Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854)

627
Sys ema ics o A ican ough-scaled liza ds, wi h
desc ip ion o wo new species om eas e n Angola
(Squama a: Lace idae: Ichno opis Pe e s, 1854)
We ne Con adie1,2,3, Chad Kea es3,4,5, Eli G eenbaum6, Ja ie Lobón-Ro i a7,8, K ys al A. Tolley9,
Max Beni o10, Ped o Vaz Pin o7,8,11, Reuben V. an B eda12, Luke Ve bu g 3,13,14
1 Po Elizabe h Museum (Baywo ld), Beach Road, Humewood, Gqebe ha, 6013, Sou h A ica
2 Depa men o Conse a ion Managemen , Na u al Resou ce Science and Managemen Clus e , Facul y o Science, Geo ge Campus, Nelson
Mandela Uni e si y, Geo ge, 6529, Sou h A ica
3 Na ional Geog aphic Oka ango Wilde ness P ojec , Wild Bi d T us , San on, 2196, Sou h A ica
4 Sou h A ican Ins i u e o Aqua ic Biodi e si y, Makhanda, 6140, Sou h A ica
5 F . Laude dale Resea ch and Educa ion Cen e , Uni e si y o Flo ida, Da ie, FL 33314, USA
6 Depa men o Biological Sciences, Uni e si y o Texas a El Paso, El Paso, TX 79968, USA
7 CIBIO, Cen o de In es igacao em Biodi e sidade e Recu sos Gene icos, InBIO Labo a o io Associado, Campus de Vai ao, Uni e sidade do
Po o, 4485-661 Vai ao, Po ugal
8 BIOPOLIS P og am in Genomics, Biodi e si y and Land Planning, CIBIO, Campus de Vai ao, 4485-661 Vai ao, Po ugal
9 Depa men o Zoology, Uni e si y o Johannesbu g, Auckland Pa k, Johannesbu g, 2006, Sou h A ica
10 Uni e sidad In e nacional Menéndez Pelayo, Calle de Isaac Pe al 23, 28040 Mad id, Spain
11 Fundação Kissama, Rua 60 Casa 560, La do Pa io a, Luanda, Angola
12 Uni o En i onmen al Sciences and Managemen , No h-Wes Uni e si y, Po che s oom, 2531, Sou h A ica
13 En i o-Insigh CC, Uni 8 Oppid aai O ice Pa k, P e o ia, 0050, Sou h A ica
14 Depa men o Zoology and En omology, Uni e si y o P e o ia, P e o ia, 0001, Sou h A ica
h ps://zoobank.o g/F811EE38-D26A-4C49-A863-D2800F54BA7B
Co esponding au ho : We ne Con adie ([email p o ec ed])
Academic edi o Uwe F i z | Recei ed 13 July 2025 | Accep ed 3 Oc obe 2025 | Published 21 No embe 2025
Ci a ion: Con adie W, Kea es C, G eenbaum E, Lobón-Ro i a J, Tolley KA, Beni o M, Vaz Pin o P, an B eda RV, Ve bu g L (2025) Sys e-
ma ics o A ican ough-scaled liza ds, wi h desc ip ion o wo new species om eas e n Angola (Squama a: Lace idae: Ichno opis Pe e s, 1854).
Ve eb a e Zoology 75 627–672. h ps://doi.o g/10.3897/ z.75.e167366
Abs ac
Ichno opis is a genus o medium-sized lace ids endemic o sub-Saha an A ica, cha ac e ised by ough head shields. The genus
cu en ly comp ises six nominal species dis ibu ed ac oss much o sou he n, cen al, and eas e n A ica. Some species a e appa en ly
ac i e a only ce ain imes o he yea , esul ing in limi ed specimen collec ions and se e ely hampe ing esea ch. This sca ci y o
ma e ial has his o ically made comp ehensi e sys ema ic e iews o he genus di icul and has led o he desc ip ion o nume ous
egional mo phological a ian s as dis inc species o subspecies. Ma e ial collec ed in ecen yea s has enabled us o p o ide a new
phylogene ic hypo hesis o Ichno opis using wo mi ochond ial genes (16S and ND4) and wo nuclea genes (c-mos and RAG-1).
Ou phylogene ic da ase includes 56 indi iduals ep esen ing i e o he six cu en ly ecognised species (excluding I. chapini).
Addi ionally, he b oad geog aphical sampling o he widesp ead I. capensis g oup has allowed us o explo e he axonomic s a us o
se e al species and subspecies wi hin he g oup. As a esul , we demons a e he monophyly o Ichno opis in ela ion o o he A i-
can lace ids and p esen he mos comp ehensi e phylogeny o he genus o da e. We also p o ide he i s phylogene ic placemen s
o I. anganicana and I. g andiceps, which allows us o alida e hei axonomic s a uses. Fu he mo e, we eco e ed a new c yp ic
species closely ela ed o I. g andiceps, and iden i ied se e al well-suppo ed clades wi hin he I. capensis g oup, all co obo a ed by
mul i-locus species delimi a ion analyses. One o hese clades is desc ibed he ein as a new species, while he emaining axa o in e -
Ve eb a e Zoology 75, 2025, 627–672 | DOI 10.3897/ z.75.e167366
Copy igh We ne Con adie e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
628
es a e discussed and highligh ed o u u e in es iga ion. Based on ou indings, we ecommend he ollowing axonomic e isions:
Ichno opis longipes and I. mac olepido a should emain synonyms o I. capensis; I. bi i a a pallida and I. capensis nig escens a e
ea ed as a junio synonyms o I. bi i a a; and I. o e lae i is conside ed a junio synonym o I. anganicana. Al hough we could no
de e mine he phylogene ic placemen o I. chapini due o he lack o gene ic ma e ial, i s head mo phology and scala ion suppo i s
eassignmen o he I. bi i a a g oup. Thus, i is e ained as a alid species pending he a ailabili y o new ma e ial o u he ax-
onomic ac ions. In conclusion, his s udy esol es se e al long-s anding axonomic issues wi hin one o A ica’s mos unde s udied
lace id gene a and lays a solid ounda ion o u u e esea ch on he genus Ichno opis.
Keywo ds
A ica, c yp ic species, ep iles, sub-Saha a, axonomy
In oduc ion
The amily Lace idae comp ises 388 ecognised spe-
cies and nume ous subspecies dis ibu ed ac oss A ica,
Eu ope, and Asia (Ue z e al. 2025). Al hough he main
species ho spo can be ound in he Palea c ic, pa icu-
la ly in a id zones such as no he n A ica and he A a-
bian Peninsula, subs an ial di e si y also occu s a he
sou h. Cen al and sou he n A ica—spanning much o
sub-Saha an A ica, om sou h o he Congo Ri e Ba-
sin o he sou he nmos ip o he con inen —ha bou s a
leas 65 ecognised species ac oss 11 gene a (Ue z e al.
2025). This accoun s o app oxima ely 17% o global
lace id di e si y, unde sco ing he egion’s signi icance
as a cen e o endemism and e olu iona y di e si ica ion
wi hin he amily. Despi e i s ichness, he lace id auna
o cen al and sou he n A ica emains compa a i ely un-
de s udied (Tolley e al. 2016), highligh ing a gap in ou
knowledge on he amily’s biogeog aphy and e olu ion-
a y his o y.
In ecen yea s, se e al phylogene ic s udies ha e
a emp ed o add ess his gap by ocussing on cen al
and sou he n A ican lace id gene a, including Adol us
(G eenbaum e al. 2011, 2018), Pedioplanis (Makokha e
al. 2007; Con adie e al. 2012; Childe s e al. 2021; Pa -
inha e al. 2021), Me oles (Edwa ds e al. 2012, 2013a),
Nuc as (Edwa ds e al. 2013b; B anch e al. 2019; Baue
e al. 2019, 2020, 2025; Bap is a e al. 2020), and He-
liobolus (Ma ques e al. 2022a). Some o hese and o he
s udies ha e also ocused on alpha axonomy (G een-
baum e al. 2011; Edwa ds e al. 2013a; Englende e al.
2013; Wagne e al. 2014). Collec i ely, hese s udies
ha e imp o ed ou knowledge o lace id sys ema ics
and di e si y, esul ing in he desc ip ion o mul iple new
species. Despi e his p og ess, de ailed phylogene ic and
phylogeog aphic s udies a e s ill lacking o ce ain gen-
e a, such as Holaspis, Ichno opis, La as ia, and T opi-
dosau a.
The A ican lace id genus Ichno opis Pe e s, 1854
comp ises se e al small o medium-sized, ough-scaled,
e es ial species ha inhabi mesic o xe ic sa annas in
cen al and sou he n A ica (B anch 1998; Spawls e al.
2018; Pie e sen e al. 2021; Beni o e al. 2025). In e es -
ingly, he genus includes se e al sympa ic species ha
a e p esumed o ha e an annual ep oduc i e s a egy,
in which hey b eed asynch onously, and his migh be
a s a egy o educe in e speci ic compe i ion (B oad-
ley 1967a, 1974, 1979; Jacobsen 1987). Howe e , ou
knowledge o his ecological phenomenon is hampe ed
by he lack o obus ecological s udies on his genus.
Knowledge o he genus is based la gely on he wo k
o Boulenge (1921) and subsequen species desc ip ions
by de Wi e and Lau en (1942), Lau en (1952), Ma x
(1956) and B oadley (1967b), as well as a ecen comp e-
hensi e syn hesis o he genus p o ided by an den Be g
(2017). Six species a e cu en ly ecognised as alid:
Ichno opis bi i a a Bocage, 1866; Ichno opis capen-
sis (Smi h, 1838); Ichno opis chapini Schmid , 1919;
Ichno opis g andiceps B oadley, 1967; Ichno opis mi-
c olepido a Ma x, 1956; and Ichno opis anganicana
Boulenge , 1917. Fu he mo e, he subspecies, I. bi i a a
pallida Lau en , 1964 is conside ed o be alid, while ad-
di ional species o subspecies (i.e., I. capensis nig escens
Lau en , 1952; I. mac olepido a Pe e s, 1864; I. longipes
Boulenge , 1902; I. o e lae i de Wi e & Lau en , 1942)
ha e been desc ibed bu a e cu en ly no conside ed
alid, o a e con o e sial (Ue z e al. 2025). Con usion
he e o e pe sis s in he li e a u e ega ding he numbe
o accep ed species, hei diagnos ic cha ac e is ics, and
synonyms ( an den Be g 2017). Thus, in he absence o
a s ong phylogene ic amewo k o Ichno opis ha can
be used o cla i y species bounda ies, and a de ailed mo -
phological e ision, his con usion is likely o con inue.
In ecen yea s, collec ions o Ichno opis ac oss i s
ange ha e imp o ed, pa icula ly due o a conce ed e -
o o ca y ou biodi e si y su eys in unde -sampled e-
gions such as Angola (Con adie e al. 2016, 2022a; Beni-
o e al. 2025), he Democ a ic Republic o he Congo
(DRC) (Kea es 2024) and Zambia (Pie e sen e al. 2017).
This has allowed o alida ion o he axonomic s a us o
some desc ibed species wi hin a phylogene ic amewo k
in he cu en s udy, as well as p o iding a mo e in o med
knowledge baseline ega ding he di e si y and e olu ion
o his g oup in cen al and sou he n A ica. Th ough his
p ocess, we aim o s abilise he axonomy o he g oup
and lay a ounda ion o u u e wo k.
Ve eb a e Zoology 75, 2025, 627–672 629
Ma e ial and Me hods
Sampling
O e he pas decade, mul iple new Ichno opis spec-
imens we e collec ed ac oss cen al and sou he n A i-
ca, especially om Angola, Democ a ic Republic o he
Congo (DRC), Mozambique, Sou h A ica and Zambia
( Table 1). Ini ial species iden i ica ions we e based on
key diagnos ic ea u es (e.g., do sola e al colou a ion,
head scala ion, sup aocula –sup acilia -p e on al con-
ac s) epo ed in he li e a u e (Boulenge 1921; Ma x
1956; B oadley 1967b; an Be g 2017), supplemen ed
by examina ion o ype and opo ypic o nea - opo ypic
ma e ial (see Mo phology below), and by conside ing
geog aphic p oximi y o he espec i e ype locali ies.
DNA samples we e collec ed om ei he li e , muscle
o ail ips and p ese ed in 99% e hanol a e which
ouche specimens we e ixed in 10% o malin and ans-
e ed o 70% e hanol o long- e m s o age a he Po
Elizabe h Museum (PEM, Sou h A ica), he Colecção
He pe ológica do Lubango (CHL, Angola), he Na ion-
al Museum Namibia (NMNW, Namibia), he Museu
de His ó ia Na u al e da Ciência–Uni e sidade do Po -
o (MHNC-UP, Po ugal), he Museo delle Scienze di
T en o (MUSE, I aly), and he Fundação Kissama Col-
lec ion (FKH, Angola). Rep esen a i e ma e ial was also
deposi ed wi h he Minis y o En i onmen , Luanda,
Angola (MINAMB), and he Museu de His ó ia Na u al
de Mapu o, Mozambique (MHNM). Fo all newly col-
lec ed specimens, geog aphic coo dina es we e eco ded
in decimal deg ees (WGS84 da um, ou decimal places)
using a handheld GPS, and ele a ion in me e s abo e sea
le el (a.s.l.).
DNA ex ac ion, ampli ica ion and
sequencing
DNA was isola ed om issue samples using a s anda d
sal ex ac ion me hod (Aljanabi and Ma inez 1997).
S anda d Polyme ase Chain Reac ion (PCR) p ocedu es
we e u ilised o ampli y one pa ial mi ochond ial ibo-
somal gene (16S RNA [16S]), one pa ial mi ochond ial
gene (NADH-dehyd ogenase subuni 4 [ND4]), and wo
pa ial nuclea genes (oocy e ma u a ion ac o [c-mos],
ecombina ion ac i a ing gene 1 [RAG-1]). Each ampli-
ica ion was conduc ed wi h a PCR mix u e o 25 µl o-
al olume, con aining 12.5 µL Taq DNA Polyme ase 2x
Mas e Mix (Ampliqon; 3 mM MgCl2, 0.4 mM dNTPs
and Ampliqon Taq DNA polyme ase), 2 µl o wa d p im-
e (10 µM), 2 µl e e se p ime (10 µM), and 8.5 µl o
ex ac ed genomic DNA (20–50 ng/µl) and wa e com-
bined. The cycling p o ile o all he genes was as ol-
lows: Ini ial dena u a ion s ep a 94°C o 5 min, ollowed
by 30–37 cycles o 94°C o 30 s, 42–58°C o 45 s, and
72°C o 45 s, wi h a inal ex ension a 72°C o 8 min.
The cycling p o ile o he genes di e ed only in he an-
nealing empe a u e and he numbe o cycles (Table S1).
The p epa ed PCR p oduc s we e sen o Mac ogen Co -
po a ion in Ams e dam, The Ne he lands, o pu i ica ion
and sequencing wi h o wa d p ime s.
Phylogene ic analyses
Fo he phylogene ic analyses, 46 Ichno opis indi idu-
als we e sequenced, supplemen ed wi h sequences om
10 indi iduals a ailable om GenBank. Six Me oles
squamulosus indi iduals we e used as ou g oup axa
( Table 1). Fo each gene, new sequences we e checked
and edi ed using BioEdi Sequence Alignmen Edi o
.7.2.5 (Hall 1999), and hen aligned wi h he GenBank
sequences in MEGA .7.0.27 (Tamu a e al. 2013), using
he Clus alW .1.6 alignmen algo i hm wi h de aul se -
ings o alignmen pa ame e s (Thompson e al. 1994).
Single gene maximum likelihood (ML) ees we e c e-
a ed in IQ-TREE .2.1.3 (Nguyen e al. 2015) o isually
check he placemen o sequences and compa e opolo-
gies. DAMBE .7.3.1 (Xia 2018) was used o es o sa -
u a ion using he indi idual as well as combined i s and
second codon posi ions o each p o ein-coding gene. As
none o he genes we e ound o be sa u a ed, he genes
we e no pa i ioned by codons. In addi ion, cong uence
be ween indi idual gene da ase s, as well as be ween mi-
ochond ial and nuclea da ase s, we e es ed using 100
eplica es o he pa i ion-homogenei y es (PHT) (Fa is
e al. 1994, 1995) in PAUP* .4.0a169 (Swo o d 2003).
All gene- ee combina ions we e cong uen , allowing
o he c ea ion o a conca ena ed da ase o 2128 base
pai s o u he phylogene ic analyses, wi h he indi id-
ual gene alignmen s joined using SequenceMa ix .1.8.2
(Vaidya e al. 2011).
The op imal pa i ion scheme and bes - i ing models
o molecula e olu ion we e selec ed using ModelFinde
implemen ed in IQ-TREE (Che nomo e al. 2016; Minh
e al. 2021). The ollowing se ings we e used: -p pa i ion
ile (each pa i ion has i s own e olu ion a e), a g eedy
s a egy and he F eeRa e he e ogenei y model exclud-
ed (only in a iable si es and Gamma a e he e ogenei y
conside ed) (Che nomo e al. 2016; Kalyaanamoo hy e
al. 2017). As M Bayes is no able o implemen all he
IQ-TREE models, he -mse m bayes command was used
o ensu e ha only models ha we e compa ible wi h
M Bayes we e selec ed. The bes - i ing model schemes
selec ed o each da ase we e as ollows: 16S: GTR+G,
ND4: GTR+I+G, c-mos+RAG-1: HKY+G.
Maximum likelihood phylogenies we e gene a ed in
IQ-TREE, using a andom s a ing ee and he bes - i -
ing model schemes selec ed o each da ase (as selec ed
abo e). The ul a as boo s ap app oxima ion (UFBoo )
me hod (Hoang e al. 2018) was implemen ed using 5000
eplica es and a minimum co ela ion coe icien o 0.99.
To ensu e accu acy, he analysis was un wice o con-
i m ha independen ML sea ches eco e ed he same
opologies.
Bayesian in e ence (BI) was un using M Bayes
.3.2.7a (Ronquis e al. 2012) on he CIPRES Science
Ga eway XSEDE (h p://www.phylo.o g; Mille e al.
2010) using he gene-pa i ioned scheme and model se-
lec ion iden i ied by ModelFinde implemen ed in IQ-
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
630
Table 1. Ma e ial used o he phylogene ic analyses, including sample numbe , museum ca alogue numbe , coun y, locali y, geog aphic coo dina es, and GenBank/ENA accession numbe s. Abb e ia-
ions: Aa on M. Baue ield numbe s (AMB), B i ish Museum o Na u al His o y (BMNH), Cali o nia Academy o Sciences (CAS), Colecção He pe ológica do Lubango (CHL), Chad Kea es ield se ies
(CKD), En i o-Insigh (EI), Fundação Kissama Collec ion (FKH). K ys al Tolley ield numbe s (KTH, RSP, WP), Ma ius Bu ge ield numbe s (MBUR), Museu de His ó ia Na u al e da Ciência - Uni-
e sidade do Po o (MHNCUP), Ninda Bap is a ield numbe s (NB), Museu de His ó ia Na u al de Mapu o (MHNM), Na ional Museum Namibia (NMNW), Museo delle Scienze di T en o (MUSE),
Ped o Vaz Pin o ield numbe s (P, L se ies), Po Elizabe h Museum (PEM), Reuben V. an B eda ield numbe s (RE), Sebas ian Ki cho ield numbe s (SK), S ua V. Nielsen ield numbe s (SVN),
Thomas B anch ield numbe s (TB), We ne Con adie ield numbe s (WC, ANG), William R. B anch ield numbe s (WRB). Missing da a o una ailable in o ma ion is indica ed as NA.
Sample No. Museum ca alogue No. Species Coun y Locali y La i ude Longi ude 16S ND4 RAG-1 c-mos
WC-4515 PEM R23530 Ichno opis b. bi i a a Angola Wes o Cui o own on Aludungo oad −12.3278 16.9067 PV357721 OZ347967 PV412867
NB0675 CHL0675 Ichno opis b. bi i a a Angola Luando In eg al Na u e Rese e −10.2772 16.9533 PV357719 PV412839 PV412865 PV390641
P1-318 FKH-0833 Ichno opis b. bi i a a Angola Cambau −10.1048 15.2182 PV357720 PV412840 PV412866 PV390642
KTH09-075 PEM R17934 Ichno opis b. pallida Angola 7 km Eas o Humpa a −14.9820 13.4352 HF547775 HF547731 HF547694
ABC2 NA Ichno opis capensis Namibia Ka ima Mulilo −17.5066 24.2688 JX962898 JX963023 JX962916
AMB-6001 NMNW Ichno opis capensis Namibia Road o Tsumkwe −19.4600 19.7200 DQ871148 HF547732 DQ871206
AMB-6067 CAS 209602 Ichno opis capensis Sou h A ica Kosi Bay, KwaZulu-Na al −26.9400 32.8200 DQ871149 HF547733 DQ871207
ANG-311 PEM R20495 Ichno opis capensis Angola 8.5 km No h o Ri o −16.6232 19.0535 PV357725 PV412844 OZ347936
BMNH 2019. 2745 BMNH 2019. 2745 Ichno opis capensis Zambia Sioma Ngwezi Na ional Pa k −16.8987 23.5985 MK464418
BMNH 2019. 2746 BMNH 2019. 2746 Ichno opis capensis Zambia Cha uma Fa m −13.0701 22.9288 MK464417
BMNH 2019. 2747 BMNH 2019. 2747 Ichno opis capensis Zambia Lukwakwa −12.6608 24.4370 MK464416
BMNH 2019. 2750 BMNH 2019. 2750 Ichno opis capensis Zambia Nanzila Plains, Ka ue Na ional Pa k −16.2814 25.9168 MK464415
BOX23-C02 PEM R24748 Ichno opis capensis Sou h A ica Tembe Elephan Pa k, KwaZulu-Na al −27.0217 32.4583 OZ347416 OZ347968
EI-0437 NA Ichno opis capensis Sou h A ica Lepalale, Limpopo −23.6391 27.5981 OZ347969 OZ347937 OZ347957
EI-0443 PEM R25370 Ichno opis capensis Sou h A ica Lepalale, Limpopo −23.6391 27.5981 OZ347970 OZ347958
EI-0444 PEM R25371 Ichno opis capensis Sou h A ica Lepalale, Limpopo −23.6391 27.5981 OZ347417 OZ347971 OZ347938 OZ347959
L-18 PEM R22069 Ichno opis capensis Angola Gambos, Fos e ‘s a m −15.8500 14.6833 PV357726 PV412845 OZ347939
MOZ14-356 NHNM Ichno opis capensis Mozambique Chiza ane, Zona B aza Lodge −25.0137 34.0376 OZ347418 OZ347972 OZ347940
MOZ14-357 PEM R21112 Ichno opis capensis Mozambique Chiza ane, Zona B aza Lodge −25.0137 34.0376 OZ347419 OZ347973 OZ347941
MOZ14-358 PEM R21113 Ichno opis capensis Mozambique Chiza ane, Zona B aza Lodge −25.0137 34.0376 OZ347420 OZ347974 OZ347942 OZ347960
NB0771 CHL0771 Ichno opis capensis Angola Bicua Na ional Pa k −15.2435 14.8915 PV357727 PV412846 PV412870 PV390644
NB0772 CHL0772 Ichno opis capensis Angola Bicua Na ional Pa k −15.2435 14.8915 PV357728 PV412847 PV412871 PV390644
NB0779 CHL0779 Ichno opis capensis Angola Bicua Na ional Pa k −15.1049 14.8403 PV357729 PV412848 PV412872 PV390644
NB1116 CHL1116 Ichno opis capensis Angola Cusseque −13.6851 17.0795 PV357730 PV412849 OZ347943 PV390647
NB1123 CHL1123 Ichno opis capensis Angola Cusseque −13.6782 17.0832 PV357732 PV412851 PV390649
NB1124 CHL1124 Ichno opis capensis Angola Cusseque −13.6782 17.0832 PV357733 PV412852
NB1138 CHL1138 Ichno opis capensis Angola Cusseque -13.6782 17.0832 PV357734
PEM R15556 PEM R15556 Ichno opis capensis Mozambique 7 km No h o Chibu o −24.6231 33.5661 OZ347975
RE211206B1 NMNW R11561 Ichno opis capensis Namibia Khaudum −18.2876 20.9897 OZ347421 OZ347976 OZ347944
RE211206B3 NMNW R11562 Ichno opis capensis Namibia Khaudum −18.2876 20.9897 OZ347977 OZ347945
SK13128 NA Ichno opis capensis Namibia Naye-Naye −19.9235 20.6976 MN015330 MN030223
Ve eb a e Zoology 75, 2025, 627–672 631
Sample No. Museum ca alogue No. Species Coun y Locali y La i ude Longi ude 16S ND4 RAG-1 c-mos
WC-3660 PEM R22021 Ichno opis capensis Zambia Ngonye Falls −16.6736 23.5969 OZ347422 OZ347978 OZ347946 OZ347961
WC-6797 PEM R27394 Ichno opis capensis Angola Quembo Ri e b idge camp −13.5275 19.2806 PV357731 PV412850 PV412873 PV390648
WC12-A191 PEM R20009 Ichno opis capensis Angola HALO Cui o Cuana ale o ice −15.1392 19.1436 PV357724 PV412843 OZ347947
WP031 NA Ichno opis capensis Namibia 50 km No h o Gobabis −22.0000 19.1400 HF547734 HF547695
WRB INH2 NA Ichno opis capensis Mozambique Inhasso o −21.7143 35.2103 OZ347423 OZ347979 OZ347948
WC-4585 PEM R23525 Ichno opis capensis Angola Quembo Ri e sou ce −13.1095 19.0061 PV357722 PV412841 PV412868 PV390643
WC-4618 PEM R23500 Ichno opis capensis Angola Quembo Ri e sou ce −13.1360 19.0453 PV357723 PV412842 PV412869
RE211206D1 NMNW R12212 Ichno opis g andiceps Namibia Khaudum −18.2876 20.9897 OZ347424 OZ347980
P3-059 MHNCUP-REP 0983 Ichno opis longico pa sp. no . Angola Cuemba −12.1707 18.2257 PV357739 PV412857 OZ347962
P9-035 MHNCUP-REP 0984 Ichno opis longico pa sp. no . Angola Mona Quimbundo −10.0583 19.8056 PV357737 PV412855 OZ347949 OZ347963
TB-44 PEM R19903 Ichno opis longico pa sp. no . Angola Camp Chi i, Miombo o es /camp −9.3969 20.4319 PV357735 PV412853 PV412874
TB-46 PEM R19905 Ichno opis longico pa sp. no . Angola Camp Chi i, Miombo o es /camp −9.3969 20.4319 PV357738 PV412856 PV412876
WC-4557 PEM R23409 Ichno opis longico pa sp. no . Angola Lungwebungu Ri e camp b idge c ossing −12.5835 18.6660 PV357741 PV412859 PV412877
WC-4560 PEM R23531 Ichno opis longico pa sp. no . Angola Sombanana illage i e −12.3071 18.6235 PV357736 PV412854 PV412875 OZ347964
WC-6291 PEM R23996 Ichno opis longico pa sp. no . Angola Lake Tchanssengwe −12.4140 18.6442 PV357740 PV412858 OZ347950
P0-44 MHNCUP-REP 0983 Ichno opis mic olepido a Angola Se a do Moco – Canjonde −12.4261 15.1478 PV357742 PV412860 PV412878 PV390651
WC-3969 PEM R23306 Ichno opis obus a sp. no . Angola 4 km ups eam om Cuana ale Ri e
sou ce
−13.0508 18.8973 PV357715 PV412835 PV412862 PV390640
WC-3994 PEM R23279 Ichno opis obus a sp. no . Angola Cuana ale Ri e sou ce −13.0903 18.8940 PV357718 PV412838 PV412864
WC-4056 PEM R23362 Ichno opis obus a sp. no . Angola d i e o Cuana ale Ri e Camp om
Samanunga illage
−13.0380 18.8298 OZ347425 OZ347981 OZ347951
WC-4816 PEM R23420 Ichno opis obus a sp. no . Angola Cuando Ri e sou ce −13.0035 19.1275 PV357716 PV41283
CKD-457 PEM R28456 Ichno opis anganicana DRC Upemba Na ional Pa k −9.0442 26.9966 OZ347426 OZ347982 OZ347952
CKD-432 PEM R28448 Ichno opis anganicana DRC Upemba Na ional Pa k −9.0442 26.9966 OZ347427 OZ347983 OZ347953 OZ347965
CKD-433 PEM R28449 Ichno opis anganicana DRC Upemba Na ional Pa k −9.0442 26.9966 OZ347428 OZ347984 OZ347954 OZ347966
CKD-442 PEM R28452 Ichno opis anganicana DRC Upemba Na ional Pa k −9.0442 26.9966 OZ347429 OZ347985 OZ347955
MTSN 9947 MUSE-VER 09947 Ichno opis anganicana DRC Kindingi, Wes o Kabobo Pla eau −5.2626 28.9076 OZ347430 OZ347956
ABH3 Me oles squamulosus Mozambique unknown JX962896 EF632221 EF632266
ABH9 Me oles squamulosus Tanzania Laela −8.7500 32.1833 JX962897 JX963022 JX962915
MBUR00872 Me oles squamulosus Sou h A ica Cle eland, Limpopo −24.0219 31.1991 LT745784 LT745812 LT745838
RSP373 Me oles squamulosus Sou h A ica Vene ia Limpopo Rese e, Limpopo −22.2661 29.3329 HF547777 HF547737 HF547699
SVN362 PEM R19626 Me oles squamulosus Sou h A ica Lapalala Game Rese e, Limpopo
Landmanslus , Limpopo
−23.8759 28.3061 HF547776 HF547736 HF547697
WP125 Me oles squamulosus Sou h A ica Rooipoo Na u e Rese e No he n Cape −28.5937 24.2100 HF547778 HF547738 HF547701

Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
632
TREE. Two pa allel uns o he MCMC we e un o 20
million gene a ions, each wi h ou independen chains,
and ees we e sampled e e y 1000 gene a ions using
BEAGLE (Ay es e al. 2019). A bu n-in o 20% was used
o gene a e he consensus ee. T ace .1.7.2 (Rambau
e al. 2018) was used o assess he e ec i e sample size
(ESS) o he un pa ame e s. The ESS alues we e ound
o be abo e 200, indica ing ha he bu n-in was adequa e.
Bo h he ML and BI ees we e gene a ed using FigT ee
.1.4.4 (Rambau 2018). Nodes wi h boo s ap suppo
(BS) ≥ 95% o ML analyses as well as pos e io p oba-
bili ies (PP) ≥ 0.95 o he BI analyses (Huelsenbeck and
Rannala 2004) we e ega ded as well suppo ed.
Species delimi a ion analyses we e pe o med o ex-
plo e species bounda ies and elucida e whe he he e
was po en ial c yp ic di e si ica ion wi hin Ichno opis.
Mi ochond ial genes (16S, ND4) we e combined o spe-
cies delimi a ion analyses, excluding he ou g oup axa,
and he sequences we e immed o 1134 bp o minimise
missing da a in he da ase s. Se e al di e en delimi a-
ion analyses we e un: Au oma ic Ba code Gap Disco -
e y (ABGD), Assemble Species by Au oma ic Pa i ion-
ing (ASAP), Poisson T ee P ocesses (PTP), Mul i- a e
Poisson T ee P ocess (mPTP), and Bayesian Poisson
T ee P ocesses (bPTP). Alignmen s we e p epa ed and
uploaded on o he ABGD Web In e ace (h ps://bioin o.
mnhn. /abi/public/abgd/abgdweb.h ml, web e sion 22
May 2023) and he ASAP Web In e ace (h ps://bioin-
o.mnhn. /abi/public/asap/asapweb.h ml, web e sion
22 May 2023) as FASTA iles. Fo ABGD, he ollowing
se ings we e used: S anda d pai wise dis ance (p dis-
ance) me ics, minimum ba code gap wid h (1), in aspe-
ci ic di e gence minimum (0.001) and maximum (0.1)
(Puilland e e al. 2012). Fo ASAP, he Simple Dis ance
(p dis ance) subs i u ion model was used (Puilland e e
al. 2021). Mul i-locus ML phylogenies we e c ea ed o
he wo conca ena ed mi ochond ial genes in IQ-TREE
as ou lined abo e. The phylogenies we e ende ed as
un oo ed nexus iles and uploaded on o he bPTP web
se e (h p://species.h-i s.o g/p p; Zhang e al. 2013) o
he PTP and bPTP analyses, and he mPTP analysis was
conduc ed using he web se e a h p://mp p.h-i s.o g/#/
ee (Kapli e al. 2016).
Unco ec ed pai wise dis ances (p dis ances) we e es i-
ma ed in MEGA X (Kuma e al. 2018) o mi ochond ial
genes (16S, ND4). Sequences we e immed o minimise
missing da a in he da ase s and p dis ances we e es ima -
ed using uni o m a es, pai wise dele ion o emaining
da a, and 500 boo s ap eplica es.
Mo phology
We examined all newly collec ed ma e ial in he collec-
ions o he Na ional Museum o Namibia (NMNW),
Windhoek, Namibia, and Po Elizabe h Museum (PEM).
Addi ional mo phological da a we e ga he ed om he
ollowing museum collec ions: PEM, Di song Na ional
Museum o Na u al His o y, P e o ia (TM); Musée Royal
de l’A ique Cen ale, Te u en, Belgium (RMCA); and
Ins i u Royal des Sciences Na u elles de Belgique, B ux-
elles, Belgium (IRSNB). The mo phological da ase was
u he supplemen ed wi h da a om p ima y li e a u e
(Boulenge 1921; Lo e idge 1933; de Wi e and Lau en
1942; Lau en 1952, 1964; B oadley 1967b; Haacke 1970)
and unpublished da a o D.G. B oadley and W.R. B anch.
The inal da ase inco po a es mo phological da a om
he ype specimens o : Ichno opis bi i a a, I. capensis
nig escens, I. chapini, I. mac olepido a, I. mic olepido a,
and I. o e lae i. Addi ionally, high- esolu ion images o
he ype specimens o I. longipes and I. anganicana we e
consul ed, as well as key specimens in he Dundo Muse-
um, Angola (DM) and Museum o Compa a i e Zoology,
USA (MCZ). Lis o ma e ial examined can be ound in
he Appendix. This ep esen s all he ele an ype ma e-
ial, excep o he ype o Ichno opis capensis, which
emains unaccoun ed o in he Na u al His o y Museum,
London (BMNH). The a ailable ma e ial enables us o
con iden ly assign ou specimens o known species and o
make in o med axonomic decisions.
Scale nomencla u e, scale coun s, and measu emen s
used in he desc ip ions ollow p e ious s udies on A i-
can Lace idae (Con adie e al. 2012; B anch e al. 2019;
Pa inha e al. 2021; Beni o e al. 2025), and we e adjus -
ed as needed o add ess he mo phology o Ichno opis.
The ollowing measu emen s we e aken in millime es
(mm) using a digi al callipe (accu acy o 0.01 mm) wi h
he aid o a Nikon SMZ1270 mic oscope: Snou – en
leng h (SVL, ip o he snou o he pos e io edge o he
cloaca); ail leng h (TAIL, ip o ail o pos e io edge o
he cloaca, measu ed only o specimens wi h comple e
o iginal ails); o al leng h (TL, combined SVL and ail
leng h); head leng h (HL, om he an e io edge o he
occipi al scale o he ip o he snou ); head wid h (HW,
wides pa o head –usually measu ed jus behind he
eye); head heigh (HH, measu ed jus behind he eyes);
snou o on o a m (S-FL, om ip o snou o an e io
inse ion o o elimb); eye diame e (ED, om op an e-
io o he pos e io edge o eye); snou o eye dis ance
(SE, om ip o snou o an e io edge o eye); eye- o-eye
dis ance (EE, om an e io edge o one eye o an e io
edge o he o he eye); ympanum leng h (Tymp-L, a
i s wides pa e ically); lowe jaw leng h (LJL, an e-
io edge o he jaw bone o ip o lowe jaw); in e -limb
leng h (ILL, dis ance be ween axilla y and inguinal e-
gions); o elimb leng h (FLL, om elbow o w is ); hind
limb leng h (HLL, om knee o heel); hind oo leng h
(HFL, om ankle o ip o ou h oe, excluding claw);
ou h inge leng h (FFL, excluding claw), ou h oe
leng h (FTL, excluding claw), leng h o an e io sup a-
ocula scale (SO), dis ance be ween an e io sup aocula
o second lo eal (SO-L, measu emen be ween he closes
poin o he an e io sup aocula o he pos e io edge o
he second lo eal), on al scale wid h (FNW, a i s wides
poin ), and on al scale leng h (FNL). All measu emen s,
excep o EE, FNW, FNL, we e aken on he igh side
o he body.
The ollowing scala ion de ails we e eco ded wi h he
aid o a dissec ing mic oscope: Numbe o sup alabials
(SL, an e io o he subocula ); numbe o in alabials
(IL); he numbe and condi ion o he nasal scales; he
Ve eb a e Zoology 75, 2025, 627–672 633
numbe and condi ion o he lo eals; scala ion condi ion
o he lowe eyelids; numbe o sup acilia ies (SC); he
condi ion o empo al scales and he elonga e empo al
pla e (scales be ween pa ie als and empo al scales); he
deg ee o head s ia ions ( idges) p esen on he do sal
head; he condi ion o he ympanum opening and he
ympanic shield; he condi ion o he on onasals (FN),
p e on als (PF, whe he he PF is in con ac wi h an e io
SO and i he PF is in con ac wi h 1s SC), on al (F),
in e pa ie al (IP), pa ie als (P) and occipi al scales (O);
numbe o sup aocula s (SO) [no e: o his s udy we e-
s ic he sup aocula s o he wo la ge scales and e e
o he clus e o 1–4 smalle scales pos e io o he 2nd
SO as he pos -sup aocula s]; numbe o smalle scales in
on o he an e io SO, ouching on al, p e on al and
1s SC; numbe o g anules in con ac wi h he wo la ge
SO and SC; numbe o pai ed chin shields (CS, and he
numbe in con ac ); midbody scale ows (MSR); longi-
udinal en al scale ows (LVSR, coun ed midway be-
ween o e- and hind limbs); ans e se ows o en als
(TVSR, coun ed om he axilla o he g oin); emo al
po es ( igh /le ); and subdigi al lamellae unde he 4 h oe
(LUFT).
To in es iga e he mo phological a ia ion be ween
Ichno opis species and o compa e hem wi h p e ious-
ly published ma e ial (Beni o e al. 2025), wo sepa a e
p incipal componen analyses (PCA) we e un on adul
specimens. We conside ed specimens o be adul s i he
SVL was la ge han 40 mm, as ha was he smalles size
a which we could clea ly obse e he hemipenal bulge
in males. Fi s , a PCA was pe o med on he ull da ase ,
which included all measu emen a iables (Da ase 1).
The ini ial analysis e ealed ha head- ela ed measu e-
men s (HH, HL, HW) accoun ed o mos o he a ia ion.
To de e mine whe he o he a iables con ibu ed no able
a ia ion, a second PCA was un on a educed da ase ha
excluded hese head measu emen s (Da ase 2). All he
a iables we e i s size-co ec ed using a linea eg es-
sion wi h body size (SVL) as he co a ia e, and he esid-
uals we e used as inpu a iables o he PCA. Va iables
wi h communali ies > 0.5 we e e ained in he analysis,
a a imax o a ion was applied, and ec o s wi h eigen-
alues > 1.0 we e ex ac ed (Tabachnick and Fidel 2019).
The esul ing p incipal componen (PC) sco es we e sa ed
and subsequen ly used as inpu o a mul i a ia e analysis
o a iance (MANOVA), wi h species as he ixed ac-
o . Di e ences be ween species we e e alua ed pos hoc
using Tukey’s HSD es . All analyses we e conduc ed in
RS udio .2023.09.1+494 (RS udio Team 2022).
In o de o explo e o he po en ial diagnos ic cha ac-
e s be ween species, we es ed he mo phological a ia-
ion in Da ase 1 and 2 ac oss di e en axa using pe mu-
a ional ANOVAs (PERMANOVAs) wi h he package
RRPP (Collye and Adams 2018) implemen ed in RS u-
dio .2023.09.1+494 (RS udio Team 2022). Va iables
we e size-co ec ed (SVL) and log- ans o med p io o
he analyses o mi iga e he e ec s o size and mul icol-
linea i y. Finally, s anda d boxplo s we e used o isually
ep esen he a iables ha we e signi ican ly di e en
be ween species.
Mapping
To enable p oduc ion o con empo a y geog aphic dis i-
bu ion maps o all Ichno opis species, obse a ion lo-
ca ions we e sou ced om published da ase s (e.g., an
den Be g 2017; Ma ques e al. 2018), museum da abases
(PEM, TM, RMCA, IRSNB), and o he ci izen science
eposi o ies (h p://www.ina u alis .o g; h p:// mus.adu.
o g.za [ eco ds we e download be o e he websi e shu
down]). Each eco d ob ained om online sou ces was
checked o diagnos ic ea u es men ioned in his s udy o
con i m species iden i ica ions. Those which could no be
con iden ly iden i ied we e excluded om he mapping
exe cise. Howe e , his o ical specimens ha we e no ex-
amined, o o which iden i ica ion could no be e i ied,
we e en a i ely mapped based on hei ini ial iden i i-
ca ion in he o iginal publica ion o museum ca alogue.
The online GeoNames gaze ee (h p://www.geonames.
o g) o he GEOLoca e Web Applica ion (h ps://www.
geo-loca e.o g/web/WebGeo e .aspx) was used o geo-
e e ence all his o ical da a lacking p ecise loca ion in-
o ma ion. Finally, all alid obse a ion eco ds we e
mapped using QGIS .3.2 (h p://qgis.o g). Da a used
o mapping can be ound a : h ps://doi.o g/10.6084/m9.
igsha e.30285421.
Resul s
Phylogene ic analyses
Bo h maximum likelihood and Bayesian in e ence ana-
lyses eco e ed iden ical opologies o he conca ena ed
da ase (Figs 1, S1, S2), wi h s ong suppo a mos majo
nodes. Al hough he mi ochond ial and nuclea gene ees
p oduce di e en opology in he placemen o I. mic o-
lepido a, hese di e ences we e no conside ed o ep e-
sen s ong phylogene ic con lic (see Figs S3, S4). Ou
phylogene ic esul s eco e ed ou dis inc clades wi hin
Ichno opis.
Ichno opis anganicana was consis en ly suppo ed as
sis e o all o he congene s in all analyses, om which
i is highly di e gen (Table 2). All species delimi a ion
me hods suppo ed i s dis inc species s a us and u he
subdi ided i in o wo lineages (Fig. 1).
Ichno opis mic olepido a was eco e ed as sis e o
he I. bi i a a clade, hough his ela ionship was no
s ongly suppo ed in he BI analysis. Ne e heless, all
species delimi a ion analyses a ou ed i s speci ic s a-
us, wi h no ably high pai wise p dis ances o 16S and
ND4 genes (Table 2). The I. bi i a a clade con ained
only ou samples and amongs hem, opo ypic ma e-
ial o I. b. pallida (KTH09-075), which exhibi ed high
in aspeci ic a ia ion. This sample di e ed ma kedly
om o he I. b. bi i a a samples, wi h di e gence al-
ues compa able o hose in he I. capensis g oup, bu
lowe han be ween o he species (Table 2). Addi ion-
ally, all species delimi a ion analyses—excep mPTP—
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
634
iden i ied each I. bi i a a lineage as a candida e
species (Fig. 1).
Ichno opis g andiceps was eco e ed as a sis-
e axon o he I. capensis g oup. Fu he mo e,
newly collec ed ma e ial om Angola was e-
co e ed as a dis inc lineage, sis e o a sample
om Namibia (RE211206D1) collec ed om
nea he ype locali y o I. g andiceps, and wi h
high di e gence in bo h mi ochond ial ma ke s
(Table 2). All delimi a ion analyses suppo ed he
dis inc i eness o he Angolan lineage as a candi-
da e new species (Fig. 1).
Wi hin he I. capensis g oup, high le els o
in aspeci ic a ia ion we e de ec ed, wi h h ee
majo clades (Clades 1–3) e u ned in he phy-
logene ic analysis. Species delimi a ion analyses
eco e ed be ween h ee and se en candida e spe-
cies among hese clades, which do no necessa ily
ag ee wi h he h ee majo clades. Sequence di-
e gence among Clade 1 wi h Clades 2 and 3 (col-
lec i ely e e ed o as I. capensis sensu la o) was
>5% o 16S and ~12% o ND4 (Table 2), com-
pa able o species-le el h esholds amongs o h-
e A ican Lace idae (~2–12% 16S and 7–23%
ND4; Con adie e al. 2012; Edwa ds e al. 2013a;
B anch e al. 2019; Pa inha e al. 2021). Fu he -
mo e, I. capensis sensu la o was consis en ly sup-
po ed as dis inc ac oss all species delimi a ion
me hods. I should be no ed ha he in a-speci -
ic a ia ion wi hin I. capensis sensu la o, while
su p isingly high, was no compa able wi h spe-
cies le el di e gence (Fig. 1; Table 2). O e all,
we p opose ha he gene ic e idence, aken wi h
o he e idence (see below) sugges s ha each o
hese g oups can be conside ed sepa a ely e ol -
ing me apopula ions unde he gene al lineage
concep o species (de Quei oz 1998; see below).
Mo phology
The wo PCAs p oduced simila esul s despi e
analysing di e en subse s o o iginal a iables.
In PCA1 (Da ase 1), PC1 (37.3%) and PC2
(13.11%) oge he explained 50.41% o he o-
al a ia ion, wi h PC1 p ima ily co ela ed o
head- ela ed a iables (HL and SE) and PC2
co ela ed o limb measu emen s (FTL and HFL;
Fig. 2; Table S2). Simila ly, in PCA2 (Da ase
2), PC1 (36.24%) and PC2 (11.41%) accoun ed
o 47.65% o he o al a ia ion, in which PC1
is also co ela ed wi h head mo phology (SE and
LJL), while PC2 co ela ed o he same a iable
as in p e ious analysis, sugges ing consis en
unde lying mo phological pa e ns ega dless o
a iable inclusion. The MANOVA showed sig-
ni ican di e ences be ween he species o only
PC2 (P = 0.000) in bo h PCAs, and o PC1 o
PCA2 (P = 0.007; Table S2). Pos hoc pai wise
compa isons (Tukey’s HSD) e ealed no signi -
Table 2. Mean sequence di e gences (unco ec ed p dis ances) be ween Ichno opis species o 16S and ND4 genes, gi en as pe cen ages. The numbe s in he diagonal g ey boxes ep esen he mean
in aspeci ic sequence di e gences and s anda d e o s, numbe s below he diagonal g ey boxes ep esen he mean in e speci ic sequence di e gences, while numbe s abo e he diagonal g ey boxes
ep esen s anda d e o s o he in e speci ic sequence di e gences. n/c – was no possible o es ima e sequence di e gences.
16S 12345678
1I. capensis sensu la o 3.0 ± 0.5 0.9 1.5 1.4 1.4 1.4 1.7 1.6
2I. longico pa sp. no . 5.9 2.6 ± 0.8 1.5 1.4 1.3 1.2 1.6 1.6
3I. obus a sp. no . 11.2 12.0 0.2 ± 0.2 1.1 1.4 1.3 1.7 1.5
4I. g andiceps 11.5 11.8 6.3 n/c 1.4 1.3 1.7 1.6
5I. b. pallida 11.6 10.5 9.9 10.3 n/c 0.9 1.7 1.5
6I. b. bi i a a 11.5 9.8 8.8 9.3 5.4 3.8 ± 0.8 1.6 1.4
7I. mic olepido a 16.4 16.1 16.4 16.1 14.4 13.9 n/c 1.8
8I. anganicana 15.8 14.5 12.9 13.8 13.1 12.2 18.4 1.8 ± 0.4
ND4
1I. capensis sensu la o 7.7 ± 0.6 0.8 1.3 1.3 1.4 1.1 1.3 1.2
2I. longico pa sp. no . 12.3 6.6 ± 0.6 1.3 1.3 1.4 1.1 1.3 1.2
3I. obus a sp. no . 22.1 19.6 1.4 ± 0.3 1.3 1.6 1.2 1.5 1.4
4I. g andiceps 22.3 20.4 15.3 n/c 1.6 1.2 1.5 1.4
5I. b. pallida 21.5 20.6 22.3 20.7 n/c 1.1 1.6 1.7
6I. b. bi i a a 19.2 17.6 20.0 19.6 12.8 9.4 ± 0.9 1.3 1.3
7I. mic olepido a 24.0 21.6 24.4 24.4 22.6 20.6 n/c 1.4
8I. anganicana 19.1 18.0 21.7 21.8 22.7 19.4 21.2 0.1 ± 0.1
Ve eb a e Zoology 75, 2025, 627–672 635
ican di e ences be ween species o PC1 (P > 0.05),
bu signi ican di e ences we e de ec ed o PC2 in bo h
analyses (Table S3). The main di e ences de ec ed we e
be ween species o di e en g oups (e.g., I. bi i a a e -
sus I. capensis; I. capensis e sus I. g andiceps) bu no
wi hin he di e en g oups (Table S3).
Figu e 1. IQ-TREE maximum likelihood consensus phylogeny o Ichno opis wi h likelihood boo s ap suppo alues (abo e) and
Bayesian pos e io p obabili ies (below) indica ed a each node (see key in op le ). The colou ed ba s o he igh o he phylogeny
summa ise he esul s o each o he species delimi a ion analyses. Inse image: I. capensis sensu s ic o. The scale ba ep esen s
subs i u ions/si e.
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
642
Ichno opis chapini Schmid , 1919
Chapin’s ough-scaled liza d
Figu e 8; Table 3
Taxonomic no e. When I. chapini was desc ibed, i was
di e en ia ed om i s congene s based on he p esence
o an an e io sup alo eal, hus ha ing wo an e io lo-
eal scales (Schmid 1919). Howe e , addi ional ma e-
ial collec ed om Ad a in no heas e n DRC does no
possess any an e io sup alo eal (de Wi e 1933; Lau en
1952). Despi e he limi ed ma e ial a ailable, he scala-
ion obse ed in he ype specimen appea s o be anom-
alous. This is u he suppo ed by he ac ha we ha e
only eco ded his condi ion (an an e io sup alo eal)
once o all o he o he Ichno opis specimens examined
(n = 432). As in he p e ious species, I. chapini was as-
signed o he I. capensis g oup based on he obse a ion
ha he p e on al is sepa a ed om he an e io sup a-
ocula (Boulenge 1921). Examina ion o high- esolu ion
pho og aphs o he holo ype (Fig. 8) and physical exam-
ina ion o addi ional ma e ial om RMCA showed ha
his species belongs o he I. bi i a a g oup, based on
he mo e ounded head (Fig. 2). This species also seems
geog aphically well isola ed om o he species in he ge-
nus, bu his migh jus be an e ec o unde -sampling.
The seasonali y o Ichno opis species makes hem ha d
o obse e ou side o he b eeding season. Consequen -
ly, coupled wi h he absence o any mode n ma e ial and
hus molecula da a, we e ain his species as alid un il
mo e da a become a ailable.
Holo ype. AMNH 10674, adul emale, collec ed om
Aba, Hau -Uele P o ince, DRC in July 1911.
Gene al desc ip ion. A medium-sized lace id wi h a o-
bus , ounded snou . Head scala ion mode a ely s ia ed.
Nos il pie ced be ween h ee nasals; he sup anasals a e
in b oad con ac behind he os al; single on onasal, as
b oad as long; pai ed p e on al scales in b oad con ac
medially; p e on al no in con ac wi h an e io sup a-
ocula and sepa a ed om he sup acilia ies by a smalle
scale; wo la ge sup aocula s, which a e sepa a ed om
he sup acilia ies by one ow o small scales (6–8) and
p eceded by a clus e o 2–3 (3 median) smalle scales;
one pos -sup aocula scale; wo lo eal scales p esen ,
which a e sepa a ed om he an e io sup aocula by wo
scales (excep in he holo ype, whe e he an e io lo e-
al is di ided o o m a sup alo eal on bo h sides and on
he le side o BE_RMCA_Ve .R.3657); subocula in
con ac wi h lip; 4–5 (mos ly 4) sup alabials in on o
subocula ; 6–7 (mos ly 6) in alabials; i e chin shields,
wi h he an e io 2–3 in b oad con ac (in he holo ype
only he i s wo chin shields a e in con ac , while in BE_
RMCA_Ve .R.3657 he hi d chin shield is in na ow
Table 3. Summa y o mo phological da a o he Ichno opis bi i a a g oup. Measu emen s a e all shown in millime es (mm).
Values a e gi en as a ange, wi h mean ± s anda d de ia ion in pa en hesis. Ju eniles we e excluded om he measu emen s, bu
we e included in he scala ion da a. Fo abb e ia ions see Ma e ials and Me hods sec ion. n = sample size.
Cha ac e s I. bi i a a I. chapini I. mic olepido a I. anganicana
n = 39 n = 3 n = 6 n = 32
SVL 42.2–75.0 (63.2 ± 8.33) 53.8–58.0 (55.6 ± 2.15) 48.7–52.0 (50.4 ± 1.45) 41.0–60.0 (53.9 ± 4.21)
TAIL 85–156 (109.7 ± 15.47) 77 69.8 55.6–107.9 (81.7 ± 12.15)
HL 12.1–15.3 (13.6 ± 1.22) 11.7–12.9 (12.2 ± 0.64) 12.6 11.2–14.2 (12.6 ± 0.79)
HW 6.0–9.9 (8.3 ± 1.08) 7.0 –8.7 (7.7 ± 0.89) 6.7–7.3 (7.0 ± 0.20) 6.1–8.7 (7.6 ± 0.51)
HH 5.4–8.7 (7.2 ± 0.90) 6.5 5.3–6.7 (5.9 ± 0.62) 5.2–8.4 (6.8 ± 0.71)
ED 4.1–4.7 (4.4 ± 0.29) 4.2–5.2 (4.7 ± 0.70) 3.6 3.9–4.5 (4.2 ± 0.18)
SE 3.9–6.8 (5.6 ± 0.90) 4.9–5.9 (5.4 ± 0.53) 4.3–5.2 (4.7 ± 0.37) 5.1–6.6 (5.8 ± 0.36)
LL 14.2–17.2 (15.6 ± 1.6) 11.5 –13.8 (12.7 ± 1.63) 13.3 12.3–16.0 (13.8 ± 1.02)
IL 23.6–33.9 (26.7 ± 3.31) 24.3–33.0 (28.8 ± 4.33) 22.5 19.4–29.4 (24.8 ± 3.0)
FLL 5.8–9.4 (7.5 ± 1.13) 6.1–6.9 (6.5 ± 0.52) 5.4 4.7–8.7 (6.5 ± 0.80)
HLL 9.0–12.4 (10.4 ± 1.28) 9.2–9.5 (9.3 ± 0.23) 7.4 7.9–10.7 (9.2 ± 0.79)
TAIL/SVL 1.3–2.4 (1.8 ± 0.28) 1.3 1.4 1.2–2.0 (1.6 ± 0.23)
HL/SVL 0.2 (0.2 ± 0.01) 0.2 (0.2 ± 0.02) 0.3 0.2–0.3 (0.2 ± 0.02)
ES/HL 0.4–0.5 (0.4 ± 0.02) 0.4–0.5 (0.4 ± 0.02) 0.4 0.4–0.5 (0.5 ± 0.01)
HW/HL 0.5–0.7 (0.6 ± 0.05) 0.3–0.7 (0.6 ± 0.10) 0.6 0.6 (0.6 ± 0.03)
MSR 29–40 34–35 43–50 28–42
LVSR 8–10 8–10 8 –10 8–10
TVSR 22–31 24–25 26–30 20–27
SL 3–6 (mos ly 4) 4–5 (mos ly 4) 4 3–5 (mos ly 4)
IL 6–9 (mos ly 6) 6–7 (mos ly 6) 6–8 (mos ly 7) 5–7 (mos ly 6)
SC 3–4 (mos ly 4) 3–5 (mos ly 4) 4 4–5 (mos ly 4)
LUFT 17–24 18–20 16–19 17–22
Femo al po es 10–14 8–9 10–13 10–15

Ve eb a e Zoology 75, 2025, 627–672 643
con ac an e io ly); 3–5 (mos ly 4) sup acilia ies; 34–35
midbody scale ows; 8–10 longi udinal ows o enla ged
en al pla es; 24–25 ans e se en al scale ows;18–20
subdigi al lamellae unde 4 h oe; 8–9 emo al po es pe
high. Size: Adul specimens a ied om 53.8–58.0 mm
(median: 55.0 mm) SVL and 77 mm TAIL (all speci-
mens’ ails missing o unca ed; his measu emen is
based on Schmid 1919). La ges emale: 58 mm SVL
(AMNH 10674 – holo ype); la ges male: 55.0 mm SVL
(BE_RMCA_Ve .R.3656 – Ad a, DRC). Colou a ion
(based on p ese ed specimens; Fig. 8): Do sal su ace
uni o mly g eyish b own, wi h sca e ed da ke b own
o black scales. A dis inc la e al whi e s ipe o igina es
a he subocula egion, bo de ed bo h do sally and en-
ally by na ow black lines; his s ipe ex ends o e he
o elimbs bu does no each he hind limbs in emales
(AMNH 10674 and BE_RMCA_Ve .R.3657), bu each-
es he hind limbs in he male (BE_RMCA_Ve .R.3656).
A second ain do sola e al line is p esen abo e he low-
e whi e s ipe, and only ex ends o jus pos e io o he
o ea ms in emales, while in he male his s ipe is mo e
p ominen and ex ends o jus abo e he hind limbs. Be-
ween hese lines a e a se ies o ans e se black spo s
in he emales, each spanning 2–3 scales in wid h and
app oxima ely hal a scale in leng h, loca ed a he ips o
he scales. In he male he space be ween he wo whi e
s ipes o ms a p ominen black band wi h sca e ed black
scales. This band ex ends on o he empo al and snou
a ea. Do sally, wo simila se ies o ans e se black
ma kings lank he e eb al egion, ex ending la e ally
o he do sola e al s ipe. Ven al scales and chin shields
a e whi e wi h sub le g ey ma gins. The wo ou e mos
en al ows a e punc ua ed wi h small b own do s in e-
males, bu o m a con inuous na ow black band be ween
he limbs in he male and ex ends on o he sup alabials.
Sup alabials and in alabials a e i egula ly mo led wi h
ligh and da k pigmen a ion. Limbs g eyish b own do -
sally, ansi ioning o a ligh e one on he en al su -
aces.
Dis ibu ion. Only known om no heas e n DRC in he
icini y o Aba (Fig. 3). Gi en his locali y’s p oximi y
o he bo de wi h Sou h Sudan (< 10 km), i is likely o
occu in he la e coun y.
Habi a and Na u al His o y. Ve y li le is known abou
his species, bu i is expec ed o ha e simila habi a e-
qui emen s o o he Ichno opis species.
Figu e 8. Holo ype (AMNH 10674) o Ichno opis chapini om Aba, Hau -Uele P o ince, Democ a ic Republic o he Congo.
Pho og aphs o body in A do sal and B en al iews, and head in C do sal, D la e al and E en al iews. Scale ba s ep esen 10
mm. Pho og aphs: Lau en Vonnahme.
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
644
Ichno opis mic olepido a Ma x, 1956
Moun Moco ough-scaled liza d
Figu es 9, 10; Table 3
Taxonomic no e. Desc ibed based on i e specimens
e ie ed om he c op o a Da k Chan ing Goshawk
(Melie ax me aba es) a he base o Se a do Moco ( he
geog aphical ea u e o Se a do Moco which includes he
highes peak in Angola a 2620 m a.s.l., is o en collo-
quially e e ed o as Moun Moco) (Ma x 1956). Pa ke
(1936) was ac ually he i s o documen his species as
I. bi i a a om Se a do Moco and alluded o i s small-
e do sal scales. Rema kably, hese smalle do sal scales
we e one o he main diagnos ic ea u es when I. mic o-
lepido a was desc ibed. Howe e , i s axonomic s a us
has been dispu ed in he pas (Maye 2013) because o i s
esemblance o I. bi i a a and he lack o p ecise local-
i y da a, gi en ha he ype se ies was ound in he c op
o a da k chan ing goshawk (Ma x 1956). No addi ional
specimens we e collec ed un il PVP collec ed a opo ypic
specimen in Oc obe 2020 a Se a do Moco (Beni o e al.
2025). Thanks o his new ma e ial, Beni o e al. (2025)
p o ided he i s phylogene ic placemen o he species,
alida ing i s axonomic s a us and demons a ing ha
his species belongs o he I. bi i a a g oup.
Holo ype. FMNH 74285, adul male, collec ed om he
‘ oo o Moun Moco’ [= Se a do Moco], Huambo P o -
ince, Angola, by Ge d Hein ich on 19 Sep embe 1954.
Pa a ypes. FMNH 74283–84 ( emales), FMNH 74286–
87 (males); same collec ion de ails as holo ype.
Figu e 9. Pho og aphs in li e o Ichno opis mic olepido a (MHNCUP-REP0983) om Se a do Moco, Huambo P o ince, Angola
(adap ed om Beni o e al. 2025). Pho og aphs in A do sola e al iew o he ull body and B la e al iew o he head. Pho og aphs:
Ped o Vaz Pin o.
Figu e 10. Ichno opis mic olepido a (MHNCUP-REP0983) specimen om Se a do Moco, Huambo P o ince, Angola (adap ed
om Beni o e al. 2025). Pho og aphs o body in A do sal and B en al iews, and head in C do sal, D la e al and E en al iews.
Scale ba s ep esen 10 mm. Pho og aphs: Max Beni o.
Ve eb a e Zoology 75, 2025, 627–672 645
Addi ional ma e ial. MHNCUP-REP0983, adul male,
collec ed a Se a do Moco, Huambo P o ince, Angola
(–12.4554°, 15.1632°, 2300 m a.s.l.), on 18 Oc obe 2020
by Ped o Vaz Pin o (Beni o e al. 2025); ju enile speci-
men collec ed a Se a do Moco, Huambo P o ince, An-
gola, 1500–1900 m a.s.l., in Ma ch 1934 by Ka l Jo dan
(Pa ke 1936).
Gene al desc ip ion. A medium-sized, obus lace id
wi h a ounded snou and s ongly s ia ed and keeled head
scales. Nos il pie ced be ween h ee nasals; he sup ana-
sals a e in b oad con ac behind he os al; single on-
onasal, as b oad as long; pai ed p e on al scales in b oad
con ac medially; p e on al in con ac wi h he an e io
sup aocula and ei he in con ac o na owly sepa a ed
om sup acilia ies by a smalle scale; wo la ge sup aoc-
ula s, p eceded by a single scale (documen ed by Ma x
1956 as a small sup aocula ); he an e io sup aocula is
in b oad o na ow con ac wi h he 1s sup acilia y an e-
io ly; he pos e io pa o he an e io sup aocula and
he pos e io sup aocula a e sepa a ed om he sup acil-
ia ies by one ow o small scales (6); one pos -sup aoc-
ula scale; wo lo eal scales p esen , which a e sepa a ed
om he an e io sup aocula by one scale; subocula in
con ac wi h lip; ou sup alabials in on o subocula ;
6–8 in alabials (mos ly se en); i e chin shields, wi h he
an e io h ee pai s in b oad con ac ; ou sup acilia ies
(Ma x 1956 eco ded i e, bu he included he pos e i-
o lo eal); 43–50 midbody scale ows; 8–10 longi udinal
ows o enla ged en al pla es; 26–30 ans e se en al
scale ows; 16–19 subdigi al lamellae unde he 4 h oe;
10–13 emo al po es pe high. Size: Adul specimens
a ied om 48.7–52.0 mm (mean: 50.4 mm) SVL and
69.8 mm TAIL (only one specimen wi h in ac ail). La g-
es emale: 51 mm SVL (FMNH 74283); la ges male:
52 mm SVL (FMNH74285, 74286). Colou a ion (Fig.
9): The do sal pa e n ea u es a ligh b own cen al band
ex ending om jus behind he head o he hind limbs.
This band is bo de ed on each side by wo b oken ows
o black blo ches. Along he la e al sides o he body, wo
c eam o yellow longi udinal s ipes un om he le el o
he ea openings pos e io ly o he hind limbs. Be ween
hese s ipes lie a se ies o pai ed whi e ocelli, each bo -
de ed ex e nally by black ings. Below he lowe la e al
s ipe he e is a con inuous ow o single whi e ocelli. The
do sal su ace o he head is b own, mo led wi h black
speckling ac oss mos scales. The mou h is bo de ed in
black, which ades o whi e along he uppe po ion o he
sup alabials and he lowe po ion o he in alabials. The
i s ow o chin shields is en i ely black, while ows wo
h ough i e a e bicolou ed—black medially and whi e
la e ally. The h oa (gula egion) is pale ed-o ange, in-
e spe sed wi h black scales and ma ked by wo dis inc
b igh yellow-o ange spo s loca ed benea h he pos e io
ends o he lowe jaws. The en al su ace is uni o mly
whi e
Dis ibu ion. Cu en ly only known om he slopes
o Se a do Moco, in he cen al Angolan highlands
(Fig. 3).
Habi a and Na u al His o y. The specimens om he
ype se ies we e p eyed upon by a da k chan ing goshawk
(Melie ax melaba es) (Ma x 1956). The specimen col-
lec ed by PVP (MHNCUP-REP0983) was ound du ing
he day on op o an exposed small ock in open mon-
ane g assland, wi h hick ege a ion co e a 2300 m
a.s.l. (Beni o e al. 2025). The mon ane habi a in Se a
do Moco is mainly o med by a hick laye o g ass and
small bushes as well as many ocks unde nea h. This ype
o habi a is likely o hinde he species de ec abili y.
Ichno opis anganicana Boulenge , 1917
Tanzanian ough-scaled liza d
Figu es 11–13; Table 3
Taxonomic no e. This species was desc ibed om he
‘Eas Coas [o ] Lake Tanganyika’ in mode n-day Tan-
zania based on a single subadul specimen ha was col-
lec ed in 1896. When Boulenge (1917) desc ibed I. an-
ganicana, he asc ibed he holo ype o a subadul male.
Howe e , a e ou examina ion o high- esolu ion pho-
og aphs o he ype specimen i was no possible o sex
i , so we ega ded i as an unsexed subadul specimen.
Since i s desc ip ion, no addi ional ma e ial has been doc-
umen ed. Howe e , due o he ague desc ip ion p o ided
by Boulenge (1917), he axonomic s a us o his species
has been ques ioned by some au ho s (Maye 2013). On
he o he hand, based on some diagnosable head scala-
ion ea u es (i.e., sup aocula s in con ac wi h sup acil-
ia ies), his species was p elimina y e ained as alid in
subsequen yea s (Spawls e al. 2002, 2018; an den Be g
2017; Ue z e al. 2025).
In his s udy, an adul emale specimen collec ed om
he mid-ele a ion Miombo woodlands wes o he Kabo-
bo Pla eau, DRC (MTSN 9947; Fig. 11E) ag eed wi h he
desc ip ion o I. anganicana based on he sup aocula
a angemen , (i.e., an e io sup aocula in di ec con ac
wi h he sup acilia ies), and he colou a ion (b onzy oli e
do sum wi h h ee ine black s ipes on nape). Howe e ,
Boulenge (1917), in his desc ip ion o he ype speci-
men a e 20 yea s o p ese a ion, did no documen he
unique, e enly-spaced blue do sola e al spo s obse ed in
he new DRC specimen (Fig. 11E). Based on his new
in o ma ion abou he do sal colou a ion, we e isi ed he
li e a u e, examined known museum specimens (p e i-
ously asc ibed o I. bi i a a in eas e n DRC and adjacen
Zambia and Tanzania) and consul ed online ci izen sci-
ence pla o ms.
O special in e es is he case o he i s specimens o
I. bi i a a om Ipemi, Udzungwa Moun ains, Tanzania,
documen ed by Lo e idge (1933). He s a es ha , in com-
pa ison o he ype, he ega ds his specimens as conspe-
ci ic wi h I. bi i a a and dis inc om I. anganicana, o
which he also examined he ype. Howe e , he p o ides
no u he de ails. In his desc ip ion o he specimens’ co-
lou , he o e ed a de ailed accoun o he colou a ion as
ollows: ‘…se ies o blo ches which is a he mo e black
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
646
han ches nu -b own ha ing he appea ance o ocelli by
eason o a blueish-whi e cen al spo in each …’. Exam-
ina ion o high esolu ion images o he wo Ipemi spec-
imens in he Museum o Compa a i e Zoology (MCZ
R30836–7) con i med he p esence o he unique blue
la e al spo s (al hough aded o whi e in p ese a i e)
and he do sal colou a ion, bu he sup aocula s we e no
in con ac wi h he sup acilia ies, as epo ed in he ype
specimen (BMNH 1946.9.3.49) o I. anganicana. This
di e ence migh ha e been he eason why Lo e idge
(1933) conside ed his ma e ial o be conspeci ic wi h
I. bi i a a a he han I. anganicana.
Addi ionally, de Wi e and Lau en (1952) again men-
ioned hese unique do sola e al blue spo s in he colou
desc ip ion o I. o e lae i: “… om his place i is some-
imes eplaced by a se ies o small blue spo s mo e o less
bo de ed wi h black, ex ending o he base o he hind
limbs; blue spo s a e also p esen on he uppe band, be-
ween he on and hind limbs.” ( ansla ion om F ench
o English). When we examined he ype specimens o
I. o e lae i a he RMCA (Fig. 13), we no only con i med
he emnan s o blue la e al spo s and he nape colou a ion,
bu we also con i med he p esence o con ac be ween
he sup aocula s and he sup acilia y scales, in ag eemen
Figu e 11. Pho og aphs in li e o Ichno opis anganicana om ac oss i s ange, depic ing he e enly spaced do sola e al blue spo s.
Pho og aphs om A Rukwa, Tanzania; B Cambua, Democ a ic Republic o he Congo; C Rumphi, Malawi (h ps://www.ina u alis .
o g/obse a ions/146895735); D Nyika Na ional Pa k, Malawi (h ps://www.ina u alis .o g/obse a ions/146684850); E Kindingi,
Lake Tanganyika (MTSN 9947), Democ a ic Republic o he Congo; and F Upemba Na ional Pa k, Democ a ic Republic o he
Congo (h ps://www.ina u alis .o g/obse a ions/249778421). Pho og aphs: Da id Lloyd-Jones; B – Colin Tilbu y, C – Ma c Hen-
ion, D – Tim B amme , E – Wandege Muninga, D – Na ali Honig.
Ve eb a e Zoology 75, 2025, 627–672 647
Figu e 12. Holo ype (BMNH 1946.9.3.49) o Ichno opis anganicana om ‘Eas coas o Lake Tanganyika’, Tanzania. Pho o-
g aphs o body in A en al and B do sal iews, and head in C do sal, D la e al and E en al iews. Pho og aphs: Pa ick Campbell.
Figu e 13. Holo ype (BE_RMCA_Ve .R.9691) o Ichno opis o e lae i om Kapanga, Hau -Ka anga P o ince, Democ a ic Re-
public o he Congo. Pho og aphs o body in A en al and B do sal (no e he e enly spaced whi e do sola e al spo s indica ed by
he a ows) iews, and head in C do sal, D la e al and E en al iews. Scale ba s ep esen 10 mm. Pho og aphs: Max Beni o.

Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
648
wi h I. anganicana. Ne e heless, his ea u e was only
p esen in he holo ype (BE_RMCA_Ve .R.9691) and
one o he o iginal pa a ypes (BE_RMCA_Ve .R.1869,
la e used as a pa a ype o I. nig escens). Howe e , he
o he pa a ype ma e ial con o med mo phologically o ei-
he I. bi i a a (see abo e) o I. capensis sensu la o (see
below).
O he published sou ces showing pho og aphs o I. bi-
i a a (sic) wi h blue spo s include de Wi e (1933: pla e
2, ig. 1) om sou heas e n DRC, Spawls e al. (2018:
202, bo om igh ) om sou hwes e n Tanzania, and
Phadima e al. (2024: 22; also on iNa u alis 146895735)
om no hwes e n Malawi. Addi ional eco ds we e
also ound on iNa u alis (191773297, 146684850,
147210660, 87417155) and Rep ileMap (169500) om
DRC, Malawi and Zambia
Based on he combined e idence, all he abo e ma e i-
al can hus be con iden ly assigned o I. anganicana. We
he e o e ake his oppo uni y o expand on he o iginal
desc ip ion o I. anganicana and synonymise I. o e lae i
wi h I. anganicana.
Synonymy. Ichno opis o e lae i de Wi e & Lau en ,
1942: 173 (new synonymy).
Holo ype. BMNH 1946.9.3.49 (96.5.14.14), collec ed
om ‘Eas Coas [o ] Lake Tanganyika’, Tanzania, p e-
sen ed o he museum by M . WH. Nu in 1896.
Gene al desc ip ion. A medium-sized lace id wi h a
obus , ounded snou . Head scala ion weakly o mod-
e a ely s ia ed. Nos il pie ced be ween h ee nasals;
he sup anasals a e in b oad con ac behind he os al;
single on onasal, as b oad as long; pai ed p e on al
scales in b oad con ac medially; p e on al mos ly in
con ac wi h he an e io sup aocula (n = 29 in con ac ,
se en no in con ac ; h ee in con ac on one side only)
and sepa a ed om sup acilia ies by a smalle scale; wo
la ge sup aocula s, which a e ei he in di ec con ac (n
= 15) o sepa a ed (n = 18) om he sup acilia ies by a
se ies o small scales; hose ha a e no in con ac a e
sepa a ed by one ow o small scales (3–9) and p eced-
ed by a clus e o 1–6 (1.7 a e age) smalle scales; one
pos -sup aocula scale; wo lo eal scales p esen , which
a e sepa a ed om he an e io sup aocula by wo scales;
subocula in con ac wi h lip; 3–5 (mos ly 4) sup alabials
in on o subocula ; 5–7 (mos ly six) in alabials; i e
chin shields, wi h he an e io h ee in b oad con ac ; 4–5
(mos ly ou ) sup acilia ies; 28–42 (a e age: 36.0) mid-
body scale ows; 8–10 (a e age: 8.4) longi udinal ows
o enla ged en al pla es; 20–27 (a e age: 22.8) ans-
e se en al scale ows; 17–22 subdigi al lamellae unde
he 4 h oe; 10–15 emo al po es pe high. Size: Adul
specimens a ied om 41.0–60.0 mm (mean: 53.9 mm)
SVL and 55.6–107.9 mm (mean: 81.7 mm) TAIL. La ges
emale: 60 mm SVL (NMZB-UM 24433 – Misuku Hills,
Malawi); la ges male: 56 mm SVL (NMZB-UM 24432
– Misuku Hills, Malawi). Colou a ion (Fig. 11): The op
o he head and he an e io pa o he body a e coppe y
ed, some imes wi h h ee clea ly de ined black s ipes
on he nape. The an e io pa o he do sum is g ey wi h
sca e ed b own pai ed blo ches wi h black edging, ex-
ending on o he ail. The lanks a e da k b own o black,
ypically wi h in e up ed whi e do sola e al s ipes. The
uppe s ipe o igina es behind he eye and ex ends on o
he neck and hen b eaks up in o smalle whi e blo ch-
es. The lowe s ipe begins an e io ly a he sup alabi-
als, acing pos e io ly h ough he ea and o e he a m,
b eaking in o smalle whi e blo ches on he an e io hi d
o he body. Be ween hese wo s ipes lies a b oad da k
b own o black band. Diagnos ic, e enly-spaced g een
o blue spo s s a abo e he a m and ex end pos e io ly
o he g oin in bo h sexes. Benea h he lowe in e up ed
whi e s ipe/blo ches lies ano he na ow band o b own
o black scales, some imes accompanied by o ange spo s
o blo ches ex ending on o he en e . Du ing he b eed-
ing season, males exhibi mo e p ominen o ange lanks,
while he lowe whi e s ipe and la e al head become i -
id yellow an e io ly. The specimen om he DRC (MTSN
9947) exhibi s a b igh o ange lowe jaw. The en e is
ypically plain whi e bu can ha e ligh g ey colou a ion.
Dis ibu ion. Known om wes e n Tanzania, sou h o
no he n Malawi, and eas wa d o no he n Zambia and
sou he n DRC (Fig. 3).
Habi a and Na u al His o y. The Lukwa i specimen
was disco e ed in g assland adjacen o B achys egia
woodland. This specimen exhibi ed peculia leg- ucking
beha iou , whe ein i aised i s body and olded i s legs o
he sides (Spawls e al. 2018; Lloyd-Jones pe s. comm.).
A g a id emale was obse ed laying eggs in Janua y
(iNa u alis 146684850). Shelled eggs in he o iduc s o
one specimen measu ed 13.5 mm × 6.5 mm (Robe son e
al. 1963). S omach con en s we e documen ed o con ain
Ac ididae, Man idae, Isop e a, and A aneae (Robe son
e al. 1963).
The Ichno opis g andiceps g oup
Phylogene ically and mo phologically, he I. g andiceps
g oup includes I. g andiceps and a candida e new species
om Angola desc ibed he e, which sha e he ollowing
mo phological ea u es: Robus , b oad and dep essed
head; he p e on al always sepa a ed om he an e io
sup aocula ; weak head s ia ions; and uni o m b own o
ed do sum. This g oup is es ic ed o he Kalaha i Basin,
om cen al Angola o no he n Namibia (Fig. 14).
Ve eb a e Zoology 75, 2025, 627–672 649
Ichno opis g andiceps B oadley, 1967
Zambezi ough-scaled liza d
Figu es 15, 16; Table 4
Taxonomic no e. This is he mos ecen ly desc ibed
species o Ichno opis. I was desc ibed om he wes e n
Zambezi Region in no h-eas e n Namibia, based on only
h ee specimens, and was dis inguished om sympa ic
I. capensis based on i s la ge size, ounded head and do sal
colou a ion (B oadley 1967b). This is a a ely documen ed
species and i is only known om he ype se ies, ou ad-
di ional specimens collec ed om no h-eas e n Namibia
(Haacke 1970), one specimen om Khaudum, Namibia
( an B eda 2023), and one specimen om wes e n Zambia
(Pie e sen e al. 2017). Con adie e al. (2022a) en a i e-
ly assigned ma e ial om eas e n Angola o his species
based on sha ed mo phology, bu phylogene ic analyses
(see Resul s) eco e i as a sepa a e sis e lineage, which
ep esen s a candida e new species desc ibed below.
Holo ype. USNM 163989, an adul male, collec ed ‘25
miles wes o Mohembo, Bo swana, on he bo de o he
Cap i i S ip (Sou h Wes A ica)’, Namibia by T.N. Li -
e sedge and S.W. Goussa d on 20 May 1967.
Pa a ypes. NMZB-UM 16278 (male) and USNM 163990
(ju enile); same collec ion de ails as holo ype.
Gene al desc ip ion. A la ge, obus lace id wi h a poin -
ed snou . Head scala ion weakly s ia ed. Nos il pie ced
be ween h ee nasals; he sup anasals a e in b oad con ac
behind he os al; single on onasal, as b oad as long;
pai ed p e on al scales in b oad con ac medially; p e-
on al sepa a ed om he an e io sup aocula by a small-
e scale (excep on he igh -side o TM 86237) and sepa-
a ed om sup acilia ies by a smalle scale (excep on he
igh -side o TM 38309); wo la ge sup aocula s, which
a e sepa a ed om he sup acilia ies by one ow o small
scales (5–9) and p eceded by a clus e o smalle scales
(3–7); 2–3 pos -sup aocula scales; wo lo eal scales p es-
en , which a e sepa a ed om he an e io sup aocula
by wo scales; subocula in con ac wi h lip; 4–5 (mos ly
i e) sup alabials in on o subocula ; 5–7 (mos ly six)
in alabials; i e chin shields, wi h he an e io h ee in
b oad con ac ; 4–5 (mos ly i e) sup acilia ies; 44–47
(a e age: 45.6) midbody scale ows; 10 longi udinal ows
o enla ged en al pla es; 27–31 (a e age: 28.3) ans-
e se en al scale ows; 20–26 subdigi al lamellae unde
he 4 h oe; 8–14 emo al po es pe high. Size: Adul spec-
imens a ied om 57.2–77.9 mm (mean: 65.6 mm) SVL
and 103.4–148.0 mm (mean: 124.9 mm) TAIL. La ges
emale: 77.9 mm SVL (RE211206D1/NMNW R12212 –
Khaudum, Namibia); la ges male: 70 mm SVL (USNM
163989 – 40 km W o Mohembo, Bo swana). Colou a ion
(in p ese a i e; Fig. 16): Abo e pale g ey-b own, wi h
da ke s ippling and a ew sca e ed da k black spo s on
he body and ail. A poo ly de ined da k b own do sola e -
al band ex ends om he neck o he g oin, whe e i b eaks
up in o a line o la e al spo s on he ail. Sides o he head
and lowe lanks whi e. Ven e whi e. In ju eniles o sub-
adul s (Fig. 15), he da k b own la e al band is eplaced
by a mus a d-colou ed band (Pie e sen e al. 2017).
Figu e 14. Reco ds o he Ichno opis g andiceps g oup, based on all li e a u e eco ds (open ci cles), examined ma e ial (closed
ci cles) and gene ically analysed ma e ial (whi e cen es). Respec i e ype locali ies a e indica ed by a ows: Ig – I. g andiceps, and
I – I. obus a sp. no .
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
650
Figu e 15. Pho og aphs in li e o ju enile Ichno opis g andiceps. Specimens pho og aphed om A Ngonye Falls (TM 86237),
Zambia and B Chi okoloki, Zambia. Pho og aphs: A – Da en Pie e sen; B – F ank Willems.
Figu e 16. Ichno opis g andiceps (RE211206D1/NMNW R12212) specimen om Khaudum, Namibia. Pho og aphs o body in A
en al and B do sal iews, and head in C do sal, D la e al and E en al iews. Pho og aphs: We ne Con adie.
Table 4. Summa y o mo phological da a o he Ichno opis g andiceps g oup. Measu emen s a e p esen ed in millime es (mm).
Values a e gi en as a ange wi h mean ± s anda d de ia ion in pa en hesis. Da a o adul s and subadul s a e p esen ed sepa a ely.
Fo abb e ia ions, see he Ma e ials and Me hods sec ion. n = sample size.
Cha ac e s I. g andiceps I. g andiceps I. obus a sp. no . I. obus a sp. no .
n = 6 (adul s) n = 4 (subadul s) n = 5 (adul s) n = 11 (subadul s)
SVL 57.2–77.9 (65.6 ± 7.52) 57.2–77.9 (65.6 ± 7.52) 71.9–78.8 (74.9 ± 3.12) 35.6–51.1 (44.6 ± 5.33)
TAIL 103.4–148.0 (124.9 ± 16.21) 38.5–50.2 (44.9 ± 5.97) 121.0–140.0 (133.0 ± 8.29) 68.0–100.9 (88.2 ± 11.76)
HL 12.7–18.2 (16.0 ± 2.11 9.9–13.5 (12.0 ± 4.558) 17.8–18.7 (18.3 ± 0.39) 9.5–04.7 (11.9 ± 1.60)
HW 7.7–12.0 (9.3 ± 1.97) 5.8–7.9 (6.9 ± 1.08) 11.1–11.8 (11.5 ± 0.31) 5.6–8.5 (7.2 ± 1.07)
HH 6.4–9.7 (7.8 ± 1.39) 5.4–6.8 (6.2 ± 0.72) 9.1–9.4 (9.3 ± 0.13) 5.8–6.8 (6.4 ± 0.41)
ED 4.9–6.0 (5.2 ± 0.50) 3.0–4.6 (4.0 ± 0.90) 2.9–3.8 (3.2 ± 0.42) 2.3–2.7 (2.6 ± 0.16)
SE 6.3–8.2 (7.0 ± 0.85) 4.6–6.0 (5.5 ± 0.73) 6.1–8.4 (8.3 ± 0.12) 5.4–6.0 (5.6 ± 0.25)
Ve eb a e Zoology 75, 2025, 627–672 651
Dis ibu ion. Known om no heas e n Namibia and
adjacen Bo swana, and om wes e n Zambia (Fig. 13).
The appa en gap in dis ibu ion be ween no heas e n
Namibia/ Bo swana and wes e n Zambia likely e lec s a
lack o sampling, and he species’ ange is belie ed o be
mo e con inuous.
Habi a and Na u al His o y. On ogene ic colou di e -
ences ha e been obse ed be ween ju eniles and adul s
( his s udy). Found in sympa y wi h I. capensis sensu
la o. Associa ed wi h Baikiaea woodland on deep Kala-
ha i allu ial sands and ha d lime- ich soils in open wood-
land (Haacke 1970; Pie e sen e al. 2021).
Ichno opis obus a sp. no .
Robus ough-scaled liza d
h ps://zoobank.o g/1E069D01-1F86-488F-9025-DEBDB-
47BEA00
Figu es 17, 18; Tables 4, 5
Ch esonymy.
Ichno opis c . g andiceps – Con adie e al. (2022a: 198); Ichno o-
pis a . g andiceps – Beni o e al. (2025: 893).
Holo ype. PEM R23420 ( ield numbe WC-4816), adul
male, collec ed om Cuando Ri e sou ce (–13.0035°,
19.1275°, 1343 m a.s.l.), Moxico P o ince, Angola by
We ne Con adie and James Ha ey on 21 No embe
2016.
Pa a ypes. 4 specimens: a) PEM R23361 ( ield numbe
WC-4063) and PEM R23362 ( ield numbe WC-4056),
adul emales, collec ed on he oad be ween Cuana ale
Ri e sou ce camp and Samanunga illage (–13.0380°,
18.8298°, 1605 m a.s.l.), Moxico P o ince, Angola by
We ne Con adie and Luke Ve bu g on 13 Ma ch 2016;
b) PEM R23421, adul male, same collec ion de ails as
holo ype; c) PEM R23482 ( ield numbe WC-4804),
adul male, collec ed om Cuando Ri e sou ce, ap 4
(–13.0016°, 19.1296°, 1372 m a.s.l.), Moxico P o ince,
Angola by We ne Con adie and James Ha ey on 15 No-
embe 2016.
Addi ional ju enile ma e ial. 12 specimens: a) PEM
R23279–80; INBAC (no numbe ), collec ed om Cua-
na ale Ri e sou ce lake (–13.0933°, 18.8940°, 1367 m
a.s.l.), Moxico P o ince, Angola by We ne Con adie on
1 Ma ch 2016; b) PEM R23299–300, g assland wes o
Cuana ale Ri e sou ce en ou e o Samanunga illage
(–13.0751°, 18.8848°, 1366 m a.s.l.), Moxico P o ince,
Angola by We ne Con adie and Luke Ve bu g on 16
Ma ch 2016; c) PEM R23303–9, ap 4 km ups eam
om Cuana ale Ri e sou ce lake (–13.0508°, 18.8973°,
1380 m a.s.l.), Moxico P o ince, Angola by We ne Con-
adie om 28 Feb ua y o 15 Ma ch 2016.
E ymology. The species name obus a is he eminine
o m o he La in adjec i e obus us, meaning ‘ obus ’ o
‘s u dy’, in e e ence o he la ge, hea y-buil adul s o
his species.
Diagnosis. Assigned o Ichno opis due o he absence o
a well-de ined colla , digi s no se a ed o inged, sub-
digi al lamellae keeled, and subocula bo de ing he lip.
A la ge Ichno opis wi h a single on onasal; subocula
bo de ing he lip; a single an e io lo eal; eebly de el-
oped head shield s ia ions; p e on als well sepa a ed
om he an e io sup aocula ; and sup acilia ies sepa a -
ed om he sup aocula s by a se ies o smalle scales.
The new species can be dis inguished om o he Ich-
no opis species based on a combina ion o he ollowing
cha ac e s: P e on als well sepa a ed om he an e io
sup aocula ( e sus mos ly in con ac in I. bi i a a, I. mi-
c olepido a and I. anganicana); high numbe o midbody
scale ows (43–48 e sus 25–42 in I. capensis sensu la o);
la ge, obus head and ounded snou ( e sus small de-
p essed head and poin ed snou in I. capensis sensu la o);
ou (46%) o i e (50%) sup alabials an e io o he sub-
Cha ac e s I. g andiceps I. g andiceps I. obus a sp. no . I. obus a sp. no .
LL 14.3–18.3 (16.1 ± 1.99) 10.5–15.0 (13.3 ± 1.96) 20.5–24.2 (21.6 ± 1.53) 1.1–17.0 (13.8 ± 1.90)
IL 24.8–38.9 (30.4 ± 5.99) 19.4–38.9 (26.1 ± 6.9) 33.4–40.1 (36.3 ± 2.38) 18.2–26.0 (22.0 ± 2.77)
FLL 6.9–9.4 (7.8 ± 1.11) 4.2–6.5 (5.3 ± 1.15) 7.2–8.6 (8.3 ± 0.62) 3.8–6.4 (5.1 ± 0.89)
HLL 10.6–13.7 (11.8 ± 1.35) 7.0–9.0 (8.3 ± 1.13) 12.8– 13.9 (13.4 ± 0.44) 6.5– 10.5 (8.7 ± 1.35)
TAIL/SVL 1.3–2.2 (1.9 ± 0.34) 2.0 (2.0 ± 0.02) 1.6–1.9 (1.8 ± 0.17) 1.8–2.1 (2.0 ± 0.09)
HL/SVL 0.2–0.3 (0.2 ± 0.01) 0.3 (0.3 ± 0.01) 0.2–0.3 (0.2 ± 0.01) 0.3 (0.3 ± 0.01)
ES/HL 0.4–0.5 (0.5 ± 0.03) 0.4–0.5 (0.5 ± 0.02) 0.4–0.5 (0.5 ± 0.01) 0.4–0.5 (0.4 ± 0.00)
HW/HL 0.5–0.7 (0.6 ± 0.05) 0.5–0.6 (0.6 ± 0.04) 0.6–0.7 (0.6 ± 0.02) 0.6–0.7 (0.6 ± 0.03)
MSR 44–47 43–48
LVSR 10 9–10
TVSR 27–31 26–33
SL 4–5 (mos ly 5) 4–6 (mos ly 5)
IL 5–7 (mos ly 6) 5–7 (mos ly 6)
SC 4–5 (mos ly 5) 4–5 (mos ly 5)
LUFT 20–26 20–26
Femo al po es 8–14 8–14
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
658
an e io sup aocula (only in con ac in 15 ou o 245
specimens examined) and sepa a ed om sup acilia ies
by a smalle scale; wo la ge sup aocula s, which a e sep-
a a ed om he sup acilia ies by one ( e y a ely wo)
ow o small scales (4–9) and p eceded by a clus e o
3–10 smalle scales; wo lo eal scales p esen , which a e
sepa a ed om he an e io sup aocula by 2–3 scales;
1–2 pos -sup aocula s; subocula in con ac wi h he lip;
3–6 (mos ly ou ) sup alabials in on o he subocula ;
5–8 (mos ly six) in alabials; i e chin shields, wi h he
an e io h ee in b oad con ac ; 3–5 (mos ly ou ) su-
p acilia ies; 25–42 (a e age: 36.7) midbody scale ows;
8–10 (a e age: 8.8) longi udinal ows o enla ged en-
al pla es; 20–31 (a e age: 25.8) ans e se en al scale
ows; 16–26 (a e age: 21.6) subdigi al lamellae unde
he 4 h oe; 6–15 emo al po es pe high. Size: Adul
specimens a ied om 40.0–67.8 mm (mean: 54.6 mm)
SVL and 69.5–149.0 mm (mean: 110.4 mm) TAIL. La g-
es emale: 65 mm SVL (NMZB-UM 9228 – Um ali,
Zimbabwe); la ges male: 67.8 mm SVL (BE_RMCA_
Ve .R.7785 – Dilolo, DRC). Colou a ion (Fig. 20): In
males, he lanks ea u e a s iking, b oad black longi u-
dinal band ha o igina es a he ip o he snou , passes
h ough he eye, and ex ends pos e io ly well beyond he
hind limbs, g adually ading owa ds he ip o he ail.
This black band is bo de ed by wo dis inc whi e s ipes:
The uppe s ipe begins jus behind he eye, while he
lowe s ipe o igina es a he os al pla e, c osses he
ympanum, and uns pa allel o he black band along he
leng h o he body. Below he lowe whi e s ipe, a i id
eddish-o ange s ipe is especially p ominen on he an-
e io lanks. A seconda y sho black line also o igina es
a he snou , uns ac oss he sup alabials along he side
o he head, and e mina es an e io o he inse ion o
he o elimbs. The main black band on he lank is o en
sca e ed wi h small whi e spo s, pa icula ly owa ds he
pos e io end o he body. In b eeding males, he whi e
s ipes on he head and neck, as well as he gula egion,
become in used wi h a b igh yellow hue. The do sal su -
ace is a ich eddish-b own, ado ned wi h sca e ed da k
b own speckling. Females exhibi a mo e subdued co-
lou a ion, wi h an o e all g ey-b own one ha is ligh e
on he en al side. A single, less p onounced da k black
s ipe o igina es a he snou , passes h ough he eye, and
con inues along he lanks, g adually ading owa ds he
ail. Ju eniles and subadul s a e o en g ey in colou a-
ion wi h a whi e do sola e al s ipe. The en e is mos ly
whi e, bu some specimens exhibi g ey colou a ion wi h
sca e ed black specks.
Dis ibu ion. Widesp ead, occu ing ac oss se e al
coun ies in sou he n A ica, including Angola, Namibia,
Bo swana, Zambia, Zimbabwe, Mozambique, and pa s
o Sou h A ica and Malawi (Fig. 19). His o ical eco ds
om no h-eas e n Angola and DRC assigned o I. capen-
sis o I. o e lae i need o be e-e alua ed in ligh o his
s udy and migh be assignable o ei he I. anganicana,
I. bi i a a o I. longico pa sp. no . (see new species de-
sc ip ion below).
Habi a and Na u al His o y. This species p e e s a id
o mesic sa anna habi a s. I is a diu nal liza d, ac i ely
Figu e 23. Syn ype (BMNH 1946.8.4.23) o Ichno opis longipes om Mazoë, Zimbabwe. Pho og aphs o body in A en al and
B do sal iews, and head in C do sal, D la e al and E en al iews. Scale ba s ep esen 10 mm. Pho og aphs: Pa ick Campbell.

Ve eb a e Zoology 75, 2025, 627–672 659
o aging o small in e eb a es such as e mi es, spide s,
bee les, and g asshoppe s. Females lay up o nine eggs
pe clu ch, ypically du ing he summe mon hs om
Oc obe o No embe . The eggs measu e app oxima ely
5.5–7.0 mm by 8.5–9.5 mm. The incuba ion pe iod ang-
es om 56 o 77 days, wi h ha chlings eme ging be ween
Janua y and Ma ch. Females may p oduce up o wo
clu ches wi hin a single b eeding season.
Ichno opis longico pa sp. no .
Long-bodied ough-scaled liza d
h ps://zoobank.o g/2D3A7A69-E837-4542-89C3-D4C-
C18A2A160
Figu es 24, 25; Tables 6, 7
Ch esonymy.
Ichno opis capensis o e lae i – Lau en (1950: 12, in pa ); Ichno -
opis capensis – Con adie e al. (2022a: 198, in pa ); Ichno opis a .
capensis – Beni o e al. (2025: 893).
Holo ype. PEM R23410 ( ield numbe WC-4558), adul
male, collec ed om Lungwebungu Ri e camp b idge
c ossing, (–12.5835°, 18.6660°, 1304 m a.s.l.), Moxico
P o ince, Angola by We ne Con adie and Luke Ve bu g
on 22 Oc obe 2016.
Pa a ypes. 6 specimens: a) PEM R23409 ( ield numbe
WC-4557), adul male, same collec ion de ails as ho-
lo ype; b) PEM R23502 ( ield numbe WC-4522), and
PEM R23531 ( ield numbe WC-4560), adul male and
emale espec i ely, collec ed om Sombanana Village
(–12.3108°, 18.6239°, 1403 m a.s.l.), Moxico P o ince,
Angola by We ne Con adie and Luke Ve bu g on 9 Oc-
obe 2016; c) PEM R23505–7 ( ield numbe WC-4543,
WC-4562 and WC-4563, espec i ely), adul males, col-
lec ed om Lake Tchanssengwe (–12.4102°, 18.6348°,
1414 m a.s.l.), Moxico P o ince, Angola by We ne Con-
adie and Luke Ve bu g on 21 Oc obe 2016.
Addi ional ma e ial. 9 specimens: a) PEM R19903 ( ield
numbe TB 44) and PEM R19905 ( ield numbe TB 46),
adul emale and male espec i ely, collec ed om Camp
Chi i, Miombo o es /camp (–9.3969°, 20.4319°, 1004 m
a.s.l.), Lunda-Sul P o ince, Angola by Tom B anch on
24 Oc obe 2008; b) PEM R23977 ( ield numbe WC-
6267), ju enile, collec ed om nea Lungwebungu T ap
2 (–12.5820°, 18.6656°, 1208 m a.s.l.), Moxico P o ince,
Angola by We ne Con adie and Alex Rebelo on 22 Ap il
2018; c) PEM R23986 ( ield numbe WC-6266), ju e-
nile, collec ed om Lungwebungu T ap 1 (–12.5801°,
18.6674°, 1298 m a.s.l.), Moxico P o ince, Angola by
We ne Con adie and Alex Rebelo on 22 Ap il 2018; d)
PEM R23996–7 ( ield numbe s WC-6291 and WC-6292,
espec i ely), ju eniles, collec ed om Lake Tchanssen-
gwe (–12.4140°, 18.6442°, 1393 m a.s.l.), Moxico P o -
Table 6. Summa y o mo phological da a o he Ichno opis capensis g oup. Measu emen s a e in millime es (mm). Values a e
gi en as a ange wi h mean ± s anda d de ia ion in pa en hesis. Ju eniles we e excluded om he measu emen s, bu we e included
in he scala ion da a. Fo abb e ia ions, see he Ma e ials and Me hods sec ion. n = sample size.
Cha ac e s I. capensis sensu la o I. longico pa sp. no .
n = 256 n = 12
SVL 40.0–67.8 (54.8 ± 5.11) 62.7–71.2 (66.4 ± 2.37)
TAIL 69.5–149.0 (110.4 ± 17.89) 117–160 (137.5 ± 15.94)
HL 10.1–14.7 (12.7 ± 0.97) 13.1–17.2 (15.1 ± 1.16)
HW 5.6–8.6 (7.2 ± 0.67) 7.8–9.4 (8.7 ± 0.46)
HH 4.5–7.5 (5.9 ± 0.62) 6.1–8.5 (7.4 ± 0.61)
ED 3.9–6.3 (4.5 ± 0.52) 2.4–5.9 (4.4 ± 0.98)
SE 4.0–7.0 (5.2 ± 0.77) 6.0–7.3 (6.9 ± 0.40)
LL 12.5–16.4 (14.2 ± 1.18) 14.4–18.7 (17.8 ± 1.27)
IL 18.5–31.6 (25.5 ± 3.29) 27.4–31.6 (28.7 ± 1.42)
FLL 5.0–8.2 (6.2 ± 0.70) 6.8–8.4 (7.5 ± 0.46)
HLL 7.4–13.2 (10.5 ± 1.25) 10.6–13.5 (12.3 ± 0.96)
TAIL/SVL 1.2–2.7 (2.0 ± 0.28) 1.9–2.4 (2.2 ± 0.22)
HL/SVL 0.2–0.3 (0.2 ± 0.01) 0.2–0.3 (0.2 ± 0.01)
ES/HL 0.4–0.5 (0.5 ± 0.02) 0.4–0.5 (0.4 ± 0.02)
HW/HL 0.2–0.7 (0.6 ± 0.06) 0.5–0.6 (0.6 ± 0.03)
MSR 25–42 34–41
LVSR 8–10 9–10
TVSR 20–31 25–31
SL 3–6 (mos ly 4) 4–5 (mos ly 4)
IL 5–8 (mos ly 6) 6–7 (mos ly 6)
SC 3–5 4
LUFT 16–26 19–24
Femo al po es 6–15 10–13
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
660
ince, Angola by We ne Con adie and Alex Rebelo on 23
Ap il 2018; e) BE_RMCA_Ve .R.17490, 17492 (adul
males) and BE_RMCA_Ve .R.17491 (adul emale),
collec ed om he Dundo egion, Lunda-No e P o ince,
Angola by Ba os Machado on 14 Decembe 1947.
E ymology. The species name longico pa is he eminine
o m o he La in adjec i e longico pus, de i ed om
longus (long) and co pus (body), e e ing o his species’
elonga e body.
Diagnosis. Assigned o Ichno opis due o he absence o
a well-de ined colla , digi s no se a ed o inged, sub-
digi al lamellae keeled, and subocula bo de ing he lip.
A slende Ichno opis wi h a single on onasal; subocu-
la bo de ing he lip; a single an e io lo eal; eebly de-
eloped head shield s ia ions, p e on als well sepa a ed
om he an e io sup aocula ; and sup acilia ies sepa a -
ed om he sup aocula s by a se ies o smalle scales.
The new species can be dis inguished om o he Ich-
no opis species based on a combina ion o he ollowing
Figu e 24. Pho og aphs in li e o Ichno opis longico pa sp. no . Pho og aphs o A adul male (PEM R23410) om Lungwebungu
Ri e camp b idge c ossing, Angola, and B adul emale (PEM R23531) om Sombanana illage, Angola. Pho og aphs: We ne
Con adie.
Figu e 25. Holo ype (PEM R23410) o Ichno opis longico pa sp. no . om Lungwebungu Ri e camp b idge c ossing, Moxico
P o ince, Angola. Pho og aphs o body in A en al and B do sal iews, and head in C do sal, D la e al and E en al iews. Scale
ba s ep esen 10 mm. Pho og aphs: We ne Con adie.
Ve eb a e Zoology 75, 2025, 627–672 661
cha ac e is ics: P e on als well sepa a ed om he an e-
io sup aocula ( e sus mos ly in con ac in I. bi i a-
a, I. mic olepido a and I. anganicana); lowe numbe
(34–41) o midbody scales ows (44–47 in I. g andiceps
and 43–48 in I. obus a sp. no .); small, dep essed head
and poin ed snou ( e sus la ge obus head and ounded
snou in I. g andiceps and I. obus a sp. no .); ou su-
p alabials an e io o he subocula ( e sus mos ly i e in
I. g andiceps and I. obus a sp. no .); dis inc i e occip-
i al scale usually ex ending pos e io ly well beyond he
le el o he pa ie als ( e sus la ge apezi o m occipi al
wedged be ween he pa ie als, no p o uding pas pa i-
e als in I. g andiceps and I. obus a sp. no .).
The new species esembles I. capensis sensu la o in
i s na ow, poin ed snou , wi h he p e on als well sep-
a a ed om he an e io subocula . I di e s in ha he
new species exhibi s black spo s on he chin shields and
gula scales ( e sus immacula e in I. capensis sensu la o)
and he absence o a clea uppe whi e do sola e al s ipe
ha sepa a es he da k black la e al band om he do sal
b own e eb al band ( e sus p esen in mos I. capensis
sensu la o).
In he phylogene ic analysis, he unco ec ed p dis-
ances show ha he new species di e s by >5.9% o
16S and >12.3% o ND4 sequence di e gence om o h-
e Ichno opis species (Table 2).
Holo ype desc ip ion (Fig. 25). Adul male measu ing
67.7 mm SVL and 160 mm TAIL (2.4 × SVL). Body
mode a ely dep essed; head dis inc ly dep essed, almos
wice as long as b oad (HL 15.8/HW 8.5 mm), i s leng h
equi alen o 23.3% o SVL, expanded in he empo al
egion and e y dis inc om he neck. Adp essed hind
limb jus eaching he an e io edge o ea opening. The
oo leng h is longe han he head leng h (FL 19.2/HL
15.8 mm).
Do sal head shields e y eebly s ia ed and keeled;
nos il pie ced be ween h ee nasals, he sup anasals in
b oad con ac behind he os al; on onasal as long as
b oad (2.2 × 2.2 mm); p e on als much longe han b oad
(2.7 × 1.5 mm), in b oad con ac medially, no eaching
he an e io sup aocula s (sepa a ed by a small keeled
scale), in con ac wi h he an e io and pos e io lo eal;
on al mo e han wice as long as i s maximum wid h be-
ween he pos e io ips o he p e on als (4.6 × 2.1 mm),
ounded an e io ly and s ongly na owed pos e io ly;
pai ed on opa ie als longe han b oad (3.0 × 2.0 mm);
pa ie als longe han b oad (3.8 × 2.6 mm), ex ending
pos e io ly, widely sepa a ed by a la ge in e pa ie al and
occipi al, he pos e io ma gin ex ending pas he pos e-
io bo de s o he pa ie als; h ee keeled empo al scales
bo de ing he pa ie al, he i s one longes , ollowed by
he second and hi d (smalles ); wo enla ged sup aocu-
la s, he an e io sup aocula sligh ly longe han he pos-
e io one and longe han i s dis ance om he pos e io
lo eal (2.4 mm s. 1.5 mm), in con ac wi h he pos e io
hal o he on al, sepa a ed om he pos e io lo eal by
wo smalle keeled scales; he an e io sup aocula s a e
p eceded by a clus e o i e smalle keeled scales, he
pos e io sup aocula is ollowed by h ee smalle keeled
Table 7. Measu emen s (in mm) and scale coun s o he ype se ies o Ichno opis longico pa sp. no . Scale coun s a e gi en as
Righ /Le . Fo abb e ia ions, see he Ma e ials and Me hods sec ion. = unca ed.
Ca alogue No. PEM R23410 PEM R23409 PEM R23502 PEM R23505 PEM R23506 PEM R23507 PEM R23531
Type S a us Holo ype Pa a ype Pa a ype Pa a ype Pa a ype Pa a ype Pa a ype
Sex Male Male Male Male Male Male Female
SVL 67.7 63.0 64.3 63.9 65.4 66.2 63.1
TAIL 160.0 146.0 149.0 92 127.0 148.0 117.0
HL 15.6 15.8 15.6 14.7 15.3 15.8 13.1
HW 8.5 8.6 9.4 8.9 8.9 9.2 7.8
HH 7.1 7.3 7.5 7.3 7.1 7.5 6.1
ED 3.1 3.6 3.6 2.4 2.8 3.8 3.0
SE 6.6 6.6 6.7 6.7 6.6 7.2 6.1
LL 17.6 18.0 18.1 16.8 17.3 18.7 15.2
IL 30.8 31.4 29.0 28.9 31.2 29.5 31.6
FL L 7.4 7.8 8.2 7.0 7.5 7.3 6.8
HLL 13.5 13.1 12.6 13.3 13.5 13.2 11.5
TAIL/SVL 2.4 2.3 2.3 1.9 2.2 1.9
HL/SVL 0.2 0.3 0.2 0.2 0.2 0.2 0.2
ES/HL 0.4 0.4 0.4 0.5 0.4 0.5 0.5
HW/HL 0.5 0.5 0.6 0.6 0.6 0.6 0.6
MSR 39 37 38 40 37 40 37
LVSR 9 9 9 9 9 10 9
TVSR 30 28 29 31 28 30 28
SL 4/4 4/4 4/4 4/5 4/4 4/4 4/5
IL 6/6 7/6 6/6 7/7 6/6 6/6 6/6
SC 4/4 4/4 4/4 4/4 4/4 4/4 4/4
LUFT 22 20 21 22 24 22 22
Femo al po es 12/10 10/10 12/12 12/11 13/12 11/10 10/10
Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
662
pos -sup aocula scales, he wo sup aocula s a e sepa a -
ed om he sup acilia ies by a single ow o nine small
keeled scales. Fi e sup acilia ies, he i s wo much lon-
ge han he o he s and o ming a long oblique su u e.
Lowe nasal in con ac wi h he os al, i s sup alabial,
and an e io lo eal (na ow con ac on le side); pos na-
sal small, in con ac wi h he o he wo nasals, an e io
lo eal, and on onasal. Two lo eals, he pos e io one
much la ge and di ided below; ou sup alabials an e io
o he subocula , whose lowe bo de on he lip is much
sho e (3×) han he uppe bo de ; h ee sup alabials
pos e io o subocula ; empo al scales s ongly keeled;
a na ow ympanic shield on he uppe an e io co ne o
he e ically elonga e ea opening. Lowe eyelid scaly
wi h a median se ies (4–5) o e ically elonga e scales.
Six in alabials; i e pai s o la ge chin shields, he i s
h ee pai s in median con ac ; gula scales imb ica e; no
colla .
Do sal scales hombic, s ongly keeled and imb ica e;
la e als smalle and eebly keeled, passing g adually in o
he smoo h, ounded en al pla es, which a e b oad-
e han long; 39 scales a ound he middle o he body;
en al pla es in nine longi udinal and 30 ans e se ows
be ween he o e- and hind limbs; p eanal scales i egu-
la ; scales on uppe su aces o limbs hombic, s ongly
keeled, and imb ica e; 12/10 emo al po es on each side;
subdigi al lamellae plu ica ina e and spinulose, 22 unde
he 4 h oe; caudal scales s ongly keeled abo e and below,
excep hose jus pos e io o he en , which a e smoo h.
Colou a ion. (In li e, b eeding colou a ion; Fig. 24A):
The do sum a ies om g ey on he head and nape o ed-
dish-b own on he do sum and g ey on he ail. The side
o he body has a da k black band ha o igina es on he
snou and un pos e io ly o he ail, whe e i disappea s
a he ip. Below his black band is a whi e s ipe ha
o igina es on he snou , uns below he eye o he on
limb, is less dis inc be ween he o e- and hind limb, and
hen ades on o he ail. Below his whi e line is ano he
black s ipe ha o igina es on he snou , unning along
he edges o he sup a- and in alabials o he on limbs.
The whi e s ipes on he sides o he head and he gula
egion a e pale yellow. Below he black band and whi e
lank s ipe ( ha appea s as sca e ed whi e and g ey spo s
in places) is an o ange band. Limbs a e b ick ed and
g ey. Chin shields and gula scales ha e sca e ed black
blo ches o a ying sizes. The en e is whi e wi h sca -
e ed black specks. Colou a ion (in p ese a i e; Fig. 25):
Abo e pale g ey-b own; a well-de ined b oad (co e ing
3–4 scales a midbody) black do sola e al band ex ends
om he ip o he snou o he g oin; below his black
do sola e al band is a na ow whi e band (co e ing one
scale a midbody) which ex ends om he ip o he snou
o jus pos e io o he on limb, ading owa ds he g oin
and ail base. Below his whi e band is ano he na ow
black band ex ending om he men al, along he edge o
he jaw (edge o sup a- and in alabials) o jus pos e io
o he o elimb inse ion. Flanks bea a ligh b own band
( wo scales wide); gula and chin shields wi h sca e ed
black spo s o blo ches; limbs do sally b own and en al-
ly whi e; 2–3 whi e spo s on he an e io su aces o he
legs; en e whi e wi h sca e ed black specks.
Pa a ype and addi ional ma e ial a ia ion. The pa a-
ypes a e in ag eemen wi h he holo ype in scala ion,
wi h only mino a ia ion: P e on al always sepa a e
om he an e io sup aocula by one scale (excep PEM
R19905 on igh side); on onasal always sepa a e om
he 1s sup acilia ies (excep in PEM R23409 and PEM
R23506); wo ( a ely h ee) scales sepa a ing an e io su-
p aocula om he pos e io lo eal; clus e o 3–9 scales
in on o he an e io sup aocula ; single ow o 6–9
scales sepa a ing he sup aocula s om he sup acilia ies;
1–4 pos -sup aocula s; ou sup acilia ies; 4–5 sup alabi-
als; 6–7 in alabials; i e chin shields, wi h i s h ee in
con ac (PEM R23505 has six chin shields, wi h he i s
ou in con ac on he le side); 9–10 ans e se en al
pla es; 25–31 longi udinal en al pla es; 34–41 midbody
scale ows; 19–24 subdigi al lamellae unde he 4 h oe;
10–13 emo al po es on each high. PEM R23409 exhib-
i s some abe an head scala ion in ha he an e io lo eal
seems o be di ided, o ming a sup alo eal ha sepa a es
he an e io lo eal om he on onasal and he pa ie al,
and he pos e io lo eal is di ided in o wo scales. Size:
Adul specimens a ied om 62.7–71.2 mm (mean: 65.3
mm) SVL and 117.0–160.0 mm (mean: 141.2 mm) TAIL.
La ges emale: 65.1 mm SVL (PEM R19905 – Camp
Chi i, Angola); la ges male: 71.2 mm SVL (BE_RMCA_
Ve .R.17492 – Dundo, Angola). Colou a ion o all males
a e in ag eemen wi h he holo ype. The pa a ype emale
(PEM R23531; Fig. 24B) is dulle in colou a ion, almos
uni o mly eddish-b own do sally and g ey la e ally, wi h
no whi e s ipes o black bands.
Dis ibu ion. Only eco ded om he headwa e s o he
Lungwebungu and Cuando Ri e s in cen al Angola,
no hwa ds o he DRC bo de (Fig. 19). Some speci-
mens om Mabwe Ri e , Upemba Na ional Pa k, DRC
(IRSNB 7895, 7897, 7907–9, 78728), exhibi he same
dis inc da k b own o black gula ma kings and migh
be assigned o his species. I con i med, his new species
could be mo e widely dis ibu ed han cu en ly hough .
Habi a and na u al his o y. This species was no ound
o be sympa ic wi h any o he Ichno opis species, bu i
occu s in close geog aphical p oximi y o I. capensis sen-
su la o and I. obus a sp. no . This species is associa ed
wi h we Miombo woodland.
Ve eb a e Zoology 75, 2025, 627–672 663
Key o he genus Ichno opis Pe e s, 1854
1a Snou dep essed and poin ed, p e on al sepa a ed om an e io sup aocula by one o wo smalle keeled scales ...2
1b Snou obus and ounded, p e on al mos ly in con ac wi h an e io sup aocula ...................5 (I. bi i a a g oup)
2a 43–47 midbody scale ows, body obus , head b oad, adul SVL > 70 mm ......................... 3 (I. g andiceps g oup)
2b 25–42 midbody scale ows, body slende , head na ow, adul SVL < 65 mm ............................. (I. capensis g oup)
3a Con ined o he d ie Zambezian Baikiaea woodlands o Comb e um-Vachellia bush eld in wes e n Zambia and he
Zambezi Region o Namibia and adjacen Bo swana; occu s below 1000 m a.s.l ................................I. g andiceps
3b Con ined o he Angolan Pla eau, which consis s o mois e Angolan Miombo woodland; occu s abo e 1300 m
a.s.l. ................................................................................................................................................I. obus a sp. no .
4a Long, slende body and head; da k black spo s/blo ches on chin shields and gula scales, no clea uppe whi e do -
sola e al line sepa a ing do sola e al black band om do sal b own e eb al band ............... I. longico pa sp. no .
4b Sho body and head; no da k black spo s/blo ches on chin shields o gula scales, clea whi e uppe do sola e al
line sepa a ing black do sola e al band om do sal b own e eb al band ............................. I. capensis sensu la o
5a ≥ 42 (42–50) midbody scale ows, known only om M Moco, Angola .......................................... I. mic olepido a
5b ≤ 42 (28–42) midbody scale ows ..............................................................................................................................6
6a P esence o do sola e al blue o yellow spo s in li e ..................................................................................................7
6b No do sola e al blue o yellow spo s, only known om no h-eas e n DRC ...............................................I. chapini
7a P e on als mos ly sepa a ed om he an e io sup aocula , which is o en in con ac wi h he 1s sup acilia y; e enly
spaced blue do sola e al spo s; occu s in sou he n DRC, Zambia, Malawi and Tanzania .................. I. anganicana
7b P e on al mos ly in con ac wi h an e io sup aocula , which is always in con ac wi h he 1s sup acilia y; closely
spaced yellow spo s abo e a m; occu s in Angola, eas e n DRC, Republic o he Congo and Gabon ....... I. bi i a a
Discussion
Al hough Ichno opis species a e locally abundan and
widely dis ibu ed ac oss sub-Saha an A ica, excluding
Wes A ica, he genus emains among he mos axonom-
ically neglec ed o A ican lace ids. This his o ical o e -
sigh is likely d i en by a combina ion o s ong seasonal
ac i i y pa e ns, which limi de ec abili y ou side o peak
ep oduc i e pe iods (B oadley 1967a, 1974, 1979; Ja-
cobsen 1987; his s udy), and a legacy o axonomic con-
usion da ing back o e a cen u y (Boulenge 1921; Lau-
en 1952, 1964; an den Be g 2017; Beni o e al. 2025).
As a esul , specimen collec ions ha e emained spa se in
many egions, and he g oup’s ue species ichness has
long been unde es ima ed.
Th ough expanded geog aphic sampling, co e ing
much o he known anges o I. bi i a a and I. capensis,
and he gene a ion o he i s gene ic da a o I. angan-
icana and I. g andiceps, we p o ided he mos comp e-
hensi e and geog aphically inclusi e phylogene ic ame-
wo k o Ichno opis species o da e. Species delimi a ion
analyses, combined wi h a subs an ially enhanced mo -
phological da ase , suppo he alidi y o mos cu en ly
ecognised species (wi h he excep ion ha we could no
alida e he phylogene ic s a us o I. chapini) and jus i-
y he o mal desc ip ion o wo new Angolan endemics:
Ichno opis obus a sp. no . and I. longico pa sp. no .
These indings unde sco e he con inued impo ance o
in eg a ing molecula and mo phological app oaches in
axonomic e isions o mo phologically conse a i e lin-
eages.
Con e sely, ou phylogene ic amewo k also acili a -
ed he eassessmen o se e al axonomically ambiguous
species and subspecies, many o which had no p e i-
ously been e alua ed using phylogene ic analyses. Fo
ins ance, al hough I. b. pallida exhibi s no able gene ic
di e gence om o he I. bi i a a popula ions, we con-
se a i ely synonymise he subspecies wi h he nominal
o m un il u he da a become a ailable. Fu he mo e,
based on ou indings, we ecommend e aining I. lon-
gipes and I. mac olepido a as synonyms o I. capensis
sensu la o, due o he absence o consis en diagnos ic
mo phological di e ences and inclusion o opo ypic
ma e ial in ou phylogene ic analyses. Addi ionally, we
p opose synonymising I. c. nig escens wi h I. bi i a a,
and ea ing I. o e lae i as a junio synonym o I. an-
ganicana, based on mo phological cong uence. Newly
collec ed Ichno opis ma e ial om wes o he Kabobo
Pla eau and Upemba Na ional Pa k in DRC allowed us
o eassess he poo ly known I. anganicana, p e ious-
ly known only om i s ype specimen and a ague ype
locali y (Boulenge 1917; Mei i e al. 2018). Ou da a
con i m he species’ axonomic alidi y, help e ise he
species desc ip ion and sugges a b oade geog aphic
dis ibu ion han p e iously ecognised. As a esul , mos
his o ical eco ds o I. bi i a a om Eas A ica a e he e
eassigned o I. anganicana.
While he e was a mo e comp ehensi e sampling o
I. capensis ela i e o I. bi i a a, he limi ed ma e ial
a ailable o I. bi i a a (n = 4, including opo ypic I. b.
pallida) exhibi ed mode a e gene ic di e gence, and spe-
cies delimi a ion analyses consis en ly iden i ied hese
lineages as dis inc . Gi en he b oad unsampled dis i-
bu ion o I. bi i a a, coupled wi h he high gene ic di-
e si y obse ed wi hin he a ailable da ase , inc eased
geog aphic sampling, especially wi hin opog aphically

Con adie W e al.: Sys ema ics o A ican ough-scaled liza ds
664
complex Angola, DRC, Republic o he Congo and Ga-
bon, may e eal addi ional c yp ic di e si y. Mo eo e ,
ou species delimi a ion analyses eco e ed mul iple pu-
a i e candida e species wi hin he I. capensis sensu la o
complex. I mus be no ed ha some o hese lineages
may co espond o p e iously desc ibed species, such as
I. longipes, bu esol ing his will equi e a ge ed sam-
pling om ype locali ies o museomics (i.e., his o ical
DNA [hDNA]) o assess po en ial synonymy o e alida-
ion (McGui e e al. 2018; Raxwo hy and Smi h 2021;
Zacho e al. 2021; Lalueza-Fox 2022; Le sch e al. 2025).
Al hough we we e unable o assess he phylogene ic
placemen o I. chapini due o he absence o gene ic da a,
we ecommend i s p o isional assignmen o he I. bi i -
a a g oup based on simila i ies in head mo phology and
scala ion. Un il u he ma e ial becomes a ailable o
molecula analysis, we p opose e aining I. chapini as a
alid species.
The species delimi a ion analyses con i med he spe-
cies s a us o bo h newly desc ibed and p e iously ec-
ognised axa, less conse a i e me hods also e ealed
po en ial c yp ic di e si y wi hin I. anganicana, I. bi i -
a a, I. longico pa sp. no ., and I. capensis sensu la o.
We cau ion ha single-locus app oaches can o e es ima e
species bounda ies by con la ing in aspeci ic a ia ion
wi h in e speci ic di e gence, especially unde condi ions
o incomple e lineage so ing o limi ed geog aphic sam-
pling (e.g., Ca s ens e al. 2013; Sukuma an and Knowles
2017). To mi iga e his isk, we employed i e indepen-
den species delimi a ion me hods, enabling c oss- ali-
da ion and allowing us o conse a i ely in e p e only
hose lineages suppo ed by mul iple lines o e idence
(mo phology, colou a ion, ecology) as candida e species
(e.g., Fujisawa and Ba aclough 2013; Zhang e al. 2013;
Kapli e al. 2016).
Mo phological analyses b oadly suppo he h ee ma-
jo clades eco e ed by he phylogene ic analyses, wi h
diagnos ic di e ences in head shape, scala ion, and b eed-
ing colou a ion. These ai s a e la gely consis en ac oss
mul iple popula ions, making hem aluable o species
iden i ica ion in he ield. The I. capensis, I. g andiceps
and I. bi i a a g oups exhibi dis inc i e mo pholog-
ical cha ac e is ics ha aid in dis inguishing hem om
each ano he , e en in he absence o gene ic da a. How-
e e , I. anganicana is an excep ion: I o ms a deeply
di e gen mi ochond ial lineage ha is sis e o all o he
Ichno opis species in he phylogeny; ne e heless, i e-
mains mo phologically simila o all membe s o he I.
bi i a a g oup, being hus included as pa o his g oup.
The ecogni ion o I. anganicana as a dis inc species
is suppo ed p ima ily by molecula di e gence, unique
geog aphic dis ibu ion, and i s dis inc i e colou a ion
– mos no ably he p esence o e enly spaced blue la -
e al spo s – unde sco ing he impo ance o in eg a ing
gene ic and pheno ypic da a in esol ing c yp ic di e si y
wi hin mo phologically conse a i e lineages.
Biogeog aphically, Ichno opis shows i s highes di-
e si y ac oss he no he n and wes e n pa s o cen al
and sou he n A ica, wi h he Kalaha i Basin eme ging as
a ho spo o species ichness o his g oup. In con as ,
eas e n A ica is ep esen ed by only a single species
(i.e., I. anganicana), while Sou h A ica ha bou s jus
one axon om he I. capensis complex. These pa e ns
a e shaped by egional habi a he e ogenei y, his o ical
ba ie s o gene low, and likely also by limi ed his o ical
sampling in la ge swa hes o sui able habi a (G eenbaum
e al. 2018).
The desc ip ion o wo new Ichno opis species om
Angola con ibu es o he wa e o ep ile species disco -
e ies in he egion o e he pas wo decades (Con adie
e al. 2012, 2020a, 2020b, 2020c; S anley e al. 2016;
B anch e al. 2019, 2021; Ma ques e al. 2019a, 2019b,
2020, 2022a, 2022b, 2023a, 2023b, 2024; Ce íaco e
al. 2020b, 2020c, 2020d, 2024; Halle mann e al. 2020;
Lobón-Ro i a e al. 2021, 2022, 2025a, 2025b; Pa inha
e al. 2021, 2025a, 2025b, 2025c; Wagne e al. 2021;
Ba es e al. 2023; Röll e al. 2024). Since 2012, Angola’s
known lace id di e si y has nea ly doubled, om 10 o
18 species, mi o ing g ow h in o he ep ile g oups such
as skinks and geckos (Con adie 2024). Gi en ha la ge
po ions o Angola emain poo ly su eyed, u he spe-
cies disco e ies a e likely o occu wi hin he e i o y.
The wo new Ichno opis species desc ibed he e we e
disco e ed h ough in ensi e ieldwo k in cen al and
sou heas e n Angola, egions long unde explo ed due o
decades o ci il con lic and he di icul access o hese
a eas (Con adie e al. 2021). Recen he pe ological su -
eys in hese a eas ha e yielded nume ous new species,
expanded aunal eco ds, and ha e p o ided comp e-
hensi e species in en o ies (e.g., Con adie e al. 2016,
2020a, 2020b, 2021, 2022a, 2022b, 2023; Nielsen e al.
2020), unde sco ing he impo ance o a ge ed ieldwo k
and sys ema ic biodi e si y assessmen s in his o ically
inaccessible a eas (Tolley e al. 2016). These wo new-
ly desc ibed Ichno opis species (I. obus a sp. no . and
I. longico pa sp. no .) a e cu en ly conside ed Angolan
endemics, hough hey may also occu in adjacen egions
o DRC and Zambia. Finally, he Angolan highlands and
adjacen Miombo woodlands appea pa icula ly signi -
ican , ha bou ing se e al endemic o nea -endemic axa
(Baue e al. 2023; Becke e al. 2023). These indings
align wi h p e ious esea ch iden i ying his egion as
a ho spo o ep ile endemism and suppo calls o en-
hanced conse a ion a en ion.
Conclusion
This s udy p esen s he mos comp ehensi e phylogene ic
and axonomic e ision o he genus Ichno opis o da e.
By in eg a ing mi ochond ial and nuclea molecula da a,
de ailed mo phological assessmen s, and b oad geog aph-
ic sampling, we e eal ha Ichno opis liza ds ha bou
mo e di e si y han p e iously ecognised. Ou indings
suppo he desc ip ion o wo new axa, indica e mul i-
ple po en ial c yp ic species and cla i ies he axonomic
s a us o se e al his o ically ambiguous names. The e-
co e ed phylogene ic s uc u e and s ong geog aphic
Ve eb a e Zoology 75, 2025, 627–672 665
pa i ioning unde sco e he e olu iona y signi icance
o cen al and sou he n A ica, pa icula ly he Angolan
highlands, as a cen e o di e si ica ion and endemism o
ep iles. This wo k no only s abilizes he axonomy o
Ichno opis, bu also lays a obus ounda ion o u u e
e olu iona y, ecological and conse a ion s udies o his
g oup ac oss i s ange.
Acknowledgemen s
This wo k was made possible h ough he collabo a ion and suppo o
nume ous indi iduals and ins i u ions. We hank he Minis y o En i-
onmen o he Republic o Angola (MINAMB), pa icula ly D . Miguel
Xa ie , Di ec o o he Ins i u o Nacional de Biodi e sidade e Conse -
ação (INBC), o hei ins i u ional collabo a ion. Ma e ial was collec -
ed and expo ed unde he ollowing pe mi s issued by MINAMB: 31/
GGPCC/2016 and 151/INBAC/MINAMB/2019. We also acknowledge
he excep ional logis ical assis ance p o ided by Fundação Kissama,
especially Vlady Russo.
WC hanks he Wild Bi d T us , which adminis e s he Na ional
Geog aphic Oka ango Wilde ness P ojec (2015–2019 Na ional Geo-
g aphic Socie y g an ), and Ch is B ooks, who o ganized he SAREP
Aqua ic Biodi e si y Su eys o he uppe Angola ca chmen s o he
Cubango-Oka ango Ri e Basin (May 2012) and he lowe Cui o and
Cuando Ri e Basins (Ap il 2013). We a e also g a e ul o he Na u-
al His o y Museum o Mapu o, which endo sed and p o ided pe mi s
(315/MHN/E.27/2014) o ca y ou pa o his wo k in Mozambique.
This wo k ecei ed inancial and logis ical suppo om se e al in-
s i u ions: The Na ional Geog aphic Socie y Commi ee o Resea ch
and Explo a ion (CRE 9281-13); he Sou h A ican Na ional Biodi e -
si y Ins i u e; he Na ional Resea ch Founda ion o Sou h A ica (G an
#92776) o he 2014 Mozambique su ey; and Khangela Sa a is o
camp logis ics in 2014. CK hanks and acknowledges Upemba Na ion-
al Pa k, Fo go en Pa ks Founda ion (DPF), Ins i u Congolais pou
la Conse a ion de la Na u e (ICCN), Uni e si y o Lubumbashi, and
Hankuzi Explo a ions o hei assis ance wi h acili a ing he collec ion
o aluable Congolese samples o his s udy.
JLR is cu en ly suppo ed by Associação BIOPOLIS CIBIO Base
FUI 2020–2023 (UIDB1 50027 i2020). This wo k was also pa ially
suppo ed by he Syn hesis+ BE-TAF P ojec 2022 G an ob ained by
JLR. We hank he CTM s a a CIBIO – especially Susana Lopes, So ia
Mou ão, and Pa ícia Ribei o – o hei dedica ed labo a o y suppo .
EG acknowledges Ana Be ancou o he Bo de Biomedical Re-
sea ch Cen e (BBRC) Genomics Analysis Co e Facili y o ech-
nical se ices and acili ies. This wo k was suppo ed by G an
5U54MD007592 om he Na ional Ins i u e on Mino i y Heal h and
Heal h Dispa i ies (NIMHD), pa o he US Na ional Ins i u es o
Heal h (NIH).
We a e g a e ul o Ga in Cael (RMCA) and Oli ie Pauwels
(RBINS) o p o iding access o ele an ma e ial om hei he pe o-
logical collec ions. Pho og aphs o key museum specimens we e gen-
e ously p o ided by: Eugen Beh ens (Museo delle Scienze, MUSE),
Lau en Vonnahme (Ame ican Museum o Na u al His o y, AMNH),
Pa ick Campbell (Na u al His o y Museum London, NHML), Joshua
Ma a (The Field Museum o Na u al His o y, FMNH), Ma hew Gage
(Museum o Compa a i e Zoology, Ha a d Uni e si y, MCZ), and
F ank Tillack (Museum ü Na u kunde Be lin, ZMB). Sebas ian Ki ch-
ho (Na u al His o y Museum Abu Dhabi, NHMAD) kindly p o ided
addi ional sequences.
We also hank he ollowing indi iduals o hei assis ance in he
ield and sample collec ion: Ninda Bap is a, Gab iella Bi encou -Sil a,
Thomas B anch, William R. B anch, Hanlie Engelb ech , James Ha ey,
Timó eo Júlio, Michele Menegon, Gö z Nee , and Alex Rebelo. Reuben
an B eda ex ends special hanks o Ed Ne he lands, Ash Bulla d, Haley
Du on, Be nie Jo daan, Be ha Buiswalelo, F ancois Becke , and Louis
du P eez o hei help in he ield, as well as Pie Bey ell and F ancois
Jacobs and hei eam o acili a ing ieldwo k in Namibia. Specimens
in Namibia we e collec ed unde he Na ional Commission o Resea ch,
Science and Technology pe mi AN20191118. We a e g a e ul o he
la e Don B oadley and Bill B anch o allowing us o inco po a e hei
unpublished da a on he genus.
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