622 results on '"Tolley, Krystal"'
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102. African Herald snakes,Crotaphopeltis, show population structure for a widespread generalist but deep genetic divergence for forest specialists
- Author
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Engelbrecht, Hanlie M., primary, Branch, William R., additional, Greenbaum, Eli, additional, Burger, Marius, additional, Conradie, Werner, additional, and Tolley, Krystal A., additional
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- 2020
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103. Taxonomic inflation due to inadequate sampling: are girdled lizards (Cordylus minor species complex) from the Great Karoo one and the same?
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Tolley, Krystal A, Telford, Nicolas S, Taft, Jody M, Bates, Michael F, Conradie, Werner, Makhubo, Buyisile G, and Alexander, Graham J
- Subjects
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LIZARDS , *INTERGLACIALS , *SPECIES , *PRICE inflation , *VICARIANCE , *BACTERIAL diversity - Abstract
The Great Karoo and Namaqualand of South Africa are home to a species complex of morphologically conserved lizards that occur in allopatry (Karoo: Cordylus aridus , Cordylus cloetei , Cordylus minor ; Namaqualand: Cordylus imkeae). However, there are negligible morphological differences and a lack of obvious physical or climatic barriers, particularly among the three Karoo species. We hypothesized that poor geographic coverage in previous studies and lack of an explicit species concept has caused taxonomic inflation. We therefore tested species boundaries by examining multiple criteria: multi-gene phylogenetics, niche distribution modelling and re-examination of diagnostic morphological features with a larger sample size. We found that C. aridus , C. cloetei and C. minor lack diagnosable differences for both genetics and morphology. Distribution modelling, ranging from present day to the last interglacial period, show connectivity has been maintained especially during cooler periods. Conversely, C. imkeae is morphologically diagnosable, genetically distinct and lacks connectivity with the other taxa. By evaluating multiple operational criteria, we conclude that the C. minor species complex comprises only two species, C. minor (with C. aridus and C. cloetei as junior synonyms) and C. imkeae , demonstrating that species defined from inadequate data and lack of an explicit species concept can lead to taxonomic inflation. [ABSTRACT FROM AUTHOR]
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- 2022
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104. Stable climate corridors promote gene flow in the Cape sand snake species complex (Psammophiidae).
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Taft, Jody M., Maritz, Bryan, and Tolley, Krystal A.
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GENE flow ,QUATERNARY Period ,GENETIC variation ,INTERGLACIALS ,SPECIES distribution - Abstract
Climate shifts during the Quaternary Period have driven changes in regional range dynamics for many species, influencing population structure of species and in some cases promoting speciation. Within southern Africa, the psammophine snakes Psammophis trinasalis and P. namibensis were historically considered subspecies of P. leightoni but were elevated to species rank based on ecological differences. Preliminary phylogenetic analyses suggested intraspecific, not interspecific genetic variation between these taxa, but this finding was based on very limited data and could not be confirmed. To assess the level of genetic differentiation within the P. leightoni species complex, we explored the evolutionary history of these snakes by combining phylogenetic analyses, species distribution modelling and an examination of morphology. We generated a comprehensive, multi‐gene phylogeny for Psammophis that included wider geographic sampling of the three species in the complex. Using this phylogeny, Bayesian and distance‐based species delimitation analyses showed intraspecific, not interspecific divergences between taxa in the complex, suggesting that they collectively represent a single taxon. Furthermore, non‐metric multidimensional scaling analysis of scalation characters showed no differences between the species. Moreover, palaeo‐modelling at three time periods since the last interglacial period suggest that there have been varying levels of connectivity between these taxa, which has likely facilitated gene flow between them. Given the evidence, we propose that the P. leightoni complex represents a single species and therefore formally synonymise the three species. [ABSTRACT FROM AUTHOR]
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- 2022
- Full Text
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105. Nucras broadleyi Branch & Conradie & Pinto & Tolley 2019, sp. nov
- Author
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Branch, William R., Conradie, Werner, Pinto, Pedro Vaz, and Tolley, Krystal A.
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Nucras ,Nucras broadleyi ,Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Lacertidae ,Taxonomy - Abstract
Nucras broadleyi sp. nov. Angolan Sandveld Lizard urn:lsid:zoobank.org:act: C82E3A75-96FF-4D2A-9B52-3ABF4B58BC2B (Figures 4–6) Chersonymy. Nucras tessellata var. taeniolata (Bocage 1895: 30), Nucras tessellata var. taeniolata (Boulenger 1910: 474), Nucras tessellata var. holubi (Boulenger 1917: 210), Nucras intertexta var. holubi (Boulenger 1920: 20), Nucras tessellata (Monard 1937: 73; Laurent 1964: 56), Nucras ornata (Broadley 1965: 23), Nucras tessellata (Broadley 1972: 30; Ceríaco et al. 2016: 56; Burger 2014: 171), Nucras aff. tessellata (Marques et al. 2018: 221; Branch et al. 2019: 317). Type material. The type series is comprised of the three most recently collected specimens, which are housed in PEM and TM. Holotype. A subadult male (PEM R 24005, AG 018), 10 km west of Lola, edge of Bentiaba River valley, Namibe Province, Angola (-14.29028, 13.53056, WGS 84, 802 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J.S. de Almeida on 2 November 2015. Paratypes (2). a) A subadult female (PEM R 24157, AG 166), 8.8 km southwest of Farm Mucungo, Namibe Province, Angola (-14.80167, 12.41917, WGS 84, 385 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J. S. de Almeida on 8 November 2015. b) An unsexed adult (TM 40392), “ 34 km S of Moçâmedes to Porto Alexandre, Angola, 1512 Ca ” (= 34 km S Namibe to Tômbwa), Namibe Province, Angola (approx. -15.48220, 12.18289). Collected by W.D. Haacke on 30 March 1971. Additional referred material: The following additional material was used to expand the description of variation within the species: a) an adult male (MD 1967, Laurent 1964), “km 34 de la route de Moçâmedes à Sa da Bandeira” (= 34 km from Namibe on Lubango road, -15.03333, 12.41667), collected 24 October 1949, b) MBL 646, 647 a, 647b (Bocage 1895: 30) from Maconjo (approx. -15.01667, 13.20000), c) BM 1970.6.29.10–11 (Boulenger 1910: 474) from Ponang Kuma (= Donguena, approx. -17.01667, 14.71667), and d) MHNC 91.0524 (Monard 1937) from Capelongo (approx. -14.88333, 15.083333), collected April 1933. Etymology. The specific epithet is a patronym in honor of Donald G. Broadley for his numerous contributions to the herpetofauna of Africa. Don (as most of us knew him) was the first to recognize the Angolan population as a separate species (Broadley 1972). The name is constructed in the masculine genitive. Diagnosis. Assigned to Nucras due to a well-defined collar (absent in Ichnotropis), toes not serrated or fringed (versus serrated or fringed in Meroles), subdigital lamellae smooth (versus keeled in Pedioplanis and Heliobolus), subocular bordering lip, the nostril is pierced between two nasals, nasal well separated from upper labial, and dorsal scales small, smooth, and juxtaposed. * anterior subocular (posterior subocular), ** total (largest number, t = truncated) The new species can be diagnosed from other Nucras species based on a combination of the following characters: series of transversely enlarged plates present under forearm (versus absent or only feebly enlarged in Nucras lalandii), a small series (0–6) of small granules present between supraciliaries and supraoculars (versus mostly absent in N. boulengeri and N. lalandii), 23–29 lamellae under 4 th toe (versus less than 22 in N. lalandii), dorsum with a series of longitudinal pale stripes (versus dark cross bands present in N. lalandii and N. scalaris or a series of pale vertebral spots, sometimes forming irregular transverse bands in N. intertexta or lack of any dorsal patterns in N. aurantiaca), four pale stripes on nape with outer stripes forming a. Bloemfontein Museum Province Province Province Province Province Province Province Province Province National Cape Cape Cape Cape Cape Cape Cape NMB – , , Northern Western, Northern,, Northern Northern, Western,, Western Kamanjab, KwaZulu-Natal Cape , Western Museum Locality South Africa Africa South South Africa South Africa South Africa GenBank Africa South South Africa Namibia, South Africa Africa South Elizabeth Port – 1 005257 951543 951540 951542 951541 632229 871208 547691 871206 871207 547718 RAG PEM HG HF HF HF HF EF DQ HF DQ DQ HF, Science 4 005232 951535 951536 751398 547725 547732 547733 547759 of ND HG — HF — HF — FR HF HF HF HF Academy 005211 951559 951556 951558 951557 080358 871152 547772 871148 871149 547800 California S 16 HG HF HF HF HF AF DQ HF DQ DQ HF CAS – : number 206725 R 18745 R 16873 R 11574 R 16872 209602 R 18376 abbreviations Museum accession CAS PEM PEM NMB PEM NA NA NMNW CAS PEM Museum. accession 5584 08 069 - 20650 21061 20687 0531 08 6001 6067 049 analysis Field ID AMB KTH MB MB MB MH GW AMB AMB SVN genetic in Samples used Species tessellata tessellata tessellata tessellata tessellata longicaudata australis australis capensis capensis suborbitalis. ) continued 2 Table (Genus Nucras Nucras Nucras Nucras Nucras Outgroup Latastia Australolacerta Australolacerta Ichnotropis Ichnotropis Meroles continuous light stripe with the outer edges of the parietals (similar to Broadley’s (1972) N. tessellata tessellata var. “ T;” differs from N. livida and N. tessellata where the outer stripes often do not form a continuous light stripe with the outer edges of the parietals; differs from N. caeiscaudata and N. ornata where there are only three longitudinal stripes present on nape and sometimes the vertebral ones are absent), well defined occipital scale separating parietals (versus reduced or absent in northern Namibia N. holubi, which is referred to as N. intertexta damarana Parker; as well as absent in N. caesicaudata), parietal foramen absent (often present in all other species except N. taeniolata), and postnasals separated (usually fused in N. taeniolata). In the phylogenetic analysis, the uncorrected p -distances show that this clade differs by>8% for 16S,>14% for ND4, and>1% for RAG 1 sequence divergence from other members of the N. tessellata clade. Description of Holotype (Fig. 4). Body relatively slender (SVL approx. 4.5 times the head length, tail truncated), with hindlimbs larger than forelimbs (femur of hind limb equal to length of tibia); head narrow and elongated (56% longer than wide) with narrow pointed but blunt snout, that is slightly longer than distance from back of eye to rear of ear opening. Rostrum protruding and visible from below. Nasals paired and in contact (0.2 mm suture length), not swollen, nostril directed backwards separating postnasals. Frontonasal single, wider than long (1.1 × 1.8 mm). Prefrontals paired and in broad median contact with one another (0.6 mm suture length), wider than long (1.1 × 1.2 mm). Frontal entire, longer than wide (2.7 × 1.9 mm). Two large rounded supraoculars, both in contact with the frontal, with anterior supraocular preceded by a single large scale in contact with prefrontal, frontonasal, and posterior loreal, with posterior supraocular bordered by a single large scale in contact with parietal and frontoparietal. Paired frontoparietal in broad contact (1.3 mm suture length), nearly as wide as long (1.7 × 1.5 mm). Parietals twice as long as wide (3.1 × 1.8 mm), fully separate by a large, pentagonal interparietal (2.5 × 1.2 mm) that is twice as long as wide, slightly shorter than frontoparietals and nearly equal to length of frontonasal and prefrontal combined. Small subtriangular occipital (0.5 × 0.7 mm). Two loreals, second much larger than first. Six supraciliaries on each side, 1 st is the longest. A single minute granule scale between supraocular and supracilliares on right side, none on left side. Four supralabials anterior to subocular and three supralabials posterior to subocular, on both sides. Subocular slightly elevated medial and bordering the lip, its lower border being shorter than the upper. Three temporal scales, first longer than others, smooth. Tympanic shield as wide as long, border of ear opening. No ear lobes. Lower eyelid with transparent brille formed by five larger scales, surrounded by numerous smaller scales. Lower eyelid separated from subocular and enlarged temporal scales by a series of 10 smaller scales. Small scale above 3 rd supralabial separating the posterior loreal and subocular. Enlarged scale bordering 1 st post subocular, supralabial, and the subocular. Six infralabials on both sides, with 3 rd being longest; four enlarged pairs of chin shields, last largest and first three in broad contact. Twenty-four gular scales in a straight line between symphysis of chin shields and median collar plate, equal in size except last 4–5 larger. Collar free, comprising seven enlarged plates (median subtriangular) and extending slightly onto side of neck as a crease, bordered by 2–3 smaller scales. Dorsal scales small, juxtaposed, granular, smooth, larger on sides toward ventrals. Midbody scales 42. Ventral plates eight longitudinal and 28 transverse rows (from collar to groin), plates of the innermost rows longer than broad, with outer row notably smaller than other rows, transverse row of ventrals across chest just behind collar longer than broad; preanal scales irregular, median ones larger. Scales on upper surface of forearm large, smooth or slightly keeled. Scales on lower surface of forearm with eight enlarged plates, at least twice the width of scales on upper forearm. Scales on upper surface of tibia rhombic, subimbricate, smooth, and much larger than dorsal scales. Tibia below with a series of large plates. Subdigital lamellae under fourth toe 23 R / 25L. Femoral pores 13 R / 15L. Dorsal scales on tail oblique, strongly keeled diagonally, and truncate behind, ventral scales on tail obtusely keeled. Coloration. Dorsum with eight pale cream to white dorsolateral longitudinal stripes, separated by dark brown to black stripes. These stripes are more boldly patterned anteriorly, fading posteriorly. No light vertebral stripe. The two pale paravertebral stripes are separated by a very narrow strip of darker scales that starts on the interparietal through the occipital scale and fades posteriorly onto body and tail. The dorsolateral stripe extending along outer borders of parietals continues onto the tail. It is followed by the upper lateral stripe extending from posterior of the eye onto the head through the mid-temporal with a brief break above the ear opening, and continues onto the tail. The lower lateral stripe starts at the subocular, through the ear opening, broken briefly above the arm, after which it continues all the way onto the tail. Ventrum white and lower limbs oblique white. Fore limbs upper surface black with scattered pale blotches. Hind limbs light brown with pale blotches. Upper surface of tail red-brown, similar to hind limbs. Scales bordering the orbit are black edged. Variation (Figs. 5–6). Meristic and escalation data are summarized in Table 1. The largest specimen examined is (BM 1907.6.29.10) 74 + 144 mm (tail regenerated). Regarding coloration, there seem to be three main variations among material examined: 1) 8–9 longitudinal stripes as in holotype (in PEM R 24005, MBL 647a, 647b, MHNC 91.0524–5), 2) 4–5 pale longitudinal stripes broken up posteriorly with flanks spotted (in BM 1970.6.29.10–11, TM 40392, MD 1967), and 3) broken paravertebral stripes, continuous dorsolateral line and barred flanks (in PEM R 24157), similar to N. intertexta. Distribution. Found only in semi-arid south-western Angola, throughout much of Namibe Province and extending onto the escarpment of southern Hu í la and Cunene Provinces (Fig. 1). Known localities include: Maconjo (Bocage 1895: 30), Ponang Kuma (=Donguena) (Boulenger 1910: 472), 34 km from Namibe on Lubango road (Laurent 1964: 56), 34 km south of Tombwa (TM 40397), 8.8 km southwest of Farm Mucungo (this study), 10 km west of Lola (this study), and Capelongo (Monard 1937: 73). The locality of Caconda (Bocage 1895) extends the species distribution further north into Huíla Province, but the specimens could not be critically evaluated by Broadley (1972) and are now presumably lost. Habitat. The species appears to be associated with mopane woodlands, dry savannas, and semi-desert shrublands (Barbosa 1970). The new material was found in sandy plains with scattered low granite outcrops, with varying degrees of short grass cover and scattered bushes. Vegetation included Colophospermum mopane, Ficus sp., Senegalia (=Acacia) mellifera, Commiphora sp., Boscia foetida, and Salvadora persica. The confirmed historical records were also obtained within the dry woodland zone, even though the possible occurrence of the species in Caconda would place the species above 1,500 m asl and well into the mesic conditions of Brachystegia habitats (Barbosa 1970)., Published as part of Branch, William R., Conradie, Werner, Pinto, Pedro Vaz & Tolley, Krystal A., 2019, Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola, pp. 82-95 in Amphibian & Reptile Conservation 13 (2) on pages 86-91, DOI: 10.5281/zenodo.3731977, {"references":["Bocage JVB. 1895. Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, Portugal. 203 p., 20 pls.","Boulenger GA. 1910. A revised list of South African reptiles and batrachians, with synoptic tables, special reference of specimens in the South African Museum, and descriptions of new species. Annals of the South African Museum 5: 455 - 538.","Boulenger GA. 1917. A revision of the lizards of the genus Nucras, Gray. Annals of the South African Museum 13: 195 - 216.","Boulenger GA. 1920. Monograph of the Lacertidae. Volume 1. Trustees of the British Museum (Natural History), London, United Kingdom. x + 352 p.","Monard A. 1937. Contribution a l'herpetologie d'Angola. Arquivos do Museu Bocage 8: 19 - 154.","Laurent RF. 1964. Reptiles et amphibiens de l'Angola (troiseme contribution). Publicacoes Culturais Com- panhia de Diamantes de Angola 67: 1 - 165.","Broadley DG. 1965. Some problems presented by sand lizards of the Nucras tessellata group. Journal of the Herpetological Association of Africa 1: 18 - 23.","Broadley DG. 1972. A review of the Nucras tessellata group (Sauria: Lacertidae). Arnoldia 20: 1 - 35.","Ceriaco LMP, de Sa SC, Bandeira S, Valerio H, Stanley EL, Kuhn AL, Marques M, Vindum JV, Blackburn DC, Bauer AM. 2016. Herpetological survey of Iona National Park and Namibe Regional Natural Park, with a synoptic list of the amphibians and reptiles of Namibe Province, Southwestern Angola. Proceedings of the California Academy of Sciences 63 (2): 15 - 61.","Burger M. 2014. Nucras tessellata (A. Smith, 1838). Pp. 171 - 172 In: Atlas and Red List of the Reptiles of South Africa, Lesotho, and Swaziland. Editors, Bates MF, Branch WR, Bauer AM, Burger M, Marais J, Alexander GJ, de Villiers MS. Suricata 1. South African National Biodiversity Institute, Pretoria, South Africa. 485 p.","Marques MP, Ceriaco LMP, Blackburn DC, Bauer AM 2018. Diversity and distribution of the amphibians and terrestrial reptiles of Angola: atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences Series 4, 65 (Supplement II): 1 - 501.","Branch WR, Baptista N, Vaz Pinto P, Conradie W. 2019. The reptiles of Angola - History, updated checklists, endemism, hot spots, and future directions for research. Pp. 283 - 326 In: Biodiversity of Angola. Science and Conservation: A Modern Synthesis. Editors, Huntley BJ, Ferrand N, Russo V, Lages F. Springer Open, Cham, Switzerland. 552 p.","Barbosa LAG. 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, Angola. 323 p."]}
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- 2019
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106. Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola
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Branch, William R., Conradie, Werner, Pinto, Pedro Vaz, and Tolley, Krystal A.
- Subjects
Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Lacertidae ,Taxonomy - Abstract
Branch, William R., Conradie, Werner, Pinto, Pedro Vaz, Tolley, Krystal A. (2019): Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola. Amphibian & Reptile Conservation 13 (2): 82-95, DOI: http://doi.org/10.5281/zenodo.3731977, {"references":["Aljanabi SM, Martinez I. 1997. Universal and rapid saltextraction of high quality genomic DNA for PCR based techniques. Nucleic Acids Research 25: 4,692- 4,693.","Arevalo E, Davis SK, Sites JJW. 1994. Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology 43: 387-418.","Barbosa LAG. 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, Angola. 323 p.","Bauer AM, Childers JL, Broeckenhoven C, Mouton PLN. 2019. A new Nucras Gray, 1838 (Squamata: Lacertidae) from the Strandveld of the Western Cape, South Africa. Zootaxa 4560(1): 149-163.","Bocage JVB. 1895. Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, Portugal. 203 p., 20 pls.","Boulenger GA. 1910. A revised list of South African reptiles and batrachians, with synoptic tables, special reference of specimens in the South African Museum, and descriptions of new species. Annals of the South African Museum 5: 455-538.","Boulenger GA. 1917. A revision of the lizards of the genus Nucras, Gray. Annals of the South African Museum 13: 195-216.","Boulenger GA. 1920. Monograph of the Lacertidae. Volume 1. Trustees of the British Museum (Natural History), London, United Kingdom. x + 352 p.","Branch WR, Bauer AM. 1995. The herpetofauna of the Little Karoo, Western Cape, South Africa, with notes on life history and taxonomy. Herpetological Natural History 3: 47-89.","Branch WR, Tolley KA. 2017 Oral presentation (Abstract). New lacertids from Angola. African Herp News 66: 11.","Branch WR. 1998. Field Guide to the Snakes and other Reptiles of Southern Africa. Struik, Cape Town, South Africa. 368 p.","Branch WR. 2016. Preface, Amphibian & Reptile Conservation special Angola-Africa issue. Amphibian & Reptile Conservation 10(2): i-iii (e128).","Branch WR, Haacke W, Vaz Pinto P, Conradie W, Baptista N, Verburgt L. 2017. Loveridge's Angolan geckos, Afroedura karroica bogerti and Pachydactylus scutatus angolensis (Sauria, Gekkonidae): new distribution records, comments on type localities, and taxonomic status. Zoosystematics and Evolution 93: 157-166.","Branch WR, Baptista N, Vaz Pinto P, Conradie W. 2019. The reptiles of Angola - History, updated checklists, endemism, hot spots, and future directions for research. Pp. 283-326 In: Biodiversity of Angola. Science and Conservation: A Modern Synthesis. Editors, Huntley BJ, Ferrand N, Russo V, Lages F. Springer Open, Cham, Switzerland. 552 p.","Broadley DG. 1965. Some problems presented by sand lizards of the Nucras tessellata group. Journal of the Herpetological Association of Africa 1: 18-23.","Broadley DG. 1972. A review of the Nucras tessellata group (Sauria: Lacertidae). Arnoldia 20: 1-35.","Burger M. 2014. Nucras tessellata (A. Smith, 1838). Pp. 171-172 In: Atlas and Red List of the Reptiles of South Africa, Lesotho, and Swaziland. Editors, Bates MF, Branch WR, Bauer AM, Burger M, Marais J, Alexander GJ, de Villiers MS. Suricata 1. South African National Biodiversity Institute, Pretoria, South Africa. 485 p.","Ceriaco LMP, Marques MP, Bandeira S, Agarwal I, Stanley EL, Bauer AM, Heinicke MP, Blackburn DC. 2018. A new earless species of Poyntonophrynus (Anura, Bufonidae) from the Serra da Neve Inselberg, Namibe Province, Angola. ZooKeys 780: 109-136.","Ceriaco LMP, de Sa SC, Bandeira S, Valerio H, Stanley EL, Kuhn AL, Marques M, Vindum JV, Blackburn DC, Bauer AM. 2016. Herpetological survey of Iona National Park and Namibe Regional Natural Park, with a synoptic list of the amphibians and reptiles of Namibe Province, Southwestern Angola. Proceedings of the California Academy of Sciences 63(2): 15-61.","Conradie W, Branch WR, Measey GJ, Tolley KA. 2012. Revised phylogeny of Sand lizards (Pedioplanis) and the description of two new species from south-western Angola. African Journal of Herpetology 60: 91-112.","FitzSimons VFM. 1943. The lizards of South Africa. Memoirs of the Transvaal Museum 1: 1-528.","Forstner MRJ, Davis SK, Arevalo E. 1995. Support for the hypothesis of Anguimorph ancestry for the suborder Serpentes from phylogenetic analysis of mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 4: 93-102.","Haacke WD. 2008. A new leaf-toed gecko (Reptilia: Gekkonidae) from south-western Angola. African Journal Herpetology 57(2): 85-92.","Haagner GV, Branch WR, Haagner AJF. 2000. Notes on a collection of reptiles from Zambia and adjacent areas of the Democratic Republic of the Congo. Annals of the Eastern Cape Museum 1: 1-25.","Huelsenbeck JP, Ronquist F. 2001. MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754-755.","Jacobsen NHG. 1989. A Herpetological Survey of the Transvaal. Ph.D. Dissertation, University of Natal, Department of Biological Sciences, Durban, South Africa. 1,621 p.","Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, et al. 2012. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28(12): 1,647-1,649.","Kumar S, Stecher G, Tamura K. 2016. MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1,870-1,874.","Laurent RF. 1964. Reptiles et amphibiens de l'Angola (troiseme contribution). Publicacoes Culturais Com- panhia de Diamantes de Angola 67: 1-165.","Marques MP, Ceriaco LMP, Bandeira S, Pauwels OSG, Bauer AM. 2019a. Description of a new long-tailed skink (Scincidae: Trachylepis) from Angola and the Democratic Republic of the Congo. Zootaxa 4568(1): 51-68.","Marques MP, Ceriaco LMP, Stanley EL, Bandeira S, Agarwal I, Bauer AM. 2019b. A new species of Girdled Lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, soutwestern Angola. Zootaxa 4668(4): 503-524.","Marques MP, Ceriaco LMP, Blackburn DC, Bauer AM 2018. Diversity and distribution of the amphibians and terrestrial reptiles of Angola: atlas of historical and bibliographic records (1840-2017). Proceedings of the California Academy of Sciences Series 4, 65(Supplement II): 1-501.","Mayer W, Pavlicev M. 2007. The phylogeny of the family Lacertidae (Reptilia) based on nuclear DNA sequences: convergent adaptations to arid habitats within the subfamily Eremiainae. Molecular Phylogenetics and Evolution 44: 1,155-1,163.","Meier R, Kwong S, Vaidya G, Ng PKL. 2006. DNA barcoding and taxonomy in Diptera: a tale of high intraspecific variability and low identification success. Systematic Biology 55: 715-728.","Miller MA, Pfeiffer W, Schwartz T. 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Pp. 1-8 In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 November 2010, New Orleans, Louisiana, USA. Institute of Electrical and Electronics Engineers (IEEE), Piscataway, New Jersey, USA. 115 p.","Monard A. 1937. Contribution a l'herpetologie d'Angola. Arquivos do Museu Bocage 8: 19-154.","Neumann O. 1900. Description of a new lizard of the genus Nucras from Usoga, British East Africa. Annals and Magazine of Natural History, including Zoology, Botany, and Geology 7(5): 56.","Posada D. 2008. jModelTest: phylogenetic model averaging. Molecular Biology and Evolution 25: 1,253-1,256.","Smith A. 1838. Contributions to the natural history of Southern Africa. Article VIII. Magazine of Natural History 2(14): 92-94.","Spawls S, Howell K, Drewes R, Ashe J. 2018. A Field Guide to the Reptiles of East Africa. 2nd Edition. Bloomsbury Natural History, London, United Kingdom. 624 p.","Stamatakis A, Hoover P, Rougemont J. 2008. A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57: 758-771 . Stamatakis A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2,688-2,690.",". Stamatakis A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2,688-2,690.","Stanley EL, Ceriaco, LMP, Bandeira S, Valerio H, Bates MF, Branch WR. 2016. A review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the description of a new species from the Pro-Namib desert. Zootaxa 4061(3): 201-226.","Tolley K, Weeber J, Maritz B, Verburgt L, Bates M, Conradie W, Hofmeyr R, Turner A, Da Silva J, Alexander G. 2019. No safe haven: protection levels show imperiled South African reptiles not sufficiently safe-guarded despite low average extinction risk. Biological Conservation 233: 61-72."]}
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- 2019
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107. CHAIRMAN'S REPORT 2022-2023.
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Tolley, Krystal
- Abstract
The Chairman's Report for the Herpetological Association of Africa (HAA) from 2022-2023 highlights the new members of the Executive Committee and their roles. The report mentions various accomplishments during this period, including the inclusion of different languages on the HAA website, finalizing tax clearances and international payment methods, and successfully hosting an in-person conference. The committee also approved new policies related to research ethics and publishing. The report discusses the establishment of minimum standards for each committee portfolio and the implementation of Standard Operating Procedures for various HAA processes. The HAA is now a registered non-profit organization with a Board of Directors. The report also mentions changes in portfolio names and the consideration of providing free membership to qualifying seniors. The committee is exploring initiatives such as setting up a publishing office and holding an Annual General Meeting online. The report acknowledges the challenges of increasing participation and membership and discusses efforts to address these challenges, including involving Early Career Researchers and Herpetologists in auxiliary panels and portfolios. The report mentions an unexpected expense related to publishing charges from the journal publisher. The report concludes with the announcement of an online election for a new committee and the Chairman's farewell, expressing confidence in the next generation of African herpetologists. [Extracted from the article]
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- 2023
108. The Biology of Chameleons
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TOLLEY, KRYSTAL A., HERREL, ANTHONY, TOLLEY, KRYSTAL A., and HERREL, ANTHONY
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- 2013
109. New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozambique inselbergs
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Conradie, Werner, Bittencourt-Silva, Gabriela B., Harith M. Farooq, Loader, Simon P., Menegon, Michele, and Tolley, Krystal A.
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Nothophryne Poynton, 1963 is a monotypic genus of frog, with the nominal species N. broadleyi found only on Mount Mulanje, in southern Malawi. Recent surveys in northern Mozambique, however, have uncovered at least four new species associated with four inselbergs (Mount Inago, Mount Namuli, Mount Ribáuè and Taratibu Hills). Previous phylogenetic analyses using mitochondrial genes suggest that each mountain isolate has an endemic species of Nothophryne. Herein we provide a rediagnosis of the genus and comparative diagnoses of four new species based on new material. ZooBank— urn:lsid:zoobank.org:pub:19C53BF3-BA52-4CAE-933F-5BBEE0AF0457
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- 2018
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110. A new species of Uroplatus (Gekkonidae) from Ankarana National Park, Madagascar, of remarkably high genetic divergence
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RATSOAVINA, FANOMEZANA M., primary, SCHERZ, MARK D., additional, TOLLEY, KRYSTAL A., additional, RASELIMANANA, ACHILLE P., additional, GLAW, FRANK, additional, and VENCES, MIGUEL, additional
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- 2019
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111. Horizon scanning for South African biodiversity: A need for social engagement as well as science
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Seymour, Colleen L., primary, Gillson, Lindsey, additional, Child, Matthew F., additional, Tolley, Krystal A., additional, Curie, Jock C., additional, da Silva, Jessica M., additional, Alexander, Graham J., additional, Anderson, Pippin, additional, Downs, Colleen T., additional, Egoh, Benis N., additional, Ehlers Smith, David A., additional, Ehlers Smith, Yvette C., additional, Esler, Karen J., additional, O’Farrell, Patrick J., additional, Skowno, Andrew L., additional, Suleman, Essa, additional, and Veldtman, Ruan, additional
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- 2019
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112. Patterns of geographic variation between mitochondrial and nuclear markers in Heaviside's (Benguela) dolphins (Cephalorhynchus heavisidii)
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GOPAL, Keshni, primary, KARCZMARSKI, Leszek, additional, and TOLLEY, Krystal A., additional
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- 2019
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113. Genetic structure associated with habitat diversification supports the independent evolution of ecomorphs inBradypodion pumilum
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Tolley, Krystal A, primary, Hopkins, Kevin P, additional, and da Silva, Jessica M, additional
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- 2019
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114. A new species of tree snake (Dipsadoboa, Serpentes: Colubridae) from ‘sky island’ forests in northern Mozambique, with notes on other members of the Dipsadoboa werneri group
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BRANCH, WILLIAM R., primary, BAYLISS, JULIAN, additional, BITTENCOURT-SILVA, GABRIELA B., additional, CONRADIE, WERNER, additional, ENGELBRECHT, HANLIE M., additional, LOADER, SIMON P., additional, MENEGON, MICHELE, additional, NANVONAMUQUITXO, CRISTÓVÃO, additional, and TOLLEY, KRYSTAL A., additional
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- 2019
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115. Ancient habitat shifts and organismal diversification are decoupled in the African viper genusBitis(Serpentes: Viperidae)
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Barlow, Axel, primary, Wüster, Wolfgang, additional, Kelly, Christopher M. R., additional, Branch, William R., additional, Phelps, Tony, additional, and Tolley, Krystal A., additional
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- 2019
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116. No safe haven: Protection levels show imperilled South African reptiles not sufficiently safe-guarded despite low average extinction risk
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Tolley, Krystal A., primary, Weeber, Joshua, additional, Maritz, Bryan, additional, Verburgt, Luke, additional, Bates, Michael F., additional, Conradie, Werner, additional, Hofmeyr, Margaretha D., additional, Turner, Andrew A., additional, da Silva, Jessica M., additional, and Alexander, Graham J., additional
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- 2019
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117. Diversifying into the branches: Species boundaries in African green and bush snakes, Philothamnus (Serpentes: Colubridae)
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Engelbrecht, Hanlie M., primary, Branch, William R., additional, Greenbaum, Eli, additional, Alexander, Graham J., additional, Jackson, Kate, additional, Burger, Marius, additional, Conradie, Werner, additional, Kusamba, Chifundera, additional, Zassi-Boulou, Ange-Ghislain, additional, and Tolley, Krystal A., additional
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- 2019
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118. Convergence and vicariance: speciation of chameleons in the Cape Fold Mountains, South Africa, and the description of three new species of BradypodionFitzinger, 1843
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Tolley, Krystal A, Tilbury, Colin R, and Burger, Marius
- Abstract
ABSTRACTThe mechanisms that underpin ecological speciation, morphological convergence and the evolution of ecological morphotypes (ecomorphs) in squamates have allowed for a better appreciation of the speciation process in chameleons. In particular, attention has been drawn to several populations of chameleons (Sauria, Chamaeleonidae, Bradypodion) from the Cape Fold Mountains, South Africa. Previous work suggested that these populations are genetically divergent, but with strong similarities in phenotype. Using an integrative taxonomic approach that accounts for genetic diversity, habitat and morphology, three of these populations are described as species. One population is from an isolated forest patch and is genetically different at the species level, but morphologically similar to Bradypodion damaranum(Boulenger, 1887) from forested areas in the Knysna region. Although not sister species, the two are in the same clade and probably diverged through vicariance of the forest. Two other populations are from fynbos habitat in adjacent mountain ranges (Tsitsikamma/Langkloof/Kouga mountains and Baviaanskloof Mountains) and are also morphologically similar, but genetically divergent at the species level. These two species are not sister taxa and are not in the same clade yet have a virtually identical phenotype presumably as the result of convergent evolution for the fynbos habitat. Within the context of morphological taxonomy, these populations have been difficult to evaluate. However, when viewed in the context of ecological speciation, convergence and morphological conservatism, the species boundaries are apparent, allowing for them to be described as new taxa.
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- 2022
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119. Rise of oceanographic barriers in continuous populations of a cetacean: the genetic structure of harbour porpoises in Old World waters
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Ridoux Vincent, Öztürk Bayram, A Öztürk Ayaka, Llavona Ángela, Jauniaux Thierry, Ferreira Marisa, Duke Sarah, Birkun Alexei, Tolley Krystal A, Ray Nicolas, Piry Sylvain, Baird Stuart JE, Fontaine Michaël C, Rogan Emer, Sequeira Marina, Siebert Ursula, Vikingsson Gísli A, Bouquegneau Jean-Marie, and Michaux Johan R
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Biology (General) ,QH301-705.5 - Abstract
Abstract Background Understanding the role of seascape in shaping genetic and demographic population structure is highly challenging for marine pelagic species such as cetaceans for which there is generally little evidence of what could effectively restrict their dispersal. In the present work, we applied a combination of recent individual-based landscape genetic approaches to investigate the population genetic structure of a highly mobile extensive range cetacean, the harbour porpoise in the eastern North Atlantic, with regards to oceanographic characteristics that could constrain its dispersal. Results Analyses of 10 microsatellite loci for 752 individuals revealed that most of the sampled range in the eastern North Atlantic behaves as a 'continuous' population that widely extends over thousands of kilometres with significant isolation by distance (IBD). However, strong barriers to gene flow were detected in the south-eastern part of the range. These barriers coincided with profound changes in environmental characteristics and isolated, on a relatively small scale, porpoises from Iberian waters and on a larger scale porpoises from the Black Sea. Conclusion The presence of these barriers to gene flow that coincide with profound changes in oceanographic features, together with the spatial variation in IBD strength, provide for the first time strong evidence that physical processes have a major impact on the demographic and genetic structure of a cetacean. This genetic pattern further suggests habitat-related fragmentation of the porpoise range that is likely to intensify with predicted surface ocean warming.
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- 2007
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120. Long-term feeding ecology and habitat use in harbour porpoises Phocoena phocoena from Scandinavian waters inferred from trace elements and stable isotopes
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Gobert Sylvie, Siebert Ursula, Tolley Krystal A, Fontaine Michaël C, Lepoint Gilles, Bouquegneau Jean-Marie, and Das Krishna
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Ecology ,QH540-549.5 - Abstract
Abstract Background We investigated the feeding ecology and habitat use of 32 harbour porpoises by-caught in 4 localities along the Scandinavian coast from the North Sea to the Barents Sea using time-integrative markers: stable isotopes (δ13C, δ15N) and trace elements (Zn, Cu, Fe, Se, total Hg and Cd), in relation to habitat characteristics (bathymetry) and geographic position (latitude). Results Among the trace elements analysed, only Cd, with an oceanic specific food origin, was found to be useful as an ecological tracer. All other trace elements studied were not useful, most likely because of physiological regulation and/or few specific sources in the food web. The δ13C, δ15N signatures and Cd levels were highly correlated with each other, as well as with local bathymetry and geographic position (latitude). Variation in the isotopic ratios indicated a shift in harbour porpoise's feeding habits from pelagic prey species in deep northern waters to more coastal and/or demersal prey in the relatively shallow North Sea and Skagerrak waters. This result is consistent with stomach content analyses found in the literature. This shift was associated with a northward Cd-enrichment which provides further support to the Cd 'anomaly' previously reported in polar waters and suggests that porpoises in deep northern waters include Cd-contaminated prey in their diet, such as oceanic cephalopods. Conclusion As stable isotopes and Cd provide information in the medium and the long term respectively, the spatial variation found, shows that harbour porpoises experience different ecological regimes during the year along the Scandinavian coasts, adapting their feeding habits to local oceanographic conditions, without performing extensive migration.
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- 2007
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121. A phylogeny and genus-level revision of the African file snakes Gonionotophis Boulenger (Squamata: Lamprophiidae)
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30998360 - Burger, Marius, Broadley, Donald G., Burger, Marius, Tolley, Krystal A., Conradie, Werner, Wishart, Sarah, 30998360 - Burger, Marius, Broadley, Donald G., Burger, Marius, Tolley, Krystal A., Conradie, Werner, and Wishart, Sarah
- Abstract
The sub-Saharan African file snake genus Gonionotophis is currently comprised of 15 species. However, the concept of this genus has been confounded by morphological and genetic differences between the constituent taxa. Due to the dearth of DNA samples, a taxonomic assessment has been impractical to date. We therefore sequenced two mitochondrial and one nuclear marker (16S, cyt b , and c-mos) from 45 samples representing ten species of Gonionotophis to construct a molecular phylogeny using Bayesian and likelihood approaches. Four divergent and well- supported clades were recovered, including: (1) grantii + brussauxi ; (2) poensis + stenophthalmus ; (3) nyassae ; and (4) capensis , chanleri , crossi , guirali and savorgnani . Based on these results and morphological data, the genus Gonionotophis is restricted to the first clade, Mehelya is resurrected for the species in the second clade, and new genera are described for the remaining two clades
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- 2018
122. African Herald snakes, Crotaphopeltis, show population structure for a widespread generalist but deep genetic divergence for forest specialists.
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Engelbrecht, Hanlie M., Branch, William R., Greenbaum, Eli, Burger, Marius, Conradie, Werner, and Tolley, Krystal A.
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COLUBRIDAE ,HABITAT partitioning (Ecology) ,GENE flow ,SNAKES ,GENETIC speciation ,SPECIALISTS - Abstract
The African colubrid snake genus Crotaphopeltis currently comprises six species and occurs throughout sub‐Saharan Africa. The most widespread of these, Crotaphopeltis hotamboeia, inhabits most biomes, aside from rainforest and hyper‐arid regions, and its catholic niche has presumably facilitated substantial gene flow. Despite this, the geographical range is large enough that ecological or physical barriers might exist, facilitating allopatric diversification. In contrast, most of the other species are habitat specialists with limited distributions (e.g., Crotaphopeltis tornieri) and would be expected to show strong genetic structure. We therefore examined species boundaries within Crotaphopeltis in a phylogenetic context using five markers (16S, cyt b, ND4, c‐mos, and RAG‐1) for four of the six species. Species delimitation methods included two coalescent‐based and one barcoding approach. Widespread geographical sampling of C. hotamboeia allowed examination of genetic structuring across its range. The species status of Crotaphopeltis barotseensis, C. degeni, and C. hotamboeia was confirmed, whereas the Afromontane species C. tornieri comprised two candidate species. Crotaphopeltis hotamboeia did not show cryptic speciation, although its phylogeographic structure corresponded with the spatiotemporal pattern of the African savanna. Our results show how the heterogeneous African environment could influence genetic partitioning of habitat specialist and generalist species at broad geographical scales. [ABSTRACT FROM AUTHOR]
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- 2020
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123. Cryptic diversity in Rhampholeon boulengeri (Sauria: Chamaeleonidae), a pygmy chameleon from the Albertine Rift biodiversity hotspot
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Hughes, Daniel F., primary, Tolley, Krystal A., additional, Behangana, Mathias, additional, Lukwago, Wilber, additional, Menegon, Michele, additional, Dehling, J. Maximilian, additional, Stipala, Jan, additional, Tilbury, Colin R., additional, Khan, Arshad M., additional, Kusamba, Chifundera, additional, and Greenbaum, Eli, additional
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- 2018
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124. Cryptic diversity in the common flap-necked chameleon Chamaeleo dilepis in South Africa
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Main, Devon C, primary, van Vuuren, Bettine Jansen, additional, and Tolley, Krystal A, additional
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- 2018
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125. A contribution to the phylogeny and taxonomy of the Pachydactylus weberi group (Squamata: Gekkonidae): a case of intraspecific colour polymorphism confounding taxonomy
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Šmíd, Jiří, primary, Engelbrecht, Hanlie, additional, Taft, Jody M., additional, Telford, Nicolas S., additional, Makhubo, Buyisile G., additional, Bauer, Aaron M., additional, and Tolley, Krystal A., additional
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- 2018
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126. A phylogeny and genus-level revision of the African file snakesGonionotophisBoulenger (Squamata: Lamprophiidae)
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Broadley, Donald G., primary, Tolley, Krystal A., additional, Conradie, Werner, additional, Wishart, Sarah, additional, Trape, Jean-François, additional, Burger, Marius, additional, Kusamba, Chifundera, additional, Zassi-Boulou, Ange-Ghislain, additional, and Greenbaum, Eli, additional
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- 2018
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127. New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozambique inselbergs
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Conradie, Werner, primary, Bittencourt-Silva, Gabriela B., additional, Farooq, Harith M., additional, Loader, Simon P., additional, Menegon, Michele, additional, and Tolley, Krystal A., additional
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- 2018
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128. Kinyongia mulyai Tilbury & Tolley, 2015, sp. nov
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Tilbury, Colin R. and Tolley, Krystal A.
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Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Kinyongia ,Kinyongia mulyai ,Taxonomy - Abstract
Kinyongia mulyai sp. nov. Synonomy: Kinyongia adolfifriderici Tilbury 2010 Holotype. PEM-R 19199 (CT 426 — Fig. 11), a sub-adult male with partially everted hemipenes, collected by Colin Tilbury and Isak Hattingh on 21 st March 2010, from path side vegetation at 1700 metre a.s.l., Mount Nzawa, Moba District, Katanga Province, Democratic Republic of the Congo (6 ° 51.07 ’ S; 29 ° 35.87 ’ E). Diagnosis. Within the genera Kinyongia and Trioceros, the ontogenetic development of body crests and rostral process’s—whilst not developed to the same degree as seen in adults—is still evident in sub-adult specimens, and can be reliably used to determine the presence or absence of these characters. Although Kinyongia mulyai sp. nov. is known from only one sub-adult specimen, it is likely that the morphological characters are sufficiently well developed to provide reliable criteria on which to base a morphological determination. Kinyongia mulyai sp. nov. is distinguishable from the closely-related species K. adolfifriderici, K. gyrolepis (Greenbaum et al. 2012) and K. excubitor (Barbour 1911) by the absence of a dorsal crest in the male. It is also distinguished from K. adolfifriderici by having a narrower and more elongate head (Fig. 10), with a head length/inter-orbital width ratio of 4 (versus 3 for K. adolfifriderici), and from both K. gyrolepis and K. adolfifriderici by having a higher number of upper labial scales (17 vs. a maximum of 15 and 14 respectively; Greenbaum et al. 2012). Distinguished from all the other Kinyongia congeners by the absence of any form of naso-rostral ornamentation in males. Description of the Holotype. A small chameleon with a snout-vent length of 44mm and tail measuring 63mm. Head length from casque tip to snout 13.7mm and the inter-orbital width at mid orbit 2.8mm. Casque low, barely elevated above the nape. Parietal crest consists of a row of 3 enlarged tubercles at the apex of the casque. This forks anteriorly into two short rows each consisting of 2 low tubercles. Supra-orbital ridges more or less smooth. The supra-orbital ridges are separated from each other across the mid-orbital point by three flattened tubercles. The canthal ridge is adorned with a row of 4 prominent rounded tubercles—the most anterior the largest. A temporal crest composed of 4 similarly sized enlarged tubercles arises from the mid posterior orbital rim; this crest ascends along the posterior rim of the casque to its apex. The top of the casque and the zone below the temporal crest are covered with similar-sized, flattened tubercles. The nares open infero-posteriorly about midway between the anterior orbital rim and the snout. A single row of tubercles separates the inferior orbital rim and the upper labials. There is no rostral process. Seventeen upper labials and 17 lower labials are present from the tip of the snout to the posterior margin of the orbital rim. Two tubercles separate the upper labials from the end of the canthal ridge. There is no trace of a gular crest, or ventral crest. The sides of the body are clad with somewhat heterogeneous, flattened polygonal tubercles, with larger tubercles showing a tendency to rosette formation on the lower flanks. A network of thin interstitial grooves extends between the tubercles. There is no row of enlarged flank tubercles. The dorsal vertebral crest is represented by a single small cone at the nape behind the casque, followed by a second much smaller cone. Thereafter the dorsal keel is smooth. The tail is likewise smooth. The outer surfaces of the limbs are covered in enlarged flattened tubercles while the inner surfaces of the limbs are clad with small regular tubercles. Colour in Life. Head—Top of head olive green with lighter green tubercles on casque. Skin of eyeballs light brown with two thin horizontal stripes running through the middle and extending posteriorly along the temporal crest and anteriorly just below the canthal ridge. The zone below the temporal crest is powder blue scattered with greenish tubercles. A small triangular area between the mouth line, nares and the orbital rim is blue green. The gular region is suffused with pale orange and yellow extending posteriorly to between the front limbs. A pale line extends from below the eye to just beyond the commissure of the mouth. Background colouration of body is olive green. Interstitial skin between the tubercle rosettes on lower flanks forms a network of dark interstitium. The dark interstitium of the flanks is broken by two broad zones of pale interstitium extending from the dorsal ridge down over the flanks to the middle of the body. There is no white line on the abdomen. Outer side of limbs with light green tubercles, skin on inner sides of limbs off white. Tail olive green with many thin dark bands. Habitat. The Holotype was collected in closed canopy Afrotemperate montane forest at 1800 metres a.s.l. from path side vegetation, perched at approximately 2.5 metres from the ground. One other chameleon (not collected) was observed occupying a sleeping perch on an exposed liana vine over 20 metres above the ground. Etymology. Named for Mr. Jules Mulya, whose energetic support and assistance with smoothing away mountains of obstacles to the progress of the Mt. Nzawa expedition led to the discovery of both new species.
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- 2015
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129. Rhampholeon (Rhinodigitum) hattinghi Tilbury & Tolley, 2015, sp. nov
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Tilbury, Colin R. and Tolley, Krystal A.
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Reptilia ,Squamata ,Animalia ,Rhampholeon ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy ,Rhampholeon hattinghi - Abstract
Rhampholeon (Rhinodigitum) hattinghi sp. nov. Synonymy: Rhampholeon boulengeri Tilbury 2010. Holotype. PEM-R 19194, an adult male with everted hemipenes, collected by Colin Tilbury and Isak Hattingh on 21 st March 2010, from path side vegetation at 1700 metres a.s.l., Mount Nzawa, Moba District, Katanga Province, Democratic Republic of the Congo (6 �� 51.07 ��� S; 29 �� 35.87 ��� E). Paratypes. PEM-R 19193, adult male; PEM-R 19195, adult male, PEM-R 19196, adult female; PEM-R 19197, adult female; PEM-R 19198, adult female; all collected on the same date and locality as the holotype. Diagnosis. Due to its markedly bicuspid claws, amelanotic parietal peritoneum and acalyculate bag-like hemipenes with dual apical horns, this taxon can be placed within the sub-genus Rhampholeon (Rhinodigitum) Matthee et al. 2004. In spite of the remarkably conserved external morphology within this genus, this new taxon is easily distinguishable from congeners by virtue of the following identifying characteristics: The consistent absence of deep mite pockets (or pits) in the inguinal flexure distinguishes this taxon from Rh. beraduccii Mariaux & Tilbury and the six species of the Rh. platyceps G��nther complex (Branch et al. 2014), including Rh. platyceps, Rh. chapmanorum Tilbury, Rh. maspictus Branch et al., Rh. tilburyi Branch et al., Rh. bruessoworum Branch et al. and Rh. nebulauctor Branch et al.; the presence of deep pits in the axillae of this taxon distinguishes it from Rh. nchisiensis Loveridge and Rh. acuminatus Mariaux & Tilbury; the relatively smooth supra-orbital and canthal crests distinguish this taxon from Rh. boulengeri, Rh. uluguruensis Tilbury & Emmrich and Rh. moyeri Menegon et al., in all of which a peaked cluster of tubercles forms distinct protuberances above the eye and the nasal aperture (Fig. 5). Description of Holotype. Snout-vent 56mm, tail 11mm. Body habitus leaf like - typical of all other Rhampholeon (Rhinodigitum) species. Head short, casque flattened, top of head shallowly concave. The lateral crests are studded with several prominent tubercles. Parietal crest indistinct, indicated by a short row of 3 marginally enlarged tubercles. The supra-optic ridge gathers into a low cluster of tubercles anteriorly above each eye, but without forming a supra-optic horn. The two supra-orbital ridges are connected to each other by a series of 18 inter-orbital tubercles arranged in a shallow V across the top of the head. The canthal ridges are formed by a row of enlarged, relatively smooth tubercles which terminate anteriorly at the base of a small, finely tuberculated stump-like process that barely projects off the anterior end of the snout. The nares open posteriorly, from a small bulge at a point roughly one third of the distance between the anterior orbital rim and the front of the maxilla. A distinct temporal crest arises from the mid post-orbital rim and consists of 5 tubercles on the right and six tubercles on the left, of which the most posterior is the largest. Three large sub-conical tubercles are spaced along the inferoposterior rim of the orbit. A prominent tubercle is situated just above the mouth in line with the posterior orbital rim. The dorsal vertebral line is weakly crenulated, fading to smooth over the pelvic region. The dorsal margin of the tail is likewise smooth. Deep axillary mite pockets (or pits) are present, whilst the inguinal flexures are smooth. The background scalation of the body, tail, limbs and belly is composed of tightly packed sub-homogeneous, stellate-edged tubercles. The gular region of the throat extending to between the front limbs is studded with many small conical tubercles. These also are found on the belly, underside of the tail and the underside and particularly on the outer-surface of the limbs. The sides of the body have regularly-spaced, enlarged conical tubercles, with three particularly enlarged cones situated in a row at midbody, the first sited above the shoulder, the second at midbody, and the last just before the pelvis. Claws are markedly biscuspid, several prominent palmar and plantar accessory spines are seen at the base of the toes, and the outer soles of the feet are relatively spinose whilst the inner soles are smooth with a cobblestone appearance. Colour in life. Adult male (Figs 6 & 7). Top of head and snout dark brown extending to cover the sides of the snout above the line of the mouth and to a lesser extent the eyeball and superior temporal zone. Eyeball with dark radiations especially prominent over the upper half of the eyeball. Gular region off-white. Body light brown mottled with olive green. Five dark blotches are spaced along the dorsal vertebral ridge, and a broad dark blotch is also located over the dorsum of the tail. Two thin dark green stripes, originating from the second and third vertebral blotches, extend infero-posteriorly over the flanks. Three dark spots are arrayed along the flank, each spot centred over a prominent conical tubercle. The inner sides of the fore and hind limbs pale orange. A prominent white spot is located on each side of the tail. Adult female stress pattern (Fig. 8). Dorsal ridge orange from the tip of the casque to the tip of the tail. Flanks speckled with small orange spots and three large white spots on the flanks, each surround an enlarged conical tubercle. Sides of head and gular region flecked in white, sometimes forming short vertical stripes. Eyeball dark and speckled with light blue tubercles. Lateral flank stripes dark but heavily flecked with light blue. Hemipenes. Short bag-like, truncus acalyculate; a pair of short, broad-based apical horns incurving above the sulcus spermaticus (Fig. 9). The upper surface of the horns is covered in closely packed thorn-like papillae, much like a pineapple skin. Fourteen papillae are arranged in a proximal or basal cluster terminating in a distal row of two papillae. Inferior to each apical horn, a prominent balloon-like membranous evagination billows anteriorly. Sexual Dimorphism. Apart from the prominent hemipenal bulges there is little sexual dimorphism evident in the type series (Table 4). In males the tail comprises between 16���23 % of the total length whilst in females it is between 14���18 % of total length. Number Sex Length Rp Casque/snout Sub-orbital Axil pit Inguinal pit (mm) (mm) rows *= Holotype, measurements in millimetres; s-v = snout-vent length; Io tub = inter-orbital tubercles; Io diam = interorbital diameter; Length Rp = length of rostral process; Sub-orbital rows = number of rows of tubercles between the inferior orbital rim and the upper labials; na = not measured. Variation. In the paratypes the parietal crest was either absent, or merely indicated as in the holotype. The supra-orbital crest is relatively smooth in all specimens, only forming a low cluster of tubercles in 2 specimens at the point where the inter-orbital tubercles terminate above the eye. The soles of the feet may vary from relatively smooth to a sub-acuminate field of tubercles. The hemipenes of PEM-R 19193 and PEM-R 19195 do not differ in any substantial way from that of the holotype. Reproduction. The parietal peritoneum in the abdominal cavity of the largest female (PEM-R 19198) was unpigmented and the fallopian tubes each held a single egg measuring 6.4mm x 5.6mm. Habitat. All specimens were found in closed canopy Afrotemperate montane forest on low vegetation alongside a path. Perch heights varied from a few centimetres up to 50 cm from the ground. Etymology. Named for Isak Hattingh, who enthusiastically but unwittingly agreed to accompany the first author on a trip which tested many boundaries. He never complained once., Published as part of Tilbury, Colin R. & Tolley, Krystal A., 2015, Contributions to the herpetofauna of the Albertine Rift: Two new species of chameleon (Sauria: Chamaeleonidae) from an isolated montane forest, south eastern Democratic Republic of Congo, pp. 345-364 in Zootaxa 3905 (3) on pages 355-359, DOI: 10.11646/zootaxa.3905.3.2, http://zenodo.org/record/240513, {"references":["Tilbury, C. R. (2010) The Chameleons of Africa, an Atlas including the chameleons of Europe, the Middle East and Asia. Edition Chimaira, Germany, 831 pp.","Matthee, C. A., Tilbury, C. R. & Townsend, T. (2004) A phylogenetic review of the African leaf chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society of London Series B, Biological Sciences, 271, 1967 - 1976. http: // dx. doi. org / 10.1098 / rspb. 2004.2806","Branch, W. R., Bayliss, J. & Tolley, K. A. (2014) Pygmy chameleons of the Rhampholeon platyceps complex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique. Zootaxa, 3814 (1), 1 - 36. http: // dx. doi. org / 10.11646 / zootaxa. 3814.1.1"]}
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- 2015
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130. Contributions to the herpetofauna of the Albertine Rift: Two new species of chameleon (Sauria: Chamaeleonidae) from an isolated montane forest, south eastern Democratic Republic of Congo
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Tilbury, Colin R. and Tolley, Krystal A.
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Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Tilbury, Colin R., Tolley, Krystal A. (2015): Contributions to the herpetofauna of the Albertine Rift: Two new species of chameleon (Sauria: Chamaeleonidae) from an isolated montane forest, south eastern Democratic Republic of Congo. Zootaxa 3905 (3): 345-364, DOI: http://dx.doi.org/10.11646/zootaxa.3905.3.2
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- 2015
131. A new species of Chameleon (Sauria: Chamaeleonidae: Kinyongia) highlights the biological affinities between the Southern Highlands and Eastern Arc Mountains of Tanzania
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Menegon, Michele, Loader, Simon, Davenport, Tim, Howell, Kim, Tilbury, Colin, Machaga, Sophy, and Tolley, Krystal
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lcsh:Zoology ,lcsh:QL1-991 - Abstract
A new species of chameleon is described from the Livingstone and Udzungwa Mountains of Tanzania. The new species is morphologically most similar to Kinyongia vanheygeni. Furthermore, a single, short rostral appendage shows the species similarity to other Eastern Arc endemic Kinyongia species (e.g. K. uthmoelleri, K. oxyrhina, K. magomberae and K. tenuis). Females of all these species lack any rostral ornamentation and are all very similar morphologically. Males of the new species, on which the morphological diagnosis is based, can be distinguished from other Kinyongia by a shorter rostral appendage that bifurcates at the tip. They are easily distinguished from K. vanheygeni, otherwise the most similar species, by differences in head scalation and the length and shape of the rostral appendage. The new species is associated with montane rainforest and is known from only four forest fragments of which two are in the Udzungwa and two in the Livingstone Mountains. Phylogenetically, the new species is sister to K. tenuis and K. magomberae, which together, form a clade that also contains K. oxyrhina. The disjunct distribution of the new species, in the Livingstone and Udzungwa mountains, stretches across the ‘Makambako Gap’ which is a putative biogeographical barrier separating the distinct faunas of the Southern highlands and Eastern Arc Mountains. Evidence from this species however, points to potentially closer biological affinities between the Livingstone and Udzungwa mountains., Acta Herpetologica, Vol 10 No 2 (2015)
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- 2015
132. Ever since Gondwana: the influence of changing climate, fragmenting forest, and spreading savanna on the biogeography of African reptiles
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Tolley, Krystal A.
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Africa -- Natural history ,Habitat destruction -- Research ,Biodiversity -- Research ,Climate change -- Environmental aspects ,Savannas -- Natural history -- Environmental aspects ,Ecological research ,Biological sciences - Abstract
Continental Africa has experienced major biome shifts throughout the Cenozoic, with a progression from humid climate and pan-African forest to a mesic/arid, savanna-dominant landscape. This shift has major implications for [...]
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- 2017
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133. Chameleons on the move: survival and movement of the Cape dwarf chameleon, Bradypodion pumilum, within a fragmented urban habitat
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Tolley, Krystal A., Raw, Robert NV, Altwegg, Res, and Measey, G. John
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Animal Science and Zoology - Abstract
Reptiles have an amazing diversity of life-history attributes ranging from the shortest- to longest-lived of vertebrate species. Estimating survival in wild populations is of key importance when understanding population dynamics and life-history evolution. However, data are lacking for a large and charismatic group of lizards, the chameleons. We conducted a Robust Design (RD) capture–mark–recapture (CMR) experiment on the Cape dwarf chameleon, Bradypodion pumilum, at two nested sites within the Cape Town Metropolitan Area, South Africa, in order to estimate survival and movement of adults in and around an isolated 3.5 ha patch of suitable habitat. Over a nine-week period, 97 individuals were identified in 379 captures from five primary capture sessions with three secondary events each. Analysis of CMR data provided evidence that smaller chameleons have a substantially lower survival per 10-day period than larger chameleons. RD analysis showed that males were more prone to temporary emigration than females, while open multi-strata analysis revealed that smaller chameleons more readily moved between the sites than larger chameleons. Our findings offer first important insights into chameleon survival and life-history dynamics, which suggest a more vagile subadult population and the possibility of male biased dispersal. Our results have implications for managing the conservation of threatened chameleon populations in highly fragmented urban habitats.African Zoology 45(1): 99–106 (April 2010)
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- 2010
134. Impact of species delimitation and sampling on niche models and phylogeographical inference: A case study of the East African reed frog Hyperolius substriatus Ahl, 1931
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Bittencourt-Silva, Gabriela B., primary, Lawson, Lucinda P., additional, Tolley, Krystal A., additional, Portik, Daniel M., additional, Barratt, Christopher D., additional, Nagel, Peter, additional, and Loader, Simon P., additional
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- 2017
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135. Conservation genetics of an endemic and threatened amphibian (Capensibufo rosei): a leap towards establishing a genetic monitoring framework
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da Silva, Jessica M., primary and Tolley, Krystal A., additional
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- 2017
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136. Genetic diversity and differentiation of the Western Leopard Toad (Sclerophrys pantherina) based on mitochondrial and microsatellite markers
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da Silva, Jessica M., primary, Feldheim, Kevin A., additional, Measey, G. John, additional, Doucette-Riise, Stephen, additional, Daniels, Ryan J., additional, Chauke, Lucas F., additional, and Tolley, Krystal A., additional
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- 2017
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137. Chameleons of Southern Africa Ed. 1
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Tolley, Krystal, Burger, Marius, Tolley, Krystal, Tolley, Krystal, Burger, Marius, and Tolley, Krystal
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Chameleons are fascinating creatures: they evoke a strong response in people, be it delight and wonder or fear. Chameleons of Southern Africa explores this interesting group of lizards and discusses their unusual and intriguing characteristics. The book presents an overview of all types of chameleons (of which there are up to 160 species), reproduction, behavior and their relationships to one another. There is a comprehensive identification guide to all southern African species ? some 25 have been identified to date ? together with distribution maps and colourful photographs of their many features and guises. For interested students, gardeners, naturalists and even just the curious, Chameleons of Southern Africa is an essential guide to these captivating lizards
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- 2012
138. Conservation status and threats for African reptiles
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Tolley, Krystal A., primary, Alexander, Graham J., additional, Branch, William R., additional, Bowles, Philip, additional, and Maritz, Bryan, additional
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- 2016
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139. The effects of substratum on locomotor performance in lacertid lizards
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Vanhooydonck, Bieke, Measey, John, Edwards, Shelley, Makhubo, Buyisile, Tolley, Krystal, Herrel, Anthony, Centre for Invasion Biology, Stellenbosch University, Evolutionary Morphology of Vertebrates, and Universiteit Gent = Ghent University [Belgium] (UGENT)
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[SDV]Life Sciences [q-bio] ,[SDV.BA]Life Sciences [q-bio]/Animal biology ,Biology ,ComputingMilieux_MISCELLANEOUS - Abstract
Locomotion is important to animals because it has direct implications for fitness through its role in predator escape, prey capture, and territory defence. Despite significant advances in our understanding of animal locomotion, studies exploring how substrate properties affect locomotor performance remain scant. In the present study, we explore how variation in substrate (sand, slate, cork) affects locomotor performance in lacertid lizards that differ in morphology. Moreover, we explore whether substrate effects are the same for different types of locomotor performance (speed, acceleration, and stamina). Our results show that the substrate affected most types of locomotor performance studied but not always in the same way. Although substrate effects were species-dependent for the maximal speed over 50cm and the distance run to exhaustion, this was not the case for acceleration capacity. These results suggest that substrate texture differentially affects burst performance vs. longer duration measures of locomotor performance. Finally, straightforward relationships between habitat use and the substrate on which performance was maximized were not observed. This suggests that the evolution of locomotor capacity is complex and that animals may show compromise phenotypes allowing them to deal with a variety of substrates in their natural environment.(c) 2015 The Linnean Society of London
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- 2015
140. Rediscovery, range extension, habitat and phylogenetic relation of the endemic Scaled Sandveld Lizard Nucras scalarisLaurent, 1964 (Sauria: Lacertidae) in the central Angolan plateau
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Baptista, Ninda L, Tolley, Krystal A, Bluhm, Marc, Finckh, Manfred, and Branch, William R
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ABSTRACTThe Scaled Sandveld Lizard Nucras scalarisLaurent, 1964is a poorly known lacertid endemic to north-eastern Angola and is only known from the type series collected more than half a century ago. The original description provided a comprehensive morphological description, but there was a lack of information regarding its evolutionary relationships and habitat associations. A recent discovery of N. scalarisfrom Cusseque, Bié Province, on the central Angolan plateau provided the opportunity to address some knowledge gaps of this species. A phylogenetic analysis confirmed its distinctiveness at the species level. Despite a limited dataset, the analysis suggests that N. scalarisis sister to N. broadleyirather than N. lalandii,the latter having been assumed to be the sister species, because of morphological similarity between them. The new record of N. scalarisrepresents a range extension of more than 350 km to the southwest and extends the elevation range to 1 570 m above sea level, compared to previous known localities at about 1 300 m above sea level (Alto Cuílo and Alto Chicapa, in Lunda Sul Province). The specimen also provides new information on live coloration, as well as the habitat association of grassy shrubland that is dominated by geoxylic suffrutices (the so-called ‘underground forests’) characteristic of the Angolan Miombo Woodlands. This new information raises the question whether poor survey data have led to an under-estimation of faunal diversity in this peculiar and overlooked vegetation type, and reinforces the need for further surveys which could highlight the importance of this habitat.
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- 2020
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141. Rhampholeon (Rhinodigitum) chapmanorum Tilbury 1992
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
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Reptilia ,Squamata ,Animalia ,Rhampholeon ,Biodiversity ,Rhampholeon chapmanorum ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Rhampholeon (Rhinodigitum) chapmanorum Tilbury, 1992 Chapmans��� Pygmy Chameleon Synonomy: Rhampholeon chapmani Tilbury 1992 Rhampholeon chapmanorum Klaver & B��hme 1997 Rhampholeon (Rhinodigitum) chapmanorum Matthee, Tilbury & Townsend 2004. Holotype: Adult male (BMNH 1988.399), collected by Colin and Sarah Tilbury. Type locality: in evergreen forest at 940 m asl., above the Chididi Police post, on top of Malawi Hills, southern Malawi. Variation in adult coloration in an adult male and an adult female are illustrated in Fig. 4 D and Fig. 4 E, respectively. Note also the pronounced nasal and supraorbital processes on the head of the male. For full details of morphology, hemipenial structure, biology and habitat see Tilbury (2010)., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on page 17, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Tilbury, C. R. (1992) A new dwarf forest chameleon (Sauria: Rhampholeon) from Malawi, Central Africa. Tropical Zoology, 5, 1 - 9. http: // dx. doi. org / 10.1080 / 03946975.1992.10539176","Klaver, C. J. J. & Bohme, W. (1997) Liste der rezenten Amphibien und Reptilien-Chamaeleonidae. Das Tierreich, 112, i - xiv, 1 - 85.","Matthee, C. A., Tilbury, C. R. & Townsend, T. (2004) A phylogenetic review of the African pygmy chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society of London, Series B, 271, 1967 - 1976.","Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp."]}
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- 2014
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142. Rhampholeon (Rhinodigitum) bruessoworum Branch, Bayliss & Tolley, 2014, sp. nov
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
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Reptilia ,Rhampholeon bruessoworum ,Squamata ,Animalia ,Rhampholeon ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Rhampholeon (Rhinodigitum) bruessoworum sp. nov. Mount Inago Pygmy Chameleon Synonymy: Rhampholeon sp. Bayliss et al. 2010, p 17. fig. 13. Etymology. The specific epithet honours the contributions of the brothers Carl and Darren Bruessow to the protection of wildlife in southern Malawi, particularly via the Mount Mulanje Conservation Trust. Types. The type series comprises three specimens, including: Holotype.- An adult female (PEM R 20375; Fig. 9 A) collected by J. Bayliss, 5 September 2009, in a small patch of wet forest at the base of a granite inselberg of Mt. Inago, Zambézia Province, Mozambique (15 ˚04' 51 ”S 37 ˚ 23 ' 37 ”E, ca 1478 m a.s.l.). Allotype. An adult male (PEM R 20376, Fig. 9 B), same collecting details as holotype. Paratype. An adult female (PEM R 20374), same collecting details as holotype. Meristics. Measurements of the type series of Rhampholeon bruessoworum sp. nov. are summarized in Table 9. Diagnosis. The Mt. Inago Pygmy Chameleon is referable to Rhampholeon (subgenus Rhinodigitum) by possessing an unpigmented parietal peritoneum, claws that are strongly bicuspid, smooth plantar surfaces, and a rostral process. It can be distinguished from most other species in Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It differs from Rh. platyceps and Rh. maspictus sp. nov. in its small size (Rh. chapmanorum in having a relatively large rostral process in males (small in both sexes in Rh. chapmanorum), and from all other members of the Rh. platyceps complex in Mozambique (i.e. Rh. maspictus sp. nov., Rh. nebulauctor sp. nov. and Rh. tilburyi sp. nov.) in having a relatively longer tail in both sexes. From all other Rhampholeon it is also genetically well differentiated, and all chameleons examined form a monophyletic clade. Description of Holotype. Adult female, viscera exposed by a single ventral incision. Head: Dorsum of head flattened, with no upward flexure of the snout; casque flat, edged with weakly-defined lateral crests that are mainly restricted to the posterior region of the casque; temporal crest weakly-developed, comprising a single, interrupted row of large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridges reduced to a few scattered enlarged scales but with a very small multiscaled process forming a ‘soft horn’ at each end of the inter-orbital ridge that passes across the crown and is composed of 10 small granular tubercles, and that demarcate the posterior edge of a slight frontal depression; inferior orbital rim with 4 (right) and 5 (left) enlarged tubercles; snout bordered on each side by moderately developed rostral crests, that fuse together at the tip of the snout which is adorned with a very small, flattened rostral process (1.6 mm) which underneath is only slightly free from the rostral, and is four small tubercular scales long and four tubercular scales wide at its base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more conical but smaller than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest very weakly developed, reduced to 9 crenulations of enlarged, but not obviously spinose scales; crenulations most strongly developed over mid-body, reduced in size over on neck and which are present on the tail in more reduced form; deep axillary and inguinal pits are present; flank scalation heterogeneous, composed of small, stellate granules with few scattered, enlarged spinose tubercles, the largest at the shoulder and in two clusters; chest, belly and lower surface of tail smooth; limb scalation more tubercular, with a few enlarged, spinose tubercles on the forearms; claws strongly bicuspid; accessory planter spines on the soles of the fore and hind feet are present, but reduced to very small, soft, spinose scales at the base of the claws; tail flattened laterally, flexing downward on the distal third. Colour in life (based on two images of holotype, JB; Fig. 9 A): Mid-body mottled brown with two narrow, oblique purple-brown lateral stripes; dorsal surface of fore-body, neck, top of head and upper surfaces of limbs darker brown; Throat lightly mottled cream extending onto chin and labials, which have a light yellow flush; belly and base of tail pale brown. Colour in preservative: Body mottled brown with two narrow oblique reddish-brown bars on mid-flanks; lower surface of neck, belly, base of tail, soles of feet, and lower limbs pale brown. Description of Allotype (as for holotype, unless noted): Adult male, incision along base of tail with hemipenal muscle removed; hemipenes not everted. Head: Parietal crest almost absent, composed of a four enlarged tubercles in the mid-line; five enlarged tubercles on both sides of the inferior orbital rim; rostral process six small tubercular scales long and four tubercles wide at base. Body: Dorsal crest very weakly developed, reduced to nine crenulations of enlarged but not obviously spinose scales, subequal in development to those of the female holotype; tail relatively long (30.5 % SVL), with a prominent hemipeneal bulge. Colour in life (based on two images of allotype; Fig. 9 B): Mid-body mottled brown with small orange brown blotches along dorsal crest, separated by 2–3 crenulations; throat, belly and lower surfaces of limbs and tail pale brown. Colour in preservative: After preservation body pale brown with vestiges of two narrow oblique stripes on the flanks; pale brown below. Paratype variation (as for holotype, unless noted): Adult female with a large ventral incision. The ‘soft horn’ on the supraorbital ridge is very small and hardly protrudes; only four enlarged tubercles on inferior orbital rim; plantar spines very reduced to small, soft, spinose scales. Size. Presumably a small species, as all three specimens in the type series were sexually mature, with turgid testes or developing ova. Largest male - PEM R 20376 (allotype) 39.0 + 17.1 = 56.1 mm; largest female—PEM R 20374 (holotype) 47.5 + 15.3 = 62.7 mm. Distribution. Restricted to the type locality; Mt. Inago, Zambézia Province, northern Mozambique. Habitat. The Inago Massif shows habitat zonation, and is surrounded by Brachystegia woodland at the base. The type series were collected at night in mid-altitude (~ 1500 m) evergreen forest at the base of a granite inselberg (Fig. 9 C). This forest type comprised relatively large trees between 20-30 m high, with the upper canopy layer composed of species such as Drypetes natalensis, Schefflera umbellifera and Newtonia buchananii, whilst the midcanopy layer (Chrysophyllum gorungosum, Myrianthus holstii, bridelia sp. and Garcinia sp. These remnant forest patches are greatly reduced in size (et al. 2010).
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- 2014
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143. Rhampholeon tilburyi Branch, Bayliss & Tolley, 2014, sp. nov
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
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Rhampholeon tilburyi ,Reptilia ,Squamata ,Animalia ,Rhampholeon ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Rhampholeon tilburyi sp. nov. Mount Namuli Pygmy Chameleon Synonomy: Rhampholeon platyceps Branch & Ryan 2001, p 282. Rhampholeon tilburyi Timberlake et al. 2010, p 57 nomen nudum Rhampholeon nov. sp. (tilburyi) Timberlake et al. 2010, p 61 nomen nudum Rhampholeon platyceps (tilburyi) Timberlake et al. 2010, p 61 nomen nudum Rhampholeon Portik et al. 2013, p 416. Etymology. The specific epithet is a patronym for our good friend Dr. Colin Tilbury for his outstanding contributions to knowledge of African chameleons, most recently exemplified in his magnum opus, the Chameleons of Africa (Tilbury 2010). For the last 30 years his travels throughout Africa in search of chameleons, combined with his stunning photography, have been a source of inspiration. His studies have done much to enhance our understanding of African chameleons, and it is with great pleasure that we name this chameleon in his honour. Types. The type series comprises eight specimens, including: Holotype. An adult female (PEM R 14921; Fig. 8 D) collected by a local guide, 30 November 1998, in the Ukalini Forest that nestles under the south face of the main Namuli peak, Namuli Massif, Zamb��zia Province, Mozambique (15 ˚ 22 'S 37 ˚04'E, ca 1550 m a.s.l.). Allotype. An adult male (PEM R 17132, Fig. 8 A) with everted hemipenes, collected by K.A. Tolley and S. van Noort, 26 May 2006, at night perched on a dead branch 50 cm above ground in a patch of forest dominated by overgrown tea (Camellia sinensis) on the SDZ Cha Sarl Tea Estate at Guru��, Namuli Massif, Zamb��zia Province, Mozambique (15 ˚ 26 ' 51.6 ���S, 37 ˚00���, 32.6 ���E, ca 839 m a.s.l.). Paratypes. Six specimens, comprising two females (PEM R 17134, Fig. 8 B, same collecting details as allotype; PEM R 17135, perched on a tea bush in an overgrown patch of tea, on SDZ Cha Sarl Tea Estate at Guru��, 15 ˚ 26 ' 42.8 ���S, 37 ˚00���, 19.2 ���E); three males (PEM R 17131, SDZ Cha Sarl Tea Estate at Guru��, 15 ˚ 26 ' 49 ���S, 37 ˚00���, 29 ���E; PEM R 17133, same collecting details as allotype; PEM R 20372, Muretha Plateau, Namuli Massif, Zamb��zia Province, Mozambique (15 ˚ 23 ' 26 ���S, 37 ˚02���,03���E, ca 1804 m a.s.l.), collected by J. Bayliss 27 May 2007; and a very small juvenile (PEM R 20373, same details as previous specimen). Meristics. Measurements for the type series of Rhampholeon tilburyi sp. nov. are summarized in Table 8. Diagnosis. The Mt. Namuli pygmy chameleon is referable to the Rhampholeon (subgenus Rhinodigitum Matthee et al. 2004) by possessing a short hemipenis that is almost bag-like, acalyculate and adorned with a pair of simple apical ���horns��� with a variable number of thorn-like papillae arranged on the outer aspect of the horn; having an unpigmented parietal peritoneum, claws that are strongly bicuspid, smooth plantar surfaces, a rostral process, and short tail (Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It differs from Rh. platyceps and Rh. maspictus sp. nov. by its smaller size (Rh. platyceps complex, including the populations from Mt Mabu, Mt Chiperone and Mt Inago, by retaining in adult males a prominent flexure (> 32 ˚) of the snout in front of the orbit (flat or Rh. platyceps and Rh. maspictus sp. nov.; Rh. nebulauctor sp. nov. and Rh. bruessoworum sp. nov.), and a narrower head (HW/HL% 49.4 %; 53.2���59.8 % in all other species). Finally, the species is also genetically well differentiated from all other Rhampholeon, and all specimens examined form a monophyletic clade from Mt. Namuli. Description of Holotype. Adult female, viscera exposed by a single ventral incision and two lateral incisions on left flank. Head: Flattened, with only a small upward flexure of the snout at the front of the orbit (angling upwards at approximately 28 ˚ from a line between tip of snout and top of casque); casque weak, flat with small, conical tubercular edges; temporal crest moderate, forming a straight ridge of 7 raised, rounded tubercles, 3 rd and last largest; lateral crests weak, composed of small conical tubercles that demarcate the edge of the casque; temporal crest moderate with a single row of interrupted large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridge well marked but without obvious clusters of tubercles (vaguely present at the posterior edge), and with a very small ���soft horn��� of enlarged tubercles at the junction with the shallow interorbital ridge; interorbital ridge composed of 12 small granular tubercles that demarcate the posterior edge of a prominent frontal depression; two enlarged tubercles on inferior orbital rim; rostral ridges very well marked, and fusing at tip of the snout which bears a rostral process (1.5 mm, in fixation flattened against rostrum), about 7 small granules long and 5-6 granules wide at base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more stellate than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest weak, composed of a double row of enlarged, rounded tubercles along the backbone; the crest undulates between 13 equally spaced clusters of enlarged, spinose tubercles that are largest on the middle of the body, flatter and smooth over the forelimb and sacral regions, and only vaguely visible on the tail; deep axillary and inguinal pits are present; flank scalation heterogenous, composed of small, stellate granules with scattered, enlarged spinose tubercles, the largest occurring over the shoulder; chest, belly and lower surface of tail smooth; limb scalation more irregular, with numerous larger, spinose tubercles on forearms; soles of feet sub-spinose; claws strongly bicuspid; accessory planter spines at base of claws very small, almost absent; tail gently tapering, dorsoventrally flattened, 21.7 % of total length, and covered with homogenous conical tubercles. Colour in preservative: Body mottled brown with reddish brown reticulation on flanks; head darker above, blotched with dark brown; belly and inner surfaces of limbs paler with darker specking; tail dark brown above and below. Description of Allotype (as for holotype, unless noted). Adult male, with everted hemipenes and viscera exposed by a single, ragged ventral incision and liver tissue excised for DNA analysis. Head: A very prominent angular flexure of the snout at the front of the orbit (approx 40 ˚ from line between tip of snout and casque); supraorbital ridge well marked with clusters of tubercles at posterior and anterior level of orbit, the latter forming a ���soft horn��� two scales high; rostral ridge fusing anteriorly with an obvious rostral process (1.3 mm), that is rounded in profile and composed of about 6 small granules long and 5-6 granules wide at base. Body: Claws strongly bicuspid; accessory planter spines at the base of claws weakly developed; tail 24.0% of total length. Coloration in life (Fig. 8 A): Head, body and tail tan to brown with orange flush along midflank and greenish flush over back and outer surfaces of hind limbs; two prominent diagonal lines on flanks, formed from three darkish spots joined by grey infusion; belly, throat, inner surfaces of limbs and underside of tail light tan; outer skin of limbs darker brown; snout with wide orange band running diagonally through nostril; skin of orbit and temporal region darker brown with scattered, intense green-blue individual tubercles; iris orange-red. Coloration after preservation: Body very dark, almost blackish; all normal coloration lost due to fixation in concentrated formalin. Paratype variation. The tail varies from 18.3���21.7 % of total length in females, and from 24.0��� 25.8 % in males; the rostral process varies from 7.9���10.4 % of head length in females, and from 15.9���16.6 % in males. In life the iris of a male paratype (R 17131) was reddish and the snout had an indistinct brown stripe rather than the orange band in the allotype. In addition, a few scattered green-blue tubercles extended onto the snout, as well as on to the orbit and temporal region. Prominent ���leaf vein��� dark lines on the flank were also well developed. Size. Largest male���PEM R 17132 (allotype) 39.6 + 12.5 = 52.1 mm; largest female���PEM R 14921 (holotype) 55.3 + 15.4 = 70.7 mm. The smallest male (SVL 33.2 mm) still has well-developed hemipenes, indicating that it is sexually mature (although testicular activity was not determined). Sexual Dimorphism. Whilst the type series is limited in number, there are indications of sexual dimorphism. Females grow considerably larger than males, and the three females in the type series have an average SVL of 51.67 mm, compared with only 34.67 mm for the three males. In addition, males have a more angular head shape (slope 39-45 ˚, Fig 9 B) than females (slope 22-28 ˚), and also proportionately longer tails and a longer rostral process (see above). However, the small number of specimens precludes statistical confirmation. More subtle difference in scalation details cannot be assessed until larger series are available. Hemipenis. (based on everted hemipenes of all three males in type series). Hemipenis short with unadorned basal and apical sections, except for a pair of curved horns that emerge prominently from the expanded, flattened crown, and which bear well developed spines on their upper surfaces; the folds of the sulcus spermaticus flare to drain into the flattened apical region. Distribution. Restricted to the type locality; Mt. Namuli, Zamb��zia Province, northern Mozambique, in both the evergreen forests on Manho and Ukalini forests on the Muretha Plateau (1804 m a.s.l.) at the base of the main peaks, respectively, and also in fallow areas of tea plantations at lower elevations (838 m a.s.l). Habitat. The Namuli Massif shows habitat zonation and is surrounded by Brachystegia woodland at the base. Grassland and scrub with forested river valleys cover the slopes, with grassy plateaus and patches of dense moist evergreen forest near the summit (Fig. 8 E). The largest surviving blocks include the Manho Forest (c. 1,000���1,100 ha) and the Ukalini Forest (c. 100 ha), the latter lodged against the base of the Namuli dome. Dominant emergent trees include Cryptocarya liebertiana, Faurea wentzeliana and Olea capensis (Timberlake et al. 2009). These forests are under high anthropogenic threat as they are cleared for potato cultivation by surrounding rural communities. The allotype and most paratypes were collected at night on perches from 30-50 cm above ground in forest adjacent to a mountain stream in a fallow areas of an operational tea estate that have overgrown into a forest. Portik et al. (2013) noted that adults and juveniles were found sheltering on branches of small trees or on the fronds of epiphytic ferns within Ukalini Forest. This contrasts with perches heights of 1.3���5 m recorded for Rh. chapmanorum (Tilbury 1992). Reproduction. The female holotype was gravid with seven almost spherical eggs (largest diameter 4.1-4.5 mm, four in the left oviduct and three in the right oviduct) that lack obvious signs of embryonic development. Diet. The stomach of the holotype contained insect fragments, including coleopteran elytra. Predation. A small chameleon, possibly referable to this species, was seen being carried by a Crowned Hornbill (Tockus alboterminatus) that was flying over the canopy of Ukalini Forest, 29 November 1998 (Branch & Ryan 2001)., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 25-28, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Branch, W. R. & Ryan, P. G. (2001) Additions to the Mozambique Herpetofauna: Two new lizards from the Namuli Massif, Mozambique. Herpetological Review, 32 (4), 281 - 282.","Bayliss, J., Monteiro, J., Fishpool, L., Congdon, C., Bampton, I., Bruessow, C., Matimele, H., Banze, A. & Timberlake, J. R. (2010) Biodiversity and Conservation of Mount Inago, Mozambique. Report produced under Darwin Initiative Award 15 / 0 36. Mulanje Mountain Conservation Trust, Malawi, 32 pp.","Portik, D. M., Travers, S., Bauer, A. M. & Branch, W. R. (2013 b) A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mount Namuli, an isolated ' sky' island of northern Mozambique. Zootaxa, 3710 (5), 415 - 435. http: // dx. doi. org / 10.11646 / zootaxa. 3710.5.2","Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp.","Timberlake, J. R., F. Dowsett-Lemaire, F., Bayliss, J., Alves, T., Baena, S., Bento, C., Cook, K., Francisco, J., Harris, T., Smith, P. & de Sousa, C. (2009) Mt. Namuli, Mozambique: Biodiversity and Conservation. Report produced under the Darwin Initiative Award 15 / 036. Royal Botanic Gardens, Kew, London, 114 pp.","Tilbury, C. R. (1992) A new dwarf forest chameleon (Sauria: Rhampholeon) from Malawi, Central Africa. Tropical Zoology, 5, 1 - 9. http: // dx. doi. org / 10.1080 / 03946975.1992.10539176"]}
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- 2014
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144. Rhampholeon (Rhinodigitum) maspictus Branch, Bayliss & Tolley, 2014, sp. nov
- Author
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
- Subjects
Reptilia ,Rhampholeon maspictus ,Squamata ,Animalia ,Rhampholeon ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Rhampholeon (Rhinodigitum) maspictus sp. nov. Mount Mabu Pygmy Chameleon Synonomy: Rhampholeon (Rhinodigitum) chapmanorum Tilbury 2010, p 179. Rhampholeon sp. Timberlake et al. 2012, p 45; Bayliss et al. 2014, p 179. Etymology. The specific epithet derives from a combination of mas (L. = man) and pictus (L. = painted), alluding to the unusual bright colours of most males, which are often retained for long periods, even when sleeping at night. Types. The type series comprises ten specimens all preserved in 50 % propanol (except for one hatchling in 90 % ethanol), including: Holotype. An adult male (PEM R 18072; Fig. 5 A, 6 A) collected by a W.R. Branch, J. Bayliss & W. Conradie, 27 May 2009, in the vicinity of the main forest base camp, Mt. Mabu, Zamb��zia Province, Mozambique (16 �� 17 ' 10.1 "S, 36 �� 24 '02.2"E; 967 m a.s.l.). Allotype. An adult female with a small ventral incision (PEM R 18061, Fig. 5 B, 5 A), same collecting details as holotype. Paratypes. Eight specimens, comprising four males (PEM R18059, 18073 (Fig. 6 B), 18074 - 75), same collecting details as holotype; three females (PEM R 18069 -70, 18076), same collecting details as holotype; and a hatchling (PEM R 18068, Fig. 6 C) Additional material (used in analysis but not forming part of the type series): 13 specimens, all from forest Camp region, Mt. Mabu, Zambezia Province, Mozambique (16 ˚ 17 ' 12 "S, 36 ˚ 24 ' 14 "E; 1000m a.s.l.); PEM R 17130 (J. Bayliss, January 2006), PEM R 17911 - 12 (J. Bayliss, 16 April 2009), PEM R 18057 -58, 18060, 18062 -67, 18071, same collecting details as holotype. Meristics. Measurements for the type series of Rhampholeon maspictus sp. nov. are summarized in Table 6. Diagnosis. The Mt. Mabu Pygmy Chameleon is referable to the Rhampholeon (subgenus Rhinodigitum Matthee et al. 2004) by possessing a short hemipenis that is almost bag-like, acalyculate and adorned with a pair of simple, curved apical ���horns��� with a variable number of thorn-like papillae arranged on the outer aspect of the horn; claws that are strongly bicuspid, smooth plantar surfaces, a rostral process, and short tail (Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It differs from all other members of the Rh. platyceps complex by the bright green male breeding coloration, including blue flanks and side of head, and yellow throat, snout and eye ring (all of which may be retained even at night). It shares with Rh. platyceps, but differs from all other populations of the complex from Mt Chiperone, Mt Namuli and Mt Inago, its large size (> 60 mm SVL) in both sexes, lack of male dwarfism, well-developed dorsal crenulations, and reduced rostral and supraocular processes; it differs from Rh. platyceps by its more dorsoventrally flattened habitus (more rounded in Rh. platyceps), and very weak or even absent accessory plantar spines (present but small in Rh. platyceps). Finally, the species is also genetically well differentiated from all other Rhampholeon, and all chameleons examined from Mt. Mabu form a monophyletic clade. Description of Holotype. Adult male, viscera exposed by a single ventral incision in the chest region; both hemipenes everted. Head: Dorsum of head flattened, with no upward flexure of the snout; casque flat with weakly define lateral crests; temporal crest moderate with a single row of interrupted large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridge present but with only few raised conical tubercles; three enlarged tubercles on inferior orbital rim; supraorbital connected in middle by shallow interorbital ridge composed of 11 small granular tubercles that demarcate the posterior edge of a prominent frontal depression; rostral ridge very well marked, forming a small bump over the nostrils and fusing anteriorly and adorned with a small rostral process (2.1 mm, in fixation flattened against rostrum); rostral process 4 small tubercular scales long and 4 tubercles wide at base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more conical than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest moderately developed, comprised of 9 weak crenulations each comprised of a cluster of enlarged spinose scales, crenulations most strongly developed over mid-body, reduced over base of neck and tail; crenulations continue onto tail comprised of 9 small groups that are largest over the distal half of the tail; deep axillary and inguinal pits are present; flank scalation heterogeneous, composed of small, stellate granules with scattered, enlarged spinose tubercles, the largest occurring over the shoulder; chest, belly and lower surface of tail smooth; limb scalation more tubercular, with a few enlarged, spinose tubercles on the forearms; claws strongly bicuspid; accessory plantar spines very reduced or almost absent. Colour in Life: Male coloration stunningly beautiful (Figs. 5 A); body, tail and limbs leaf green, slightly mottled and also darker along the dorsal crest of body, tail and upper surfaces of fore- and hindlimbs; central region of flanks light blue, with two oblique, broad yellow-green stripes; light blue of flanks extends onto the side of head and around eye, where it becomes darker and more intense; front of chest, throat, labials, rostral process and upper surface of snout to the level of the orbit, dirty yellow; bright yellow eye ring. When stressed (e.g. in artificially close proximity to another chameleon) the body coloration changes dramatically, the body darkening to greenbrown, and the blue flank region becomes paler and infused with brown blotches (Fig. 6 A); the green orbital region, yellow eye ring and dirty yellow throat colour are retained, albeit in subdued form. Colour in preservative: Body dark brown with 3 faint oblique bars of lighter brown on flanks running from backbone to lower flank; belly, tail and limbs uniform dark brown, except light brown on inner surface of forelimbs. Description of Allotype (as for holotype, unless noted): adult female, with viscera exposed by a single, ragged ventral incision in the belly, and liver tissue excised for DNA analysis. Head: Temporal crest weakly developed comprised of a disrupted row of slightly enlarge tubercles; parietal crest absent; subocular ridge weakly pronounced with few enlarged tubercles; interorbital ridge with 13 weakly enlarged tubercles Body: Dorsal crest very weakly developed, comprising 7 crenulations of tubercles that are only slightly enlarged; crenulations most strongly developed over mid-body and almost absent on the tail; flank scalation heterogeneous, composed of small, stellate granules with very few scattered, enlarged spinose tubercles, the largest at the shoulder. Coloration in Life: Less intensely coloured than male holotype (Figs. 5 B); sides of body dorsally dull green with two vague, oblique, dark olive stripes that extend from dorsal crest, where they are broader, onto lower flank; upper flanks with large, irregular light brown blotches that coalesce and cover upper surface of tail; lower flanks brighter green with scattered pale blotches, extending as pale green onto lower surface of tail; upper surfaces of limbs olive, paler green below; upper surface of casque pale brown, extending at a narrow, almost white line along the dorsal crest; front and sides of snout mottled in light green and brown giving a dull green-brown appearance; throat cream with numerous light blue tubercles that coalesce to form irregular blotches; orbital area olive green with scattered pale blue tubercles and a dull red eye ring. When stressed the irregular light brown blotches on the upper flanks become paler, and the sides of head become blotched, especially in the labial region, giving an appearance of alternating pale and dark bands radiating from the eye (Fig. 6 A). Colour in preservative: Body and head uniform dark brown above and below. Paratype variation. All paratype males have well-everted hemipenes. In life the orbital scalation of a male paratype (R 17130) was light green, with a conspicuous yellow ring around the iris; the pale brown body had two prominent rust red ���leaf vein��� narrow lines that ran from the dorsal crest obliquely to the lower flank; there is a prominent enlarged, pale yellow-cream spinose tubercle on the fore-flank in the shoulder region; the crown of the casque and dorsal crest on the body and tail was medium brown (Fig. 6 B). A very small specimen (PEM R 18068, SVL 19.6 mm) has a mottled reddish brown head, body and limbs, with two vague, narrow oblique red-brown stripes that run from the dorsal crest to the lower flanks; the dorsal crest forms a narrow cream stripe that extends onto the casque, where it becomes disrupted into small blotches; the limbs are darker brown with scattered bright yellow tubercles of the upper surfaces; the orbital region is dull brown with scattered small blue tubercles and a dull red eye ring (Fig. 6 C). Size. Largest male���PEM R 18074 (paratype) 65.2 + 20.1 = 85.3 mm; largest female���PEM R 18069 (paratype) 64.4 + 16.2 = 80.6 mm. The smallest specimen���PEM R 18068 (paratype, unsexed) 19.9 + 4.9 = 24.8 mm) appears to be newly hatched. Sexual Dimorphism. The average total size of males (65.3 mm, n = 11) and females (64.4 mm, n = 11) is similar and suggests no sexual size dimorphism. However, males do have proportionately longer tails (TL as = % of SVL 24.0% in males, but only 18.6 % in females). Sexual dichromism in pygmy chameleons remains poorly known, and has not been studied in the Rh. platyceps complex. Tilbury (1992) noted the emergence of bright colours (pale powder blue head and neck, and bright white eyelids) in male Rh. chapmanorum in male-male interactions. The bold colours of large male Rh. maspictus sp. nov. (Fig. 5 A) was noted in three different males at the type locality, including two specimens sleeping at night on low vegetation. No female was observed showing the bright green body and blue and yellow head of adult males. Hemipenis. (based on everted hemipenes of five males in the type series). Hemipenis short with unadorned basal and apical sections, except for a pair of curved horns that form the lateral edges of a prominent, flat-topped crown; 3-4 weakly-developed spines occur on the upper surfaces of the horns; the folds of the sulcus spermaticus flare to drain into the flattened apical region. Distribution. Known only from the type locality; Mt. Mabu, Zamb��zia Province, central Mozambique, but expected to occur throughout much of the extensive forest (+ 7880 ha) remaining on the massif (Bayliss et al. 2014). Habitat. The Mabu massif is a complex of granitic inselbergs formed of syenite, an igneous intrusion of the younger Precambrian Namarroi series dating from 850 ���1100 Mya (Instituto Nacional de Geologia 1987). Although the Mabu Massif shows vegetational zonation and a significant extent (+ 7880 ha) of mid-altitudinal forest, it lacks the large area of upland plateau that occurs on Mt. Namuli. Mt. Mabu is surrounded by Brachystegia woodland at the base, much of which is replaced to the south by the Cha Madal tea estate. The paratype series were collected at night on perches from 30-100 cm above ground in forest clearings and adjacent to a mountain stream (Fig. 6 D). No specimens were collected on perches as high as 1.3 m to 5 m noted for Rh. chapmanorum (Tilbury 1992). This may be related to the presence of an additional chameleon (N. baylissi) which utilizes this forest stratum on Mt. Mabu, and that has not been recorded on Mt. Chiperone. Reproduction. The female allotype was gravid with seven almost spherical eggs (largest diameter 4.1-4.5 mm, four in the left oviduct and three in the right oviduct) that lack obvious signs of embryonic development., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 17-22, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp.","Timberlake, J. R., Bayliss, J., Dowsett-Lemaire, F., Congdon, C., Branch, B., Collins, S., Dowsett, R. J., Fishpool, L., Francisco, J. Harris, T., Kopp, M. & da Sousa, C. (2012) Mount Mabu, Mozambique: Biodiversity and Conservation. Report produced under the Darwin Initiative Award 15 / 036: Monitoring and Managing Biodiversity Loss in South-East Africa's Montane Ecosystems. Royal Botanic Gardens, Kew, London. pp. 94.","Bayliss, J., Timberlake, J., Alves, T., Branch, W. R., Bruessow, C., Collins, S., Congdon, C., De Sousa, C., Dowsett-Lemaire, F., Fishpool, L., Harris, T., Herrmann, E., Giogiardis, S. Liggitt, B., Monadjem, A., Patel, H., Spottiswoode, C., Taylor, P., Wilcocks, S., Ribeiro, D. & Smith, P. (2014) The discovery, biodiversity, and conservation of Mabu forest-the largest mid-altitude rainforest in southern Africa. Oryx, 48 (2), 177 - 185. http: // dx. doi. org / 10.1017 / s 0030605313000720","Tilbury, C. R. (1992) A new dwarf forest chameleon (Sauria: Rhampholeon) from Malawi, Central Africa. Tropical Zoology, 5, 1 - 9. http: // dx. doi. org / 10.1080 / 03946975.1992.10539176"]}
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- 2014
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145. Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique
- Author
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
- Subjects
Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Branch, William R., Bayliss, Julian, Tolley, Krystal A. (2014): Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique. Zootaxa 3814 (1): 1-36, DOI: http://dx.doi.org/10.11646/zootaxa.3814.1.1
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- 2014
146. Rhampholeon (Rhinodigitum) nebulauctor Branch, Bayliss & Tolley, 2014, sp. nov
- Author
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
- Subjects
Reptilia ,Squamata ,Animalia ,Rhampholeon ,Rhampholeon nebulauctor ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Rhampholeon (Rhinodigitum) nebulauctor sp. nov. Mount Chiperone Pygmy Chameleon Synonomy: Rhampholeon champmanorum (sic.) Timberlake et al. 2007, p 20. Etymology. The specific epithet is a noun in apposition and derived from nebula (L. = cloud, mist) and auctor (L. = maker), i.e. ���cloudmaker���, alluding to the ���Ciperoni���, the local name for the cold drizzle that comes to the Shire Highlands of southern Malawi as moist air from the Indian Ocean is forced to rise over Mt. Chiperone. Types. The type series comprises five specimens, including: Holotype.- An adult female (PEM R 17278) collected by J. Bayliss, 1 December 2008, in the shrub understorey of evergreen forest on the southeast slopes of Mt. Chiperone Massif, Zamb��zia Province, Mozambique (16 ˚ 30 ' 25.9 ���S, 35 ˚ 43 ' 33.4 ���E, ca 1000 m a.s.l.). Allotype. An adult male (PEM R 17281), collected by J. Bayliss on 27 November 2008; same locality details as holotype. Paratypes. Three specimens, comprising an adult female (PEM R 17277) and two subadult females (PEM R 17279 - 80), all collected by J. Bayliss between 26 November and 3 December 2008, same locality details as holotype. Meristics. Measurements for the type series of Rhampholeon nebulauctor sp. nov. are summarized in Table 7. Diagnosis. The Chiperone Pygmy Chameleon is referable to the Rhampholeon (subgenus Rhinodigitum Matthee et al. 2004) by having an unpigmented parietal peritoneum, claws that are strongly bicuspid, smooth plantar surfaces, a rostral process, and short tail (Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It is geographically closest to Rh. chapmanorum, but differs from that species by its smaller size, the presence of a relatively large rostral process in males, and accessory planter spines that are very poorly developed in both sexes. It is well differentiated from Rh. platyceps and Rh. maspictus sp. nov. by its smaller size (SVL Rhampholeon, and all chameleons examined from Mt. Chiperone form a monophyletic clade. Description of Holotype. Adult female, viscera exposed by a large ventral incision. Head: Dorsum flattened, with no upward flexure of the snout; casque flat, edged with weakly-defined lateral crests that are mainly restricted to the posterior region of the casque; temporal crest weakly-developed, comprising a single, interrupted row of large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridges reduced to a few scattered enlarged scales but with a very small multi-scaled process forming a ���soft horn��� at each end of the inter-orbital ridge that passes across crown and is composed of 11 small granular tubercles, and that demarcate the posterior edge of a slight frontal depression; inferior orbital rim with 4 (right) and 4 (left) enlarged tubercles; snout bordered on each side by moderately developed rostral crests, that fuse together at the tip of the snout which is adorned with a very small, flattened rostral process (1.5 mm), and is three small tubercular scales long and three tubercular scales wide at its base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more conical but smaller than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest very weakly developed, reduced to 7 crenulations of enlarged, but not obviously spinose scales; crenulations most strongly developed over mid-body, reduced in size over on neck and tail; crenulations continue onto tail but in more reduced form; deep axillary and inguinal pits are present; flank scalation heterogeneous, composed of small, stellate granules with few scattered, enlarged spinose tubercles, the largest at the shoulder and in a single cluster; chest, belly and lower surface of tail smooth; limb scalation more tubercular, with a few enlarged, spinose tubercles on the forearms; claws strongly bicuspid; accessory planter spines on the soles of the fore and hind feet are present, but reduced to a few very small, soft, spinose scales at the base of the claws; tail flattened laterally, flexing slightly downward on the distal third. Colour in life (Fig. 7 A, B): Mid-body mottled brown with two vague oblique lateral stripes; fore-body, neck, head and upper surfaces of limbs with extensive bright green tubercles, with scattered light blue tubercles on temporal region of head and throat and on front of belly; scales around orbit dark blue, with a red rim to the iris; tip of casque and dorsal crest cream; enlarged scales of the dorsal crest crenulations orange, that is more extensive and intense along the top of the tail. Colour in preservative. Body mottled brown with no obvious lateral stripes; lower surface of neck, belly, base of tail, soles of feet, and lower limbs pale brown, except on throat and sides of head which are dark brown. Description of Allotype (as for holotype, unless noted): adult male, very large ventral incision; hemipenes not everted. Head: Supraorbital ridge reduced to a few scattered enlarged scales but with a distinct multi-scaled process forming a ���soft horn��� at the end of the inter-orbital crest that is slightly more pronounced than in female holotype; interorbital ridge, shallow and composed of 12 small granular tubercles; rostral process six tubercular scales long and five wide at base. Body: Dorsal crest very weakly developed, reduced to eight crenulations of enlarged but not obviously spinose scales, which are subequal in development to those of the female holotype; tail relatively long (26.5 % SVL), with a prominent hemipenal bulge; distal third of tail flexes strongly downward. Colour in preservative: Body mottled brown with no obvious lateral stripes on the flanks. Pale brown below. Paratype variation. In the largest female paratype (PEM R 17277) the ���horn��� on the supraorbital ridge is reduced to a very small cluster of about 5 spinose tubercular scales; rostral process is very small, 3 scales long and 4 wide at base. Size. Presumably a small species, as all except the smallest female were reproductively active. Largest male���PEM R 17281 (allotype) 32.3 + 11.6 = 43.9 mm; largest female���PEM R 17277 (paratype) 48.9 + 13.2 = 62.1 mm. Reproduction. The three adult females contain enlarged ova, albeit without signs of embryonic development: PEM R 17277, Four ova ca. 3.4 mm dia.; PEM R 17278, three ova largest measuring 5.0 mm x 3.5 mm; PEM R 17279, two small ova (ca. 3.1 mm dia.). The smallest female, PEM R 17280, has small developing follicles. The only male (PEM R 17281) has well-developed testes despite its small size (32.3 mm SVL). Distribution. Restricted to the type locality; Mt. Chiperone, Zamb��zia Province, northern Mozambique. Habitat. The Chiperone Massif shows habitat zonation and is surrounded by Brachystegia woodland at the base. Chameleons were found on the southeast side of the mountain at 1000 m in wet forest dominated by Khaya anthotheca, Strombosia schefflerii, Rawsonia burtt-davyi, and Drypetes arguta (Fig. 7 C.), Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 22-24, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Timberlake, J. R., Bayliss, J., Alves T., Baena, S., Francisco, J., Harris, T. & da Sousa, C. (2007) The Biodiversity and Conservation of Mount Chiperone, Mozambique. Report produced under the Darwin Initiative Award 15 / 036: Monitoring and Managing Biodiversity Loss in South-East Africa's Montane Ecosystems. Royal Botanic Gardens, Kew, London. pp. 33."]}
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- 2014
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147. Rhampholeon (Rhinodigitum) platyceps Gunther 1892
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
- Subjects
Reptilia ,Squamata ,Animalia ,Rhampholeon ,Biodiversity ,Rhampholeon platyceps ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Rhampholeon (Rhinodigitum) platyceps G��nther, 1892. Mount Mulanje Pygmy Chameleon Synonomy: Brookesia platyceps Loveridge 1933 Brookesia platyceps platyceps Loveridge 1953 Brookesia platyceps carri Loveridge 1953 Rhampholeon (Rhinodigitum) platyceps Matthee, Tilbury & Townsend 2004. Holotype: Adult female (BMNH 1946.8.21.73), collected by A. White. Type locality (by subsequent designation, Stevens 1974): Mt. Mulanje, southern Malawi. Variation in adult coloration of an adult male (Fig. 4 A) and adult female (Fig. 4 B), and evergreen forest habitat in the Lichenya plateau region, are shown (Fig. 4 C). For full details of morphology, hemipenial structure, biology and habitat see Tilbury (2010)., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 15-16, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Gunther, A. (1892) Report on a collection of reptiles and batrachians transmitted by Mr H. H. Johnson from Nyasaland. Proceedings of the Royal Society of London, 1982, 555 - 558.","Loveridge, A. (1953) Zoological results of a fifth expedition to East Africa. III Reptiles from Nyasaland and Tete. Bulletin of the Museum of Comparative Zoology, 110, 141 - 322.","Matthee, C. A., Tilbury, C. R. & Townsend, T. (2004) A phylogenetic review of the African pygmy chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society of London, Series B, 271, 1967 - 1976.","Stevens, R. A. (1974) An annotated check list of the amphibians and reptiles known to occur in southeastern Malawi. Arnoldia (Rhodesia), 6 (30), 1 - 22.","Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp."]}
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- 2014
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148. Genetic structure associated with habitat diversification supports the independent evolution of ecomorphs in Bradypodion pumilum.
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Tolley, Krystal A, Hopkins, Kevin P, and da Silva, Jessica M
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HABITATS , *GENE flow , *GENETIC markers , *ECOLOGY , *MITOCHONDRIAL DNA , *NATURAL selection - Abstract
There are notable examples of repeated evolution of ecomorphs within groups of closely related species, these are typically viewed as remarkable cases of natural selection. In most cases, directional selection for certain phenotypic traits under specific (and differing) environmental conditions is implicated. Differing ecomorphological forms for closely related species (or for populations within a species) have been documented for several lineages of South African dwarf chameleons (Bradypodion), although this pattern is not ubiquitous across the genus. Within one species, Bradypodion pumilum, two ecomorphs have been observed, these differ in terms of morphology, performance, ecology and habitat, yet genetic differences (mtDNA) were not apparent, with shared haplotypes across the ecomorphs. We investigated whether more sensitive genetic markers (eight microsatellite loci) would show population level structure between ecomorphs, which could indicate lack of gene flow across the ecological (habitat) boundary. Our results suggest that the two ecomorphs are distinct at a population level, implying that gene flow is absent or reduced and that the two ecomorphs are on separate evolutionary trajectories. Given that mtDNA could not detect population structure, we infer that the ecological boundary between ecomorphs is relatively recent in evolutionary history. The two ecomorphs are distinctive in terms of their morphology, ecology, habitat and functional traits, these differences are likely driven and maintained by reproductive isolation across the habitat boundary. [ABSTRACT FROM AUTHOR]
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- 2019
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149. Anuran 3D models reveal the relationship between surface area‐to‐volume ratio and climate.
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Ferreira Amado, Talita, Moreno Pinto, Marta Gloria, Olalla‐Tárraga, Miguel Ángel, and Tolley, Krystal
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BODY size ,WATER conservation ,LENGTH measurement ,CLIMATE change ,CLIMATOLOGY - Abstract
Aim: The relationship size–environment has been continuously debated for ectotherms. Surface‐to‐volume ratios are recurrently suggested as the cause of body size responses to climate, but most inferences on organismal size clines stem from single linear measurements of body length. Here, we illustrate how new photogrammetric techniques can be applied to characterize amphibian morphologies with 3D models. Using the natterjack toad (Epidalea calamita) as model organism, we compared the performance of climatic models in accounting for body size (measured as snout‐to vent length, SVL) and body geometry (surface area‐to‐volume ratios) variation across macroclimatic gradients. Location: Palaearctic. Taxon: Epidalea calamita (former genus Bufo). Methods: We used photogrammetry techniques to reconstruct 104 museum specimens of Epidalea calamita into 3D models and to collect geometric data (volume and surface area) to calculate surface area‐to‐volume ratios (SA:V). We then apply spatial and non‐spatial regression to examine the relationships between SA:V, SVL and a set of bioclimatic variables. Results: In accordance with the water conservation hypothesis, SA:V decreased towards more arid environments, a pattern that remained consistent after accounting for spatial autocorrelation effects. Only mean annual temperature was significant correlated with SVL. The contrast between both sets of models indicates that more traditional body size metrics (SVL) and body geometry (SA:V) do not operate the same constraints. Main conclusions: We show that the variation in SA:V can be explained by the geographical variation in aridity. Evaporative water loss in wet‐skinned ectotherms is highly dependent on the surface area of the skin exposed to the air. In contrast, SVL models detected that body length was more associated with mean annual temperature. This finding stresses the importance of selecting different metrics of body geometry and body size when examining variation across climatic gradients. [ABSTRACT FROM AUTHOR]
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- 2019
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150. Ancient habitat shifts and organismal diversification are decoupled in the African viper genus Bitis (Serpentes: Viperidae).
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Barlow, Axel, Wüster, Wolfgang, Kelly, Christopher M. R., Branch, William R., Phelps, Tony, and Tolley, Krystal A.
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BITIS ,HABITATS ,VICARIANCE ,ALLELES ,OLIGOCENE Epoch ,MIOCENE Epoch - Abstract
Aim: The expansion of open habitats during the mid‐Miocene has been hypothesized as a driver of allopatric speciation for many African taxa. This habitat‐dependent mode of diversification has been implicated in the shift from C3 (e.g. forest/woodland) to C4 dominated systems (i.e. open savanna, grasslands) in a number of African squamates. We examined this hypothesis using a genus of African viperid snakes (Bitis) with both open habitat and forest‐dwelling representatives. Location: Africa. Methods: A comprehensive multilocus dataset was used to generate a calibrated species tree using a multispecies coalescent model. Individual gene trees and patterns of nuclear allele sharing were used to assess species monophyly and isolation. To test the habitat‐dependent evolution hypothesis, we generated an ancestral character state reconstruction for open and closed habitats using the dated phylogeny. This was related to the timing of open habitat expansion and forest/woodland contraction in Africa. Results: The genus Bitis originated in the Oligocene, with species level diversification in the late Miocene/Pliocene. Four well‐supported clades correspond to the recognized subgenera Bitis, Keniabitis, Macrocerastes and Calechidna. Several previously unrecognized lineages potentially represent cryptic species. Main conclusions: Habitat‐dependent evolution does not appear to have been a main driver for generic level viperine diversification: the ancestral state for Bitis was open habitat and at least one clade moved into forest in the Miocene, long after forest had contracted and fragmented. Forest‐dependent species diversified only in the late Miocene, presumably as forest became further reduced in extent, fitting an allopatric model of speciation. Although our results do not favour a general pattern of habitat‐dependent diversification in Bitis, cladogenesis within the subgenus Calechidna for "arenicolous" species (Bitis caudalis complex) and "rupicolous" species (B. atropos‐cornuta complex), corresponds to the aridification of southwest Africa. This suggests there are subtleties not captured in the broad open habitat category, which are relevant for understanding the role of habitat‐dependent evolution. [ABSTRACT FROM AUTHOR]
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- 2019
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