Research Article |
Corresponding author: Michael F. Bates ( michaelfrancisbates@gmail.com ) Academic editor: Uwe Fritz
© 2023 Michael F. Bates, Javier Lobón-Rovira, Edward L. Stanley, William R. Branch, Pedro Vaz Pinto.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bates MF, Lobón-Rovira J, Stanley EL, Branch WR, Vaz Pinto P (2023) A new species of green-eyed Cordylus Laurenti, 1768 from the west-central highlands of Angola, and the rediscovery of Cordylus angolensis (Bocage, 1895) (Squamata: Cordylidae). Vertebrate Zoology 73: 599-646. https://doi.org/10.3897/vz.73.e95639
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Cordylus angolensis (Bocage, 1895) was described 128 years ago on the basis of a single specimen collected at Caconda in the west-central highlands of Angola. Additional specimens referred to this species were collected at ‘Mombolo’ (also in the central highlands) during the Vernay Angola Expedition in 1925. As the holotype was apparently destroyed in the fire of 1978 at the Museu Bocage in Lisbon and no additional specimens have been collected, its taxonomic status and phylogenetic relationships has remained uncertain. The species has eluded all efforts aimed at its re-discovery in the vicinity of the type locality, with a single specimen from near Condé, north of Mombolo—collected in 1970 by Wulf Haacke—the only other specimen of Cordylus known from west-central Angola. Recent field work in the Angolan highlands resulted in the collection of a series of specimens from Taqueta Mountain (west of Caconda), Monte Verde (Sandula, ‘Mombolo’) and Uassamba (Vondo). A phylogenetic analysis, using three mitochondrial and six nuclear genes, indicated the existence of two distinct species-level lineages in the Angolan highlands. These two species are allopatric and morphologically distinct, differing especially in terms of their colour patterns, eye colour and certain scalation characteristics. We therefore confirm that C. angolensis is a valid species and designate a neotype, and describe a new species, Cordylus momboloensis sp. nov.
Africa, Cordyliformes, distribution, Sauria, taxonomy
Cordylus Laurenti, 1768 is the most speciose and geographically widespread genus of girdled lizards (Cordylidae), comprising 21 species of mainly rupicolous lizards that occur from the south-western tip of South Africa northwards through Botswana and Namibia to Angola, and elsewhere to Mozambique, Zimbabwe, Zambia, Malawi, south-eastern Democratic Republic of the Congo, Tanzania and Kenya, with an apparently isolated population in southern Ethiopia (
Cordylus originally encompassed all heavily armoured members of the family Cordylidae, but a phylogenetic analysis by
Despite some early reptile and amphibian collecting in Angola during the colonial era (e.g.,
Four species of Cordylus have been recorded from Angola, namely C. angolensis and three closely related species restricted to the south-western parts of the country, namely C. machadoi, C. namakuiyus and C. phonolithos.
Cordylus vittifer machadoi was described by
Live specimens of various species of Cordylus: A C. machadoi (PEM R25218 – 6 km north of Humpata, Huíla Province, Angola; photo: William R. Branch); B C. namakuiyus (PEM R18005 – between Namibe and Omahua lodge, Namibe Province, Angola; photo: William R. Branch); C C. phonolithos (CAS 263581, holotype – vicinity of N’Dolondolo, Serra da Neve, Namibe Province, Angola; photo: Luis M. P. Ceríaco); D C. rhodesianus (near Chimoio, Mozambique; photo: David Maguire); E C. tropidosternum (Kayalekera, Malawi; photo: Luke Verburgt); F C. jonesii (Umbabat Private Nature Reserve, Mpumalanga Province, South Africa; photo: Darren Pietersen); G C. vittifer (NMB R8523 – Farm: Dipka 220, Vrede district, Free State Province, South Africa; photo: Michael F. Bates); H C. cordylus (NMB R8540 – Ha Sehlaba, Lesotho; photo: Michael F. Bates). Representatives of all four Cordylus clades recovered by the current genetic analysis are included.
Material from “Damaraland” (north-western Namibia), identified as Zonurus griseus Cuvier, 1829 by Peters (1869) and Z. cordylus by
A molecular phylogeny conducted by
A third species in the C. machadoi group, namely C. phonolithos, was described by
Fortunately, the holotype of C. angolensis was examined at the Museu Bocage in 1968 by the late Dr Donald Broadley (Natural History Museum, Bulawayo) before it was apparently destroyed in the fire a decade later. Broadley’s scale counts agree with data presented by
Recent field work in central-western Angola resulted in the collection of a series of specimens from Taqueta Mountain (c. 100 kms west of Caconda), Monte Verde (Sandula, ‘Mombolo’) and Mount Uassamba (Vondo). In order to examine the evolutionary relationships between these populations, we conducted a phylogenetic analysis using three mitochondrial and six nuclear genes. A detailed morphological evaluation was also conducted, indicating the existence of two distinct species, one of which we refer to C. angolensis, while the other represents a new species (initially referred to as C. ‘Mombolo’) which we describe below.
Between 2016 and 2021, specimens of Cordylus were collected across Angola, focusing on the central highlands. A total of 25 specimens, comprising 12 C. angolensis-like specimens and 13 specimens of the C. machadoi group, were collected as vouchers. Specimens were euthanised by injection of tricaine methanesulfonate (MS222) (
Phylogenetic analysis was used to provide a phylogenetic context and support the morphological findings. For that purpose we generated 213 new Cordylus sequences from 24 individuals from across Angola following
Cordylidae samples included in this study with museum voucher numbers/field numbers and GenBank accession numbers for nine genes. Voucher specimens are indicated with museum accession numbers:
Species | Catalogue Number | Field Number | Country/Locality | 16S | 12S | ND2 | PRLR | Kif24 | c-mos | RAG-1 | MYH2 | BDNF | Source |
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Cordylus angolensis (Neotype) |
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NB 0612 | Angola/Taqueda Mtn | OQ869891 | OQ869867 | OQ869522 | OQ869474 | OQ869428 | OQ867122 | OQ869498 | OQ869452 | OQ869544 | This study |
C. angolensis | CHL 611 | NB 0611 | Angola/Taqueda Mtn | OQ869890 | OQ869866 | OQ869521 | OQ869473 | OQ869427 | OQ867121 | OQ869497 | OQ869451 | OQ869543 | This study |
C. angolensis | CHL 613 | NB 0613 | Angola/Taqueda Mtn | OQ869892 | OQ869868 | OQ869523 | OQ869475 | OQ869429 | OQ867123 | OQ869499 | OQ869453 | OQ869545 | This study |
C. angolensis |
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NB 0614 | Angola/Taqueda Mtn | OQ869893 | OQ869869 | OQ869524 | OQ869476 | OQ869430 | OQ867124 | OQ869500 | OQ869454 | OQ869546 | This study |
C. machadoi | — | KTH 09059 | Angola/Humpata | HQ167204 | KT941132 | KT941260 | KT941311 | KT941234 | KT941205 | KT941369 | HQ167424 | KT941168 |
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C. machadoi |
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KTH 09056 | Angola/Humpata | KT941143 | KT941129 | — | KT941308 | KT941231 | KT941202 | KT941366 | KT941245 | KT941165 |
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C. machadoi |
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KTH 09057 | Angola/Humpata | KT941145 | KT941131 | KT941259 | KT941310 | KT941233 | KT941204 | KT941368 | KT941247 | KT941167 |
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C. machadoi |
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KTH 09058 | Angola/Humpata | KT941144 | KT941130 | KT941258 | KT941309 | KT941232 | KT941203 | KT941367 | KT941246 | KT941166 |
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C. machadoi |
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KTH 09006 | Angola/Humpata | KT941142 | KT941128 | — | KT941307 | KT941230 | KT941201 | KT941365 | KT941244 | KT941164 |
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C. machadoi |
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KTH 09005 | Angola/Humpata | KT941141 | KT941127 | KT941257 | KT941306 | KT941229 | KT941200 | KT941364 | KT941243 | KT941163 |
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C. machadoi | CHL 698 | NB 0698 | Angola/Tundavala | OQ869894 | OQ869870 | OQ869525 | OQ869477 | OQ869431 | OQ867125 | OQ869501 | OQ869455 | OQ869547 | This study |
C. machadoi | CHL 699 | NB 0699 | Angola/Tundavala | OQ869895 | OQ869871 | OQ869526 | OQ869478 | OQ869432 | OQ867126 | OQ869502 | OQ869456 | OQ869548 | This study |
C. machadoi | CHL 700 | NB 0700 | Angola/Tundavala | OQ869896 | OQ869872 | OQ869527 | OQ869479 | OQ869433 | OQ867127 | OQ869503 | OQ869457 | OQ869549 | This study |
C. machadoi | CHL 701 | NB 0701 | Angola/Tundavala | OQ869897 | OQ869873 | OQ869528 | OQ869480 | OQ869434 | OQ867128 | OQ869504 | OQ869458 | OQ869550 | This study |
C. machadoi |
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NB 0702 | Angola/Tundavala | OQ869898 | OQ869874 | — | OQ869481 | OQ869435 | OQ867129 | OQ869505 | OQ869459 | OQ869551 | This study |
C. machadoi | WRB 0040 | — | Angola | JQ389806 | JQ389799 | — | JQ389851 | JQ389812 | — | — | — | — |
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C. machadoi | WRB 0041 | — | Angola | JQ389807 | JQ389800 | — | — | — | — | — | — | — |
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C. momboloensis sp. nov. | FKH 0129 | P9-17 | Angola/Vondo | OQ869904 | OQ869880 | OQ869534 | OQ869487 | OQ869441 | OQ867135 | OQ869511 | — | OQ869557 | This study |
C. momboloensis sp. nov. (Paratype) | FKH 0125 | P9-13 | Angola/Vondo | OQ869900 | OQ869876 | OQ869530 | OQ869483 | OQ869437 | OQ867131 | OQ869507 | OQ869460 | OQ869553 | This study |
C. momboloensis sp. nov. (Paratype) |
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P9-14 | Angola/Vondo | OQ869901 | OQ869877 | OQ869531 | OQ869484 | OQ869438 | OQ867132 | OQ869508 | OQ869461 | OQ869554 | This study |
C. momboloensis sp. nov. (Paratype) | FKH 0127 | P9-15 | Angola/Vondo | OQ869902 | OQ869878 | OQ869532 | OQ869485 | OQ869439 | OQ867133 | OQ869509 | OQ869462 | OQ869555 | This study |
C. momboloensis sp. nov. (Paratype) | FKH 0128 | P9-16 | Angola/Vondo | OQ869903 | OQ869879 | OQ869533 | OQ869486 | OQ869440 | OQ867134 | OQ869510 | OQ869463 | OQ869556 | This study |
C. momboloensis sp. nov. (Holotype) |
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AG16.36 | Angola/Mt Verde | OQ869899 | OQ869875 | OQ869529 | OQ869482 | OQ869436 | OQ867130 | OQ869506 | — | OQ869552 | This study |
C. momboloensis sp. nov. (Allotype) |
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P9-140 | Angola/Sandula | OQ869905 | OQ869881 | OQ869535 | OQ869488 | OQ869442 | OQ867136 | OQ869512 | OQ869464 | OQ869558 | This study |
C. momboloensis sp. nov. | — | P1-279 | Angola/Bocoio | OQ869914 | — | — | — | — | — | — | — | — | This study |
C. namakuiyus |
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— | Angola/near Pico do Azevado | KT941137 | KT941123 | KT941253 | KT941302 | KT941225 | KT941194 | KT941358 | KT941239 | KT941157 |
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C. namakuiyus |
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— | Angola/near Pico do Azevado | KT941138 | KT941124 | KT941254 | KT941303 | KT941226 | KT941195 | KT941359 | KT941240 | KT941158 |
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C. namakuiyus |
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— | Angola/near Caraculo | KT941135 | KT941121 | KT941251 | KT941300 | KT941223 | KT941192 | KT941356 | KT941237 | KT941155 |
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C. namakuiyus |
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— | Angola/near Caraculo | KT941136 | KT941122 | KT941252 | KT941301 | KT941224 | KT941193 | KT941357 | KT941238 | KT941156 |
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C. namakuiyus |
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— | Angola/ near Pico do Azevado | KT941139 | KT941125 | KT941255 | KT941304 | KT941227 | KT941196 | KT941360 | KT941241 | KT941159 |
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C. namakuiyus |
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— | Angola/ near Pico do Azevado | KT941140 | KT941126 | KT941256 | KT941305 | KT941228 | KT941197 | KT941361 | KT941242 | KT941160 |
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C. namakuiyus |
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KTH 09212 | Angola/between Namibe and Omauha Lodge | KT941148 | KT941134 | — | KT941313 | KT941236 | KT941207 | — | KT941250 | KT941170 |
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C. namakuiyus | FKH 0262 | P9-169 | Angola/Chapeu Armado | OQ869908 | OQ869884 | OQ869537 | OQ869491 | OQ869445 | OQ867139 | OQ869515 | OQ869467 | OQ869561 | This study |
C. namakuiyus | CHL 609 | NB 0609 | Angola/Mariquita | OQ869907 | OQ869883 | — | OQ869490 | OQ869444 | OQ867138 | OQ869514 | OQ869466 | OQ869560 | This study |
C. namakuiyus | CHL 616 | NB 0616 | Angola/Meva | OQ869906 | OQ869882 | OQ869536 | OQ869489 | OQ869443 | OQ867137 | OQ869513 | OQ869465 | OQ869559 | This study |
C. phonolithos | AMB 10272 | — | Angola/Serra da Neve | MN342160 | — | QGN01202 | — | — | — | — | — | — |
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C. phonolithos |
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— | Angola/Serra da Neve | MN342159 | — | QGN01201 | — | — | — | — | — | — |
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C. phonolithos | CHL 848 | NB 0848 | Angola/Serra da Neve | OQ869909 | OQ869885 | OQ869538 | OQ869492 | OQ869446 | OQ867140 | OQ869516 | OQ869468 | OQ869562 | This study |
C. phonolithos |
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NB 0849 | Angola/Serra da Neve | OQ869910 | OQ869886 | OQ869539 | OQ869493 | OQ869447 | OQ867141 | OQ869517 | OQ869469 | OQ869563 | This study |
C. phonolithos |
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NB 0850 | Angola/Serra da Neve | OQ869911 | OQ869887 | OQ869540 | OQ869494 | OQ869448 | OQ867142 | OQ869518 | OQ869470 | OQ869564 | This study |
C. phonolithos | CHL 851 | NB 0851 | Angola/Serra da Neve | OQ869912 | OQ869888 | OQ869541 | OQ869495 | OQ869449 | OQ867143 | OQ869519 | OQ869471 | OQ869565 | This study |
C. phonolithos | CHL 852 | NB 0852 | Angola/Serra da Neve | OQ869913 | OQ869889 | OQ869542 | OQ869496 | OQ869450 | OQ867144 | OQ869520 | OQ869472 | OQ869566 | This study |
C. beraduccii | JB 6 | — | Tanzania | KT941403 | KT941400 | KT941393 | KT941390 | JQ389811 | KT941187 | KT941351 | KT941386 | KT941150 |
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C. beraduccii | JB 7 | — | Tanzania | KT941404 | KT941401 | KT941394 | KT941391 | — | KT941188 | KT941352 | KT941387 | KT941151 |
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C. beraduccii | WRB 0037 | — | Tanzania/Mtera | HQ167172 | HQ167061 | KT941395 | HQ167501 | HQ167283 | KT941189 | KT941353 | HQ167392 | KT941152 |
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C. cordylus | AMB 8168 | — | S. Africa/E. Cape | HQ167182 | HQ167071 | HQ166970 | HQ167511 | HQ167293 | — | — | HQ167402 | — |
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C. cordylus |
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— | S. Africa/E. Cape | HQ167190 | HQ167079 | HQ166978 | HQ167519 | HQ167301 | — | — | HQ167410 | — |
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C. cordylus |
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— | S. Africa/E. Cape | HQ167189 | HQ167078 | HQ166977 | HQ167518 | HQ167300 | — | — | HQ167409 | — |
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C. cordylus |
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— | S. Africa/E. Cape | HQ167232 | HQ167121 | HQ167015 | HQ167561 | HQ167343 | — | — | HQ167452 | — |
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C. imkeae | MBUR 01795 | — | S. Africa/N. Cape | HQ167197 | HQ167086 | HQ166985 | HQ167526 | HQ167308 | — | — | HQ167417 | — |
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C. imkeae | MBUR 01796 | — | S. Africa/N. Cape | HQ167198 | HQ167087 | HQ166986 | HQ167527 | HQ167309 | KT941191 | KT941355 | HQ167418 | KT941154 |
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C. jonesii | AMB 8310 | — | S. Africa/Limpopo | HQ167199 | HQ167088 | HQ166987 | HQ167528 | HQ167310 | KT941198 | KT941362 | HQ167419 | KT941161 |
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C. jonesii | AMB 8396 | — | S. Africa/Limpopo | HQ167200 | HQ167089 | HQ166988 | HQ167529 | HQ167311 | KT941199 | KT941363 | HQ167420 | KT941162 |
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C. macropholis | AMB 8873 | — | S. Africa/W. Cape | HQ167206 | HQ167095 | HQ166993 | HQ167535 | HQ167317 | — | — | HQ167426 | — |
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C. macropholis | AMB 8874 | — | S. Africa/W. Cape | HQ167207 | HQ167096 | HQ166994 | HQ167536 | HQ167318 | KT941208 | KT941371 | HQ167427 | KT941171 |
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C. marunguensis | EBG 2994 | — | DRC/Marungu Mountains | JQ389803 | JQ389798 | KT941396 | JQ389849 | JQ389810 | KT941209 | KT941372 | JQ389846 | KT941172 |
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C. marunguensis |
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— | DRC/Marungu Mountains | JQ389803 | JQ389798 | — | JQ389849 | JQ389810 | — | KU298675 | — | — |
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C. marunguensis |
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— | DRC/Marungu Mountains | JQ389802 | JQ389797 | — | JQ389848 | JQ389809 | — | — | JQ389846 | — |
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C. mclachlani | AMB 8855 | — | S. Africa/W. Cape | HQ167208 | HQ167097 | HQ166995 | HQ167537 | HQ167319 | KT941210 | KT941373 | HQ167428 | KT941173 |
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C. mclachlani | CmclSU 1 | — | S. Africa/W. Cape | HQ167209 | HQ167098 | HQ166996 | HQ167538 | HQ167320 | — | KT941373 | HQ167429 | — |
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C. meculae |
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— | Mozambique/Mecula | HQ167210 | HQ167099 | — | HQ167539 | HQ167321 | KT941413 | KT941411 | HQ167430 | KT941416 |
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C. meculae |
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— | Mozambique/Mecula | HQ167211 | HQ167100 | — | HQ167540 | HQ167322 | KT941414 | KT941412 | HQ167431 | KT941417 |
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C. meculae |
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— | Mozambique/Mecula | HQ167233 | HQ167122 | — | HQ167562 | HQ167344 | KT941415 | — | HQ167453 | KT941418 |
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C. meculae |
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— | Mozambique/Mecula | HQ167234 | HQ167123 | — | HQ167563 | HQ167345 | — | — | HQ167454 | — |
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C. minor | CminSU | — | S. Africa/N. Cape | HQ167212 | HQ167101 | HQ166997 | HQ167541 | HQ167323 | — | — | HQ167432 | — |
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C. minor |
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— | S. Africa/W. Cape | HQ167169 | HQ167058 | HQ166958 | HQ167498 | HQ167280 | KT941186 | — | HQ167389 | KT941149 |
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C. minor |
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— | S. Africa/W. Cape | HQ167170 | HQ167059 | HQ166959 | HQ167499 | HQ167281 | — | — | HQ167390 | — |
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C. niger | AMB 8875 | — | S. Africa/W. Cape | HQ167216 | HQ167105 | HQ166999 | HQ167545 | HQ167327 | — | — | HQ167436 | — |
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C. niger | CnigSU 1 | — | S. Africa/W. Cape | HQ167217 | HQ167106 | HQ167000 | HQ167546 | HQ167328 | KT941211 | KT941374 | HQ167437 | KT941174 |
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C. oelofseni | AMB 8851 | — | S. Africa/W. Cape | HQ167218 | HQ167107 | HQ167001 | HQ167547 | HQ167329 | — | — | HQ167438 | — |
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C. oelofseni | AMB 8860 | — | S. Africa/W. Cape | HQ167221 | HQ167110 | HQ167004 | HQ167550 | HQ167332 | KT941212 | KT941375 | HQ167441 | KT941175 |
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C. oelofseni | AMB 8862 | — | S. Africa/W. Cape | HQ167222 | HQ167111 | HQ167005 | HQ167551 | HQ167333 | — | — | HQ167442 | — |
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C. oelofseni | CoelSU 1 | — | S. Africa/W. Cape | HQ167219 | HQ167108 | HQ167002 | HQ167548 | HQ167330 | KT941213 | KT941376 | HQ167439 | KT941176 |
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C. oelofseni | CoelSU 2 | — | S. Africa/W. Cape | HQ167220 | HQ167109 | HQ167003 | HQ167549 | HQ167331 | — | — | HQ167440 | — |
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C. rhodesianus | ELSPET 4 | — | Pet Trade | HQ167230 | HQ167119 | HQ167013 | HQ167559 | HQ167341 | KT941214 | KT941377 | HQ167450 | KT941177 |
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C. rhodesianus | ELSPET 5 | — | Pet Trade | HQ167231 | HQ167120 | HQ167014 | HQ167560 | HQ167342 | — | — | HQ167451 | — |
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C. rivae | TJC 564 | — | Ethiopia | KM242128 | KM242126 | — | KM242122 | KM242118 | — | — | KM242124 | — | Nielsen and Colston (2014) |
C. rivae | TJC 565 | — | Ethiopia | KM242129 | KM242127 | — | KM242123 | KM242119 | — | — | KM242125 | — | Nielsen and Colston (2014) |
C. tropidosternum | JB 8 | — | Tanzania | KT941405 | KT941402 | KT941397 | KT941392 | KT941389 | KT941215 | KT941378 | KT941388 | KT941178 |
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C. tropidosternum | WRB 0038 | — | Tanzania | HQ167236 | HQ167125 | — | HQ167565 | HQ167347 | — | — | HQ167456 | — |
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C. tropidosternum | WRB 0042 | — | Tanzania | HQ167235 | HQ167124 | KT941398 | HQ167564 | — | KT941216 | KT941379 | HQ167455 | KT941179 |
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C. ukingensis | WRB 0039 | — | Kenya/Uzungwe Mts | HQ167237 | HQ167126 | KT941399 | HQ167566 | HQ167348 | KT941217 | KT941380 | HQ167457 | KT941180 |
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C. vittifer | AMB 6073 | — | S. Africa/ Mpumalanga | HQ167241 | HQ167130 | HQ167019 | HQ167570 | HQ167352 | KT941218 | KT941381 | HQ167461 | KT941181 |
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C. vittifer | AMB 8274 | — | S. Africa/Limpopo | HQ167242 | HQ167131 | HQ167020 | HQ167571 | HQ167353 | KT941219 | KT941382 | HQ167462 | KT941182 |
|
C. vittifer | AMB 8603 | — | Eswatini | HQ167243 | HQ167132 | HQ167021 | HQ167572 | HQ167354 | KT941220 | KT941383 | HQ167463 | KT941183 |
|
Namazonurus campbelli |
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— | Namibia | HQ167174 | HQ167063 | HQ166962 | HQ167503 | HQ167285 | KT941221 | KT941384 | HQ167394 | KT941184 |
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Ouroborus cataphractus | MBUR 01792 | — | S. Africa/N. Cape | HQ167177 | HQ167066 | HQ166965 | HQ167506 | HQ167288 | KT941222 | KT941385 | HQ167397 | KT941185 |
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Pseudocordylus melanotus subviridis |
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— | S. Africa/KwaZulu-Natal | HQ167268 | HQ167157 | HQ167046 | HQ167597 | HQ167379 | — | — | HQ167486 | — |
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P. m. subviridis |
|
— | S. Africa/KwaZulu-Natal | HQ167267 | HQ167156 | ADO13379 | HQ167596 | HQ167378 | — | — | HQ167485 | — |
|
Gene | Primer | Length (bp) | Sequence | Reference |
16S | 16Sa | 581 | 5’ CGCCTGTTTATCAAAAACAT 3’ |
|
16Sb | 5’ CCGGTCTGAACTCAGATCACGT 3’ | |||
12S | 12sf700 | 459 | 5’ AAACTGGGATTAGATACCCCACTAT 3’ |
|
12sr600 | 5’ GAGGGTGACGGCGGTGTGT 3’ | |||
ND2 | L4437-R | 1025 | 5’ AAGCTTTCGGGCCCATACC 3’ |
|
H5540-F | 5’ TTTAGGGCTTTGAAGGC 3’ | |||
R102 | 5’ CAGCCTAGGTGGGCGATTG 3’ |
|
||
RAG-1 | RAG1_f1a | 1040 | 5’ CAGCTGYAGCCARTACCATAAAAT 3’ |
|
RAG1_r2 | 5’ CTTTCTAGCAAAATTTCCATTCAT 3’ | |||
c-mos | G73 | 407 | 5’ GCGGTAAAGCAGGTGAAGAAA 3’ |
|
G74 | 5’ TGAGCATCCAAAGTCTCCAATC 3’ | |||
BDNF | BDNF_f | 732 | 5’ GACCATCCTTTTCCTKACTATGGTTATTTCATACTT 3’ |
|
BDNF_r | 5’ CTATCTTCCCCTTTTAATGGTCAGTGTACAAAC 3’ | |||
PRLR | PRLRf1 | 532 | 5’ GACARYGARGACCAGCAACTRATGCC 3’ |
|
PRLRr1 | 5’ GACYTTGTGRACTTCYACRTAATCCAT 3’ | |||
Kif24 | Kif24f | 572 | 5’ WGGCTGCTGRAAYTGCTGGTG 3’ |
|
Kif24r | 5’ SAAACGTRTCTCCMAAACGCATCC 3’ | |||
MYH2 | MYH2f | 765 | 5’ GAACACCAGCCTCATCAACC 3’ |
|
MYH2r | 5’ TGGTGTCCTGCTCCTTCTTC 3’ |
Bayesian inference (BI) and maximum likelihood (ML) analyses were performed with sequence data for 85 specimens using separate analyses for each gene, as well as different analyses for concatenated sets of three mitochondrial genes (ND2, 16S and 12S) and six nuclear genes (RAG-1, c-mos, BDNF, PRLR, MYH2 and Kif24), and all genes combined. The partitioning schemes were determined using PartitionFinder2 (
The identity of an individual from Morro do Pundo, Bocoio (P1-279 in Table
This study was based on material in the herpetological holdings of the following museums/collections: American Museum of Natural History, New York (
Specimens were examined using binocular dissecting microscopes (up to 40 or 48 times magnification). Measurements were performed using digital calipers (0.01 mm) or a ruler (1 mm), often under magnification. Apart from five
Measurements were performed on the right side of the body unless damaged. The following were measured: snout-vent length (SVL, from tip of snout to anterior margin of vent, with lizard on its back and flattened); tail length (posterior margin of vent to tail tip, original tails unless indicated); head length (tip of snout to posterior margin of lateral temporals); head width (widest part in the temporal area); head height (midpoint of eye from top of head to bottom of lower jaw); ear length (greatest distance more-or-less dorso-ventally); eye length (excluding small scales around the eye); nostril-eye distance (shortest distance between); eye-ear distance (shortest distance); snout-eye length (tip of snout to anterior margin of eye, excluding small scales around the eye); internarial distance (shortest distance between nostrils); inter-orbital distance (measured between the middle of the eye sockets); snout-arm length (from tip of snout to anterior margin of forelimb); axilla-groin distance (posterior edge of forelimb insertion to anterior edge of hindlimb insertion); forearm (inner part of elbow fold to tip of claw of fourth (longest) finger; 4th (longest) toe of hindlimb.
Scale counts were, unless indicated, performed on both sides of each lizard. The following counts were taken: supralabials (all scales bordering the upper lip, excluding the rostral, but including the scale at the corner of the mouth; the second last supralabial is the largest); infralabials (all scales bordering lower lip, excluding the mental, the posterior one situated at the corner of the mouth below and partly behind the posterior supralabial); sublabials (large scales in contact with the infralabials, the posterior one situated near the ear opening below and partly behind the posterior infralabial); number of chin shields in contact with anterior sublabials; supraoculars (large scales); supraciliaries (narrow and elongate scales above the eye and in contact with the much larger supraoculars, excluding tiny granules and postocular scales that may be in contact with the posterior supraocular); loreals; suboculars (all scales bordering eye and in contact with the supralabials, but excluding the preocular and any postoculars; when present, a fourth subocular is invariably situated near the back of the eye); preoculars; rows of gulars between posterior angles of jaws, excluding tiny granules; dorsals, transverse rows (from immediately behind occipitals to vent); dorsals, longitudinal rows (counted midway between fore- and hindlimbs, excluding granular or minute scales on either side if present); ventrals, transverse rows (from axilla to groin; the most anterior row curves anteriorly and the most posterior row curves posteriorly); ventrals, longitudinal rows (counted midway between fore- and hindlimbs; the most lateral row consists of scales generally at least half as wide as the adjacent inner ones; when this row is not clearly distinguishable, the formula ‘12 + 2’ is used, indicating a possible extra row on either side); subdigital lamellae on 4th (longest) toe (the most basal scale counted is fully present on the digit); femoral pores; differentiated femoral scales (generation glands).
The interpretation of ‘occipital’ requires explanation. In the genus Cordylus, occipitals are typically defined as the scales posterior to, and in contact with, the pair of posterior parietals, plus those scales in contact with the posterior upper temporals [on either side of the posterior parietals] (e.g.,
Arrangement of occipitals (red circles) and post-occipitals (yellow triangles) in A Cordylus angolensis (CHL 611), B C. momboloensis sp. nov. (
All material referable to C. angolensis, as well as that of C. ‘Mombolo’, was compared to one another and samples of other Angolan Cordylus, namely C. machadoi, C. namakuiyus and C. phonolithos (Fig.
In the diagnoses and diagnostic key below, comparative morphological details were derived from:
For osteological descriptions of the west-central Angolan cordylid material, microComputed Tomography (CT) datasets of C. ‘Mombolo’ (
We follow a lineage-based species concept whereby a species is represented by an independently evolving metapopulation lineage (
Both the BI and ML analyses were largely concordant, yielding the same well-supported topology. Phylogenetic analyses demonstrate that Angolan Cordylus is sister to the C. jonesii–C. rhodesianus group (PP: 0.99, BS: 87) (Fig.
Maximum likelihood (ML) phylogenetic tree for Cordylus based on concatenated dataset, with Bayesian inference (BI) support overlaid. Numbers above the key nodes indicate BI posterior probability (≥ 0.95 were considered supported), while below the key nodes indicate ML bootstrap values (≥ 95% were considered supported). Major Angolan clades are highlighted in different colours.
Angolan Cordylus represents a well-supported monophyletic group (PP: 1, BS: 100), divided into a northern clade, which includes C. angolensis from the central highlands of Angola and an undescribed sister species, C. ‘Mombolo’, from slightly further north (PP: 0.99, BS: 88), and a southern clade that includes the three species of the C. machadoi group (C. machadoi, C. namakuiyus and C. phonolithos). The Taqueta material is referable to C. angolensis on the basis of its similarity to the holotype (see below). These clades show large genetic ND2 p-distance between them (~10.00%). The uncorrelated ND2 p-distance found between C. angolensis and C. ‘Mombolo’ was 9.2% (Table
ND2 divergences (uncorrected pairwise distances) between cordylid taxa, including all Cordylus from Angola. Bold values depict intraspecific divergences.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | |
1. Namazonurus campbelli | — | ||||||||
2. Pseudocordylus m. subviridis | 22.46 | 7.7 | |||||||
3. Cordylus vittifer | 22.12 | 20.82 | 11.7 | ||||||
4. Cordylus jonesii | 22.02 | 20.75 | 16.54 | 2.8 | |||||
5. Cordylus namakuiyus | 22.07 | 19.50 | 16.05 | 16.43 | 4.3 | ||||
6. Cordylus machadoi | 23.19 | 20.27 | 16.38 | 16.82 | 7.57 | 2.4 | |||
7. Cordylus phonolithos | 21.56 | 18.90 | 16.77 | 15.60 | 9.40 | 8.94 | 0.4 | ||
8. Cordylus ‘Mombolo’ | 22.74 | 21.25 | 16.60 | 16.70 | 9.85 | 11.59 | 12.29 | 1.1 | |
9. Cordylus angolensis | 22.22 | 19.82 | 16.55 | 15.15 | 10.85 | 11.97 | 11.92 | 9.22 | 0.2 |
Length of scales in the first transverse row of dorsals (Figs
Dorsal views of the head in Angolan Cordylus. Cordylus angolensis—A CHL 611, B CHL 615, C
Presence or absence of a loreal scale (Figs
Nasal scales (Figs
Position of the nostril (Figs
The position of the nostril relative to the suture between rostral and first supralabial is also informative. In C. angolensis and C. ‘Mombolo’ the entire nostril is situated posterior to the suture, as is the case in C. tropidosternum and C. vittifer, whereas in all three species of the C. machadoi group at least some part of the nostril is situated above the suture, which is often also the case with C. rhodesianus and C. jonesii (Fig.
Texture of sublabial scales (Figs
Lateral views of the head in Angolan Cordylus. Cordylus angolensis—A CHL 611, B CHL 615, C
Ventral views of the head in Angolan Cordylus. Cordylus angolensis—A CHL 611, B CHL 615, C
Arrangement of the nasal shield, nostril and scales of the snout in Cordylus: A C. angolensis (CHL 615 – Taqueta Mountain, Angola); B C. momboloensis sp. nov. (
Relative size of the frontonasal scale (Fig.
Proximity of the frontonasal to the frontal scale (Fig.
Occipitals and post-occipitals (Figs
Anterior parietals (Fig.
Femoral pores and differentiated femoral scales (generation glands) (Table
Comparative meristic data for Cordylus from the Angolan clade and various species in the East African clade (including C. jonesii–C. rhodesianus). The first row of data refers to the range of values, the second row is the mean, and the third row is the standard deviation. For C. angolensis, data for the (now lost) holotype, as recorded by
Species | Sample Size | Gulars in contact with 1st pair chin shields | Gulars btwn angle of jaws | Transverse rows of dorsals | Longitudinal rows of dorsals | Transverse rows of ventrals | Longitudinal rows of ventrals | Lamellae under 4th toe | Lamellae under 4th finger | Femoral pores (total both legs) | Differentiated femoral scales (total both legs) |
C. angolensis | 5 (4 M, 1 F) | 5–6 5.2 ±0.447 | 20–21 [26] 20.2 ±0.447 | 24–25 24.4 ±0.548 | 20–23 [26] 21.2 ±1.30 | 22–24 [27] 22.8 ±0.837 | 14 [16] | 14 | — | M 11–12 F 11 | M 38–50 F 0 |
C. momboloensis sp. nov. | 12 (5 M, 7 F) | 4–5*–1 4.6 ±0.515 | 16–26 20.2 ±2.66 | 22–24 22.9 ±0.900 | 18–23 20.7 ±1.56 | 22–27 23.9 ±1.78 | 11–14 12.3 ±0.754 | 11–15–2 13.1 ±1.10 | — | M 10–15 F 8–12 | M 53–72 F 0 |
C. machadoi | 12 (7 M, 5 F) | 3–5 4.1 ±0.793 | 18–24–1 20.4 ±1.69 | 23–27 24.8 ±1.14 | 22–27 24.8 ±1.66 | 22–26 23.5 ±1.45 | 11–18 13.8 ±2.09 | 10–15 13.7 ±1.44 | 9–13 11.2 ±1.03 | M 11–14 F 12–14 | M 20–48 F 0 |
C. cf. machadoi (Namibia) | 1 (1 J) | 5 | 24 | 23 | 22 | 23 | 14 | 14 | 10 | J 9 | J 0 |
C. namakuiyus | 26 (11 M, 15 F) | 3–5 3.6 ±0.697 | 15–18–1 17.3 ±0.737 | 22–26 23.6 ±0.852 | 19–24 22.0 ±1.37 | 19–23–1 21.5 ±0.770 | 10–18 13.1 ±1.48 | 11–15 12.8 ±1.05 | 9–12 10.7 ±0.962 | M 8–12 F 8–12 | M 19–36 F 0 |
C. phonolithos | 4 (2 M, 1 F, 1 J) | 3–5** 4.4 ±0.787 | 17–18 17.3 0.500 | 24–26 25.0 0.816 | 22–23 22.5 0.577 | 23–2 | 16–17 16.5 0.577 | 14–1 | 12–3 | M 14–15 F 14 J 0 | M 33–42 F 0 J 0 |
C. tropidosternum | 6 (3 M, 3 F) | 4 | 23–25 23.5 0.837 | 27–28 27.3 0.516 | 19–23 21.0 1.55 | 23–28 25.3 1.86 | 12–14 12.3 0.816 | 13–14 13.3 0.516 | 12–13 12.3 0.516 | M 12 F 0–13 | M 16–29 F 0 |
C. rhodesianus | 5 (1 M, 4 F) | 2–4 3.00 ±0.707 | 22–24 23.0 ±0.707 | 26–28 27.0 ±1.00 | 22–25 23.6 ±1.14 | 22–25 22.8 ±1.30 | 12–14 13.2 ±1.10 | 12–13 12.2 ±0.447 | 10–11 10.2 ±0.447 | M 14 F 11–12 | M 49 F 0 |
C. jonesii | 5 (2 M, 3 F) | 3–4 3.2 ±0.447 | 20–22 20.8 ±0.837 | 23–24 23.6 ±0.548 | 19–24 21.4 ±1.95 | 21–25 23.4 ±1.52 | 12–14 12.4 ±0.894 | 13–14 13.4 ±0.548 | 10–12 11.6 ±0.894 | M 11–12 F 8–10 | M 10–13 F 0 |
C. vittifer | 72 (35 M, 31 F, 6 J) | 2–5–1 3.2 ±0.873 | 15–24–1 20.6 ±1.63 | 23–27 25.0 ±0.731 | 20–27 22.4 ±1.37 | 21–27 24.2 ±1.40 | 14–18 15.1 ±1.11 | 12–16–1 13.5 ±0.734 | 10–13–1 11.5 ±0.733 | M 10–18 F 0–19 J 0–12 | M 17–62 F 0–10 J 0–18 |
Ventral scale rows longitudinally (Table
Other scale counts (Table
Colour pattern (in life; Figs
Colour of the iris of the eye (Fig.
Texture of gular scales (Fig.
Cranial skeleton (Fig.
Osteoderms (Figs
The morphological and genetic differences mentioned above, together with allopatry, support the recognition of two different species in the C. angolensis clade. We take the opportunity below to describe the C. ‘Mombolo’ lineage as a new species.
Squamata: Sauria: Cordylidae
Zonurus angolensis Bocage, 1895: 25. Type locality. Caconda, Angola.
Cordylus cordylus angolensis
–
Cordylus angolensis
–
In 1968 Dr Donald G. Broadley (in litt., 18 January 2012) examined the holotype (MBL 429) of C. angolensis at the Museu Bocage in Lisbon and recorded the following data: nasals in broad contact, loreal absent (fused with preocular); frontal and frontonasal in contact, suboculars 3, supraciliaries 3, gulars 26, dorsals in 24 transverse and 26 longitudinal rows, ventrals in 27 transverse and 16 longitudinal rows, femoral pores 6 on each thigh; outer three rows of ventrals keeled and mucronate; SVL = 78 mm; tail length 74 mm tail (broken). As mentioned earlier,
According to
The new specimens from Taqueta Mountain were collected about 100 km west of Caconda and are assigned to C. angolensis on the basis of their similarity to the holotype as described by
The holotype (MBL 429) is no longer in existence, and therefore we propose and describe one of the new specimens as neotype:
Adult males:
A medium to large rupicolous Cordylus with a moderately depressed head and body. Referred to Cordylus (rather than any other species of Cordylidae) by the following combination of characters: head distinct from body, two pairs of large and well developed limbs (body serpentiform, head indistinct from body, and limbs rudimentary in Chamaesaura Schneider, 1799), scales on back large and keeled (granular in Platysaurus Smith, 1844, partly granular in Pseudocordylus Smith, 1838 and Hemicordylus Smith, 1838), non-spinose occipitals (spinose in Smaug
Cordylus angolensis is distinguishable from other members of its genus by the following combination of characters: (1) back dark brown with a paravertebral series of pale markings; (2) top of head plain brown or with occasional pale blotches; (3) iris of the eye brown; (4) scales of the first transverse row of dorsals similar in appearance to those of the row behind; (5) loreal shield absent; (6) nostril pierced in the posterior part of a large nasal, situated behind the suture of rostral and first supralabial, usually well separated from both the first supralabial and the preocular; (7) a regular row of six enlarged, non-spinose occipitals; (8) Frontonasal separated from the frontal by a pair of prefrontals (each of which usually exceeds it in size) or in contact; (9) Anterior pair of parietals usually in contact anteriorly; (10) dorsolateral and lateral scales may be weakly to moderately spinose; (11) tail spinose, but more weakly so distally; (12) dorsal scale rows transversely 24–25; (13) dorsal scale rows longitudinally 20–23; (14) ventral scale rows transversely 22–24; (15) ventral scale rows longitudinally 14; (16) subdigital lamellae on 4th toe 14; (17) femoral pores per thigh 5–6 in males and females; (18) differentiated femoral scales [generation glands] per thigh in males 19–25; (19) premaxillary teeth 7.
Its status as a distinct species is supported by monophyly with high levels of support from a suite of three mitochondrial and six nuclear markers (see above); and it differs from C. ‘Mombolo’ (see below), the most similar species genetically and morphologically, by an uncorrelated ND2 p-distance of 9.22% (Table
It differs from most other Cordylus (except C. ukingensis [Loveridge, 1932], C. macropholis [Boulenger, 1910], and C. vittifer [which occasionally has a loreal]) in lacking a loreal. It differs from C. ukingensis and C. macropholis by virtue of its smooth (rarely with a few weak keels on a few scales) versus strongly keeled (even spinose in C. macropholis) gulars, as well as by having its nostril pierced near the middle of the posterior part of the nasal scale (versus infero-posteriorly). Differs from C. tropidosternum by having smooth versus keeled gulars, and having the nostril well separated from the first supralabial (not in contact or near-contact); from C. rhodesianus by having distinctly rugose versus finely rugous to smooth upper head shields, 24–25 versus 25–29 transverse rows of dorsals, and a straight versus curved sulcus dividing the posterior part of the nasal; from C. jonesii by having its nostril pierced halfway up the posterior edge of the nasal rather than towards the centre, and a straight versus curved sulcus dividing the posterior part of the nasal; and from C. marunguensis which has the nostril pierced centrally on the lower margin of the nasal. Distinguished from C. vittifer and C. machadoi, C. namakuiyus and C. phonolithos by always lacking a loreal scale, having most or all scales of the first transverse row of dorsals of similar length (rather than longer) than those of the next row, and by having a pair of paravertebral rows of pale greenish-cream spots or blotches versus a lack of these. Most similar to C. ‘Mombolo’ (see below).
Meristic data for scalation characters in Cordylus angolensis and Cordylus momboloensis sp. nov. from Angola. When different, the number of scales on the left and right sides of the head are separated by a slash symbol (/). For occipitals, when the row is medially interrupted by post-occipitals, the number of scales on the left and right sides of the head respectively are separated by a plus sign (+).
Cordylus angolensis | Cordylus momboloensis sp. nov. | |||||||||||||||||
|
|
CHL 611 | CHL 613 | CHL 615 |
|
|
FKH 0127 (Paratype) | FKH 0128 (Paratype) | AMN R47333 (Paratype) |
|
FKH 0125 (Paratype) | AMN R47331 (Paratype) | AMN R47332 (Paratype) | AMN R47334 (Paratype) | AMN R47335 (Paratype) |
|
FKH 0129 | |
Sex | Male | Male | Male | Male | Female | Male | Male | Male | Male | Male | Female | Female | Female | Female | Female | Female | Female | Juvenile |
Supralabials | 6 | 6 | 6 | 6 | 6 | 6/5 | 5/6 | 6 | 6 | 6/5 | 6 | 6 | 5 | 6 | 5 | 5 | 5/7 | 6 |
Infralabials | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | – | 6 | 6 | 5/4 | 6/5 | 5 | 5 | 6 | 7/6 |
Sublabials | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 | 5 |
Supraoculars | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 |
Supraciliaries | 3 | 3 | 3 | 4 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 |
Suboculars | 3 | 3 | 4/3 | 3/4 | 3/4 | 4/3 | 3 | 3 | 3 | 3 | 2/3 | 3 | 4 | 3 | 3 | 3 | 3 | |
Occipitals* | 6 | 6 | 6 | 6 | 6 | 2+3 | 2+2 | 3+3 | 3+2 | 2+2 | 2+2 | 2+3 | 2+2 | 2+2 | 2+2 | 2+2 | 2+2 | 2+2 |
Post-occipitals | — | — | — | — | — | 6 | 6 | 6 | 5 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 |
Gulars transversely | 20 | 20 | 20 | 20 | 21 | 26 | 19 | 19 | 20 | 18 | 19 | 21 | 22 | 21 | 16 | 18 | 23 | |
Gulars contacting 1st pair sublabials | 5 | 5 | 6 | 5 | 5 | 5 | 5 | 5 | 4 | — | 4 | 5 | 5 | 5 | 4 | 4 | 4 | 5 |
Dorsals transversely | 25 | 25 | 24 | 24 | 24 | 22 | 24 | 23 | 23 | 22 | 24 | 24 | 22 | 22 | 23 | 22 | 24 | |
Dorsals longitudinally | 20 | 20 | 21 | 22 | 23 | 23 | 22 | 21 | 20 | 18 | 22 | 22 | 21 | 21 | 20 | 18 | 20 | |
Ventrals transversely | 22 | 23 | 22 | 23 | 24 | 22 | 26 | 26 | 25 | 23 | 27 | 25 | 23 | 23 | 23 | 22 | 22 | |
Ventrals longitudinally | 14 | 14 | 14 | 14 | 14 | 12 + 2 | 12 | 12 | 11 | 14 | 12 | 12 | 12 + 2 | 12 + 2 | 13 | 13 | 12 + 2 | |
Scales under 4th toe | 14 | 14 | 14 | 14 | 14 | 13/14 | 13 | 12 | 15 | 14 | 13 | 13 | 11 | 13 | 13 | |||
Femoral pores | 6/5 | 6/5 | 6/5 | 6 | 6/5 | 7 | 8/7 | 8/7 | 7/6 | 5 | 4 | 5 | 5 | 6 | 0 | 5 | 7/5 | |
Differentiated femoral scales | 21/20 | 19 | 21/19 | 25 | 0 | 26/27 | 30/26 | 37/35 | 30/29 | 30/25 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Mensural data and body ratios of Cordylus angolensis and Cordylus momboloensis sp. nov. from Angola (r = regenerated tail; w = width; l = length).
Cordylus angolensis | Cordylus momboloensis sp. nov. | ||||||||||||||||
|
|
CHL 611 | CHL 613 | CHL 615 |
|
|
FKH 0127 (Paratype) | FKH 0128 (Paratype) |
|
|
FKH 0125 (Paratype) |
|
|
|
|
|
|
Sex | Male | Male | Male | Male | Female | Male | Male | Male | Male | Male | Female | Female | Female | Female | Female | Female | Female |
SVL | 88.37 | 84.99 | 88.54 | 82.89 | 103.07 | 85.06 | 90.53 | 88.68 | 88.21 | 76.14 | 89.53 | 88.83 | 87.06 | 87.11 | 77.3 | 60.7 | 105.97 |
Tail length | 62.78r | 66.19 | 73.64 | 79.54 | 84.14 | 77.48r | 88.36 | 84.43 | 66.65 | 66.60r | 85.44 | 85.54 | 75r | 76.59 | 72.3 | 56.2 | 113 |
Head length | 24.61 | 21.93 | 23.24 | 20.54 | 25.72 | 25.3 | 23.07 | 23.32 | 22.54 | 22.34 | 22.30 | 22.36 | 24.64 | 23.98 | 20.63 | 17.10 | 27.74 |
Head width | 20.86 | 19.80 | 22.08 | 20.01 | 22.34 | 21.35 | 21.33 | 19.50 | 19.75 | 20.70 | 20.53 | 20.52 | 20.29 | 20.07 | 19.6 | 14.50 | 22.19 |
Head height | 10.56 | 9.76 | 10.92 | 10.71 | 10.33 | 9.22 | 11.93 | 11.34 | 9.37 | 9.28 | 11.80 | 11.39 | 10.40 | — | 8.15 | 7.61 | 10.41 |
Ear length | 7.82 | 7.14 | 7.50 | 5.53 | 6.96 | 6.78 | 6.99 | 6.96 | 6.10 | 6.06 | 8.80 | 6.92 | 6.52 | 6.65 | 5.96 | — | 5.98 |
Orbital length | 4.00 | 4.89 | 4.62 | 4.22 | 4.16 | 4.60 | 4.47 | 4.29 | 4.33 | 4.47 | 4.41 | 4.45 | 3.20 | 4.89 | 4.06 | — | 4.05 |
Nostril to eye | 3.19 | 3.07 | 3.79 | 3.92 | 4.15 | 4.67 | 4.03 | 3.91 | 4.31 | 3.07 | 2.93 | 3.58 | 5.15 | 4.56 | 4.46 | — | 5.41 |
Eye to ear | 11.65 | 10.32 | 11.03 | 11.13 | 12.08 | 11.37 | 10.80 | 9.26 | 8.46 | 9.15 | 11.34 | 9.71 | 11.12 | 10.12 | 10.04 | — | 11.90 |
Snout to eye | 7.15 | 8.51 | 8.16 | 7.25 | 7.30 | 9.90 | 7.99 | 7.14 | 7.65 | 9.14 | 6.06 | 6.66 | 9.82 | 9.33 | 8.24 | 7.16 | 10.85 |
Internarial | 6.55 | 4.54 | 4.92 | 4.40 | 4.73 | 4.95 | 5.40 | 4.77 | 5.02 | 5.08 | 5.11 | 4.94 | 5.11 | 4.29 | 4.84 | — | 5.64 |
Interorbital | 10.90 | 10.99 | 9.04 | 8.43 | 8.88 | 10.6 | 8.63 | 8.24 | 8.78 | 8.67 | 10.51 | 9.50 | 10.74 | 9.96 | 10.63 | — | 11.76 |
Snout to arm | 32.14 | 29.80 | 32.91 | 30.99 | 32.47 | 32.33 | 35.04 | 34.13 | 36.57 | 31.85 | 31.28 | 29.91 | 32.73 | 33.99 | 32.60 | 25.95 | 36.72 |
Axilla to groin | 38.23 | 40.04 | 35.90 | 35.40 | 51.37 | 36.09 | 40.15 | 38.97 | 35.43 | 35.01 | 44.29 | 44.53 | 42.84 | 43.80 | 33.32 | 29.66 | 53.50 |
Forearm length | 15.65 | 13.05 | 16.48 | 14.20 | 16.84 | 19.8 | 18.58 | 17.90 | 17.57 | — | 17.73 | 17.67 | 20.90 | 19.01 | 19.00 | — | 23.30 |
Fourth toe | 10.28 | 11.39 | 10.38 | 11.41 | 12.58 | 10.22 | 11.03 | 12.66 | 12.65 | 12.02 | 11.31 | 12.16 | 10.75 | 10.40 | 11.11 | 8.74 | 11.93 |
Tail length/SVL | 0.710 | 0.779 | 0.832 | 0.960 | 0.816 | 0.912 | 0.976 | 0.952 | 0.756 | 0.875 | 0.954 | 0.963 | 0.861+ | 0.879 | 0.935 | 0.926 | 1.066 |
Head length/SVL | 0.278 | 0.258 | 0.262 | 0.248 | 0.250 | 0.297 | 0.255 | 0.263 | 0.256 | 0.293 | 0.249 | 0.252 | 0.283 | 0.275 | 0.267 | 0.282 | 0.262 |
Head width/SVL | 0.236 | 0.233 | 0.249 | 0.241 | 0.217 | 0.251 | 0.236 | 0.220 | 0.224 | 0.272 | 0.229 | 0.231 | 0.233 | 0.230 | 0.254 | 0.239 | 0.209 |
Head w/Head l | 0.848 | 0.903 | 0.950 | 0.974 | 0.869 | 0.845 | 0.925 | 0.836 | 0.876 | 0.927 | 0.921 | 0.918 | 0.823 | 0.837 | 0.950 | 0.848 | 0.800 |
Orbital/Head l | 0.163 | 0.223 | 0.199 | 0.205 | 0.162 | 0.182 | 0.194 | 0.184 | 0.192 | 0.200 | 0.198 | 0.199 | 0.130 | 0.204 | 0.197 | — | 0.146 |
Forearm/SVL | 0.177 | 0.154 | 0.186 | 0.171 | 0.163 | 0.233 | 0.205 | 0.202 | 0.199 | — | 0.198 | 0.199 | 0.240 | 0.218 | 0.246 | — | 0.220 |
Dorsal scales rectangular, rugose, moderately keeled (less so medially), seldom spinose or mucronate, not serrated at the posterior edges; laterals oval, juxtaposed, rugous, sharply keeled and moderately spinose, no additional spines on the free end of scales; dorsals plus laterals in 25 transverse rows and 20 longitudinal rows (vertebral scales slightly smaller); on the central part of the belly the mesial ventrals are rectangular (transversely) and larger than others, the next row on either side with slightly rectangular scales, including the next two rows on either side; ventrals mostly smooth, but 2–3 lateral rows on either side with some obtusely keeled scales; ventrals in 22 transverse and 14 longitudinal rows (plus a row of oval, keeled scales on either side); a pair of enlarged and somewhat oval pre-cloacal plates is followed anteriorly (before the ventrals) by two transverse rows of much smaller irregular scales.
Scales on limbs above are large, strongly keeled and spinose; scales under fourth (longest) toe 14 on left and right feet; femoral pores 6 on left thigh, 5 on right, with distinct plugs of yellowish secretion; differentiated glandular femoral scales on thigh 21 on left, 20 on right. Tail with whorls of large, elongate, strongly keeled, spinose (spines directed backwards and longest superolaterally), weakly serrated, scales; supracaudals strongly keeled throughout most of their length, subcaudals basally distinctly keeled mainly on the distal half. Scales on palms of hands and soles of feet moderately to obtusely keeled; supradigital scales of hands smooth to weakly keeled, of feet smooth to moderately keeled; subdigital scales of hands weakly keeled, of feet moderately keeled.
Colour: The colour pattern of the neotype in life is similar to that described for the holotype (see above). The back is a mixture of light and dark brown, with some of the darker markings forming ill-defined transverse bands, and there is a series of greenish-cream blotches, mostly elongated, arranged in close proximity paravertebrally (about eight pairs). Top of the head is a mixture of pale and dark brown, with an ill-defined pale median band longitudinally. The upper parts of the tail are brown, with some keels dark brown. Belly is dirty white and the throat cream.
Cranial skeleton (Fig.
Postcranial skeleton: Segmented mesh file of
Osteoderms: (Fig.
(Figs
Colour: Similar to that described for the neotype, but CHL 611 and CHL 615 have a few scattered cream markings on the top of the head, and none have a pale median band.
Size: Largest male (CHL 611, Taqueta Mountain): SVL 88.5 mm, tail length 73.6 mm (original), total length 162.1 mm, head length (HL) 23.2 mm, head width (HW) 22.1 mm, head height (HH) 10.9 mm. Largest female (CHL 615, Taqueta Mountain): SVL 103.1 mm, tail length 84.1 mm (original), total length 187.2 mm, HL 25.7 mm, HW 22.3 mm, HH 10.3 mm.
Although no specific habitat information was available in
Although the original type locality is given as Caconda (Huíla Province), efforts over more than one century to rediscover the species at or near that locality have failed. The fact that Anchieta was based at Caconda for more than 20 years (
Geographical distribution of Cordylus angolensis (dark blue) and C. momboloensis sp. nov. (green) in Angola. The distribution of the three species of the C. machadoi group (C. machadoi [red], C. namakuiyus [pale blue], C. phonolithos [yellow]) is also indicated. Type localities are indicated by stars; for C. angolensis, the original type locality of Caconda is indicated by a black star, while the neotype locality is indicated by a dark blue star. Locality records are from the present study (see species accounts and Appendix 1) as well as
Angola: Caconda (–13.7199°, 15.0648°), Taqueta Mountain (–13.7778°, 14.1794°).
This species is known from only one locality, and little is known about its biology or population trends. Nevertheless, it should be noted that montane habitats in Angolan highlands are currently under anthropogenic pressure due to widespread excessive burning, cutting of remnant forests, and overgrazing of grasslands, all of which may negatively affect the species. The actual range of this species is difficult to infer and threats cannot be fully assessed at this time, but a Data Deficient categorisation, based on IUCN (2022a) Standards and Petitions Committee criteria, may apply.
It was not possible to locate the specimen of ‘Zonurus cordylus’ from Aruwimi (presumably Aruwimi River, a tributary of the Congo River) in northern Democratic Republic of the Congo as documented by
Cordylus angolensis not Bocage, 1895:
Allotype
:
(Where variation in additional material falls outside the range of type material, this is indicated in parentheses.) A medium to large rupicolous Cordylus with a moderately depressed head and body. Referred to Cordylus (rather than any other species of Cordylidae) by the following combination of characters: head distinct from body, two pairs of large and well developed limbs (body serpentiform, head indistinct from body, and limbs rudimentary in Chamaesaura), scales on back large and keeled (granular in Platysaurus, partly granular in Pseudocordylus and Hemicordylus), non-spinose occipitals [and post-occipitals] (occipitals spinose in Smaug), 23–24 transverse dorsal scale rows (40–43 in Ninurta; 31–46 in Karusasaurus; 15–16 in Ouroborus); loreal absent (present in Smaug, Ninurta, Ouroborus, Karusasaurus, and Namazonurus).
Cordylus momboloensis sp. nov. is distinguishable from other members of its genus by the following combination of characters: (1) back dark brown with a paravertebral series of pale markings; (2) top of head with pale blotches; (3) iris of the eye mostly pale green (blue-green to some eyes), with a brown ring around the pupil; (4) scales of the first transverse row of dorsals similar in appearance to those of the row behind; (5) loreal shield absent; (6) nostril pierced in the posterior part of a large nasal, situated behind the suture of rostral and first supralabial, always well separated from the first supralabial, and usually separated from the preocular; (7) an interrupted row of non-spinose occipitals consisting of 2–3 scales on either side of head; (8) a row of 5–6 non-spinose post-occipitals, the median scales of which are in contact with the posterior parietals and separate the occipitals; (9) Frontonasal separated from the frontal by a pair of prefrontals, each of which exceeds it in size; (10) Anterior pair of parietals usually in contact anteriorly; (11) dorsolateral and lateral scales weakly to moderately spinose; (12) tail spinose, more weakly so distally; (13) dorsal scale rows transversely 22–24; (14) dorsal scale rows longitudinally 18–23; (15) ventral scale rows transversely 22–27; (16) ventral scale rows longitudinally 12–13 [rarely 11 or 14]; (17) subdigital lamellae on 4th toe 11–15; (18) femoral pores per thigh 5–8 in males, 4–6 in females [rarely 0; 7 on one side in a non-type specimen,
Its status as a new species is supported by monophyly with high levels of support from a suite of three mitochondrial and six nuclear markers (see above); and it differs from C. angolensis, the most similar species genetically and morphologically, by an uncorrelated ND2 p-distance of 9.22% (Table
The new species differs from all other Cordylus in having nine (rather than seven, rarely six) premaxillary teeth, and from most other congeners (except C. angolensis, C. ukingensis, C. macropholis, and C. vittifer [occasionally has a loreal]) in lacking a loreal. It differs from C. ukingensis and C. macropholis by virtue of its weakly keeled or smooth versus strongly keeled (even spinose in C. macropholis) gulars, as well as by having its nostril pierced near the middle of the posterior part of the nasal scale (versus infero-posteriorly). Differs from C. tropidosternum by having smooth or weakly keeled versus keeled gulars, and having the nostril well separated from the first supralabial (not in contact or near-contact); from C. rhodesianus by having distinctly rugose versus finely rugose to smooth upper head shields, 22–24 versus 25–29 transverse rows of dorsals, and a straight versus curved sulcus dividing the posterior part of the nasal; from C. jonesii by having its nostril pierced halfway up the posterior edge of the nasal rather than towards the centre; and a straight versus curved sulcus dividing the posterior part of the nasal; and from C. marunguensis which has the nostril pierced centrally on the lower margin of the nasal. Distinguished from C. vittifer, C. machadoi, C. namakuiyus and C. phonolithos by always lacking a loreal scale, having most or all scales of the first transverse row of dorsals of similar length (rather than longer) than those of the next row, and by having a pair of paravertebral rows of pale greenish-cream spots or blotches versus a lack of these.
The new species is most similar to C. angolensis, but distinguished from it as follows: in live specimens the iris of the eye is largely pale green to blueish-green versus brown; top of head distinctly marked by pale blotches versus mostly plain brown; throat usually dirty white versus cream; a medially-interrupted row of occipital scales (2–3 on either side of head) is present, never forming a continuous row, each scale usually smaller than a post-occipital versus a continuous row of 6 occipitals across the back of the head; lower numbers of ventral scale rows longitudinally (usually 12–13 vs 14); relatively longer forearms (forearm/SVL 0.20–0.25 versus 0.15–0.19); higher numbers of generation glands in males (25–37 per thigh versus 19–25); and higher numbers of premaxillary teeth (nine versus seven).
Dorsal scales closely-set but often juxtaposed, rectangular, rugose, moderately keeled (less so on middle of back), some are also weakly spinose dorso-laterally, occasionally weakly serrated at their posterior edges; laterals occasionally subimbricate, often oval, rugose, sharply keeled and moderately spinose, no additional spines or serrations on the free ends of scales; dorsals (including laterals) in 22 transverse rows (excluding one half-row) and 23 longitudinal rows (including row of small vertebral scales which is interrupted at parts); on the central part of the belly the paired rows of mesial ventrals are rectangular (transversely), the next row on either side with only slightly transversely rectangular scales, the others being squarish, with two rows of lateral ventrals on either side consisting of longitudinally-rectangular scales; ventrals mostly smooth, but the outermost lateral row on either side, and an additional row of ‘pseudo-ventrals’ on either side, with some obtusely keeled scales; ventrals in 22 transverse and 12 longitudinal rows (excluding a row of oval, keeled scales on either side best considered ‘pseudo-ventrals’); a pair of enlarged pre-cloacal plates is followed anteriorly (before the ventrals) by 2–3 transverse rows of much smaller plates.
Scales on upper parts of hindlimbs are large, strongly keeled and strongly spinose; scales of forelimbs large, moderately keeled and moderately spinose; scales under fourth (longest) toe 13 on left foot, 14 on right; scales under fourth (longest) finger 10 on left hand, 11 on right; femoral pores 7 on each thigh, of moderate size with distinct plugs of yellowish secretion; differentiated glandular femoral scales on thighs 26 on left, 27 right. Tail whorled, dorsally with large, elongate, rugose, strongly keeled, spinose, weakly serrated scales, with spines directed backwards and longest superolaterally; supracaudals strongly keeled throughout most of their length, subcaudals basally distinctly keeled mainly on the distal half of tail. Scales on palms of hands obtusely keeled, on soles of feet moderately keeled; supradigital scales of hands weakly keeled, of feet moderately keeled; subdigital scales of hands and feet weakly keeled or smooth.
Colour of holotype (in life): The upper parts of the head, back, flanks, limbs and tail were mostly dark brown, with irregular pale or light greenish to blueish-green markings on either side of the midline (about six pairs) and on top of the head; venter whitish to cream; throat dirty white (Fig.
Holotype (
(n = 10). (Figs
External morphology: All paratypes are similar to the holotype, but differ as follows: SVL 76.1–90.5 mm (males), 60.7–89.5 mm (females). Original tail shorter than SVL (87.5–97.6%, n = 9). Head 1.05–1.21 times as long as wide, head height 0.416–0.582 times head width. Nasals in narrow contact in FKH 0125,
Dorsals plus laterals in 22–24 transverse rows and 18–22 longitudinal rows; ventrals in 22–27 transverse and 12–13 [11 in FKH 0128, 14 in
Scales under fourth (longest) toe 11–15, under fourth (longest) finger 10–12 [
Colour of paratypes (in life): Colouration of the Sandula and Vondo paratypes was similar to that of the holotype. The belly had a slightly blueish tinge.
Colour of paratypes (in preservative):
Cranial skeleton: (Fig.
Postcranial skeleton: Segmented mesh files of
Osteoderms (Fig.
(Figs
External morphology:
FKH 0129 (juvenile). Data for head only, see Figs
Colour of additional material (in life).
Largest male (
The name ‘momboloensis’ derives from the local name (“m’bolo”) for Mombolo which in Umbundo – main language in central Angola – is applied to bread or staple food. It is a reference to the fact that the region in question consists of a fertile and relatively large highland plateau (surrounded by mountains, in Luanza Sul Province). Most of the available specimens, including the holotype as well as the first series of specimens of this species collected during the Vernay Angola Expedion in 1925, were collected in Mombolo.
Although no specific habitat information is available for the material housed at the
This species was collected at Sandula (Monte Verde) and Vondo (Mount Uassamba), both in Cuanza Sul Province on high mountain peaks at 1975–2095 m a.s.l., and an individual was briefly observed at Morro do Pundo (–12.4464°, 13.9171°) in Benguela Province, at approximately 1000 m elevation near the town of Bocoio (Fig.
The
Little is known about the biology of this species, but habitat at the four sites where it has been found appears to be relatively undisturbed. Given the current knowledge, if an assessment was to be done, it is likely to result in a classification of Least Concern based on IUCN (2022a) Standards and Petitions Committee criteria.
1a | Loreal present; nostril situated above the suture between rostral and first supralabial; scales of the first transverse row of dorsals mostly elongated | 2 |
1b | Loreal absent; nostril situated posterior to suture between rostral and first supralabial; scales of the first transverse row of dorsals not elongated, similar to those of subsequent rows | 4 |
2a | Interparietal small, not completely separating the anterior parietals; ventral osteoderms absent | C. machadoi |
2b | Interparietal large and usually in contact with the frontoparietals, separating the anterior parietals; ventral osteoderms present | 3 |
3a | Femoral pores in adults 4–6 per leg | C. namakuiyus |
3b | Femoral pores in adults 7–8 per leg | C. phonolithos |
4a | Iris of the eye brown; occipitals in a regular row of six scales; ventral scales in 14 rows longitudinally; generation glands in males 19–25 per leg; forearm/SVL 0.15–0.19 | C. angolensis |
4b | Iris of the eye pale green or blue-green; occipitals 2–3 on either side of head, interrupted medially by a row of 5–6 post-occipitals; ventral scales in 12–13 (rarely 11 or 14) rows longitudinally; generation glands in males 25–37 per leg; forearm/SVL 0.20–0.25 | C. momboloensis sp. nov. |
The most recent phylogeny of the genus Cordylus indicated a distinct Angolan clade consisting of three species: C. machadoi, C. namakuiyus and C. phonolithos (
In the present study we analysed lizards from the west-central highlands of Angola and placed them in a phylogenetic context with their congeners, including the three species in the C. machadoi group. Cordylus angolensis and C. momboloensis sp. nov. form a sister clade to the C. machadoi group. Within the latter, C. machadoi (high elevation escarpment) and C. phonolithos (Serra da Neve inselberg) group together, rather than C. machadoi + C. namakuiyus (arid lowlands) as indicated by
Cordylus angolensis and C. momboloensis sp. nov. are superficially similar species of comparable size, colour pattern and scutellation. Nevertheless, we show that they differ with regard to the arrangement of occipitals/post-occipitals, numbers of ventral scale rows longitudinally, relative forearm length, numbers of generation glands in males, and in terms of eye colour – a character not usually mentioned in cordylid species comparisons (see Diagnosis above). There are also differences between C. angolensis and C. momboloensis sp. nov. with regard to the skeleton and osteoderms: 22–23 maxillary teeth on either side versus 18–21 respectively; premaxillary teeth 7 versus 9; and in C. angolensis the posterior process of the parietal is much longer compared to C. momboloensis sp. nov. Osteoderm distributions of C. momboloensis sp. nov. differ between the two specimens scanned, but greater sampling is needed to ascertain whether this is a result of ontogenetic, geographic or stochastic variation.
Our phylogeny agrees largely with that of
Recent studies on other rupicolous lizards in the south-western quarter of Angola have indicated similar distribution patterns to Cordylus. In both Angolan Afroedura and Cordylus, the main divide splits one clade in the west-central highlands and one clade widely distributed along the south-western coastal plain and escarpment. The six species of Angolan flat geckos (
These similar distribution patterns strongly suggest that the same evolutionary forces may have shaped the speciation history of these genera in Angola. The Angolan west-central highlands, corresponds to the marginal mountain chain or ancient massif (
Despite several recent studies on the herpetofauna of Angola (see references in Introduction), more surveys will be required to allow an evaluation of the true distribution of both C. angolensis and C. momboloensis sp. nov., and to investigate the possibility of additional congeners occurring in the country. The highlands of central Angola are still relatively poorly known biologically and barely explored, but it is likely that future work will reveal similar speciation patterns in other rupiculous taxa, which may significantly contribute to a better interpretation of Angolan biogeography in general. Our findings underline the importance of the Angolan west-central highlands in functioning as a regional reservoir of biodiversity, and highlight the need to conserve its highly threatened montane habitats.
Our evaluation of genetic and morphological data indicates that two species of allopatric, rupicolous Cordylus, C. angolensis and C. momboloensis sp. nov., occur in the west-central highlands of Angola. Cordylus angolensis was until now known only from the holotype collected 128 years ago. We also show that these two species form a clade most closely related to the three species of the C. machadoi complex in south-western Angola, and together the Angolan assemblage of Cordylus is a distinct clade most closely related to east African congeners, especially C. jonesii + C. rhodesianus.
We thank the following individuals: Agnes Phindane (National Museum, Bloemfontein) for measuring non-Angolan lizards and assisting with calculations of body size; Werner Conradie (Port Elizabeth Museum) for a few measurements on the holotype of C. momboloensis sp. nov.; Lauretta Mahlangu (Ditsong National Museum of Natural History, Pretoria), David Kizirian (American Museum of Natural History, New York), and Seth Eiseb and Mathilde Awases (National Museum of Namibia, Windhoek) for loans of Angolan and Namibian Cordylus specimens; Afonso Vaz Pinto and Ninda Baptista (Centro de Investigação em Biodiversidade e Recursos Genéticos, Portugal) for field support to collect specimens for this study; the late Wulf Haacke (Pretoria) for information about Angolan collecting localities and for supplying photographic images of Angolan specimens; Frank Tillack (Museum für Naturkunde der Humboldt-Universität, Berlin) for information about the holotype of C. vittifer and for supplying photographs of this specimen; Aaron Bauer (Villanova University, Villanova) for providing copies of old literature; René Navarro (University of Cape Town) for preparing the C. vittifer map; Luis Verissimo (Catholic University of Angola, Luanda) for information on locality names, habitat and elevations in Angola; Werner Conradie (Port Elizabeth Museum, Bayworld), Krystal Tolley (South African National Biodiversity Institute, Cape Town) and John Measey (Stellenbosch University, Stellenbosch), field companions of the late WRB in Angola; Alberto Sanchez-Vialas for providing detailed photographs of the neotype of C. angolensis and other material deposited at Museo Nacional de Ciensias Naturales, Madrid; Luis Ceriaco for photographs of the holotype of C. phonolithos; David Maguire, Darren Pietersen and Luke Verburgt for photographs of live cordylids; and Fatima Linares (Centro de Instrumentación Científica of Granada) for CT scanning of the neotype of C. angolensis. JLR was supported by Fundação para a Ciência e Tecnologia (FCT) contract PD/BD/140808/2018 and BIOPOLIS 2022–18. Comparative CT material was produced as part of the NSF oVert TCN (NSF:1701714).
Specimens of Cordylus examined for this study: extralimital species and species in the C. machadoi group. Localities are listed alphabetically by country or province; for South Africa the farm name is followed by number and magisterial district; museum catalogue numbers are followed in parentheses by mapping co-ordinates or quarter-degree grid cell codes (usage explained in
Cordylus beraduccii
KENYA. 20.4 km North of Kajiado, Rift Valley Province –
Cordylus jonesii
SOUTH AFRICA. Limpopo Province: Doorenwaard 313 Vhembe district –
Cordylus machadoi
ANGOLA. 5 km SSE of Chibemba, Huíla Province –
Cordylus cf. machadoi
NAMIBIA. Baynes Mountains, Kunene Region –
Cordylus marunguensis
DEMOCRATIC REPUBLIC OF CONGO. Marungu Mountains, Katanga Province –
Cordylus namakuiyus
ANGOLA. “Angola” –
Cordylus nyikae
MALAWI. Nyika Plateau, Northern Region –
Cordylus phonolithos
ANGOLA. Namibe Province: Serra da Neve, N’Dolondolo –
Cordylus rhodesianus
ZIMBABWE. Manicaland Province: Nyanga district: Chinaka farm –
Cordylus rivae
ETHIOPIA. East of Negelle, Oromia Region – TJC 0564*.
Cordylus tropidosternum
MALAWI. Chitipa district, near Jembya Forest Reserve, Northern Region –
Cordylus ukingensis
TANZANIA. Tandala, Ukinga Mountains, Njombe Region –
Cordylus vittifer variety A (Free State and Gauteng provinces) (1 in Fig. S3)
SOUTH AFRICA. Free State: Annie’s Rust 763, Heilbron district –
Cordylus vittifer variety B (Free State Province) (2 in Fig. S3)
SOUTH AFRICA. Free State, Vrede district: Berlin 497 –
Cordylus vittifer variety B (Limpopo Province) (3 in Fig. S3)
SOUTH AFRICA. Limpopo Province: Bloemhof 4, Potgietersrus district –
Cordylus vittifer variety B (Mpumalanga Province and Eswatini) (4 in Fig. S3)
SOUTH AFRICA. Mpumalanga: De Goedeverwachting 57, Carolina district –
Cordylus vittifer variety C (Free State Province) (5 in Fig. S3)
SOUTH AFRICA. Free State, Harrismith district: Tafelberg ‘A’ 1312 –
Cordylus vittifer (KwaZulu-Natal Province) (6 in Fig. S3)
SOUTH AFRICA. KwaZulu-Natal: Dewar 2400, Dundee district –
Figure S1
Data type: .tif
Explanation note: Paratype of Cordylus machadoi (
Figure S2
Data type: .tif
Explanation note: Holotype (ZMB 10762) of Zonurus vittifer. A–D Lateral, ventral and dorsal aspects of the head; E, F: dorsal and ventral aspects of the body; G, H femoral region of the right and left legs, respectively. Photos: Frank Tillack.
Figure S3
Data type: .png
Explanation note: Geographical distribution of Cordylus vittifer in South Africa and Eswatini. Shaded areas depicting the species overall distribution are based on
Figure S4
Data type: .jpg
Explanation note: Ventral aspect of Cordylus machadoi (
Figure S5
Data type: .pdf
Explanation note: Allometry plot of relative osteoderm volume (osteoderm-postcranial skeleton volume ratio) against snout-vent length (mm) in Angolan Cordylus. 1) Cordylus machadoi
Table S1
Data type: .xlsx
Explanation note: Scan settings and Morphosource DOIs for CT datasets of Angolan Cordylus studied in this description. Scan location codes: NRF-UF = Nanoscale Research Facility, University of Florida, USA. MIF-AMNH = Microscopy Imaging Facility, American Museum of Natural History, New York, USA. GE-SC = General Electric Inspection Technologies, LP Technical Solutions Center, San Carlos, California, USA. CIC = Centro de Instrumentación Científica of Granada, Spain.
Table S2
Data type: .xlsx
Explanation note: Comparative meristic data for Cordylus from the Angolan clade and various species and populations in the East African clade (including C. jonesii–C. rhodesianus). The first row of data refers to the range of values, the second row is the mean, and the third row is the standard deviation. For C. phonolithos, data is for the holotype and paratype (
Table S3
Data type: .xlsx
Explanation note: Premaxillary, maxillary and dentary teeth counts for Angolan and east African Cordylus.