iVl iul s e tul iml oi L ounparatti we /^oology
US ISSN 0006-9698
4 November 2011
A NEW DWARF AG AM A (SAURIA: AGAMIDAE) FROM ETHIOPIA
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: email@example.com
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.
(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
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).
A NEW DWARF AG AM A FROM ETHIOPIA
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 AND METHODS
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
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
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 1922.214.171.124, 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
A NEW DWARF AG AM A FROM ETHIOPIA
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 19126.96.36.199); C,
Agama rueppelli (BMNH 19188.8.131.52, syntype); D, Agama per similis (BMNH 19184.108.40.206, holotype); E, Agama
gracilimembris (MNHN 1904.0115, syntype); F, Agama weidholzi (MCZ R-44311, paralectotype).
lucyae sp. n.
34 t7 7
5U.67 59 58
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
A NEW DWARF AG AM A FROM ETHIOPIA
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
19220.127.116.11 (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
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 
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 ). 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
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.
A NEW DWARF AG AM A FROM ETHIOPIA
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
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
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.
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
A NEW DWARF AG AM A FROM ETHIOPIA
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.
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
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
A NEW DWARF AG AM A FROM ETHIOPIA
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.
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A NEW DWARF AGAMA FROM ETHIOPIA
Table 2. Measurements and pholidosis of the type series of A. lucyae sp. NOv. a
MCZ R- 146689
MCZ R- 146687
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
B RE VI OR A
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
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.
A NEW DWARF AG AM A FROM ETHIOPIA
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 1918.104.22.168).
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 1922.214.171.124 [new:
19126.96.36.199], 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 19188.8.131.52, 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 19184.108.40.206); Shendam (BMNH
1962.1570); Wukari (BMNH 19220.127.116.11-9);
Zaria (NHMW 18994); Zonkwa (BMNH
Agama hartmanni. Eritrea. ("Urn Ager
[=Omhajer], Missione Corni, Calciati," type
locality of A. cornii). South Sudan. Taufikia
Agama persimilis. Ethiopia. Bohodle
(BMNH 1918.104.22.168-73, 1922.214.171.124-60).
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 126.96.36.199).
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
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 . 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
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
Nigeria. The Herpetological Journal 1: 23-25.
Grandison, A. G. C. 1968. Nigerian lizards of the
genus Agama (Sauria: Agamidae). Bulletin of the
British Museum of Natural History (Zoology) 17:
. 1969. Agama weidholzi of West Africa and
its relationship to A. gracilimembis. Bulletin de
l' Institute Fondamental d'Afrique Noire (Ser. A) 31:
Guibe, J. 1954. Catalogue des Types de Lezards du
Museum national d'Histoire naturelle. Bayeux,
Joger, U. 1990 . The herpetofauna of the Central
African Republic, with description of a new species
of Rhinotyphlops (Serpentes: Typhlopidae), pp.
85-102. In G. Peters, and R. Hutterer eds.
Vertebrates in the Tropics. Bonn, Museum A.
Lanza, B. 1978. On some new or interesting East
African amphibians and reptiles. Monitore Zoolo-
gico Italiano N.S. , Supplement 10: 229-297.
A NEW DWARF AG AM A FROM ETHIOPIA
. 1983. A list of the Somali amphibians and
reptiles. Monitor e Zoologico Italiano N.S., Supple-
ment o 18: 193-247.
Largen, M., and S. Spawls. 2006. Lizards of Ethiopia
(Reptilia Sauria): an annotated checklist, bibliog-
raphy, gazetteer and identification key. Tropical
Zoology 19: 21-109.
, and . 2010. The Amphibians and Reptiles
of Ethiopia and Eritrea. Frankfurt am Main,
Moody, S. M. 1980. Phylogenetic relationships and
historical biogeographical relationships of the
genera in the family Agamidae (Reptilia: Lacer-
tilia). Unpublished PhD Thesis. Ann Arbor, Uni-
versity of Michigan.
Parker, H. W. 1942. The lizards of British Somaliland.
Bulletin of the Museum of Comparative Zoology 91:
Peters, W. C. H. 1869. Eine Mittheilung uber neue
Gattungen und Arten von Eidechsen. Monatsber-
ichte der Koniglich Preussischen Akademie der
Wissenschaften Berlin 1869: 57-66.
Scortecci, G. 1929. Rettili dell' Eritrea esistenti nelle
Collezioni del Museo Civico de Milano. Atti delta
Societd Italiana di Scienze Naturali, e del Museo
Civico di Storia Naturale Milano 67(3-4): 290-339.
Spawls, S., K. M. Howell, R. Drewes, and J. Ashe.
2002. A Field Guide to the Reptiles of East Africa.
London and San Diego, Academic Press.
Ullenbruch, K., O. Grell, and W. Bohme. 2010.
Reptiles from southern Benin, West Africa, with the
description of a new Hemidactylus (Gekkonidae),
and a country-wide checklist. Bonn Zoological
Bulletin 57: 31-54.
Vaillant, M. L. 1882. Reptiles et Batraciens, pp. 1-10.
In G. Revoil ed. Mission dans le pays Comalis,
Faune et Flore. Faune et Flore des Pays Comalis,
Afrique Oriental. Paris, Challamel Aine Editeur.
Wagner, P. 2010. Diversity and distribution of African
lizards. Unpublished Ph.D. Thesis. Bonn, Ger-
many, University of Bonn.
. 2011. Book review: the amphibians and rep-
tiles of Ethiopia and Eritrea. Salamandra 47: 61-
, W. Freund, D. Modry, A. Schmitz, and W.
Bohme. 2011. Studies on African Agama IX. New
insights into Agama finchi Bohme et al., 2005
(Sauria: Agamidae), with the description of a
new subspecies. Bonn zoological Bulletin 58: 25-
, I. Ineich, A. Leache, T. M. Wilms, S. Trape,
W. Bohme, and A. Schmitz. 2009. Studies on
African Agama VI. Taxonomic status of the West
African Agama (Sauria: Agamidae) with prominent
tail crests: Agama boulengeri Lataste 1886, Agama
insularis Chabanaud, 1918 and Agama cristata
Mocquard, 1905. Bonner zoologische Beitrdge 56:
Wermuth, H. 1967. Liste der rezenten Amphibien und
Reptilien: Agamidae. Das Tierreich 86: 1-127.
Wettstein, O. 1932. Eine neue Eidechse aus Senegam-
bien. Zoologischer Anzeiger 99: 303-305.