32 results on '"Tolley, Krystal"'
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2. Isolation of novel microsatellite loci in dwarf chameleons from KwaZulu-Natal province, South Africa and their cross-amplification in other Bradypodion species
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Feldheim, Kevin A., da Silva, Jessica M., and Tolley, Krystal A.
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- 2012
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3. Out on a limb: female chameleons (Bradypodion pumilum) position themselves to minimise detection, whereas males maximise mating opportunity.
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Rebelo, Alexander D, Altwegg, Res, Katz, Eric M, and Tolley, Krystal A
- Abstract
Although reproductive strategies are poorly documented in chameleons, sperm storage has been confirmed among some species for which male reproductive success may be maximised by increasing the number of females inseminated and/or preventing subsequent female copulation with rival males. Behaviour, such as mate searching, may enable males to maximise the number of mating opportunities with different partners and mate guarding may increase the likelihood that a male will sire offspring with a particular female. In this study, we investigated behavioural differences in movement (distance travelled) and perch choice between sexes of the Cape Dwarf Chameleon, Bradypodion pumilum, to offer data for a better understanding of mate searching and mate guarding behaviours. Wild chameleons were radio-tracked in 2010 (n = 8) and 2014 (n = 5) for approximately two weeks. The geographic location and perch height above ground were recorded multiple times per day to capture both the scale of movement and their perch use. Linear mixed effect models were used to assess differences between sexes. Our results indicated that females perched lower than males and moved shorter distances, offering support of a mate searching strategy in males. Ad hoc field observations revealed behaviours consistent with mate guarding by some males, and mate avoidance by some unreceptive females. Our results could indicate that males increase their reproductive fitness by using higher perches in both a mate guarding and mate searching context and moving further distances in search of receptive females. [ABSTRACT FROM AUTHOR]
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- 2022
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4. 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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5. 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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6. 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
- Author
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Tilbury, Colin R. and Tolley, Krystal A.
- Subjects
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
7. 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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8. Rhampholeon (Rhinodigitum) bruessoworum Branch, Bayliss & Tolley, 2014, sp. nov
- Author
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
- Subjects
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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9. Rhampholeon tilburyi Branch, Bayliss & Tolley, 2014, sp. nov
- Author
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
- Subjects
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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10. 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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11. 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
12. 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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13. Rhampholeon (Rhinodigitum) platyceps Gunther 1892
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Branch, William R., Bayliss, Julian, and Tolley, Krystal A.
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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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14. Linking microhabitat structure, morphology and locomotor performance traits in a recent radiation of dwarf chameleons
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Silva, Jessica, Herrel, Anthony, Measey, G. John, Vanhooydonck, Bieke, Tolley, Krystal, Higham, Timothy, Evolutionary Morphology of Vertebrates, and Universiteit Gent = Ghent University [Belgium] (UGENT)
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Morphometrics ,Species complex ,Ecology ,[SDV]Life Sciences [q-bio] ,[SDV.BA]Life Sciences [q-bio]/Animal biology ,Morphology (biology) ,15. Life on land ,Biology ,biology.organism_classification ,Grip strength ,Chemistry ,Evolutionary biology ,biology.animal ,Adaptation ,Parallel evolution ,Chamaeleonidae ,human activities ,Ecology, Evolution, Behavior and Systematics ,ComputingMilieux_MISCELLANEOUS ,Bradypodion - Abstract
Summary 1. Evidence that morphological traits associated with particular environments are functionally adapted to those environments is a key component to determining the adaptive nature of radiations. Adaptation is often measured by testing how organisms perform in diverse habitats, with performance traits associated with locomotion thought to be among the most ecologically relevant. 2. We therefore explored whether there are relationships between morphology, locomotor performance traits (sprint speed, forefoot and tail grip strength on broad and narrow dowels) and microhabitat use in five phenotypic forms of a recent radiation of dwarf chameleon – the Bradypodion melanocephalum–Bradypodion thamnobates species complex – to determine whether morphological differences previously identified between the forms are associated with functional adaptations to their respective habitats, which can be broadly categorized as open or closed-canopy vegetation. 3. The results showed significant differences in both absolute and relative performance values between the phenotypic forms. Absolute performance suggests there are two phenotypic groups – strong (B. thamnobates and Type B) and weak (B. melanocephalum and Types A and C). Relative performance differences highlighted the significance of forefoot grip strength among these chameleons, with the closed-canopy forms (B. thamnobates, Types B and C) exceeding their open-canopy counterparts (B. melanocephalum, Type A). Little to no differences were detected between forms with respect to sprint speed and tail strength. These results indicate that strong selection is acting upon forefoot grip strength and has resulted in morphological adaptations that enable each phenotypic form to conform with the demands of its habitat. 4. This study provides evidence for the parallel evolution of forefoot grip strength among dwarf chameleons, consistent with the recognition of open and closed-canopy ecomorphs within the genus Bradypodion.
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- 2014
15. Chamaeleo Laurenti 1768
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Tilbury, Colin R. and Tolley, Krystal A.
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Chamaeleo ,Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Genus Chamaeleo Laurenti 1768 Type species: Chamaeleo chamaeleon (Linnaeus 1758) Generic synonyms: Phumanola Gray 1864. Type species Chamaeleo namaquensis Smith 1831 Calyptosaura Gray 1864. Type species Chamaeleo calyptratus Dumeril & Bibron 1851 Erizia Gray 1864. Type species Chamaeleo senegalensis Daudin 1802. Species content: africanus, anchietae, arabicus, calcaricarens, calyptratus, chamaeleon, dilepis, gracilis, laevigatus, monachus, namaquensis, necasi, senegalensis, zeylanicus. This genus has a wide ranging pan African distribution extending into Europe, the Middle East, Arabia and the Indian sub-continent (Fig. 2). One species is confined to the island of Socotra. The distribution areas of the mainland species tend to be large and continuous except for Ch. anchietae which appears to have population pockets restricted to highland plateaux. Although some species may penetrate into lowland forest, or high altitude grasslands, the species of the genus generally occupy moist and dry woodland savannahs, thorn scrub, semi desert and in one species, true desert. Apart from occipital lobes in some species and prominent parietal crests in others, they have little other head ornamentation. None of them possess horns or any form of rostro-nasal or pre-orbital projections. A gular-ventral crest of single cones is found in all species being more or less developed in the various forms from very prominent in Ch. calyptratus to almost indiscernible in Ch. namaquensis. None of the species of this genus demonstrate a temporal crest. The casque is edged in a lateral parietal crest originating as a posterior continuation of the supra-orbital ridge which delineates the posterior ramus of the squamosal bone. The temporal zone is undivided. The background scalation of the flanks is generally composed of relatively homogeneous to finely heterogeneous closely packed granular tubercles. The tail of all species within this group is smooth. The plantar surfaces are smooth and claws simple. This is the only genus where the presence of tarsal spurs is seen in several of the species (Ch. arabicus, Ch. monachus, Ch. chamaeleon, Ch. necasi, Ch zeylanicus, Ch. dilepis, Ch. gracilis, Ch. calyptratus, Ch. africanus). These tend to be best developed in males and usually absent or much reduced in females. Tarsal spurs may be a synapomorphy for the genus Chamaeleo. The basic internal lung morphology consists of two large septae arising from the region of the hilum of the lung which end freely within the lung, dividing it into three chambers viz a small dorsal, a large middle and a small ventral chamber. All species possess a gular pouch and in the lung - a membrano-fibrous diaphragm that partially separates off a small dorso-cranial compartment. Many species also have several small partial septae that arise from the dorsal wall of the lung near the cranial end. The lungs are invariably adorned with varying numbers of diverticulae that trail from the inferior and posterior margins of the lung. The diverticulae vary in length and number and may be branched (Klaver 1973, 1977, 1981). Hemipenes are calyculate, with a multi rotulae arrangement of between three to five pairs of denticulated rotulae except for Ch. arabicus and Ch. namaquensis which have retained the plesiomorphic four rotulae (two pairs) configuration (Klaver & B��hme 1986). The genus is oviparous with a cyclic reproductive strategy ��� usually a single brood but up to three clutches of eggs per year in some species in ideal conditions. These species tend to have relative longevity. Females are usually sexually mature within one year and over the next few years will produce at least one clutch of eggs annually. The parietal peritoneum is unpigmented. The documented karyotype of the several species so far examined (Matthey 1931, 1957, Matthey & van Brink 1956) is 2 n= 24 = 12 M+ 12m and appears to be restricted to this genus as a synapomorphic character (Klaver & B��hme 1986), Published as part of Tilbury, Colin R. & Tolley, Krystal A., 2009, A re-appraisal of the systematics of the African genus Chamaeleo (Reptilia: Chamaeleonidae), pp. 57-68 in Zootaxa 2079 on pages 63-65, DOI: 10.5281/zenodo.187324, {"references":["Klaver, C. J. J. (1973) Lung anatomy: aid in chameleon taxonomy. Beaufortia, 20 (269), 155 - 177.","Klaver, C. J. J. (1977) Comparative lung morphology in the genus Chamaeleo Laurenti, 1768 (Sauria: Chamaeleonidae) with a discussion of taxonomic and zoogeographic mplications. Beaufortia 25 (327), 167 - 199.","Klaver, C. J. J. (1981) Lung morphology in the Chamaeleonidae (Sauria) and its bearing on phylogeny, systematics and zoogeography. Zeitschriftfuer zoologishe Systematik Evolutionsforsch, 19, 36 - 58.","Klaver, C. J. J. & Bohme, W. (1986) Phylogeny and classification of the Chamaeleonidae (Sauria) with special reference to hemipenis morphology. Bonner Zoologische Monographien, 22, 1 - 64.","Matthey, R. (1931) Chromosomes de reptiles saurians, ophidians, chelonians. L'evolution de la formule chromosomiale chez les saurians. Revue Suisse de Zoologie 38 (9), 146 - 147.","Matthey, R. (1957) Cytologie comparee et taxonomie des Chamaeleontidae (Reptilia - Lacertilia). Revue Suisse de Zoologie 64, 709 - 732.","Matthey, R. & van Brink, J. M. (1956) Note preliminaire sur la cytologie chromosomique comparee des cameleons. Revue Suisse de Zoologie 63, 241 - 246."]}
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- 2009
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16. Bradypodion ngomeense Tilbury & Tolley, 2009, sp. nov
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Tilbury, Colin R. and Tolley, Krystal A.
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Bradypodion ,Reptilia ,Bradypodion ngomeense ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Bradypodion ngomeense sp. nov. Holotype: PEM-R 16621, an adult male collected by Devi Stuart-Fox and Adnan Moussalli in the Ngome Forest, KZN 27 �� 49 ���S, 31 �� 25 ��� E, on 6 th January 2004. Paratypes: PEM-R 16617 adult female, PEM-R 16612 adult male, PEM-R 16613 adult male, PEM- R 16615 adult female, PEM-R 16618 adult male, PEM-R 16620 adult female, PEM-R 16622 adult female, all collected by Devi Stuart-Fox and Adnan Moussalli with same locality and date as holotype; PEM-R 16842 sub-adult male, PEM-R 16843 adult female collected by Colin Tilbury 05/01/ 2002 same locality as the holotype; PEM-R 5689 adult male & PEM-R 5690 adult female, collected by Colin Tilbury 08/01/ 1999, Ngome Forest; PEM-R 17518, PEM-R 17519, PEM-R 17520, PEM-R 17521 all collected by Krystal Tolley 14 / 11 / 2005, locality, Ngome Forest. Diagnosis: A moderately large-bodied, long-tailed species of the genus Bradypodion demonstrating the characteristics of the genus (Tilbury et al. 2006). This species differs from its various congeners in the following respects: the adults have a relatively tall casque angled at (usually) over 30 degrees or more to the supra-orbital line in keeping with 6 other South African species - viz: B. damaranum (Boulenger 1887), B. dracomontanum Raw 1976, some B. ventrale (Gray 1845), B. thamnobates Raw 1976, B. transvaalense (Fitzsimons 1930) and B. nemorale Raw, 1978 (Qudeni ecomorph). It differs from B. ventrale which has a tail that is less than 50 % of its snout /vent length; from B. thamnobates which has prominent enlarged plate-like flank tubercles; from B. damaranum which has areas of ���naked��� interstitium around the axilla and along the anterior para-vertebral zone; from B. nemorale which has a relatively much reduced gular crest, and a dorsal crest of cones larger than the diameter of the eye-opening; and from B. transvaalense (sensu stricta) where most adult specimens usually have dorsal cones larger than the eye opening and in which the superior temporal zone is pale coloured and the black mid-temporal stripe continues uninterrupted over the upper and mid flank (Figure 3). Description of Holotype: Adult male, Snout/vent length 65mm, Tail 81mm. Casque sharply elevated posteriorly. Pre-orbital, supra-orbital, and temporal crests rugose and composed of thick conical and subconical tubercles. A low median parietal crest extends forwards from the apex of the casque to terminate just before the mid-orbital region. The temporal crests originate from the mid post-orbital rim, are prominent and composed of a row of 6 more or less equally enlarged tubercles on the right side and 5 on the left. Posteriorly, the temporal crest then rises almost vertically as the squamosal crest to run parallel and adjacent to the lateral parietal crest to the apex of the casque. A prominent gular crest is present, composed of 14 granulated composite lobes, numbers 2 ��� 6 overlapping and numbers 2 ��� 4 are broader than long. The free distal edges of the lobes have a denticulated fringe. Five to six thin white gular interstitial grooves are present on each side of the throat, enclosing chains of enlarged rounded tubercles. A prominent dorsal crest is present, composed of 26 conical tubercles roughly triangular in outline, extending from the nuchal area to the sacrum and then extending along the proximal third of the tail. The tubercles of the dorsal crest are highest over the anterior to mid vertebral line and lowest over the sacral region, none of the cones are taller than the diameter of the eye opening. Background scalation of heterogeneous, rounded to polygonal tubercles with no obvious interstitial venation between the tubercles. A short row of slightly enlarged flattened tubercles skirts the upper mid-flank. Enlarged tubercles are scattered along the para-vertebral zone. Belly tubercles arranged in pallisades of more or less homogeneous rounded tubercles. Variation in the Paratypes: The largest specimen, a female (PEM-R 16617) has a total length of 166mm (snout/vent 77mm + tail 89mm). Considerable variation in the gular crest is present among the types. The gular crest varies from 11 to 19 individual lobes or tubercles. The length and width of the lobes is variable, in some specimens all the lobes are longer than wide, in others the length / width may be similar and in some the overlapping lobes may be broader than long. Although overlap of between 2 to 7 lobes is common, in several specimens there is no overlap of any of the lobes. In 9 / 12 specimens, the first gular projection is a simple conical tubercle, and in all specimens, the last gular projection is a simple cone. Up to the last 3 gular ornaments may be simple cones. In most specimens the flanks appear to have no interstitial web, but in 2 / 14 specimens, an indication of interstitial webbing was present. The dorsal crest in the type series consisted of between 19 ��� 26 tubercles extending between the nuchal fold to the mid-sacrum. It may additionally extend for a variable distance along the tail from being almost imperceptible to easily discernable up to half of the length of the tail. The tail is always longer than the snout/vent length in both sexes. Sexual dimorphism: Males tend to have a more rugose scalation, better developed body and head crests, and a prominent hemipenal bulge at the base of the tail. Tail length differs between the sexes, being 53���57 % of the total length in males compared to 53���55 % in females. Colour differences between the sexes are discussed below. Colour in life: Male���Head above the upper labials mostly black ��� labials and all cranial crests pale yellow. Gular region and gular crest white. Nuchal fold - white to pale yellow. Skin of eyeball, heavily flecked with red around the periphery and a single dark horizontal stripe transects the eye. Background colour of the body is blue-green as is the dorsal crest. A bright red to orange flash over the mid flank is fully enclosed within a black flank patch which terminates abruptly about 2 / 3 rds of the way along the flank. The colour of the flash is variable from white to various shades of red. Belly, front legs and tail blue-green. Female ��� Background pale green. The superior and inferior temporal zones have white patches. The superior half of the anterior 2 / 3 rds of the flank is dark green, with scattered lighter tubercles (Figures 4 & 5). Hemipenes: None of the specimens currently present in any museum collection have fully everted hemipenes so the structure of these organs remains un-described for the present. Habitat: Northern Afrotemporate ���Mist Belt��� forest between 500 and 1360 meters a.s.l. characterized by a tall, but multi-layered canopy up to 30 meters with both Afrotemperate species and sub-tropical elements (Mucina & Geldenhuys 2006). The forest occupies the deeply incised southeast facing slopes of the Ntendeka cliffs. The largest trees in the Ngome Forest are specimens of the forest waterwood (Syzygium gerrardii) reaching up to 30 meters. Other trees include Podocarpus latifolius, Octea kenyensis, Faurea macnaughtonii, Olinia radiata and Trichocladus grandiflorus. The forest is rich in species of epiphytic orchids and hydrophilous ferns. Most specimens collected in the type series were found along forest paths and the road verges in bushes and trees at perch heights between 1���10 meters from the ground. Conservation: The entire known range of this species is restricted to the Ngome Forest. It is the largest fragment of Mist Belt forest in KwaZulu-Natal with an estimated area of 3360 Ha (36.6 km 2 - Mucina & Rutherford 2006.). Between 1876 and 1899, the forest was heavily logged and most of the larger Podocarpus and other hard wood species were removed or depleted. The forest has been protected since 1905. Much of the forest (2,636 Ha) is managed by the South African Department of Water Affairs and Forestry (DWAF) and protected within the Ntendeka Wilderness Area. Although some of the adjacent scarp area has been given over to pine plantations, there is no pine within the Wilderness Area. Etymology: The species is named after the Ngome Forest - the only forest known to harbor this taxon., Published as part of Tilbury, Colin R. & Tolley, Krystal A., 2009, A new species of dwarf chameleon (Sauria; Chamaeleonidae, Bradypodion Fitzinger) from KwaZulu Natal South Africa with notes on recent climatic shifts and their influence on speciation in the genus, pp. 43-57 in Zootaxa 2226 on pages 47-50, DOI: 10.5281/zenodo.190207, {"references":["Raw, L. R. G. (1976) A survey of the dwarf chameleons of Natal, South Africa, with the description of three new species (Sauria: Chamaeleonidae). Durban Museum Novitates, 11, 139 - 161.","Raw, L. R. G. (1978) A further new dwarf chameleon from Natal, South Africa (Sauria: Chamaeleonidae). Durban Museum Novitates, 11, 265 - 269.","Mucina, L. & Rutherford, M. C. (eds) (2006) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria."]}
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- 2009
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17. Bradypodion Fitzinger 1843
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Tilbury, Colin R., Tolley, Krystal A., and Branch, William R.
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Bradypodion ,Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Bradypodion Fitzinger 1843 Type Species: Chamaeleo pumilus Daudin 1802 Composition: B. caffer (Boettger), B. damaranum, B. dracomontanum Raw, B. gutturale (Smith), B. kentanicum (Hewitt), B. melanocephalum, B. nemorale Raw, B. occidentale, B. pumilum, B. setaroi Raw, B. taeniabronchum, B. thamnobates Raw, B. transvaalense (Fitzsimons), B. ventrale, and several as yet undescribed species (Tolley et al. 2005; Branch et al. 2006). Characterization: The monophyly of the South African Bradypodion is established by a suite of nuclear and mitochondrial genes and can also be defined by three characteristics, namely; independently derived viviparity and the associated pigmentation of the parietal peritoneum, and a specific cranial structure with a broad roof-like parietal (interpreted as a retained symplesiomorphy) bearing supra-temporal processes (interpreted as a secondary character reversal Rieppel & Crumley 1997). External morphological features that are common to all Bradypodion include the presence of heterogeneous background scalation, a midline gular crest consisting in most species of composite lobes and cones, and the absence of a ventral crest. Rostronasal processes are absent in all species. The hemipenes are calyculate with a plesiomorphic 4 -rotulae apical ornamentation. All species are viviparous. The genus may also be characterized by lung morphology, comprising simple, adiverticulate, sac-like lungs with small ridge-like septae on the cephalic, dorsal and ventral walls and with an accessory gular pouch (Beddard 1997, Klaver 1973, 1881). However, not all Bradypodion species have been assessed. Distribution: Restricted to South Africa, ranging into adjacent Swaziland, and possibly Lesotho and southern Mozambique, with introduced populations in Namibia, and occupying a wide variety of habitats., Published as part of Tilbury, Colin R., Tolley, Krystal A. & Branch, William R., 2006, A review of the systematics of the genus Bradypodion (Sauria: Chamaeleonidae), with the description of two new genera, pp. 23-38 in Zootaxa 1363 on page 34, DOI: 10.5281/zenodo.174715, {"references":["Fitzinger, L. J. (1843) Systema Reptilium Auctore Leopoldo Fitzinger. Vindobonnae (Vienna).","Tolley, K. A., Tilbury, C. R., Branch, W. R. & Matthee, C. A. (2005) The dwarfs of Africa: taxonomy, distribution and diversity of dwarf chameleons (Bradypodion sensu lato). In Abstracts, Fifth World Conference of Herpetology, Stellenbosch, South Africa, 2005, pp. 156.","Branch, W. R., Tolley, K. A., & Tilbury, C. R. (2006) A new Dwarf Chameleon (Sauria: Bradypodion Fitzinger 1843) from the Cape Fold Mountains, South Africa. African Journal of Herpetology, in press.","Rieppel, O. & Crumley, C. (1997) Paedomorphosis and skull structure in Malagasy chamaeleons (Reptilia: Chamaeleoninae), Journal of Zoology (London), 243, 351 - 380.","Klaver, C. J. J. (1973) Lung anatomy: aid in chameleon taxonomy. Beaufortia, 20, 155 - 177."]}
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- 2006
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18. Nadzikambia
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Tilbury, Colin R., Tolley, Krystal A., and Branch, William R.
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Reptilia ,Squamata ,Animalia ,Nadzikambia ,Biodiversity ,Chordata ,Chamaeleonidae ,Taxonomy - Abstract
Nadzikambia genus nova Type Species: Chamaeleo mlanjensis Broadley 1966 Composition: Nadzikambia mlanjense Characterisation: The monophyly of Nadzikambia is established on the basis of a suite of mitochondrial and nuclear genetic characters and a unique hemipenis. The latter is rather short and stout with a short pedicel and shallow calyces. There are no rotulae on the apex, which are replaced with a pair of large fleshy, papillate lobes with scalloped edges. At the sulcal base of each lobe are one or two pedunculated papillae (Klaver & B��hme 1986). The lungs show a structure similar to that of Kinyongia, with two pairs of long diverticulae trailing from the inferior and posterior surfaces of the lung. A series of four small septae alternate with five larger septae across the dorsal wall with several smaller septae arising from the ventral and cephalic walls. There is no gular pouch (Klaver 1977). The parietal peritoneum is unpigmented and reproduction oviparous. The external morphology is largely conservative, and gular and ventral crests are absent. There is a weak dorsal crest, finely heterogeneous scalation that forms rosettes of tubercles on the lower flanks, and a low casque. Although cranial morphology remains undescribed, the external appearance of the cranial crests suggest that the cranium is constructed in the same manner as Kinyongia. Distribution: Only known from sub-montane forest habitats in a few scattered localities on the Mulanje Massif in southern Malawi, Central Africa. Etymology: The name is derived from ���Nadzikambe���, the name for chameleon in ChiChewa, the language used by the tribe that lives in the area around Mulanje Mountain in southern Malawi. The name Nadzikambe is Latinised by terminating it with the suffix ia thus giving it a feminine gender., Published as part of Tilbury, Colin R., Tolley, Krystal A. & Branch, William R., 2006, A review of the systematics of the genus Bradypodion (Sauria: Chamaeleonidae), with the description of two new genera, pp. 23-38 in Zootaxa 1363 on pages 35-36, DOI: 10.5281/zenodo.174715, {"references":["Klaver, C. J. J. & Bohme, W. (1986) Phylogeny and classification of the Chamaeleonidae (Sauria) with special reference to hemipenis morphology. Bonner zoologische Monographisce 22, 1 - 64."]}
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- 2006
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19. Kinyongia
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Tilbury, Colin R., Tolley, Krystal A., and Branch, William R.
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Reptilia ,Squamata ,Animalia ,Biodiversity ,Chordata ,Chamaeleonidae ,Kinyongia ,Taxonomy - Abstract
Kinyongia genus nova Type Species: Chamaeleo fischeri fischeri Reichenow 1887. Composition: K. adolfifriderici, K. carpenteri, K. excubitor, K. fischeri fischeri, K. fischeri multituberculatum (Nieden), K. fischeri uluguruense (Loveridge), K. tavetanum, K. tavetanum boehmei (Lutzman & Nečas), K. uthmoelleri, K. xenorhinum, K. oxyrhinum, and K. tenue Characterization: The monophyly of Kinyongia is established on the basis of a suite of nuclear and mitochondrial genetic characters. No morphological synapomorphy is known to define all members. Cranial structure has only been studied in K. fischeri (Rieppel & Crumley 1997). The parietal is reduced to a narrow posteriorly projecting sagittal process that meets the ascending squamosal processes at the apex of the casque to completely enclose the temporal fossa. This derived condition is similar to that found in the genera Chamaeleo, Furcifer and Calumma (Rieppel 1981, 1987, Rieppel & Crumley 1997). Scalation is generally of finely heterogeneous granules or flattened polygonal tubercles. In those species that are characterized by a head ornamentation of fused rostronasal projections (carpenteri, xenorhinum, tenue and oxyrhinum), the scalation is generally an unordered heterogeneous mix of tubercles. In species with paired rostronasal projections (fischeri, tavetanum, uthmoelleri) the flanks are adorned with tubercles clustered into “rosettes”, especially on the lower flanks. This rosetting is also seen in the hornless species excubitor. Plantar surfaces are smooth and claws are simple. None of the species have midline gular or ventral crests, occipital lobes, or annulated horns. Cranial ornamentation in some species, e.g. paired rostronasal blade-like horns in fischeri and tavetanum, and fusion of the canthal ridges into a single vertically flattened process in carpenteri, xenorhinum, tenue and oxyrhinum, are similar to features found in the Malagasy genera Calumma and Furcifer. Lung structure is relatively plesiomorphic. They are similar to those of Bradypodion and Malagasy Calumma and Furcifer, being generally simple with a number of small septae on the dorsal, cephalic and ventral walls. The lungs of Kinyongia appear to lack the accessory gular pouch and usually have trailing diverticulae from the posteroinferior surface of the lung (tavetanum, fischeri, tenue, and adolfifriderici) although these are lacking in K. xenorhinum (Klaver 1977, 1981). The lungs in the rest of the species of Kinyongia have not as yet been described. The hemipenes are calyculate with a plesiomorphic 4 rotulae apical ornamentation, and all the species are oviparous. Distribution: Distributed in East Africa with the most westerly species, K. adolfifriderici, extending into the eastern DRC, and K. excubitor reaching as far north as Mount Kenya. They are confined to tropical/sub-tropical forest biomes, often in relict montane or sub-montane forests (Fig. 1). Etymology: This genus is largely confined to the three countries that make up the central east African region namely Kenya, Tanzania and Uganda. The lingua franca for this region is Swahili. The name derives from the generic Swahili name for chameleon “Kinyonga” and identifies it as a genus that is largely confined to Swahili speaking countries. The name is Latinized by terminating the name spelling with the letters ia giving it a feminine gender. Thus the specific names remain unaltered.
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- 2006
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20. Diversification through ecological opportunity in dwarf chameleons.
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da Silva, Jessica Marie and Tolley, Krystal A.
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DWARFISM , *CHAMELEONS , *BIODIVERSITY , *MITOCHONDRIAL DNA , *MICROSATELLITE repeats , *BAYESIAN analysis - Abstract
Aim To investigate whether morphological variation in a continental radiation of lizards (Chamaeleonidae) was driven by ecological opportunity. Location KwaZulu-Natal, South Africa. Methods Chameleons from all morphotypes were genotyped ( n = 279) using 10 microsatellite and one mitochondrial marker ( n = 130). Genetic spatial structure was examined in a Bayesian framework and compared to ecomorphological forms, to identify distinct chameleon groups using a total evidence approach. Ecological niche modelling projected back to the Last Glacial Maximum and the Last Interglacial was used to make inferences regarding the processes that may have facilitated diversification through ecological opportunity. Results For the three inland groups, genetic and morphological evidence were in agreement suggesting these are candidate species. However, there was discordance in evidence for populations from coastal regions, although the different time-scales of the data allow us to make some inferences regarding this disparity. The microsatellite data suggest at least three independent origins of coastal chameleons, although all three clusters show the same ecomorph, which indicates convergence of phenotype in a similar environment. The species distribution models revealed high niche stability among inland groups, suggesting the presence of refugia for these chameleons; whereas the coastal niche is more dynamic and likely resulted in recent origins of these groups. Main Conclusions A dynamic climatic niche and corresponding vegetation shifts are likely to have shaped contrasting ecomorphological and genetic patterns within this continental radiation by creating ecological opportunity for the coastal chamleons. In contrast, inland groups are in climatically stable regions, and show concordance between lines of evidence, suggesting that niche stability has allowed for these chameleons to diverge genetically and phenotypically. [ABSTRACT FROM AUTHOR]
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- 2017
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21. Does diet drive the evolution of head shape and bite force in chameleons of the genus Bradypodion?
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Dollion, Alexis Y., Measey, G. John, Cornette, Raphaël, Carne, Liza, Tolley, Krystal A., Silva, Jessica M., Boistel, Renaud, Fabre, Anne-Claire, and Herrel, Anthony
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ANIMAL feeding behavior ,CHAMELEONS ,LIZARD evolution ,LIZARD morphology ,ANIMAL sexual behavior ,ANIMAL species - Abstract
The head is a complex integrated system that is implicated in many vital functions. As such, its morphology is impacted by different and sometimes conflicting demands. Consequently, head shape varies greatly depending on the environment and dietary ecology of an organism. Moreover, given its role in territory defence and mating in lizards, it is also subjected to strong sexual selection in these animals., We investigated the relationships between head shape, bite performance and diet in 14 of the 17 extant Bradypodion species to determine whether variation in diet can explain the observed diversity in bite force and head shape in this genus. We also evaluate differences between sexes in terms of the relationships between head shape, bite force and diet and predict tighter relationships in females given that the head in this sex is principally under natural selection., Our results show that there is indeed a correlation between head shape, diet and bite force, but the direction and magnitude are sex-dependent. Whereas we observed a correlation between absolute bite force and head shape in both sexes, size-corrected bite force was correlated with mandible and quadrate shape in females only. Despite strong correlations between bite force and prey hardness, and between prey hardness and head shape, we did not find any relationship between head shape and prey evasiveness., These data suggest that the cranial system in chameleons of the genus Bradypodion evolves under natural selection for the ability to eat large or hard prey. Moreover, significant differences in the ecomorphological relationships between the two sexes suggest that sexual selection plays a role in driving the evolution of bite force and head shape. These data suggest that ecomorphological relationships may be sex-dependent. [ABSTRACT FROM AUTHOR]
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- 2017
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22. 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 P., Davenport, Tim R. B., Howell, Kim M., Tilbury, Colin R., Machaga, Sophy, and Tolley, Krystal A.
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CHAMELEONS ,MOUNTAINS ,UPLANDS ,GENETICS ,BIOLOGY - 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. [ABSTRACT FROM AUTHOR]
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- 2015
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23. Multiple paternity and sperm storage in the Cape Dwarf Chameleon ( Bradypodion pumilum ).
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Tolley, Krystal A., Chauke, Lucas F., Jackson, Jennifer C., and Feldheim, Kevin A.
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MICROSAURA pumila , *CHAMELEON behavior , *LIZARD reproduction , *ANIMAL breeding & the environment - Abstract
Although multiple paternity and sperm storage are common in squamates, neither has been conclusively demonstrated in the chameleons (Chamaeleonidae). To examine whether these traits are present in chameleons, the Cape Dwarf Chameleon (Bradypodion pumilum) was investigated. This viviparous species inhabits a temperate climate in southern Africa, has aseasonal breeding, and can produce multiple clutches annually. Given this reproductive schedule, we hypothesised that females may mate opportunistically and store sperm, and that single clutches may have multiple sires. To examine this, six females and their clutches were genotyped using microsatellite markers, and assessed for multiple paternity. In addition, 46 females were assessed for sperm storage structures and presence of sperm in the oviduct. Multiple paternity was found in all six clutches, based on the presence of multiple alleles at all loci that could not be assigned to the mother. No evidence of specialised sperm storage structures was found, but sperm was present in the lumen of the posterior oviduct in 74% of females. While sperm storage cannot be concluded with finality, it seems likely to occur, rather assuming that three out of four of the females in the sample had mated just prior to capture. Furthermore, two post-partum females retained sperm in the oviducts after the delivery of a clutch, but in the absence of additional copulation. Given that this species has aseasonal reproduction, if sperm is indeed stored long-term, this combination of traits would be advantageous given the lack of defined breeding season and presumably low rates of mate encounter. [ABSTRACT FROM AUTHOR]
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- 2014
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24. Temporal changes in allelic variation among Cape Dwarf Chameleons, Bradypodion pumilum , inhabiting a transformed, semi-urban wetland.
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Katz, Eric M., Tolley, Krystal A., and Bishop, Jacqueline M.
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MICROSAURA pumila , *CHAMELEON behavior , *POLYMORPHIC transformations , *ANIMAL ecology - Abstract
The Cape Dwarf Chameleon,Bradypodion pumilum, is threatened by extensive habitat loss and transformation in the Western Cape Province of South Africa. As a result, the species exists as a collection of populations inhabiting an increasingly fragmented landscape within a critically endangered ecosystem. In this study we monitored microsatellite genetic variation in one of these populations ofB. pumilumacross a three-year period. From the results of a temporary site-vacancy of adult chameleons during the study, we anticipated a significant shift in microsatellite allele frequencies. Using data from eight highly polymorphic loci we detected differences in the allelic composition of chameleons inhabiting the study site between pre- and post-site-vacancy periods (AMOVARST= 0.017;P= 0.019). Results suggest that recolonisation by immigrant chameleons, from an adjacent vegetation fragment connected to the study site by corridors, caused the recorded shifts in allelic frequencies. Pairwise tests of year by year comparisons revealed significant shifts in allelic frequencies between years one and two and between one and three, but not between years two and three. Our findings highlight the susceptibility of small vertebrates to stochastic changes in the allelic composition of populations in a fragmented landscape, and can be useful for the development of biodiversity management in an increasingly fragmented habitat mosaic. [ABSTRACT FROM AUTHOR]
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- 2014
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25. Survival and abundance of Cape dwarf chameleons, Bradypodion pumilum, inhabiting a transformed, semi-urban wetland.
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Katz, Eric M., Tolley, Krystal A., and Altwegg, Res
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MICROSAURA pumila , *MARK & recapture (Population biology) , *DEMOGRAPHIC surveys , *SURVIVAL , *CHAMELEONS - Abstract
The article discusses the study that investigated the temporal dynamics of local demographic parameters for a population of Cape dwarf chameleon, Bradypodion pumilum, in Noordhoek, South Africa, using robust design (RD) capture-mark-recapture (CMR) models for population demographics estimation. The study found that a fluctuating abundance of individuals is supported by the site and higher 30-day and annual survival rates for larger chameleons, regardless of sex.
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- 2013
26. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot.
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Tolley, Krystal A., Tilbury, Colin R., Measey, G. John, Menegon, Michele, Branch, William R., and Matthee, Conrad A.
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BIODIVERSITY , *FORESTS & forestry , *CHAMELEONS , *HABITATS , *RADIATION - Abstract
Aim East Africa is one of the most biologically diverse regions, especially in terms of endemism and species richness. Hypotheses put forward to explain this high diversity invoke a role for forest refugia through: (1) accumulation of new species due to radiation within refugial habitats, or (2) retention of older palaeoendemic species in stable refugia. We tested these alternative hypotheses using data for a diverse genus of East African forest chameleons, Kinyongia. Location East Africa. Methods We constructed a dated phylogeny for Kinyongia using one nuclear and two mitochondrial markers. We identified areas of high phylogenetic diversity (PD) and evolutionary diversity (ED), and mapped ancestral areas to ascertain whether lineage diversification could best be explained by vicariance or dispersal. Results Vicariance best explains the present biogeographic patterns, with divergence between three major Kinyongia clades (Albertine Rift, southern Eastern Arc, northern Eastern Arc) in the early Miocene/Oligocene (> 20 Ma). Lineage diversification within these clades pre-dates the Pliocene (> 6 Ma). These dates are much older than the Plio-Pleistocene climatic shifts associated with cladogenesis in other East African taxa (e.g. birds), and instead point to a scenario whereby palaeoendemics are retained in refugia, rather than more recent radiations within refugia. Estimates of PD show that diversity was highest in the Uluguru, Nguru and East Usambara Mountains and several lineages (from Mount Kenya, South Pare and the Uluguru Mountains) stand out as being evolutionarily distinct as a result of isolation in forest refugia. PD was lower than expected by chance, suggesting that the phylogenetic signal is influenced by an unusually low number of extant lineages with long branch lengths, which is probably due to the retention of palaeoendemic lineages. Main conclusions The biogeographic patterns associated with Kinyongia are the result of long evolutionary histories in isolation. The phylogeny is dominated by ancient lineages whose origins date back to the early Miocene/Oligocene as a result of continental wide forest fragmentation and contraction due to long term climatic changes in Africa. The maintenance of palaeoendemic lineages in refugia has contributed substantially to the remarkably high biodiversity of East Africa. [ABSTRACT FROM AUTHOR]
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- 2011
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27. Eastward from Africa: palaeocurrent-mediated chameleon dispersal to the Seychelles islands.
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Townsend, Ted M., Tolley, Krystal A., Glaw, Frank, Böhme, Wolfgang, and Vences, Miguel
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CHAMELEONS ,PHYLOGENY ,MAXIMUM likelihood statistics ,STATISTICAL bootstrapping ,HABITATS ,PHYSIOLOGY - Abstract
The article discusses results of a multi-locus phylogenetic study of chameleons. The study material and methods are discussed, including taxon sampling and maximum-likelihood (ML) best-tree and boostrap analyses. The study showed that Seychelles-endemic Calumma tigris is the sister taxon of a southern African clade and requires accommodation in its own genus as Archaius tigris. It also noted that Calumma tigris differs from its closest relatives, Rieppeleon in microhabitat usage, tail length and general appearance.
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- 2011
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28. 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 N.V., Altwegg, Res, and Measey, G. John
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CHAMELEONS , *REPTILES , *BRADYPODION , *MICROSAURA pumila , *LIZARDS , *HABITATS - 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. [ABSTRACT FROM AUTHOR]
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- 2010
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29. A new species of chameleon (Sauria: Chamaeleonidae: Kinyongia) from the Magombera forest and the Udzungwa Mountains National Park, Tanzania.
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MENEGON, MICHELE, TOLLEY, KRYSTAL A., JONES, TREVOR, ROVERO, FRANCESCO, MARSHALL, ANDREW R., and TILBURY, COLIN R.
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ANIMAL species , *CHAMELEONS , *PHYLOGENY ,MAGOMBERA Forest Reserve (Tanzania) - Abstract
The article presents a study that deals with a new species of chameleon Kinyongia magomberae sp. nov. or the Magombera chameleon found in the Magombera forest and the Udzungwa Mountains National Park in Tanzania. Phylogenetic analyses using mitochondrial and nuclear markers helped determine the specie status. A single rostral appendage made the new chameleon similar with K. tenuis and the more widespread Eastern Arc endemic K. oxyrhina. It is determined as sister to K. tenuis and part of a clade that also contains K. oxyrhina.
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- 2009
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30. Biogeographic patterns and phylogeography of dwarf chameleons ( Bradypodion) in an African biodiversity hotspot.
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TOLLEY, KRYSTAL A., BURGER, MARIUS, TURNER, ANDREW A., and MATTHEE, CONRAD A.
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BIOGEOGRAPHY , *PHYLOGEOGRAPHY , *CHAMELEONS , *BRADYPODIDAE , *BIODIVERSITY , *CLIMATE change , *BIOCLIMATOLOGY , *MITOCHONDRIAL DNA , *ANIMAL genetics - Abstract
The southern African landscape appears to have experienced frequent shifts in vegetation associated with climatic change through the mid-Miocene and Plio–Pleistocene. One group whose historical biogeography may have been affected by these fluctuations are the dwarf chameleons ( Bradypodion), due to their associations with distinct vegetation types. Thus, this group provides an opportunity to investigate historical biogeography in light of climatic fluctuations. A total of 138 dwarf chameleons from the Cape Floristic Region of South Africa were sequenced for two mitochondrial genes (ND2 and 16S), and resulting phylogenetic analyses showed two well-supported clades that are distributed allopatrically. Within clades, diversity among some lineages was low, and haplotype networks showed patterns of reticulate evolution and incomplete lineage sorting, suggesting relatively recent origins for some of these lineages. A dispersal-vicariance analysis and a relaxed Bayesian clock suggest that vicariance between the two main clades occurred in the mid-Miocene, and that both dispersal and vicariance have played a role in shaping present-day distributions. These analyses also suggest that the most recent series of lineage diversification events probably occurred within the last 3–6 million years. This suggests that the origins of many present-day lineages were founded in the Plio–Pleistocene, a time period that corresponds to the reduction of forests in the region and the establishment of the fynbos biome. [ABSTRACT FROM AUTHOR]
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- 2006
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31. Phylogenetics of the southern African dwarf chameleons, Bradypodion (Squamata: Chamaeleonidae)
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Tolley, Krystal A., Tilbury, Colin R., Branch, William R., and Matthee, Conrad A.
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PHYLOGENY , *TAXONOMY , *MICROSAURA pumila - Abstract
The taxonomic relationships within the dwarf chameleons (Bradypodion) of southern Africa have long been controversial. Although informal phenotypic groups have been suggested, the evolutionary relationships among the 15 recognised species in southern Africa have not been previously investigated. To investigate the relationships among species within this genus, fragments of two mitochondrial genes (16S ribosomal RNA and ND2) were sequenced and analysed using maximum parsimony, maximum likelihood and Bayesian inference. All analyses showed congruent topologies, revealing at least 5 well-supported clades distributed across distinct geographic regions. The mtDNA gene tree indicated that in many instances, geographic location has played a role in shaping the evolution of this group, and that the previously suggested phenotypic groupings do not adequately reflect evolutionary relationships. Furthermore, it appears that some of the currently recognised species (described on morphology) are polyphyletic for mitochondrial sequences, most notably those occurring in the isolated forest patches of north-eastern South Africa, near the Drakensberg Escarpment. [Copyright &y& Elsevier]
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- 2004
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32. Large-scale phylogeny of chameleons suggests African origins and Eocene diversification.
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Tolley, Krystal A., Townsend, Ted M., and Vences, Miguel
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CHAMELEONS , *PHYLOGENY - Abstract
An abstract of the study "Large-Scale Phylogeny of Chameleons Suggests African Origins and Eocene Diversification," by Krystal A. Tolley, et al. is presented.
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- 2013
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