Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 (Squamata: Agamidae)

2009·1.3 MB·

by  Wagner, Philipp

© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
Bonnerzoologische Beiträge Band 56 Heft4 Seiten 215-223 Bonn, November2009
Studies on African Agama V.
On the origin oíLacerta agama Linnaeus, 1758 (Squamata: Agamidae)
Philipp Wagner'*, Thomas M. Wilms-, Aaron Bauer^^ & Wolfgang Böhme'
' Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-531 13 Bonn, Germany
[email protected]; w.boehme(a;uni-bonn.de
-Zoologischer Garten Frankfurt, Bemhard-Grzimek-AUee 1, D-60316 Frankfurt
thomas.wilms@,stadt-frankñirt.de
3 Villanova University, 800 LancasterAvenue, Villanova, Pennsylvania 19085, US
*con"esponding author
Abstract. Herein we present our strategyto preserve the nomenclatural stability ofthe widespread and common
Afrotropical lizardAgamaagama(Linnaeus, 1758). Werecognizedseveralproposedsyntypes, belongingtoava-
riety ofspecies, i.e.,Agama agama. Agama atra and Tropidiinisplica. But we have shown that these specimens
do not belong to the type material. However, ifthey were included in the type material the selection ofthe e.g.
Americansyntype, Tropidiinisplica-atropidurineiguanid,aslectotypewouldresultintaxonomicchaos,asAga-
ma agama is an African species and includes in its synonymy the type species ofthe type genus ofthe family
Agamidae.The illustratedtypesin Seba(1734)arerecognizedbyusasavarietyofagamidlizardswithatype lo-
cality encompassing theNew andthe Old World and not simply 'America", as given by Linnaeus.
Morphologically,Agama agama ishighlyvariablemaking it impossibleto assignthe syntypethatagreeswiththe
currentconceptofAgamaagamatoageographicpopulation,particularlyastheonlycharacter,viz.thecolourpat-
tern,isnolongerdiscernibleinthisspecimen.ThisendangersthestabilityoftheknownsubspeciesofAgamaaga-
ma. We therefore designate a neotype forLacerta agama Linnaeus, 1758 herein.
Keywords.Nomenclature;taxonomy;neotype; Squamata;Agamidae;Agamaagama;Africanrainbowlizard;type
locality;Africa.
Introduction
ThegenusAgama Daudin, 1802 is one ofthe mostwide- species groups (Boulenger & Power 1921, Grandison
lydistributedlizardgenerawithinAfrica, althoughthere- 1956, 1968) and most workers have considered it point-
lationshipsbetweenthe species inthegenusarestillpoor- lesstodesignate lecto-orneotypesto stabilizethe noinen-
ly known. Agama is a highly taxonomically complex clatureofthetaxa.Asanexample,Grandison(1968)des-
genus and most ofthe taxa are very similar and highly ignated a lectotype ofAgama bemieensis Monard, 1951
variable inpholidosis andonly identifiableby thenuptial (today recognized as a subspecies of Agcmia doriae
colouration ofmales(McLachlan 1981,Jacobsen 1992, Boulenger, 1885; seeMoody&Böhme 1984),therebysta-
Wagner 2007). Therefore, it is often not possible to as- bilizing anotherAgama species because several individ-
signa single specimen with colouration lacking to apar- uals of the type series were identified as A. paragama
ticular species or subspecies. This situation causes prob- Grandison, 1968. However, thesituationhaschangedand
lems in the determination oftype material to assess the several authorsarenowworkingonabroadscalephyloge-
validity ofthe different taxa. Thus, in many species and nyoftheentiregenus(Wagner& Leache, owndata).For
subspecies validity is doubtful (e.g., Agama bocowti this analysis it is essential to clarifythe taxonomic status
Rochebrune, 1884;/íg^amaco//;//Scortecci, \92^;A.pic- ofAgamaagama(Linnaeus, 1758)becausemanytaxaare
ticaudaPeters, 1877;A. congicaPeters, 1877). Pastgener- assignedtothis species assynonymsorsubspecies. Even
ic reviews have only been attempted for some inrecentyearsseveral fonuersubspeciesofA. agamahave
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
216 PhilippWagneret al: Studies onAfricanAgama V. On the origin ofLacerta agama Linnaeus,
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
Bonnerzoologische Beiträge 56 217
14, LAGERT A caudatcretí longa, pedibus pentada- ainp!,ibin. LACERTA caiidä icrcú longa.,
¿tylis, dorfo antice denticulato, coUo capite- pcdibiispeinadaííyin dorjo an-
que pone aculeate ,
tice denticulato, collo capitajite
Salamandra americ. Lacertoe aemula altera. Seb.
thef.i.piyo. t.ioy.f.j. Salamandraamer. am- poneacnleatis.Amcen.Acad.i.p.zSS
phibia. Seb. thef.i. p.i6ß. t.ioy.f.1.,2. ^\ß. nut. j6\n. iz.
CAPUT ovatum, fquamis anteriora verfus imbri- Salamandra americana amphibia.
catum, pone juxta aures aculéis insequalibus Sel',iluf.t.cCñ). i. loy.f.1,2.
fpinofum, quod fmgulare inhac fpecie. Col- Salamandra amer. lacerta £imda
lum
altera. Sch. ¡luf.i.p.ijo. t.wj.fi.
Habitat la AMERICA.
Fig. 2. Descriptionof"Lacertaamphibia"inthe 'Amoenitates Fig. 3. "Lacertaamphibia"published inthe 'MuseumAdolphi
academicce'by Linnaeus (1749, Vol. 1, p. 288). Friderici'(Linnaeus 1754,p.44)withthefirstmentioninofthe
locality 'Habitat inAmerica'.
been recognized as species. Böhme et al. (2005) have el-
evated the Agama lionotus Boulenger, 1896 complex 'West-lndischeSalamander (=West Indian salamander),
{Agama lionotus lionotus Boulenger, 1896, Agama I. el- halfsalamander, halflizard. Thespecimens shown in fig-
gonis Lönnberg, 1922, Agama I. dodomae Loveridge, ures two and three ofthe same plate (herein fig. 1) were
1923,Agama I. ufipae Loveridge, 1932) leaving the tme also recognized as Salamandra atnericana but the figure
Agamaagamatoñill speciesrank,withthefollowingsev- captions were 'Wyfie van den voorgaanden Salamander"
eral nominal subspecies ofits own: Agama agama aga- (= the female ofthe above mentioned salamander) (fig-
ma Linnaeus, 1758; Agama a. africana Hallowell, 1844 ure two) and 'Andersoort van West-Indischen Salaman-
(type locality: Liberia); Agama a. boensis Monard, 1940 derals de voregf (= another species as the previous one
(type locality: Madina Boé, Guinea-Bissau) andAgama ofthe west Indian salamander) (figure three).
a. mucosoensis Hellmich, 1957 (type locality: Mucoso
near Dondo, Angola). Agama a. savattieri Rochebrune, All thesubsequentdescriptionsofthespeciesrefertothis
1884 [type localities: Casamance (Senegal?), Mélacorée plate and the text mentioned. The name 'Salamander
(Guinea?),Albréda(TheGambia),Bathurst(=Banjul,The atnericana" is mentioned later by many authors (see be-
Gambia] has been synonymized withA. agama africana low).
by Grandison 968
(1 ).
However, confusions among agamids, geckos and sala-
Even the status ofthese subspecies remains unclear and manderswerecommonduringthe 18"' andearly 19"icen-
is dependent on the definition oíA. agama. Therefore, it turies. As an example, a gecko, today known as Hemi-
isessentialtoconsulttheoriginaldescriptionbyLinnaeus dacty'lusplatyurus, was described in an agamid genus as
(1758), the type specimens and previous descriptions of Stellioplatyurusby Schneider(1792). Inthe same work
the species. The results ofthis search are herein present- Schneider (1792) also described Stellio fimhriatus from
ed. Madagascar,todayknownagainasamemberofthegeck-
onid genus Uroplatus. Also Trapeliis savignyi Audouin,
1827, described in an agamid genus but in a gecko sec-
1. Pre-Linnaeen (before 1758) names tion ofthe text is today known as synonym ofthe gecko
Stenodactylussthenodacty'lus(Lichtenstein, 1823) (Wag-
'Salamandra americana Seba, 1734' ner & Crochet 2009).
In his Thesaurus Seba (1734) published a plate (thes. I,
p. 170, t. 107) depicting three lizards (see fig. 1) which ^Lacerta amphibia Linnaeus, 1749'
he described as 'Salamandra Americana, posteriore
parteLacertum referens, amphibia; mas'.The Thesaurus In the 'Amoenitates academicce', Linnaeus' first impor-
is published in two separate editions, one in French and tantcontributionsascollectedthesesin sevenvolumeson
Latin, the other one in Dutch and Latin. We herein rec- systematics, he (Linnaeus 1749, Vol. 1, p. 288; see fig.
ognize the Dutch edition because this would be the text 2) described the lizard which was shown in Seba (1734)
that probably most accurately reflects Seba's own lan- as 'Lacertacauda teretilonga,pedibitspentadactylis. dor-
guage. Inthe Dutchpartoftheaccompanyingtexthe de- so anticedenticulato, collo capitequeponeaculeato' lat-
scribedthe species shown in figureoneoftheplate asthe er known as 'Lacerta amphibia' and synonymised the
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
218 Philipp Wagneretai.: Studies onAfricanAgama V. On the originoiLacerta agama Linnaeus, 1758
Andersson (1900) examined these specimens and iden-
Agsnia. 30. L. caiulH tereti longa, eolio' fupra cspiteque pofticc tified two as 'true Agama colonorum Daudin, 1802' and
acukaio.
ylitiie». itcud. 1. p. i88. the last as 'Uraniscodonplica Linnaeus, 1758'. Arecent
M«J'. Ad. Fr. T. p. 44. examinationbyusofthethreespecimens(MuseumAdol-
¿V¿ mitj'. I. t. 107, f. I. i. 3. NRM
tiahtiai, in Amciii-'a. phi Friderici: 107, 108, 112; see fig. 5 herein) re-
Citlur corporis palliJus. AodoMtn miitimeßriaturn.
vealed that they represent three different valid species:
Agama cf agama (LiNNAEUS, IISS),AgamaatraDaudin,
1802 and Tropidurusplica (Linnaeus, 1758). But two of
Fig. 4. Description ofLacerta agama in the nomenclaturally
binding 10"^ edition ofLinnaeus 1758 (p. 207). these specimens are without doubt not identical with the
individuals illustrated in Seba(1734).Asitisobvious,the
lizards shown in figures one and two clearly have seg-
mented tails, which are not only lacking inthethree mu-
seum vouchers, but also in the genus Agama in general.
name 'Salamaiuiraamericana'. InnaminghisspeciesLin- These segmented tails, which are arranged inwhorls, are
naeusrefen-edtotheplatepublishedbySeba (1734,plate typical for the species of the genera Acanthocercus
107; see fig. 1 herein)andobviouslyfollowedthe sugges- Fitzinger, 1843 and Landakia Gray, 1845 (both Agami-
tions ofSeba (1734), considering this species halfsala- dae) occuiringinAfrica,AsiaandEurope. Onlythespec-
manderandhalflizard. ButLinnaeus (1749)alsorefen-ed imen offigure three shows a tail typical forAgama, but
tothedifferencesbetweenthespecimensshown inthefig- also forthe SouthAmericangenus Tropidurus. Thesedif-
ures in Seba (1734) as he synonymised figure three as ferences were also recognized by Duméril & Bibron
Salamandra americana and figures one and two as ( 1837) who referred the specimens shown in figures one
Salamandraamericanaamphibia. Severalyears later, Lin- and two ofplate 107 in Seba (1734) to Stellio vulgaris
naeus(1754, p. 44)publishedhis "MiiseimjAdolphiFrid- Sonnini & Latreille, 1802 (todayknownasLaudakiastel-
erici\ Herein, he added a type locality as 'Habitat in lio vulgaris) and eiToneously synonymised Lacerta am-
America' and again he referred to two taxa (Salamandra phibia withAgama colonorum, as they only referred fig-
americana amphibia, fig. 1 & 2; Salamandra americana ure three to Agama colonorum.
lacerate aemiila, fig. 3).
NRM
However, it isobviousbycomparingthe specimens
withtheremainingimagein Seba(1734)thatitisnotiden-
2. Linnaeus (1758) tical with one ofthe specimens. The atra specimen has a
very different colouration, whereas theplica one has the
Lacerta agama Linnaeus, 1758 mouthclosed,butsimilarinscalationstothefiguredspec-
imen.Thecf agama specimen isnotidentical inheadsca-
In the 10"i and currently still binding edition ofhis guid- lation and proportion.
ing work, Linnaeus (1758, p. 207) again re-named "Lac-
erta atnphibia'to the currently validLacerta agama Lin- Consequently, theNRM specimens arenotidentical with
naeus, 1758 (see fig. 4). Incontrasttothedescriptionsbe- the lizards shown in Seba (1734) and are not type speci-
fore, he referred all the three images ofplate 107 in Se- mensofLacertaagama Linnaeus, 1758.As anomenclat-
ba (1734) to this monotypic speciesandmentionedagain ural stabilization ofthisname is indispensable forfurther
'America'' as type locality. In his description, Linnaeus research, a neotype is designatedby us (see below).
(1758) did not referto other images orvouchers ofa mu-
NRM
seum collection. Therefore, the individuals which are the However,even ife.g. themostsimilar voucher(the
basis ofthe illustrations in Seba (1734) are the syntypes, T.plica-specimen)totheremainingimagewerethename-
becatisethe 'International CodeofZoological Nomencla- bearingtypeofLacertaagama. itwouldrequiresuppres-
ture'(ICZN 1999) doesnotrecognize 'iconotypes'andno sionbecause itwoulddestabilisetwowellknownspecies,
other types are mentioned. However, Quensel, director two genera and two families.
ofthe Swedish Academy ofScience from 1799 to 1806,
alsomentionedthreespecimens(Quensel 1802, Quensel Thus Tropidurusplica(ascuixentlyknown)wouldbecome
in Andersson 1900) from the collection ofthe Museum Lacertaagama. Wo\Ne\er,Agamacolonorum,asynonym
Adolphi Friderici in Drottningholm and noted 'that they ofLacerta agama today known as Agama agama, is al-
are identical with Linnaeus'sLacertaamphibia'(Ander- so the type species of the genus Agama. Therefore,
sson 1900, p. 1 1). However, theuseof'identical'hereon- Tropidurusplica and closely related taxa like T. lumaria
lymeans identicaltospeciesnot identicaltype specimens. and T. umbra would become members ofthe genusAga-
ma.
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
Bonner zoologische Beiträge 56 219
Further, the genus Agama is the type genus ofthe lizard dition but completely lacking colouration. As previously
familyAgamidae. This wouldmean that these Old World noted, species ofthe genus Agama, and especially Aga-
lizards would lose their name to a New World genus, maagama,arecharacterizedbyahigh variabilityofscale
whichwouldcauseataxonomicchaosandisthereforeob- counts (BouLENGER & POWER 1921, Thys van den Au-
viously unacceptable. DENAERDE 1963, Grandison 1968, Moody & Böhme
1984, Böhme etal. 2005)andthereforenearlyall taxaare
Accordingtoarticle75.6oftheICZNthisapproachshould detennined by aspects of colouration of adult males
not be taken into consideration because it endangers the (McLachlan 1981, Jacobsen 1992, Wagner 2007).
stabihtyofthewellknownspecies T.plica,thegenusAga- Thus, it is not possible to assign this voucherto a specif-
ma and the familyAgamidae. ic Agama agama population or subspecies and even not
clearly to Agama agama itself
NRM
Remarks on the identity ofthe specimens
Remarks on the erroneous type locality 'America' &
As shown above, two ofthe specimens are not identical the nomen Agama
withAgama agama as they are identified as A. atra and
Tropidurusplica. Theremainingvoucher(fig. 5c)isatrue Today it is obvious that the type locality 'in America' is
Agama species, and probably Agama agama itself. It is erroneous and does not coiTespond with the actual distri-
obvious from figure 5c that the specimen is in good con- bution of the species, genus and family. Suiprinsingly,
Agama colonoriim Daudin, 1802, the tust available syn-
onym of Agama agama, was also described from
T'Ameriqueméridionale', 'Cuba'and 'Jamaique'. Daudin
(1802) suggestedthisname forLacerta agama Linnaeus,
1758 and mentioned several names (e.g. Lacerta agama;
Iguanacordylina Laurenti, 1768; 'Salamandraamericana
Seba, LacertusmajorSloane; GuanalizardBrown;
Blue lizard Edwards; L'agame Daubenton; L'agame ou
caméléon de Mexique Stedman) as synonyms. Addition-
ally, Daudin (1802)alsomentionedaspectsofthe linguis-
tic usage of the name 'agama' and mentioned that the
name is used for a variety ofdifferent lizard species in
SouthAmerica. However,the sourceofthename 'agama'
is from the Ewe language ofthe Kwa group, spoken in
Togo, Benin and Ghana (Baumann 1936, Westermann
1954). DuMÉRiL& BiBRON (1837) werethe firstwho rec-
ognized that the type locality is in errorand mentioned a
distribution ofthe species 'sur la cote de Guinée et au
Sénégal' (= at the coast ofGuinea and in Senegal).
However, the consideration ofthe type locality as 'in er-
ror',thesimplestsolution,doesnotsolvetheproblem.Ac-
cording to article 76 ofthe code, the 'type locality ofa
nominal species-grouptaxon is the geographical place of
capture, collection or observation of the name-bearing
type; ifthere are syntypes and no lectotypehasbeen des-
ignated, thetype localityencompassesthe localitiesofall
ofthem'. As the syntypes are not available, the type lo-
calitycanonlybeconcludedfromtheidentifiedlizardgen-
era shown on the plate in Seba (1734).
Mertens (1938)alsorecognizedtheproblemand restrict-
Fig. 5. FormerlysupposedsyntypesofLacertaagamaLinna- ed itto 'Cameroon', but without the designation ofa lec-
eus, 1758 in the Museum Adolphi Friderici (NRM 107, 108, to- or neotype. According to article 76 of the code,
112). A= Tropidurusplica (Linnaeus, 1758); B= Agama atra Mertens' (1938) restricfion was invalid and the type lo-
Daudin, 1802; C=Agama cf agama (Linnaeus, 1758). cality thus still not clarified.
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
220 Philipp Wagner et al.: Studies onAfricanAgama V. On the origin ofLacerta agama Linnaeus, 1758
Fig. 6b. Neotype ofLacerta agama Linnaeus, 1758 (ZFMK 15222): details ofhead scalation.
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
Bonnerzoologische Beiträge 56 221
Fig. 6c. Neotype ofLacerta agama Linnaeus, 1758 (ZFMK 15222): SEM images ofdorsal scales. Location ofthe scales is indi-
cated by an arrow in fig. 6a.
Description of Lacerta agama Linnaeus, 1758 1884 AgamaagamasavattieriRochebrune,(synonym of
A. a. africana fide Grandison 1968) Faune
Agama agama (Linnaeus, 1758) Sénégambie, Rept.: 89.
1896 Agama smithii Boulenger, (fide Largen & Spawls
1734 Salamandraamericana Seba,NaturaliumThesauri, 2006), Proc. zool. Soc. London 1896: 213.
thes. l,Tome 1, Pi. 107, fig. 1-3. 1940 Agama boensis Monard,Arq. Mus. Bocage, Lisboa
1749 Lacertaamphibia Linnaeus,Amoenitatesacademi- 11: 155. Type locality: Madina Boé, Guinea.
cs, vol. 1: 288. 1957 Agama agama miicosoensis Hellmich, Veröff Zo-
1754 Lacertaamphibia Linnaeus, MuseumAdolphi Fri- ol. Staatsamml. München5: 44. Typelocality: Don-
derici: 44. do, Angola.
1758 LacertaagamaLinnaeus, Syst.Nat.,Ed. 10, 1: 207.
1802 Agama colononim Daudin, Hist. Nat. gén. Pail. Neotype. ZFMK 15222: Mokolo, Margui-Wandala
Kept. 3: 358. Province, northern Cameroon, leg. W. Böhme & W.
1831 Agama occipitalis Gray, in Griffith,Animal King- Hartwig, 14.11.1974 (figs 6a-c).
dom Cuvier, 9 Synops. Spec: 56.
1844 Tropidolepisafricanus Hallowell. Proc.Acad. Nat. Diagnosis.Alargespeciesofthegenus. Thenuptial male
Sei. Philadelphia 1844: 171.Type locality: Liberia. oíAgama agama is characterized by a yellow head, ared
1877 Agama colonorum var. congica Peters, M. Ber. K. throatand a tri-coloured (yellow, red and black) tail. The
preuß. Akad. Wiss. Beriin. 1877: 612. subordinate males, females, and adolescents possess an
1877 Agamapicticauda Peters, M. Ber. K. preuß. Akad. olive-green head and an olive-green to brown body with
Wiss. Beriin, 1877: 612. yellow dots, ocelli and stripes.
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
222 Philipp Wagner et al.; Studies onAfricanAgama V. On tlie origin oíLacerta agama Linnaeus, 1758
Description oftheneotype ofLacertaagama Linnaeus, colouration is as differenttoA. a. agama asA.planiceps,
1758 A. lionotiis and manymore well diagnosed species ofthe
genus. Therefore it is very doubtfulthatmucosoensis isa
mm
Adult male, measuring 290.1 1 in length (snout-vent subspecies ofagama.
length 128.93 mm,tail 161.18 mm,head length33.49mm,
head width 23.61 mm, head height 14.07 mm). Acknowledgements. We are grateful to PierreAndre Crochet
for his advises using the International Code on Zoological
Nostril slightly above the canthus rostralis, pierced in the NomenclatureandtoSvenKullanderforusingtheimagesofthe
middle ofa large vaulted nasal scale, directed obliquely NRMspecimens. CorneliusdeHaanandArievanderPastrans-
posterodorsally. Between the nostrils is a longitudinal, lated the Dutch part ofthe text in Seba.
finely keeledshield, followedby finelykeeledto smooth,
regularly arrangedscales intheinterorbitalregionandon
References
the remaining upper side ofhead. Supraocular scales are
smooth. Parietal shield more or less pentagonal, parietal Andersson,L.G. (1900): CatalogueofLinneantype-specimens
organ slightly visible. Nine supralabial scales on the left, ofLinnaeus's Reptiliainthe Royal Museumin Stockliol. Bi-
eight on the right side, nine infralabial hangtill K. SvenskaVet.-Akad. Handlingar26: 1-29.
Baumann, H. (1936): Schöpfung und Urzeitdes Menschenim
scalesonboth sides. Thescalesoriginatingfromboth sides MythosderafrikanischenVölker DietrichReimer&Andrews
ofthe parietal midline have their imbrications anteriorly SteinerVerlag, Berlin, 435 pp.
directed; thekeels in the uppertemporal region havetheir Böhme, W., Wagner, R, Malonza, R, Lötters, S. & J. Köh-
mucronesattheanteriormargins. Ear-opening large, about ler(2005):AnewspeciesofthsAgamaagamagroup(Squa-
thesamesizeasthe eye; tympanumvisible; anteriormar- mata:Againidae)fromwesternKenya,EastAfrica,withcom-
ginbeingcomposedofspiny, mucronatescales. Fourspiny ments on Agama lionotus Boulenger, 1896. Russian Journal
tufts on the left side and five on the right, one before the BoHuelrepnegteorl,ogGy.A1.2:&83J.-9H0..Power(1921):ArevisionoftheSouth
earopening, one above, one to two below and one to two African agamas allied toAgama hispida andA. atra. Trans-
behind. Nuchal crest low, consisting of 13 weakly differ- actions Royal Society SouthAfrica 9: 229-287.
entiatedlanceolatescales. Guiarscalesflat,juxtaposedand Daudin. F. M. (1802): Histoire Naturelle, générale et parti-
becoming smaller towards the guiar fold. Dorsal scales culiéredesreptiles,ouvragefaisantsuite,aThistoirenaturelle,
keeled, becoming larger from neck and tail to the middle générale et particuliére composée par Leclerc de Buffon, et
otafilthaenabondgyc;diinn5w8easckallye bruotwsnoanreotuhnedlemsisddbiosdtiyn.ctSlcyaldiefsfeorn- Dliormuee.driHigilés,etoapi.arreMCN..atCSu..r&eSlolGne.niCBnoiim,bprlvooelnt.e{31d.8eF3s.7R)De:upftEairrlpteé,st.oPlVaoroilgs.i.e4.GLéinbérr.aElne-;
entiated whorls thatconsistofthree scalerings each, dor- cyclopédique Roret, Paris, 570 pp.
sally strongly, ventrallyweaklykeeledto smooth. Ventral Grandison, G. C. (1956): OnacollectionoflizardsfromWest
scales smooth, slightly to strongly imbricate at theirpos- Africa. Mémoires de V Institut francais d'Afrique noir 18:
terior margin; in 80 rows between throat and vent. One 224-245.
row of 10 preanal pores. Scales on forelimb with strong- Grandison. G. C. (1968): Nigerian lizardsofthegenusAgama
(Sauria: Agamidae). Bulletin ofthe British Museum ofNat-
ly keeled scales, on the upper aim twice as large as the
ural Histoty ofZoology 17: 67-90.
dorsals, becoming smallertowards the underside and the Jacobsen, N. H. G. (1992): The status oíAgama aculeata ár-
manus. 3"íand 4''^ finger longest, 4'hslightly longer, dig- mala Peters, 1854 (Reptilia: Agamidae). Journal ofthe Her-
ital length decreasing 2-5-1, subdigital lamellae keeled. petologicalAssociation ofAfrica41: 30-34.
Hindlimbsalsocoveredbydistinctlykeeledscales,onthe Linnaeus,C.(1749):Amoenitatesacademicseseudissertationes
uppertights twice as large as the dorsals, becoming larg- varisephysicas,medicsbotanicieantehacseorsimeditzenunc
er towards the lower tights and smaller towards the un- collectaeetauctaecum tabulisaeneis.Vol. 1, Holmiee, Lipsias,
563 pp, tab. I-XVII.
derside. Linnaeus,C. (1754):AdolphiFridericiRegisSvecorum.Gotho-
rum. Vandalorumquae &c.in quoAnimalia Rariora Imprimis
Colouration in life. Head and nape reddish, limbs and etExotica:Quadrupedia.Aves.Amphibia,Pisces,Insecta,Ver-
body blue, base oftail white, followed by red and then mes Describuntur et Detenninatur, Latine et Suetice cum
black. Iconibus.Typ. Regia. Holmiae, Stockholm, I-XXX, 1-96pp.
+ 7 pp.
Linnaeus,C. (1758): Systemanaturceperregnatrianaturae,se-
Remarlis.Therankofthetwo subspeciesboemisandmii- cundumclasses, ordines,genera, species, cumcharacteribus,
cosoensis is questionable. Grandison (1968) mentioned differentiis,synonymis.locis. Editodecima.TomusI.Lauren-
boensis as a large-scaled Agama species and therefore ti Salvi. Holmiae. Stockholm, 823 pp.
some vouchers from Guinea could be identical with that McLachlan,G. R.(1981):TaxonomyoíAgamaliispicia(Sauria:
taxon and in this consequence boensis should be elevat- Agamidae) in southernAfrica. CimbebasiaA5: 219-227.
edto speciesrank(Wagner& Ineichunpubl. data).Aga- Mertens, R. (1938): Heipetologische Ergebnisse einer Reise
nach Kamemn. Abhandlungen der senckenbergschen natur-
ma a. imicosoeiisis, with its striking blue and yellow forschenden Gesellschaft442: 1-52.
© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at
Bonnerzoologische Beiträge 56 223
Moody. S. & W. Böhme (1984): Merkmalsvariationen und ta- NaturiECuriosorumCoUegaXenocratesdictus; SocietatisRe-
xonomischeStellungvonAgamadoriaeBoulenger, 1885und giaeAnglicans,etInstituti Bononicnsis.sodalis.Tomus1,Jans-
Agama hemieensis Monard, 1951 (Reptilia: Agamidae) aus sonio-Waesbergios & J. Wetstenium & Gul. Smith, Amster-
dem Sudangürtel Afrikas. Bonner Zoologische Beiträge 35: dam, 33, 178 pp. Ill pis.
107-128. Thysvan denAudenaerde, D. F. E. (1963): LesAgamidaedu
QuENSEL, C. (1802): Catalogue ofthe collections ofthe Royal Congo: les espéces et leurdistribution géographique. Revue
Museum. Stockholm. zoologique etbotaniqueAfricain 68: 203-250.
Schneider.J. G. (1792):Amphibiorum PhysiologiaeSpecimen Wagner, P. (2007): Studies in African Agama I: On the taxo-
Alterum Historiam et Species Generis Stellionum seu nomic status of Agama lionotiis usamharae Barbour &
Geckonum Sistens. Traiecti ad Viadnim (Frankfurt / Oder), Loveridge, 1928(Squamata.Agamidae). Heipetozoa20(1/2):
C. L. F. Aitzi 2: 54 pp. 69-73.
Seba,a.(1734): LocupletissimiRemmNaairaliumThesauriAc- Wagner, P. & P.-A. Crochet(2009): The status ofthe nomina
curataDescriptio.etIconibusArtificiosissimisExpressio.per Trapelits savignvi Audouin, 1827 and Agama savigiiii
UniversamPhysicesHistoriam. Opus, cui, in hoc RerumGe- Dumeril & BiBRON, 1837 and the valid nomen ofthe Savi-
nere,NullumparExstitit. ExTotoTerrarumOrbeCollegit,Di- gny'sAgama (Sauria: Agamidae). Zootaxa 2209: 57-64.
gessit,Descripsit.etDepingendumCuravitAlbertusSeba,Et- Westermann, D. (1954): Wörterbuch der Ewe-Sprache. Aka-
zela Oostfrisius, Academia Caesarea Leopoldino Carolinae demie Verlag, Berlin, 795 pp.
ZOBODAT - www.zobodat.at
Zoologisch-Botanische Datenbank/Zoological-Botanical Database
Digitale Literatur/Digital Literature
Zeitschrift/Journal: Bonn zoological Bulletin - früher Bonner Zoologische Beiträge.
Jahr/Year: 2009
Band/Volume: 56
Autor(en)/Author(s): Wagner Philipp, Wilms Thomas M., Bauer Aaron M., Böhme
Wolfgang
Artikel/Article: Studies on African Agama V. On the origin of Lacerta agama Linnaeus,
1758 (Squamata: Agamidae) 215-223