Studies on African Agama VII. - A new species of the Agama agama-group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehelyi Tornier, 1902

2009·1.9 MB·

by  Wagner, Philipp

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Bonnerzoologische Beiträge Band 56 Flcft 4 Seiten 285-297 Bonn, November 2009
Studies on African Agama VIL
A new species of the Agama agama-group (Linnaeus, 1758)
&
(Sauria: Agamidae) from Cameroon Gabon,
with comments on Agama mehelyi Tornier, 1902
Philipp Wagner*', Michael F. Barej' & Andreas Schmitz-
' Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Gemiany.
- Muséum d'histoire naturelle, CP. 6434, CH-121 1 Geneva 6, Switzerland.
* Corresponding author: philipp.wagner.zfmk@,uni-bonn.de.
Abstract. In the course ofrecent taxonomic studies in theAfricanAgamidae the WestAfrican species were examined
andnew species havebeen identified. In this publication anew species ofthe genusAgama Daudin, 1802 is described
fromCameroon andGabon. Malesofthenew species differfrom all otherknownAgama in the uniquecombination of
the reticulate colouration ofthe throat and the blue tip oftail. It is comparedwith the other Cameroonian members of
the genus and several oihtvAgama species fromAfrica. Additionally, the status oíAgama mehelyiTornier, 1902, only
known from its holotype, is discussed.
Keywords. Reptilia: Sauria.Agamidae,Agama sp. n.,Agama mehelyi,Africa, Cameroon, Gabon, morphology, taxon-
omy.
Introduction
AfterresearchactivitiesonthegenusAgama Daudin, 1802 who restricted the eiToneous type locality 'America' of
in the last few years many species were described, syn- Agamaagama toCaiueroon but failedto fixthisdecision,
onymised or revalidated (Padial 2005, Wagner 2007, according to article 76 ofthe hiternatio-nal Code ofZo-
Wagneretal2008a,Wagneretal. 2008b). Currently,the ologicalNomenclature(ICZN 1999)bydesignatinga lec-
genuscontains 35 recognisedspecies,butpreliminaryre- to- orneotype. Now, aftertheproperde-finition ofaneo-
sults(especially fi^omtheWestAfricancladeofthegenus) type(Wagneretal. 2009b), ataxo-nomicrevisionofthis
show several cryptic taxa within theAgama agama (Lin- species group is possible and the alreadyrecognised new
naeus, 1758) coiTiplex. Therefore, many populations are species can be described.
under investigation and especially vouchers from
Cameroonwereofspecialinteresttotheauthors,because Chirio & LeBreton (2007) listed seven species ofAga-
the type locality ofAgama agama has been restricted to ma for Cameroon: Agama a. agama (Linnaeus, 1758);
this country. Agama doriae bemieensis Monard, 1951; Agamagrácil-
imembrisChabanaud, \9\'&\AgamamehelyiTornier, 1902;
However, until recently it was not possible to character- Agamaparagama Grandison, 1968; Agama sankaranica
izethe 'true'ZacertoagamaLinnaeus, 1758,becausethe Chabanaud, 1918 and Agama sylvanus MacDonald,
syntypes illustrated in Seba (1734) are not available and 1981. In the adjoining countries ofCameroon, no other
the proposed type material in the Museum Adolphi Fri- than the above mentionedAgama species occur (Wagn-
derici collectionoftheNaturhistoriskaRiksmuseet(=The er, unpubl. data) but Chirio & LeBreton (2007) also
SwedishMuseumofNaturalHistory) is notidenticalwith identified four probable new Agama species from
the illustratedspecimens. Wagner etal. (2009b)clarified Cameroon and one ofthem (Agama sp. 1) turned out to
this situation by designating a neotype (ZFMK 15222) beidenticalwiththenewspeciesrecognizedbyusbefore,
from Cameroon. The authors followed Mertens (1938) and we describe it herein.
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286 Philipp Wagner et al.: Studies onAfricanAgama VII. New species oftiieAgama agama-group
Material & Methods plify aportion ofthe mitochondrial 6S ribosomal RNA
1
gene. PCR cycling procedure was as described in
The type material ofthe new species is deposited in the Schmitz et al. (2005). PCRproducts were purified using
Zoologisches Forschungsmuseum A. Koenig, Bonn, Zo- Quiaquickpurificationkits(Quiagen). Sequenceswereob-
ologisches Museum für Naturkunde, Berlin and the tainedusinganautomatic sequencer(ABI 377). Sequences
Museum d'histoirc naturelle, Geneva. It was compared were aligned using ClustalX (Thompson et al. 1997; de-
withvouchersfromCameroonandotherAfricancountries fault parameters) and manually checked using the origi-
housed in those collection (see appendix) and with data nal Chromatograph data in the program BioEdit (Hall
from literature (Grandison 1968, Chirio & LeBreton 1999);thisresultedinatotal of505 bp forthechosen sec-
2007). tion ofthe 16S gene. Phylogenetic trees were calculated
in theprogram Paup* 4.0bl0(Swofford2002)usingthe
Measurements were taken with a digital calliper to the neighbor-joining algorithm (NJ) and bootstrapping with
nearest of0.1 mm. Measurements and scale counts were 20000pseudo-replicates to estimatenode supportaswell
done following Grandison (1968), Moody (1980) and aswithamaximumparsimonyapproach(heuristicsearch
Moody & Böhme (1984). with TBR branch swapping, stepwise random addition
with 100replicates,bootstrappingwith2000pseudo-repli-
Moleculardata were collected to examine sequence vari- cates). Paup* 4.0b10wasalso usedtocomputetheuncor-
ation between the new species and its morphologically rected pairwise distances for all sequences.
closestrelatives.Aportionofthemitochondrial 16S rRNA
gene ofthe holotype ofthe new species (ZFMK 87698; The following measurements and scale counts werepart-
GenBankaccession number: GU133316), aseriesofoth- ly used to compare the different species (see table 1):
er specimens ofthe new species (no voucher; GenBank Snout-vent length (SVL): measured from mental scaleto
accession number: GUI33315; ZFMK 73241; GenBank cloaca; tail length (TL): measured from cloaca to tip of
accession number: GUI33317; ZFMK 75376; GenBank tail; head length (HL): measured from jugale to rostral
accession number: GUI33318; ZFMK 83762; GenBank scale;headheight(HH): measuredatthejugal-postorbital
accession number: GUI33319; ZFMK 83766; GenBank region; headwidth (HW): measuredacrossthejugal-pos-
accession number: GUI33320), as well as the proposed torbital regionjust anterior to the external auditory mea-
neotype ofLacerta agama (ZFMK 15222; GenBank ac- tus; Midbody scales (MS): scale rows around midbody;
cession number: GUI33323) from northern Cameroon, vertebralscales (VS): numberofmidlinevertebral scales,
Agama .sankaraiiica (ZFMK 84992; GenBank accession counted from midpoint ofpectoral region to midpoint of
number: GUI33327) from an unknown locality, Agama pelvic region; dorsal scales (DS): numberofmidline lon-
paragama (ZFMK 15244; GenBank accession number: gitudinaldorsal scales, countedfrom midpointofpectoral
GUI33321)fromCameroon,Waza, LogoneetChari,Aga- region to cloaca; cloacal pores (CP).
ma agama africana (ZFMK 73845, ZFMK 73846; Gen-
Bank accession numbers: GUI33311, GUI33312) both
from Senegal, Dakar-Bel Air, Agama agama ssp. (MH- Results & Discussion
NG 2689.53; GenBank accession number: GUI33310)
from Benin, Agama hoiieti (ZFMK 80057) from Mauri- Species ofthe genusAgama, unlike chameleons, the sis-
tania, 30km NW of Rosso, and Agama fiiichi (ZFMK ter taxon ofthe Agamidae (e.g. Macey et al. 2000), are
83652; GenBank accession number: GUI33314) from very conservative in body form, without developing or-
Kenya, Malaba(type locality)were sequenced. Sequences naments like e.g. horns orear laps. Only in some species
for Agama planiceps (GenBank accession number: body ornamentations likeenlargednuchal ortailcrestare
AF355476) andAgama castroviejoi(GenBank accession present. Ifone takes into consideration the results from
number: AY522929) were added to the dataset from al- the chameleons (Ziegler & Böhme 1997; Böhme &
readypublishedsequences (Matthee & Flemming 2002, Ziegler 2008), where morphologically similar species
Padial 2005). Acanthocerciis atricoHis (ZFMK 41748; showhighvariations inhemipenisstructures,asimilarsit-
GenBank accession number: GUI33322 Botswana, uation intheAgamidaecouldbeexpected. Incontrast,pre-
Gaborone) was chosen as outgroup. liminai-yresults(Böhme 1988; Wagneretal.,unpubl. da-
ta) showahigh levelofhomoplasyandconservativemor-
DNA was extracted using QuiAmp tissue extraction kits phology in the hemipenis structures in the genusAgama.
(Quiagen) or a modified Chelex-Protocol (Walsh ct al. Consequently, beside from body fonn and hemipenis
1991, Schmitz 2003). The primers 16sar-L (light chain; structure, there must be another mechanism of species
5'-CGC CTG TTTATCAAAAACAT-3') and 16sbr- recognition. A pre-mating mechanism was identified by
H (heavy chain; 5' - CCG GTC TGA ACT CAG ATC several authors (Loveridge 1933, Thys van den Aude-
ACG T- 3') ofPalumbi et al. (1991) were used to am- NAERDE 1963, McLachlan 1981, Böhme et al. 2005,
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Bonnerzoologische Beiträge 56 287
Acanthocercus atricollisZFMK41748
Agama sankaranica ZFMK 84992
AgamaplanicepsAF355476
82
53
-Agama finchiZFMK 83652
Agama agama ssp. MHNG 2689,53
Agama agama africanaZFMK 73845
100
94
Agama agama africanaZFMK 73846
—
Agama agama agama ZFMK 15222 Neotype
-AgamaparagamaZFMK 15244
100
4<>]- Agama lebretonino voucher
100
100 Agama lebretoniZFMK 87698 Holotype
100
Agama lebretoniZFMK 73241
Agama lebretoniZFMK 75376
Agama lebretoniZFMK 83762
Agama lebretoniZFMK 83766
-Agama bouetiZFMK 80057
100
100
Agama castroviejoiAY522929
0.1
Fig. 1. Phylogenetictreebasedon 505 bpofthemitochondrial 16S rRNAgene. Valuesabovethenodesrepresentneighbour-joi-
ningbootstrap (20000 pseudo-replicates) values inpercentwhile the values blow the nodes are the correspondingmaximum par-
simony supports. Significantly supported values are in bold. Values below 50% are not shown. The branch length from the out-
group (Acanthocercus)tothe ingroup (Agama) has been shortened forvisual purposes.
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288 Philipp Wagneret al.: Studies onAfricanAgama VIL New species oftheAgama agama-group
—
Wagner 2007, Wagner et al. 2008a, 2008b) as the !^ ts<
colouration ofhead, throat, forelimbs and tail ofnup-
tial males. In conclusion, the colouration ofdominant »q »Q !>q »Q Oq Cí(3 CK3 »q »Q »q »5 CK; »5 CK3 I 85
malesand, sometimes, ofpregnantfemalescanbeused ^
fortheidentificationofthedifferentspecies inthegenus
Agama, but not for analysing phylogenetic relation- N N -2
ships. Oneofthemaincharacters isthecolourationpat- > N V ? •p T1 n
tern ofthe throat (Jacobson 1992; McLachlan 1981; 3 3 ?^ 5 S S
Wagner2007; Wagneretal. 2008a,2008b). Dominant N-n N-n N-n N-n N-0 N N2^
mhaavlieosurparlesceonnttetxhetirtothortohaetr,ahdeualdt maanldefocroemlpiemtbistoirnsaanbde,- 7oU^o) o7o^ ro-o oc ro 27^ 7^ - 00 7s
especially, females. Consequently, these characters are -OOJNN -OtJNo u~O<Jn t4Ji 00
useful to distinguish different species and to compare
the herein described new species with closely related
taxa. Therefore, we compare the new species not only
in pholidosis and morphometries with other Agama
species, chiefly with that ones occurring in Cameroon,
but we turn our attention especially to the colouration o — NO o NO NO NO NO O ^ O O U00) oo o nO
K) ro
ofdominant males.
o o o o o o — — o o
o o
-J — Ko) too
Genetics: Forthegenetic datasetbothphylogeneticcal- OO oo ttoo 0^0 lo o o
culation methods completely agree in the topology of pö pö oö oö oö o o oÖ o— o— oÖ oÖ
the recovered phylogenetic tree (flg. 1), and only dif- L0»0J '0.>0J OC' 00 00 00 — ^ — o 4^
fer in the degree ofsupport recovered for the individ- C—
ual nodes. to to o—c Öoo Öto>cj Öttoo ^o o 'o— Öo
On '.>J
O O O o o o o o o o o
Thephylogenetictree isratherwell resolvedwiththree 4t:OiOLO*OJUOJOtO>G.>OUOJOLOoUOJttOoO^
major groups apparent. The first and basal fully sup-
portedcladeincludes^gö/nabouetiandtheonlyrecent- o c —
lydescribed^. castroviejoi(Hh 100/MP: 100).Thepo- —
00
siintgioonfoofthtehrisecxltardeemecoWrersets-poonrdNsowretlhlAwfirtihcatnhetapxoasiitniotnh-e NpÖO o'—— o—oo —ooo o o o oo oo too
00
more complete phylogeny of Leaché et al. (2009), oooocoooo
therefore it is likely that these two taxa are part of a o
On to 00 00 oo 00 o
more comprehensive north-western clade within the 4i ji.
genus Agama. The second clade is only well support- O O O
ed by the neighbour-joining analysis and is represent-
ed in this tree by a single species (Agama sankarani- o o co o o
o o o
ca), andrepresents the "Sahel Radiation"-clade identi- oo oo oo
fied by Leaché et al. (2009).
o o o o o o
Thethirdand largest clade includes a basal unresolved
polytomybecausethephylogeneticpositionoíA.plan-
iceps andA.fiuchicould notbe resolvedwith the used
gene fragment. But here the three well supported sub-
o p p p
cladesareofimportance: thefirstsubcladeincludesthe Ö Ö Ö Ö
proposed neotype oíAgama a. agama and the closely
relatedA. paragama (NJ: 84/MP: 80), the second sub-
clade includes the two Senegalese vouchers ofAgama 00
NO to
a. africana as well as a specimen from Benin (NJ: o o
84/MP: 62), whilethethirdclade fully supportedclade Ö Ö
(NJ: 100/MP: 100) contains all the included vouchers
oi'^Agcmia sp. n.".
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Bonnerzoologische Beiträge 56 289
Fig. 2. ImagesofAgama¡ehretonisp.n.alive:A. PregnantfemaleoíAgamalehretonisp.n. fromMt.Nlonako,Cameroon(ZFMK
ISild) B. Non-pregnant female oíAgama lehretoni sp. n. from Nyasoso, Mt. Kupe, Cameroon C. Close-up ofthe head ofthe
livinghoIlotypeofAgama lehretonisp. n. from northeastofMamfe, Mukwecha,Amebisu, Cameroo| n. D. Holotype ofAgama le-
|
hretoni sp. n. in life
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290 Philipp Wagner et al.: Studies onAfricanAgama VII. New species oftheAgama agama-group
Fig. 3. Preserved holotype ofAgama lebretoni sp. n. (ZFMK 87698) from nordeast ofMamfe. Mukwecha, Amebisu in Came-
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Bonnerzoologische Beiträge 56 291
Table 1 summarizes the unconected p-distances for the ZFMK51686: Cameroon, Magba, leg. F. Schütte, Febru-
used gene fragment. The neotype ofAgcuna a. agama ary 1990.
shows only a comparatively low genetic difference ZFMK 75376: Cameroon, Mt. Nlonako, Nguengue, leg.
(2.96%) to its sister taxonA. paragama, while it differs H.-W. Herrmann & A. Schmitz, January 2000.
fromitswestern 'subspecies'/Ío-í777;<3a. africanaby4.41% ZMB 55709, 37061: Cameroon, Makum.
and fromAgama sp. n. by 3.81%. The latter two species MHNG 2713.29 (field number AMC-360): Mofako
are separatedbyanequal genetic distance(3.81%). These Balue, Rumpi Hills, Cameroon, leg. locals people, July
distances are ofsimilar magnitude than to other well es- 2009.
tablishedspecieswithinthislargeclade{A. a. agama [neo- MHNG 2713.30(fieldnumberAMC-248): Big Massaka,
type]-A.planiceps4.72%;A. a. agama [neotype]-.-i. //7;c/?/ Rumpi Hills, Cameroon, leg. M. Barej &A. Schmitz, Ju-
3.17%;A.finchi-A. planiceps 3.59%). Regarding the ge- ly 2009.
netic distances of vouchers which we morphologically MHNG 2713.31 (tleld number AMC-009): Foyer du
confirmedtobemembersofthe sametaxonwe findthem Marin, Douala, Cameroon, 01.07.2009, leg. M. Barej &
to be genetically identical to each other; this is true both A. Schmitz.
for the two Agama a. africana specimens as well as for MHNG2713.32 (fieldnumberAMC-178): BigMassaka,
all six vouchers ofin the third subclade (Agama sp. n.). Rumpi Hills, Cameroon, 09.07.2009. leg. by locals.
The identified genetic structure (low to non-existent in- Diagnosis. A fairly large species ofAgama (total length
tra-cladedistancesandmoderatelyhigherbutmoreorless ofadult male above 25 cm), which is characterized by a
equidistant inter-clade distances) is typical for rather reticulated pattern on the throat, a bright vertebral stripe
young radiations. Our genetical analyses therefore sup- and a deep blue tail tip in adult males. Males ofthe new
ports the idea of previous studies (e.g. Wagner 2007; species are distinguishable from all other/Ígí7»?rt-species
Wagner et al. 2008a, 2008b) that many ofthe previous- by the combination of throat and tail colouration. The
lyrecognisedspecies inthe genusAgama in reality com- throat colouration (fig. 4.1) is a reticular pattern ofred
priseofseveraltaxa,whichareoftenverydifficulttosep- lines, which is so faronlyknown fromAgamaparagama
arate on the basis ofexternal morphology alone, and the (fig. 4.2), Agama sylvanus (fig. 4.3) and an undescribed
same now holds tme forAgama agama sensu lato. species from Uganda (fig. 4.5). The tail colouration is
unique within the genus because ofthe blue tip ofa tri-
Theinclusionofmorphologicallywellestablishedspecies colouredtail (see fig. 2d). Females are similarto those of
{A.planiceps.A. finchi,A.paragama), which shownear- other species oftheAgama agama species complex, and
ly equal genetic distances to the neotype ofA. a. agama can only be safely determined by their genetics.
prove that the uncovered distance ofAgama sp. n. to all
othercloselyrelated congeners confirmsthe specific dis- From southernandeasternAfrican speciesA. lebretonisp.
tinctness ofAgama sp. 1 and together with morphologi- n. differs as follows:
cal evidence (see below) warrants the formal recognition
ofthis species. FromA.planiceps,A. turuensis^A. m\vanzae,A. kaimosae
andA. atra the new species differs in their unique throat
colouration, the blue colouration ofthe tip ofthe tail and
Agama lebretoni sp. n. in having a vertebral stripe. From A. knobeli the new
species differs additionally in not possessing a tail crest.
Holotype (figs. 2c-d, 3, 4.1) FromA. miicosoensis the new species differs in having a
redheadinsteadofayellow.A. lebretonisp. n. isverydif-
ZFMK 87698 (fieldnumberMM075): Cameroon, north- ferent in tail colouration (tri-coloured instead ofbanded
east of Mamfe, Mukwecha, Amebisu, N 05°53.866' E nan-ow white and blue) fromA. lionotus and A. kirkii.
009°33.495', leg. J. Wurstner & M. Barej, September,
2007. Agama species occurring in Cameroon and Gabon differ
fromy4. lebretoni sp. n. as follows:
Paratypes
ZFMK 87694 - 697, 87699 (paratopotypes): Cameroon, Agamaagama differs inhavingablackcolourationofthe
Mamfe region, Mukwecha, Amebisu, leg. J. Wurstner & taalisloddpififnesrtienadchaarbalcuteerosneofitnhtrhoeatnceowlosupreactiieosn.:BAogtahmsapeacgiae-s
M. Barej, September. 2007.
ZFMK 87700: Cameroon, Mamfe, leg. J. Wurstner& M. 4m.a4),hawshiacmhoirsediosrsilmeislsarsttroitahteedrettoicuunliaftoednnthir-eodattohfrotahte(nfeigw.
Barej, September, 2007.
ZFMK 61243: Cameroon, Komp, Mundemba, leg. C. species. A. agama is also lacking the typical pale verte-
bral stripe which is present in nuptial coloured males of
Wild, February 1989.
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292 Philipp Wagner et al.: Studies onAfricanAgama VII. New species oftheAgama agaw^o-group
Fig. 4. Throatpatterns ofsome WestAfrican adult males oíAgama species:
1=Agama lebretonisp. n. (holotype, ZFMK 87698); 2=Agamaparagama (ZFMK 15244); 3=Agamasylvanus (ZFMK40252);
4=Agama agama (neotype, ZFMK 15222); 5=Agama sp. n. (ZFMK 88809); 6=Agama a. africana (ZFMK 20125); 7=Agama
finchi(NMK L2716); 8=Agamaplaniceps (ZFMK 55062); 9=Agama liouotus lionotus(ZFMK 83624); 10=Agama lionotusel-
gonis (ZFMK 82065); 11=Agama lionotus dodomae (ZFMK 83706); 12=Agama kaimosae (NMK L2715); 13= Agama doriae
hemieensis(ZFMK29615); \4=Agamadoriaedoriae(ZFMK55545); \5=Agamasankaranica(ZFMK40468); \6=Agamagra-
cilimemhns (ZFMK 33719).
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Bonnerzoologische Beiträge 56 293
the new species. In contrast, Aganici againa has a slight- large occipital scale, which is as large as one and halfdi-
ly lowercount ofscale rows around midbody. According ameter ofthe tympanum.
to Grandison (1968) A. agama from Nigeria has 59 to Agama lebretoni sp. n. differs clearly from Agama syl-
77rows,whereas^, lebretonisp. n. has 73 to 80rows [23
vanus in having a vertebral stripe instead ofa complete-
specimens].
lybluebody inA. sylvanus,butboth species show a sim-
From Agama doriae beniieensis the new species differs ilarpattern ofthroat colouration in adult males (figs 4.1,
4.3).
inhavingadifferentthroatpattern in dominantmales: A.
doriaewithaprominentblackdotonthebaseofthethroat
(fig. 4.13, 4.14), instead ofa reticulated throat in A. le- Agama lebretoni sp. n. differs from the holotype ofAga-
bretonisp. n.Additionally, doriaebenueensis isoneof mamehelyi(forcomments seebelow) inhavingahomog-
the fewAgama species which have the nostril below the enous body scalation.
canthusrostralis insteadofonthecanthus like inthemost
Agama'^. Fromthethreeotherproposednew speciesmentionedby
Chirio& LeBreton (2007)thenewspeciesdiffers asfol-
Agamagracilimembrisismuchsmallerinsize(100to 120 lows: from Agama sp. 2 in the larger size; from Agama
mm
inA. gracilimembris in difference to more than 250 sp. 3 in having the nostril on the canthus rostralis; from
mminy4. lebretonisp. n.)anddoesnothavespinosescales Agama sp. 4 in having a vertebral stripe and completely
aroundtheearopeningasinthemostotherAgama species. blue forelimbs instead ofred and blue coloured ones in
Again, also this species differs in the colouration ofthe Agama sp. 4.
throat ofadult males: striated inA. gracilimembris (fig.
4.16) and reticulated in the new species. In morphology,
A. gracilimembris differs in possessing strongly keeled Comparison ofA. lebretonisp. n. with probably valid
head scales and it is also one ofthe few Agama species synonyms ofAgama agama
which have the nostril below the canthus rostralis. How-
ever, both species are similar in the count ofscale rows Agcmiacolonorum var. congica Peters, 1877 wasdescribed
aroundmidbody(70 to 85 in^. gracilimembris and 73 to from Chinchoxo, Cabinda, Angola. One syntype (ZMB
80 in the new species). 9196) resembles in colouration moreAgama agama. The
typical characteristics ofA. lebretoni sp. n. (reticulated
From Agama paragama the new species differs in pos- throat, white speckled body and blue tip oftail) are lack-
sessing a lowernuchal crest, in having a red instead ofa ing in this specimen. The other syntype (ZMB 67193) is
yellow-whitish head and in having a higher number of lackingcolourationbutshowsarelativelylargebodysca-
scale rows around midbody (Wagner, unpubl. data). Ad- lafion in difference toA. lebretoni sp. n.. It is most prob-
ditionally, adult males ofA. paragama show a black in- ablethatAgamacolonorum congica isavalidspeciesbut
stead of blue tail tip. Grandison (1968) described the further investigations are needed.
colouration ofthe throat ofA. paragama (similar to the
herein described new species) as 'a dark network on a Agamapicticauda Peters, 1877 was describedby a series
cream ground which takes the forni of isolated, round, ofsix syntypes fromAdaFoáh inGhana(Adaferin Mau-
cream spots'(fig. 4.2) andasdissimilartoAgama agama ritaniafide Loveridge 1957, which isobviouslyinerror),
which has a 'longitudinal arrangement ofdarker lines or fromAcera in Guinea (= Accra in Ghana?) and from an
blotches'. However, Grandison (1968) also mentioned, unknown localityin Cameroon. Denzeretal. (1997)des-
thatthetypicalthroatcolourationof^.paragama ispres- ignated the specimen ZMB 403 from Ada Foáh as lecto-
ent inboth sexes, which is not comparable withA. lebre- type.Thissubadultspecimenresembles incolourationand
toni sp. n.. A similar situation exists in Agama tiiriiensis morphologyAgama agama. Nevertheless, the paralecto-
and Agama lionotus elgonis which have a very similar typeseriesmistakenlyincluded anadultmaleoíA. lebre-
colouration ofthe throatto each other(see fig. 4) but are toni sp. n. (ZMB 8299) from Cameroon. Therefore, be-
recognized as disfinct taxa by Wagner et al. (2008a). cause ofthe chosen lectotype, this taxon must be recog-
nizedassynonymoíAgamaagama. Differencesbetween
Agama sankaranica is completely different in body A. agama and A. lebretoni sp. n. have already been dis-
colouration, butalso in otheraspects ofmoiphology: this cussed above.
species is much smaller in size and it is one ofthe few
Agama lizards which have the nostril below the canthus Description ofthe holotype (figs 1 c-d, 2, 4.1). Habitus
rostralis. Additionally, A. sankaranica has a fewer count stout, snout-ventlength(SVL) 140.0mm, tail length (TL)
ofscalerowsaroundmidbody(64-78 [69.6] insankaran- 199.6 mm, head length (HL) 36.0 mm, head with (HW)
ica versus 73 to 80 [76.2] in lebretoni sp. n.) and a very 26.0 mm, head height (HH) 15.8 mm.
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
294 Philipp Wagner et al.: Studies onAfricanAgama VII. New species oftheAgama agama-group
Fig. 5 Distribution ofAgama lelvetoni sp. n. Cameroon. 1= Makum (ZMB 55709, 37061); 2= Magba (ZFMK 51686. 54906-
907); 3= Metchum, Wuni(ZFMK 15194 - 15200); 4=Amebisu [=Amebcsu] [type locality]; (ZFMK 87694 - 699); 5= Mamfe
(ZFMK87694-699,87700);6=Nguengue, MtNlonako(ZFMK69017);7-9=Rumpi Hills: MofakoBalue,BigMassaka(MHNG
2713.29- 30, 2713.32); 10= Mundemba (ZFMK 61243); 11= Douala (MHNG 2713.31); 12= Limbe (Victoria) (ZFMK 18891 -
894). Equatorial Guinea. 13= Bioko Island (Fernando Poo), San Carlos (ZFMK 9353 - 359). Gabon 14= Ngouassa (IRSNB
15686- 687); 15= Fougamou (ZFMK 73239 -245).