in 1895 without a documented 
destination, and Brygoo (1988) was unaware 
of its present whereabouts. Boulenger (1895) 
in his description of A. vaillanti mentioned 
that he was "able to compare the specimen 
with one of the types of A. rueppelli'' and 
obviously this type is the one donated or 
exchanged from the Paris collection. 

Agama bottegi Boulenger, 1898 [1897] 
incertae sedis 

Holotype. MSNG 28548, adult male, from 
"Lugh" (03°47'46N, 42°32'32E), Somalia, 
collected by Capt. U. Ferrandi (V. Bottego 

Diagnosis (fide Boulenger 1898 [1897]). A 
large Agama with a maximum length up to 
355 mm (SVL 120 mm). Head moderately 
convex, body scarcely depressed, hind limbs 
strong. Gular pouch absent. Tail twice as 
long as SVL. Head convex, snout not flat as 
in persimilis. Head scales moderately large, 
smooth, with the occipital scale enlarged. 
Nostril tubular, directed backward; pierced 
in the posterior part of a small nasal scale, 
situated on the canthus rostralis. Two scales 
between the nasal and the first supraciliary 
scale. Supraciliary scales smooth. Strong 
nuchal crest present in males, with conical 
scales equal to the diameter of the ear 
opening. Low crest extending down the back 
and to the tail. Ear opening larger than the 
eye. Tufts of spinose scales near the ear and 
on the sides of the neck present, longest 
spines about two-third the diameter of the 


No. 527 

ear opening. Body scales large, homoge- 
neous, imbricate, strongly keeled, and 
strongly mucronate; in 53 rows around 
midbody. Ventral scales keeled, smaller than 
the dorsals. Fourth and third fingers equal, 
fourth toe slightly longer than third, fifth not 
extending beyond first. Tail slightly com- 
pressed, covered with strongly keeled scales, 
which are larger than the body scales. One 
row of precloacal pores. 

Distribution. Only known from the type 
locality and two other localities (see Fig. 2). 

Remarks. This species is clearly distinct 
from the MCZ series from the Omo River 
Valley in aspects of size, scalation characters 
(e.g., large crests, strongly mucronate scales 
and tufts with long spinose scales), and 
coloration, but it is similar in some aspects 
(e.g., strong nuchal crest, tufts of long 
spinose scales) to Agama spinosa Gray, 1831. 

Agama gracilimembris Chabanaud, 1918 

Syntypes. MNHN 04.114-115, male and 
female, from "Dahomey [= Benin]," col- 
lected by Eugene Megy in 1904. 

Diagnosis (partly fide Grandison, 1968). A 
dwarf Agama with females (SVL 48-57 mm) 
longer than males (SVL 40^17 mm). Body 
depressed, hind limbs long. No gular pouch 
in males or females, but a marked lateral fold 
is present. Tail one and a half times longer 
than SVL. In males, base of the tail with 
prominent hemipeneal bulge. Head convex. 
Upper head scales large, strongly keeled. 
Occipital scale enlarged, greatest width as 
large as the diameter of the ear opening. 
Nostril directed posteriorly, pierced in the 
posterior part of the nasal scale which is 
below the canthus rostralis. Usually with one 
scale between the nasal and the first supra- 
ciliary scale. Supraciliary scales smooth. 
Nuchal and dorsal crests absent. Ear opening 
as large as the eye, tympanum superficial and 
exposed. Tufts of long, spinose scales around 
the ear and the sides of the neck lacking. 

Single, short, conical scales close to the 
border of the ear opening are present. Body 
scales small, heterogeneous, keeled, and 
nonmucronate, in 70-85 scale rows around 
midbody. Enlarged body scales irregularly 
intermixed on the lateral regions of the body. 
Vertebral scales 30-46. Gular and ventral 
scales strongly keeled, ventrals about the 
same size as the dorsal scales. Hind limbs 
relatively long, reaching the tympanum. 
Lamellae 13-14 under fourth toe. Males 
with one row of 8-12 precloacal pores. 

Distribution. Ghana, Benin, Nigeria, Ca- 
meroon, Central African Republic (see 
Fig. 2 for details). 

Agama weidholzi Wettstein, 1932 

Lectotype. NHMW 18318, from "Urwald 
Sumpfgebiet, Tabadienke, 30 km S Dialla- 
koto, Senegambien [= Senegal]," collected by 
Weidholz in 1930. 

Diagnosis (partly fide Grandison, 1969). A 
dwarf Agama species With females (SVL 56- 
65 mm) longer than males (SVL 54-62 mm). 
Body depressed, hind limbs long, reaching the 
area between ear opening and eye. No gular 
pouch. Tail one and a half times longer than 
SVL. Head convex. Upper head scales large, 
keeled; occipital scale enlarged, as large as ear 
opening. Nostril directed dorsolateral^, 
pierced in the posterior part of a long, slightly 
oval, swollen nasal scale below the canthus 
rostralis. Usually with one scale between the 
nasal and the first supraciliary scale. Supra- 
ciliary scales smooth. Nuchal and dorsal crests 
absent. Ear opening as large as the eye, 
tympanum superficial and exposed. Tufts of 
spinose scales absent from sides of neck. 
Sometimes one or two conical scales behind 
the ear hole. Body scales small, homogeneous 
(but tendency for rows of enlarged vertebral 
scales), keeled and nonmucronate, in 68-82 
rows around midbody. Vertebral scales 32-^48, 
16-21 lamellae under fourth toe, and males 
with 6-10 precloacal pores usually in one row. 



Distribution. Senegal, The Gambia, Guin- 
ea-Bissau, and Mali (for details see Fig. 2; 
Wagner et al., 2009). 

Agama lucyae, new species 

Holotype. MCZ R- 146689, young adult 
male. Ethiopia: "Wollega" [probably Shewa 
or Kefa province; see remarks in Distribu- 
tion section] today either Oromiya or South- 
ern Nation, Nationalities and Peoples' Re- 
gion, Omo River, 58 km NW of Soddo 
[=Sodo], about 1,300 m above sea level 
(a.s.L). Coordinates: 7.27085°N, 37.37891 °E 
[subsequently georeferenced, see remarks in 
Distribution section]. Collected: 21. VII. 1964, 
by T. Monath. 

Paratopes. MCZ R-146687, adult female; 
MCZ R-146685, subadult female; MCZ R- 
146688, subadult male; MCZ R-146686, 
juvenile male. All with same collecting data 
as holotype. 

Diagnosis. A small Agama lizard (body 
length of males up to 53.5 mm SVL, females 
to 58.5 mm SVL), with a short depressed 
head, smooth head scalation, a large occip- 
ital scale, homogeneous body scalation, and 
strongly keeled but non- or only slightly 
mucronate dorsal scales. The nasal scale is 
on the canthus rostralis. Ear hole surrounded 
by single short conical scales; few tufts of 
spinose scales behind the ear and on the 
lateral side of the neck are present, tympa- 
num superficial. Short neck crest present in 
both sexes. Gular and ventral scales smooth. 
Tail not arranged in distinct whorls. 

Differential Diagnosis. A. lucyae sp. nov. is 
distinct from Acanthocercus, Xenagama, and 
Uromastyx in lacking distinct whorls or 
regular rings on the tail and from Acantho- 
cercus by its large occipital scale. It differs 
from Pseudotrapelus by its smaller size (58.5 
versus 88 mm SVL), its relatively larger body 
scales, the tufts and single conical scales 
around the ear and on the side of the neck 
versus the absence of these scales, and the 

relatively shorter hind limbs (38.5 versus 
77 mm). It is distinct from African Trapelus 
species by its large ear hole, the visible 
tympanum, and the tufts and single conical 
scales around the ear and on the side of the 

The new taxon is smaller than most other 
Agama species occurring in the Horn. It is 
distinct from Agama robecchii in its homo- 
geneous body scalation and larger dorsal 
body scales; it differs from Agama doriae, A. 
finchi, Agama lionotus, A. bottegi, and A. 
spinosa by possessing a smaller number of 
tufts of shorter spinose scales behind the ear 
and on the sides of the neck, and nonmu- 
cronate body scales; it further differs from A. 
spinosa and A. bottegi by a much smaller 
nuchal crest and from the latter in not 
possessing a body and tail crest. It is very 
clearly distinct from all these species in color 
pattern, which is characterized by a vertebral 
stripe and, in males, longitudinal stripes and 
transverse bands. 

Agama lucyae sp. nov. differs from other 
dwarf or medium-sized Agama lizards as 
follows: from the slightly longer A. rueppelli 
by its non- or only slightly mucronate dorsal 
body scales versus mucronate and spiny 
scales, by the development of scale tufts 
behind the ear opening and on the sides of 
the neck (few short tufts versus many long 
tufts), a higher number of scale rows around 
midbody (66-69 versus 54-64), a lower 
number of vertebral scales (47-52 versus 
59-66), and the presence of a nuchal crest; 
from A. persimilis by having an acuminate 
snout (versus a rounded), nuchal crest 
present, higher numbers of scale rows 
around midbody (66-69 versus 52-57) and 
vertebral scales (47-52 versus 26-31), few 
tufts of shorter spinose scales (versus tufts of 
long scales around the ear), and a distinct 
coloration (neither the dark patches on either 
side of the vertebral stripe nor the large 
brown patch on the anterior flanks of A. 



No. 527 

persimilis are obvious in A. lucyae sp. nov.); 
from A. hartmanni by its smaller size (53.5- 
58.5 versus 87-92 mm SVL), nuchal crest, 
tufts of spinose scales behind the ear opening 
and on the neck, and lower numbers of scale 
rows around midbody (66-69 versus 76-81) 
and vertebral scales (47-52 versus 61-68); 
from A. weidholzi by having the nasal scale 
on the canthus rostralis (versus below the 
canthus rostralis), the upper head scales 
smooth, tufts of spinose scales behind the 
ear opening and on the neck, a nuchal crest, 
a slightly overlapping range but higher mean 
number of vertebral scales (47-52 versus 32- 
48), a lower mean number but slightly 
overlapping range of scale rows around 
midbody (66-69 versus 68-82), and the 
remarkable lack of a white bordered black 
patch on the shoulders; and from A. graci- 
limembris by its homogeneous body scala- 
tion, nasal scale on the canthus rostralis 
(versus below the canthus rostralis), having 
the upper head, ventral and gular scales 
smooth, possessing tufts of spinose scales 
behind the ear opening and on the neck, neck 
crest present, and a lower number of scale 
rows around midbody (66-69 versus 70-85), 
but a higher number of vertebral scales (47- 
52 versus 30-46). 

Description of the Holotype. MCZ R- 
146689 (see Fig. 3), young adult male, tail 
broken 19 mm from the base. 

Measurements. SVL: 53.5 mm, HH: 
7.5 mm, HW: 12.2 mm, HL: 16.6 mm, length 
of left forelimb: 25.5 mm, length of left hind 
limb: 38.5 mm. 

Description. Head and body depressed. 
Nostril tubular, directed laterally and slight- 
ly posterodorsally, pierced in the posterior 
part of a large convex, smooth, pear-shaped 
nasal scale which is situated on the canthus 
rostralis. Nasal scale partly visible from 
above, directly in contact with the first 
canthal scale. The first two canthal scales 
not in contact with the eye. Scales on the 

head smooth, somewhat rugose, interorbital 
scales as large or larger than the supraorbital 
scales; imbrications of temporal scales not 
uniformly directed, partly ventrally, others 
posteriorly. Occipital large, its greatest width 
equal to the diameter of the tympanum, 
pierced by a visible pineal foramen in the 
center. 10 (left)— 8 (right) upper and 9 (left)-8 
(right) lower labial scales. Ear opening as 
large as the eye, surrounded at its border by 
several single short conical scales, one tuft 
above, and one tuft of short spinose scales 
behind the ear opening, three of these tufts 
on the dorsolateral parts of the neck. Spinose 
scales of the tufts short, often consisting of 
scales of the same size, sometimes one longer 
scale in the center. Gular fold present, but 
gular pouch absent. Short nuchal crest 
present, composed of nine short erect scales. 
Dorsal scales homogeneous, in 67 scale rows 
around midbody, in 52 dorsal scales along 
the vertebrate and 68 ventral scales along the 
belly between the anterior border of the 
shoulders and cloaca. One row of eight 
precloacal pores. Dorsal body scales keeled, 
with a keel extending along the entire scale, 
not or only uncommonly very slightly 
mucronate and erect. Gular scales smooth, 
ventral scales smooth, becoming feebly 
keeled toward the lateral side of the body. 
Hind limb and fourth toe long, reaching to 
the posterior part of the tympanum when 
adpressed. Lamellae 16 under the left fourth 
finger, 17 lamellae under the left fourth toe. 
Relative length of digits of left manus 4 > 3 
> 5 > 2 > 1 ; relative length of digits of left 
pes 4>3 = 5>2>1; Tail depressed at its 
base, broken 19 mm behind cloaca. Large 
hemipeneal pockets absent. Dorsal tail scales 
strongly keeled, slightly mucronate, some- 
what larger than the body scales, and not 
arranged in whorls. 

Coloration (after formalin fixation and 
ethanol preservation). Upper parts of the 
body and head brownish, a pale vertebral 




Figure 3. Holotype of Agama lucyae sp. nov. (MCZ R-146689). A, Dorsal view; B, ventral view; C, lateral view; 
D, ventral view of head and chest; E, dorsal view of head and chest; F, lateral view of head. 



No. 527 

stripe from just behind the occiput along the 
body and extending on the tail is visible. At 
the neck, the pale vertebral stripe is bordered 
by a dark band on each side. A small dark 
patch under both ear openings is visible. A 
faintly juvenile coloration is present, with 
four dark transverse bands between the 
limbs. On these bands, several pale, darkly 
bordered ocelli are visible. Throat whitish, 
with a dark reticulated pattern that extends 
to the entire belly and partly to the underside 
of the hind limbs. The pattern is somewhat 
darker at the base of the throat, forming a 
dark patch. 

Variation of Paratypes. Adult female. 
Gravid, containing a minimum of five eggs 
(see X-ray in Fig. 4), but resembles in most 
characters the described holotype (see 
Fig. 4). Length of left forelimb 29.2 mm; 
length of left hind limb 36.4 mm (for 
additional measurements see Table 2). Nasal 
also tubular, but directed more laterally and 
only slightly dorsally. Nasal scale on the 
canthus rostralis, and partly visible from 
above. Between ear opening and eye, a round 
slightly swollen patch consisting of several 
scales is present. Gular fold and a small gular 
pouch present. Neck crest more strongly 
developed than in holotype, also consists of 
nine scales, but longer. Single conical scales 
framing the ear opening longer, tufts of 
spinose scales behind the ear and on the side 
consist of more scales. Occipital scale large, 
but bordered by a broad elongated scale on 
each side. Dorsal scales partly mucronate. 
Tail short and thin, only about one third 
longer than the snout-vent length; base of 
the tail not notably broadened. Entire dorsal 
surface of body, tail, and limbs brownish 
with obscure irregularly placed lighter and 
darker shades. Belly and underside of the tail 
dirty whitish. Throat dirty whitish, becoming 
finely marbled posteriorly, uniform dark 
patch at its base present. Subadults. Both 
resemble the holotype but characters are not 

as distinct. Nasal scale slightly above the 
canthus rostralis, visible from above. In 
MCZ R- 146688, the same swollen patch as 
in the female is recognizable. Neck crests 
present in both specimens but weakly devel- 
oped, consisting of 10 and 11 erect scales. 
Coloration as in other juvenile Agama 
lizards; four dark crossbars between the 
limbs, becoming broad at the vertebral area, 
showing a pale, darkly framed, diamond- 
shaped patch at each crossband. In MCZ R- 
146685, a pale vertebral stripe is visible. 
Throat coloration resembles the holotype, 
extending to the anterior part of the belly. 
Juvenile. MCZ R-146686 resembles the 
holotype, but characters are only rudimen- 
tarily developed. Occipital scale large, cov- 
ering about two thirds of the back parts of 
the head. Coloration similar to the described 

Etymology. The entire Omo River basin is 
an important area both geologically and 
archaeologically and several hominid fossils 
have been excavated there. Reflecting the 
authors' interest in the evolution of early 
humans, this new species is named after 
"Lucy," the famous hominid fossil of Aus- 
tralopithecus afarensis Johanson, White, and 
Coppens, 1978, found in the Hadar area (the 
northern offshoot of the Rift System) of 

Ecology. Based on catalogue data, the 
entire MCZ type series was collected on the 
same day. The series contains two adults, 
two subadults and one juvenile specimen. 
Gartshore (1985) mentioned both A. weid- 
holzi and A. gracilimembris as annual species 
because she found gravid females between 
April and May, but rarely observed adults 
after June and none after the end of August; 
the first juveniles were found thereafter. So, 
adult and juvenile specimens were never 
collected at the same time. In Agama lucyae 
sp. nov., a gravid female was collected in July 
(gravid females of A. persimilis have been 




Figure 4. Adult female paratype of Agama lucyae sp. nov. (MCZ R-146687). A, Lateral view of head; B, dorsal 
view of head; C, ventral view of head; D, dorsal view; E, X-ray, ventral view; F, ventral view. 



No. 527 
















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Table 2. Measurements and pholidosis of the type series of A. lucyae sp. NOv. a 

MCZ R- 146689 

MCZ R- 146687 

MCZ R-146686 

MCZ R-146685 

MCZ R-146688 





































10, 8 

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9, 10 

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a See Material and Methods for abbreviations. For SL and IL the first numbers refer to the left and the second to 
the right side of the specimen. 

found in August; see Spawls et al., 2002), 
together with a relatively freshly hatched 
juvenile (MCZ R-146686) and two subadults 
(MCZ R-146685, 146688). Thus, A. lucyae 
sp. nov. should not be an annual lizard like 
the other two dwarf species (A. weidholzi, A. 
gracilimembris), as three different age phases 
were collected on the same day. 

Distribution. There is some confusion 
about the exact location of the type locality. 
The original collection label shows only 
"Wollega" [=Welega or Wellega] as the 
province, whereas the MCZ online register 
gives both Wollega and Shewa. However, the 
border of Wollega is approximately 200 km 
distant from the type locality (58 km NW of 
Soddo). If the correct locality is actually in 
the Omo Gorge, then it would lie within 
either Kefa [=Kaffa] Province (west bank) or 
Shewa [=Shoa or Showa] Province (east 
bank). These provincial divisions have, since 
1996, been replaced by ethnically based 
administrative regions (kililoch) with differ- 
ent boundaries. Today the west bank of the 
Omo Gorge lies within Oromiya [=Oromia, 
also called "Region 4"], whereas the east 
bank is in the Southern Nation, Nationali- 
ties, and Peoples' Region [also called "South- 
ern Peoples' Region"]. The coordinates given 

are not part of the original label and have an 
error of 29.205 km according to the MCZ 

Agama lucyae sp. nov. is currently only 
known from its type locality, which lies to 
the west of the Rift Valley. Two distribu- 
tional scenarios are possible, (a) A distribu- 
tion further west to Sudan and northwestern 
Kenya, more likely than to Somalia, and (b) 
a restricted distribution in the Central 
Ethiopian highlands. However, A. lucyae 
sp. nov. is the only dwarf species collected 
within the Ethiopian highlands (Fig. 2) and 
presumably is a rock- or rocky-ground- 
living species rather than a dweller on sandy 
surfaces. The Ethiopian Plateau is a volcanic 
system of 3,000-m-deep basalt rocks on 
underlying marine rocks, whereas the sur- 
rounding arid areas are sandy grassland, 
savannas, and deserts (Largen and Spawls, 
2010). Therefore, (b) is perhaps the more 
likely distribution scenario, although numer- 
ous lowland species are known to move 
upstream in the deeply dissected gorges, 
sometimes even extending onto the plateau 
lands (Stephen Spawls personal communica- 

The types of Agama lucyae sp. nov. were 
collected near the Omo River. This river 



No. 527 

arises in an alpine environment and passes 
through rainforest, open savannah and 
finally through desert and is the sole 
Ethiopian feeder of Lake Turkana (the Kerio 
and Turkwell rivers feed the lake within 
Kenya). The locality is in the central part of 
the river system, which based on the exam- 
ination of satellite images, is primarily rocky, 
semi-arid, open bushland lacking dense 
vegetation. However, the terrain in this area 
changes dramatically. The northwest side of 
the Omo Gorge drops down from more than 
3,000 m altitude to less than 1,000 m in only 
about 16 km, and the gorge itself has steep 
cliffs. Therefore, the specimens could have 
been collected in highland or lowland and in 
any of several different vegetation types 
because parts of the gorge are quite densely 
vegetated, although there are also extensive 
sand flats. 


Small species are not rare in Agama (e.g., 
there is a species radiation with about seven 
species in southern Africa), but most often 
small size is associated with solitary living, 
rather than with harem groups as seen in the 
large rock agamas (Branch, 1998; Spawls et 
al., 2002). However, solitary agamas are 
often found in lower densities and the 
collection of a series of all age classes could 
indicate a group living structure. 

A new agamid species from the Horn of 
Africa is not surprising. The area is both one 
of the African hotspots for arid-adapted 
reptiles and one of the most poorly studied 
areas of the continent (Largen and Spawls, 
2010). This is certainly true of agamid 
lizards. In most species, many aspects of 
ecology, habitat preferences, and behavior 
are unknown, and several (e.g., A. hartmanni, 
A. persimilis) are only known from a handful 
of voucher specimens from two or three 
localities. However, on the basis of known 

distributions (see Fig. 4), these species prefer 
lowland and sandy savannas or semideserts, 
whereas A. lucyae sp. nov. was found in the 
Central Ethiopian highlands. The highlands 
of the Rift Valley in Ethiopia seem to be a 
distribution barrier for ground-living agamas 
within the Horn. Agama persimilis is distrib- 
uted in eastern Ethiopia and Somalia (like A. 
robecchii, not shown), southward to Kenya, 
whereas A. hartmanni is known from low- 
lands of the western side of the Rift in 
Sudan, South Sudan, and Eritrea. Only A. 
rueppelli occurs on both sides. The Ethiopian 
Rift Valley and its southern offshoots could 
also be the contact zone between two 
important species radiations, the West Afri- 
can Agama agama group and the East 
African A. lionotus group, as represented, 
for example, by A. finchi, a member of the 
former, and A. lionotus, a member of the 
latter group, neither of which crosses the 
Valley (Bohme et al., 2005), but A. doriae 
(not closely related to either group) is known 
from the Omo Gorge near the A. lucyae sp. 
nov. collection locality (Stephen Spawls, 
personal communication). On the other 
hand, species of the agamid genus Acantho- 
cercus occur on both sides of the Rift but are 
distinct in habitat preferences to Agama. In 
contrast, A. lucyae sp. nov. was found in the 
center of an area not previously known to 
support ground-living Agama lizards. 

The relationships of A. lucyae sp. nov. 
remain unclear. On the basis of morphology 
and zoogeography, it could be closely related 
to A. hartmanni, but a relationship to A. 
gracilimembris, A. persimilis, or A. rueppelli 
is also feasible. On the basis of zoogeogra- 
phy, a membership in the A. agama or A. 
spinosa (including A. hartmanni, unpublished 
data) radiations is more likely than in the A. 
lionotus radiation, given its occurrence west 
of the Rift Valley (Wagner et al., 2011). 
However, the relations of these species are 
also currently poorly resolved. 





We are grateful to Jose Rosado and 
Jonathan Losos for access to the MCZ 
collection and for the loan of the specimens 
and to Wolfgang Bohme (ZFMK), Michael 
Barej and Frank Tillack (ZMB), Ivan Ineich 
(MNHN), Andreas Schmitz (MHNG), and 
Heinz Grillitsch (NHMW) for contribution 
of images or access to their respective 
collections. The manuscript was improved 
by insightful comments from Stephen Spawls 
and Wolfgang Bohme. We are grateful to the 
Museum of Comparative Zoology, Harvard 
University, for permission to publish photo- 
graphs of their specimens. All original 
photography of MCZ specimens is copyright 
©President and Fellows of Harvard College. 
This research was supported in part by the 
Lemole Endowed Chair in Integrative Biol- 
ogy Fund at Villanova University. 

Appendix 1. Material Examined 

Agama bottegi. Somalia. Lugh (MSNG 
28548, holotype); Karin, 10°58'26.4" N, 
49°12'49.2" E (CAS 227496). 

Agama gracilimembris. Benin. Without 
locality (MNHN 04.114-5; syntypes). Camer- 
oon. Benue NP (ZFMK 33717); Boki (ZFMK 
15257-259) Central African Republic. Koum- 
bala (ZFMK 33718-721); between Kotissako 
and Sakoumba (ZFMK 33722-725). 

Agama hartmanni. South Sudan. Bahr el 
Ghazal, Meschrael Rek (ZFMK 2587); Bahr 
el Zeraf (ZFMK 2586); Boma Hills (ZFMK 
27598); 20 mi N Faschoda [=Kodok] 
(BMNH 1901.7.31.1); Rejaf (MCZ R- 
29639). Sudan. Dongola (ZMB 4355 [3 ex., 
type series]); Singa (BMNH 1913.9.16.9). 

Agama lucyae sp. nov. Ethiopia. Shewa 
Province, Wollega, Omo River, 58 km NW 
of Soddo, about 1,300 m a.s.l., 7.27085°N, 
37.37891 °E (MCZ R146685-89). 

Agama persimilis. Ethiopia. Haud 
(45°50'E, 8° N, BMNH 1937.12.5.64 [new: 

1946.8.24.51], holotype). Kenya. East of 
Wajir at Wajir Bor (MHNG 2236.20). 

Agama rueppelli. Kenya. Kanapoi (MCZ R- 
128427); Kerio River Camp (MCZ R- 140858); 
Lake Turkana 2°18'N; 36°03'E (ZFMK 
30694-695); Lodwar (MCZ R-39013, 84292); 
Lokori (MCZ R-84286); Loperot Kalabata 
(MCZ R-97175); Marsabit, 60 mi N (MCZ R- 
84290); Mbunyi near Taveta (MCZ R- 18280); 
Voi (MCZ R-41003, 41005, 41008); Voi (MCZ 
R- 18281, syntype A. r. septentrionalis); South 
Turkana (MCZ R-97168-71). Somalia. Buran 
district (MCZ R-49120). South Sudan. Bahr- 
el-Gebel, Redjaf [=Rejaf] (ZFMK 2588-89); 
(?) Bender-Meraya (BMNH 1946.8.28.5, syn- 
type of A. r. rueppelli). 

Agama weidholzL Mali. Between Negala 
and Kassaro (ZFMK 20059); Fatao, 9 km N 
(ZFMK 20063-065); Kassaro, 8 km E 
(ZFMK 20060); Kita, 5 km E (ZFMK 
20061); Kita, 20 km E (ZFMK 20062). 
Senegal. Between Tiara und Mantiankani 
(ZFMK 20072); Casamance: 13 km SW 
Kolda (ZFMK 20073-078); Darsalam, 5 km 
SE (ZFMK 20066-068); Kounkane, 12 km 
W (ZFMK 20070-071); Medina Gounas, 
20 km S (ZFMK 20069); Tabadienka, 30 km 
S Diallakoto (NHMW 18318; 18319:1-2, 4- 
8; MCZ R-4431 1-312 type series). 

Appendix 2. Material Cited for 
Distributional Information Only 

Agama gracilimembris. Central African 
Republic. Ndelle (MNHN 17.191). Nigeria. 
Igbetti (ZMC R36654, R36701, R36263); 
Kano (BMNH 1961.2067-8, 1962.566); La- 
fia (BMNH 1938.3.1.47); Shendam (BMNH 
1962.1570); Wukari (BMNH 1938.3.1.48-9); 
Zaria (NHMW 18994); Zonkwa (BMNH 
1961.949, 1962.1569). 

Agama hartmanni. Eritrea. ("Urn Ager 
[=Omhajer], Missione Corni, Calciati," type 
locality of A. cornii). South Sudan. Taufikia 
(NHMW 17039). 


No. 527 

Agama persimilis. Ethiopia. Bohodle 
(BMNH 1937.12.5.68-73, 1946.8.27.55-60). 
Kenya. Kitui Dist.: 7.7 km S of Ngomeni, 
Tolotwa (CAS 161295); Mandera Dist: ca. 
1 mi. NE. Mandera Dist.: Elev. 1,200 ft. 
(CAS 130307). Somalia. Bud Bud (MF 5331- 
32); Dusa Mareb, 55 km N-NE of (MF 
5459); El Bur [=Ceel Buur] (MF 2431); 
Geriban (MF 3865-66); Har Addei (MF 
5456); Iesomma (MF 5189). 

Agama rueppellL Ethiopia. Dabanac 
(MSNG 28905); Gumboworen (ZMB 
27387); Sassabana (BMNH 
Kenya. Kajiado, 20.8 km N of (CAS 
198918); Kakuma (CAS 130877); Lake Bar- 
ingo, Kampi-Ya-Samaki (CAS 131599); Lo- 
kichoggio (CAS 131247); Lokitaung (CAS 
131281); Loyengalani (CAS 123106). Soma- 
lia. Boorama, Awdal Region (MVZ 241339); 
Buq Village, 27 km NE by Baargaal Rd. 
(MVZ 242741); Hargeysa Airport, 6 km E by 
Bander Wanaaq Rd. (CAS 225501). South 
Sudan. Lokomarinyang (CAS 131530). 

Appendix 3. Localities Obtained from 
Literature Used for Distributional Information 

Agama gracilimembris. Benin. Pendjari NP 
(Ullenbruch et al., 2010). Central African 
Republic. Ndelle, 8°25'N, 36'E (Grandison 
1968). Ghana. Wa (Spawls in Gartshore, 
1985). Nigeria. Ado-Awaiye, Anara, Fal- 
gore, Hung, Kura, Pandam, Ringum, Yan- 
kari (all Gartshore, 1985). 

Agama rueppellL Ethiopia. Confluence of 
the Web and Ganana [Ganale] (Largen and 
Spawls, 2006). 


Bauer, A. M., T. R. Jackman, E. Greenbaum, V. Giri, 
and A. De Silva. 2010. South Asia supports a 
major endemic radiation of Hemidactylus geckos. 
Molecular Phylogenetics and Evolution 57: 343-352. 

Bohme, W. 1975. Zur Herpetofaunistik Kameruns, mit 
Beschreibung eines neuen Scinciden. Bonner zoo/o- 
gische Be it rage 26: 2^18. 

, P. Wagner, P. Malonza, J. Kohler, and S. 

Lotters. 2005. A new species of the Agama agama 
group (Squamata: Agamidae) from western Kenya, 
East Africa, with comments on Agama lionotus 
Boulenger, 1896. Russian Journal of Herpetology 12: 

Boulenger, G. A. 1895. Rettili e Batraci, pp. 9-18. In G. 
Doria, and R. Gestro eds. Esplorazione del Giuba e 
dei suoi Afjluenti compiuta dal Cap. V. Bottego 
durante gli Anni 1892-93 sotto gli auspicii della 
Societa Geografica Italiana. Annali del Museo 
Civico di Storia Naturale Giacomo Doria, vol. 2, 
no. 15. Genoa, Museo civico di storia naturale. 

. 1898 [1897]. Concluding report on the late Capt. 

Bottego's collection of reptiles and batrachians 
from Somaliland and British East Africa. Annali 
del Museo Civico di Storia Naturale di Genova 
2(18): 715-723. 

Branch, W. R. 1998. Field Guide to Snakes and Other 
Reptiles of southern Africa, 3rd ed. Sanibel Island, 
Florida, Ralph Curtis Books. 

Brygoo, E. R. 1988. Les types d'agamides (reptiles, 
sauriens) du Museum national d'Histoire naturelle. 
Catalogue critique. Bulletin de la Museum nationale 
d'Histoire naturelle Paris, Section A. Zoologie, 
Biologie et Ecologie Animates 10(3)(Suppl.l): 1-56. 

Chabanaud, P. 1918. Etude complementaire de deux 
Agama de l'Afrique occidentale et description de 
quatre especes nouvelles de reptiles de la meme 
region. Bulletin de la Museum nationale d'Histoire 
naturelle Paris 24: 104-112. 

Gartshore, M. E. 1985. Agama gracilimembris in 
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