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Brookesia micra Frank Glaw & Jörn Köhler & Ted M. Townsend & Miguel Vences 2012, sp. n.
No full textBrookesia micra sp. n. ZooBank LSID: urn:lsid:zoobank.org:act:D1A239D6-93E8-4C34-A428-F79A2C8B6405 Remark.—This species has been considered before as Brookesia sp. ‘‘Nosy Hara’’ [7]. Holotype.— ZSM 2181/2007 (FGZC 1271), adult male (hemipenes everted), collected on Nosy Hara island, 12°14'40"S, 49°0 0'30"E, ca. 10–20 m a.s.l., Antsiranana Province, northern Madagascar, on 7 March 2007 by H. Enting, F. Glaw and J. Köhler. Paratypes.— ZSM 2180/2007 (FGZC 1270), ZSM 2182/2007 (FGZC 1275), juveniles, ZSM 2183/2007 (FGZC 1278), ZSM 2185/2007 (FGZC 1280), adult males (hemipenes everted), ZSM 2184/2007 (FGZC 1279), ZSM 2186–2187/2007 (FGZC 1281– 1282), adult females, all with same data as holotype; UADBA uncatalogued (FGZC 1830), male, ZSM 1507 /2008 (FGZC 1831), male (not examined morphologically), UADBA uncatalogued (FGZC 1833), female, ZSM 1509 /2008 (FGZC 1834), adult female, ZSM 1510 /2008 (FGZC 1832), juvenile, all collected at small stream on Nosy Hara, 12°14'59"S, 49°0 0'28"E, 12 m a.s.l., Antsiranana Province, northern Madagascar, on 22 February 2008 by F. Glaw and J. Köhler. Diagnosis.— A member of the Brookesia minima group based on small body size (SVL 7.2% to all other species of the B. minima group. Description of holotype.— Adult male in good state of preservation (Fig. 7; Supporting Information S 1). Both hemipenes everted. Measurements in Table 2. Head with lateral crest starting at median level at the posterior edge of eye; prominent orbital crests; a weak crest at the posterior edge of the head, that forms a weakly developed dorsal helmet; a pair of low indistinct parasagittal crests slightly converging before terminating at the lateral crest; three similar-sized pointed tubercles on each side of posterior crest, one at termination point of lateral crest, one at termination point of parasagittal crest, and a one between parasagittal and lateral crests; one pointed tubercle on lateral surface of head, below lateral crest in temporal region; orbital crest denticulated; no supraocular cone recognizable; supranasal cone small; head longer (4.5 mm) than wide; chin and throat without longitudinal rows of slightly enlarged tubercles. Dorsal surface of body without a vertebral ridge or keel; 11 dorsolateral pointed tubercles form a complete longitudinal line on the body; most posterior (11th) pointed dorsolateral tubercle being largest, above insertion point of hindlimb; pointed dorsolateral tubercles almost equally spaced, pointing out almost perpendicularly from body; slightly enlarged, rounded tubercles form curved transversal crests on either side of the vertebral line between 1 st and 10th dorsolateral pointed tubercles; slightly enlarged scales forming four parallel longitudinal rows on dorsal surface of tail; no well-defined dorsal pelvic shield in sacral area; scattered enlarged rounded tubercles on the lateral surface of body; venter without enlarged tubercles; no pointed tubercles around cloaca; no enlarged tubercles on lateral or ventral surfaces of tail. After almost four years in ethanol, back, flanks, dorsal surfaces of limbs and anterior part of tail uniformly dark grey-brown; posterior part of tail beige; neck grey; throat and venter light grey. Variation.— For morphological measurements and proportions see Table 2 and Supporting Information S1. In preservative, ZSM 2185/ 2007 and 2188 /2007 have a greyish dorsal surface of head and neck and a slightly lighter vertebral region. The throat is uniformly dark brown (ZSM 2187/ 2007 and 2184 /2007) or beige with brown spots (ZSM 2185/2007, 2188/2007, and 1509/2008). The tail base of the males is only slightly more thickened than in the females. In life, most individuals with dorsal colouration of head, dorsum and tail light grey, lateral parts of the tail more yellowish becoming orange posterior to the tail base. Lateral parts of the body brown, with few dark brown spots; limbs almost uniformly dark brown. This pattern is likely to refer to a stress colouration. When unstressed, most parts of the body dark brown, except a beige area on the head anterior to the eyes. Tail base dark brown, orange in the middle and yellowish posteriorly. Colour and pattern of juveniles resemble those of adults and they can also show the stress colouration, but tails are slightly less colourful than in adults and usually do not show bright orange colour. Genital morphology.— Everted hemipenes of this species are available for three specimens (holotype ZSM 2181/2007, paratypes ZSM 2183/2007, ZSM 2185/2007). Each hemipenis is an elongated, relatively wide tubular structure. Apex and truncus are devoid of any ornaments. In the holotype, hemipenis length is 2.4 mm and maximum hemipenis width is 0.9 mm. When fully everted, the most characteristic feature is the apex which is a flat surface distally forming a symmetrical comb of six large, rounded papillae, of which the two inner ones are largest, followed by the two intermediate ones, whereas the outer ones are merely two slightly elevated knobs, recognizable only in fully everted organs. In general, this comb of papillae is not visible in hemipenes that are not fully everted, different from the spines of, e.g., B. peyrierasi that typically are also visible in incompletely everted organs. All three examined specimens have one hemipenis fully everted, the second one only partially everted. The organs are largely transparent, and the central retractor muscles can be easily seen through the outer hemipenial integument (Fig. 6). Etymology.— The species epithet is a latinized derivation from the Greek word ‘‘MIKRóz’’ (mikros), meaning ‘‘tiny’’ or ‘‘small’’ and referring to the extremely diminutive body size. It is used as an invariable noun in apposition. Distribution.— Only known from two sites (see localities in type series) on the small island of Nosy Hara, northern Madagascar (e.g. Fig. 8 D). Remarkably, no Brookesia species was recorded during intensive herpetological surveys of Nosy Hara, nearby islands and the adjacent mainland [40], suggesting that B. micra might be difficult to record. Natural History.— B. micra was found during the day active on the ground in a mosaic of eroded limestone boulders and dry forest leaf litter, and at night roosting on branches in very low vegetation (ca. 5–10 cm above the ground). In contrast to B. tristis and B. confidens its occurrence was not remarkably patchy.Published as part of Frank Glaw, Jörn Köhler, Ted M. Townsend & Miguel Vences, 2012, Rivaling the World's Smallest Reptiles: Discovery of Miniaturized and Microendemic New Species of Leaf Chameleons (Brookesia) from Northern Madagascar, pp. 1-24 in PLoS ONE 7 (2) on pages 16-17, DOI: 10.1371/journal.pone.0031314, http://zenodo.org/record/89404
Brookesia confidens Frank Glaw & Jörn Köhler & Ted M. Townsend & Miguel Vences 2012, sp. n.
No full textBrookesia confidens sp. n. ZooBank LSID: urn:lsid:zoobank.org:act:CB2A9146-0161-42B7-A145-6DED579F1F21 Remark.—This species has been considered before as conspecific with Brookesia sp. ‘‘Montagne des Francais’’ [7]. Holotype.— ZSM 2150/2007 (FGZC 1196), adult male (hemipenes incompletely everted), collected on trail to the ‘‘Petit Tsingy and Grotte des Chauves-Souries’’, 12°57'25"S, 49°0 7'0 6"E, 90 m a.s.l., Ankarana National Park, Antsiranana Province, northern Madagascar, on 1 March 2007 by P. Bora, H. Enting, F. Glaw, A. Knoll, and J. Köhler. Paratypes.— UADBA uncatalogued (FGZC 1194–1195), two specimens; ZSM 2151/2007 (FGZC 1197), probably male, ZSM 2152/2007 (FGZC 1198), adult female, ZSM 2153/2007 (FGZC 1199), adult female, all with same data as holotype; UADBA uncatalogued (FGZC 1608), UADBA uncatalogued (FGZC 1610), ZSM 1511/2008 (FGZC 1607), adult female, ZSM 1512 /2008 (FGZC 1609), adult male, all collected at ‘‘ Petit Tsingy’ ’, Ankarana National Park, Antsiranana Province, northern Madagascar, on 12 February 2008 by N. D’Cruze, M. Franzen, F. Glaw and J. Köhler. Diagnosis.— A member of the Brookesia minima group based on small body size (SVL 9% to all other species of the B. minima group. Description of holotype.— Adult male in good state of preservation (Fig. 7; Supporting Information S 1). Both hemipenes everted. Measurements in Table 1. Head with lateral crest starting at median level at the posterior edge of eye; prominent orbital crests with distinctly enlarged pointed tubercle at posterior level slightly below lateral crest, and a crest at the posterior edge of the head, that form a weakly developed dorsal helmet; a pair of very short straight parasagittal crests that transform posteriorly into a broad area of elevated scales which almost cover the entire posterior head surface; three pointed tubercles on each side of posterior helmet crest, a prominent one at termination point of lateral crest, an even larger one at the imaginary termination point of parasagittal crest, and a weakly developed one between parasagittal and lateral crests; one pointed tubercle on lateral surface of head, below lateral crest in temporal region; orbital crest denticulated; no supraocular cone recognizable; supranasal cone not clearly recognizable; head longer (5.3 mm) than wide; chin and throat without longitudinal rows of slightly enlarged tubercles. Dorsal surface of body without a vertebral ridge or keel; 13 dorsolateral pointed tubercles form a complete longitudinal line on the body; most posterior (13th) pointed dorsolateral tubercle being largest, above insertion point of hindlimb; pointed dorsolateral tubercles almost equally spaced, pointing out almost perpendicularly from body; slightly enlarged, pointed tubercles form curved transversal crests on either side of the vertebral line between 2nd and 10th dorsolateral pointed tubercles; slightly enlarged tubercles on dorsal surface of tail, without any reconizable pattern; no well-defined dorsal pelvic shield in sacral area; few irregularly scattered enlarged rounded ubercles on the lateral surface of body; venter without enlarged tubercles; scattered, soft-pointed tubercles on limbs; no pointed tubercles around cloaca; longitudinal row of slightly enlarged pointed tubercles lateral on tail, forming a longitudinal row from tail base to two-thirds of tail length; no enlarged tubercles on ventral surfaces of tail. After almost four years in ethanol, all surfaces pale grey to beige, with most pointed tubercles being brown; upper surface of head pale brown. Variation.— For morphological measurements and proportions see Table 1 and Supporting Information S1. In ZSM 2153/2007, the ground colour is dark brown with a whitish vertebral stripe that starts at posterior half of body and extends onto the tail which is entirely beige with scattered brownish spots. The posterior dorsal surface of head is grey. The body of ZSM 1512/2008 is light grey whereas the head is brown with a greyish transverse line between the nostrils. The tail base of the females is less thickened than in the males. In life, all individuals with dorsal colouration of head, dorsum and tail light grey or pale beige, lateral parts of the body brown, with indistinct greyish marbling lateroventrally and with few small dark brown spots around the flank tubercles; limbs almost uniformly dark brown. This pattern is likely a stress colouration. Genital morphology.— For this species, everted hemipenes are available only for the holotype (ZSM 2150/2007). On both sides, the organs are single tube-like structures of low diameter which decreases from the apex to the distal part of the truncus. The left hemipenis is not fully everted whereas the one on the right side (length 1.3 mm; width at the pedicel 0.6 mm, width at the apex 0.4 mm) might be fully everted, but this cannot be verified without examination of further material. There are no ornaments on the pedicel and truncus. The sulcus spermaticus is not recognizable on most of the truncus but becomes more distinct in the apical region. The apical region is characterized by a slight increase in diameter but not bilobed, and is covered by relatively large but poorly defined pustular papillae (Fig. 6). The hemipenis is devoid of any further prominent structures such as the spiny apical papillae of B. peyrierasi (which are visible also in incompletely everted organs). Etymology.— The species epithet is an adjective derived from the Latin ‘‘confidens’’ meaning ‘‘confident’’, ‘‘trusting’’. The known range of the species is supposedly a well protected nature reserve with apparently limited habitat destruction. Furthermore, this area might benefit from natural protection by the tsingy limestone formations which are difficult to access, thus giving hope for the species’ survival. Distribution.— Only known from a single locality within Ankarana National Park. Natural History.— Most individuals were found roosting at night on thin branches about 5–20 cm above the leaf litter in deciduous dry forest close to a small forest trail within a small area, surrounded by tsingy outcrops. At this locality, the species was relatively abundant, whereas it was not found at similar localities nearby, suggesting a patchy distribution and a preference for certain microhabitats. This hypothesis is also supported by the fact that earlier herpetological surveys in Ankarana [38,39], did not record any species of the Brookesia minima group. When stressed, individuals can quickly change colour and display a broad pale vertebral stripe contrasting with the darker flanks.Published as part of Frank Glaw, Jörn Köhler, Ted M. Townsend & Miguel Vences, 2012, Rivaling the World's Smallest Reptiles: Discovery of Miniaturized and Microendemic New Species of Leaf Chameleons (Brookesia) from Northern Madagascar, pp. 1-24 in PLoS ONE 7 (2) on pages 15-16, DOI: 10.1371/journal.pone.0031314, http://zenodo.org/record/89404
Brookesia tristis Frank Glaw & Jörn Köhler & Ted M. Townsend & Miguel Vences 2012, sp. n.
No full textBrookesia tristis sp. n. ZooBank LSID: urn:lsid:zoobank.org:act:56F36D4D-1F94-49C4-AC30-6E894B6AA998 Remark.—This species has been considered before as Brookesia sp. ‘‘Montagne des Francais’’ [7] and as Brookesia sp. nov. [37]. Holotype.— ZSM 1704 /2004 (no field number), adult male (hemipenes everted), collected at Montagne des Français, 12°19'S, 49°20'E, ca. 150 m a.s.l., Antsiranana Province, northern Madagascar, on 23 February 2004 by F. Glaw, M. Puente, R. D. Randrianiaina and guides of the hotel ‘‘ King’s Lodge’ ’. Paratypes.— UADBA uncatalogued (FGZC 477–478), ZSM 357/2004 (FGZC 654) [not examined morphologically], and ZSM 876/2010 [no field number, not examined morphologically], adult males, ZSM 354 /2004 (FGZC 651), adult female, same data as holotype; ZSM 1705 /2004–1707/2004 (no field numbers), three adult females, most likely same data as holotype. UADBA uncataloged (FGZC 1187, 1189, 1191); ZSM 2146–2149 /2007 (FGZC 1188, 1190, 1192, 1193), adult females, all collected at Montagne des Français, near remains of French Fort, 12°19'S, 49°20'E, 250–300 m a.s.l., Antsiranana Province, northern Madagascar, on 27 February 2007 by P. Bora, H. Enting, F. Glaw, A. Knoll and J. Köhler; ZSM 2018 /2008 (FGZC 1733 = MgF 061) and UADBA uncatalogued (FGZC 1734 = MgF 061), collected at Montagne des Français, canyon 2 km [air distance] W of Andavakoera, 12°19,838'S, 49°20,941'E, 250 m a.s.l., on 24 January 2006 by E. Randriamalala; UADBA uncatalogued (FGZC 1657), female, ZSM 1505 /2008 (FGZC 1656), adult male (hemipenes everted), both collected ca. 1.5 km southwest of Andavakoera (‘‘ Frontier base camp’’), 12°19'59.2"S, 49°21'20.6"E; 140 m a.s.l., Montagne des Français, Antsiranana Province, northern Madagascar, on 16 February 2008 by M. Franzen, F. Glaw, J. Köhler and Z.T. Nagy. Diagnosis.— A member of the Brookesia minima group based on small body size (SVL 6.9% to all other species of the B. minima group. Description of holotype.— Adult male in good state of preservation (Fig. 7; Supporting Information S 1). Both hemipenes everted. Measurements in Table 1. Head with lateral crest starting at median level at the posterior edge of eye, prominent orbital crests, and a crest at the posterior edge of the head, that form a weakly developed dorsal helmet; a pair of curved parasagittal crests that start above the eyes and begin to converge before terminating at the posterior crest; between the parasagittal crests there is a second pair of parallel longitudinal crests; three pointed tubercles on each side of posterior helmet crest, one at termination point of lateral crest, one at termination point of parasagittal crest, and one between parasagittal and lateral crests; one pointed tubercle on lateral surface of head, below lateral crest in temporal region; orbital crest denticulated; no supraocular cone recognizable; supranasal cone does not project beyond snout tip; head longer (5.1 mm) than wide; chin and throat without longitudinal rows of slightly enlarged tubercles. Dorsal surface of body without a vertebral ridge or keel; 11 dorsolateral pointed tubercles form a complete longitudinal line on the body; 10th dorsolateral tubercle least pointed, small; most posterior (11th) pointed dorsolateral tubercle being largest, above insertion point of hindlimb, very slightly projecting backwards; pointed dorsolateral tubercles almost equally spaced, 3rd to 6th tubercle slightly larger than others, pointing out almost perpendicularly from body; slightly enlarged, rounded tubercles form curved transversal crests on either side of the vertebral line between 1 st and 10th dorsolateral pointed tubercles; dorsal surface of tail with slightly enlarged rounded tubercles forming separated rounded crests that continue from tailbase to two-thirds down the tail; no well-defined dorsal pelvic shield in sacral area; lateral surface of body with evenly spaced enlarged rounded tubercles, mainly arranged in four longitudinal rows; venter without enlarged tubercles; scattered, soft-pointed tubercles on limbs; no pointed tubercles around cloaca; longitudinal row of slightly enlarged pointed tubercles lateral on tail, forming a longitudinal row from tail base to tail tip; no enlarged tubercles on ventral surfaces of tail. In life, colouration of head, body, limbs and tail brown to beige. Faint brown bars radiate from eye to lateral surfaces of head; chin and throat brown; ventral surfaces of body and tail pale brown. After almost seven years in ethanol, all surfaces pale grey to beige, with most pointed tubercles being brown; upper surface of head pale brown. Variation.— For morphological measurements and proportions see Table 1 and Supporting Information S1. In preservative, most paratypes are dorsally darker than the holotype, ranging from brown to dark brown. Their ventral surface is greyish except for the region of the chest, which shows brown mottling. In life several specimens got a light grey vertebral stripe when kept together with conspecifics in a bag, suggesting that this might be a stress colouration. The life colouration of juveniles generally resembles that of adults. The tail base of the females is distinctly less thickened than in the males. In life, two adult specimens had a weight of 0.18 g and 0.206 g, respectively. Genital morphology.— Only two specimens with everted hemipenes were available for examination: ZSM 1505/2008 and ZSM 1704/2004 (holotype). In both specimens, one hemipenis is clearly incompletely everted whereas the second one appears fully everted, although this cannot be stated with complete certainty. In ZSM 1505/2008, hemipenis length is 2.5 mm and maximum hemipenis width is 1.7 mm when fully turgid. The hemipenis is an irregular tubular structure without any ornaments, which in fully turgid state is somewhat balloon-shaped, widest at the central part of the truncus and more narrow at the pedicel and apex. Near the apex, two very small lobe-like structures are visible, corresponding to attachments of the retractor muscle, and a few very small spinelike papillae. However, it is not clear whether the hemipenis could possibly become further everted and then may become more strongly bilobed terminally, with a more prominent exposure of the spine-like papillae. There are no ornaments on pedicel and truncus, and no recognizable ornamental structures on apex (although apex might not be fully everted; Fig. 6). Etymology.— The species epithet is an adjective derived from the Latin ‘‘tristis’’ meaning ‘‘doleful’’, ‘‘sad’’, ‘‘sorrowful’’, and refers to the fact that the entire known range of this species (Montagne des Français) suffers from severe deforestation and habitat destruction [37] despite recently being declared as a nature reserve. Distribution.— Only known from the Montagne des Français limestone massif. Natural History.— Most individuals of B. tristis were found roosting at night on small branches about 5–50 cm above the leaf litter within a limestone massif with deciduous dry forest. By far most specimens were found in a few small areas where the species was abundant, whereas it was rare in other, apparently similar habitats. One of the females laid two large eggs (5.9 6 3.7 mm resp. 5.8 6 3.5 mm diameter) in February. The two juveniles hatched 64 and 69 days after egg deposition at temperatures between 20– 26°C. They measured 14 mm in total length. Eight days after hatching the weight of the older juvenile was 0.03 g.Published as part of Frank Glaw, Jörn Köhler, Ted M. Townsend & Miguel Vences, 2012, Rivaling the World's Smallest Reptiles: Discovery of Miniaturized and Microendemic New Species of Leaf Chameleons (Brookesia) from Northern Madagascar, pp. 1-24 in PLoS ONE 7 (2) on pages 12-15, DOI: 10.1371/journal.pone.0031314, http://zenodo.org/record/89404
Brookesia desperata Frank Glaw & Jörn Köhler & Ted M. Townsend & Miguel Vences 2012, sp. n.
No full textBrookesia desperata sp. n. ZooBank LSID: urn:lsid:zoobank.org:act:C62B456C-CDA8- 4DDE-AE0A-66DF43C57F7E Remark.—This species has been considered before as Brookesia sp. aff. karchei ‘‘Ambre’’ [7] and as Brookesia sp. nov. [41]. Holotype.— ZSM 2170/2007 (FGZC 1250), adult male (hemipenes everted), collected at Forêt d’Ambre Special Reserve, ca. 5 km southwest of Sakaramy village, 12°28'0 0"S, 49°13'37" E, 470 m a.s.l., Antsiranana Province, northern Madagascar, on 12 March 2007 by F. Glaw, J. Köhler and A. Razafimanantsoa. Paratypes.— ZSM 2171/2007 (FGZC 1251), ZSM 2175/2007 (FGZC 1258), ZSM 2176/2007 (FGZC 1260), ZSM 2177/2007 (FGZC 1263), ZSM 2179/2007 (FGZC 1269), adult males (all with everted hemipenes), ZSM 2172–2174 /2007 (FGZC 1252– 1254), ZSM 2178/2007 (FGZC 1265), adult females, all with same data as holotype; ZSM 1506/2008 (FGZC 1880, female), and UADBA uncatalogued: FGZC 1700, FGZC 1879 (male), FGZC 3111–3112, collected at Forêt d’Ambre Special Reserve, ca. 5 km southwest of Sakaramy village, 12°28'S, 49°13'E, 550 m a.s.l., Antsiranana Province, northern Madagascar, on 27 February 2008 by N. D’Cruze, F. Glaw and J. Köhler; ZSM 791 /2009 (ZCMV 13040), adult male (not examined morphologically), collected at Forêt d’Ambre Special Reserve, 12°28'18.0"S, 49°13'56"E, 438 m a.s.l., Antsiranana Province, northern Madagascar, on 16 November 2009 by A. Crottini, S. Hauswaldt, A. Lima, F. M. Ratsoavina and E. Rajeriarison. Diagnosis.— A member of the Brookesia minima group based on small body size (SVL 25–30 mm) and molecular phylogenetic relationships. Brookesia desperata is distinguished from all other species in the group by the presence of three enlarged tubercles on lateral head surface (versus 0–2). In addition it differs as follows: from B. confidens by a larger adult body size (male SVL 25.0–26.7 vs. 18.3–20.1 mm), supranasal cone present (vs. absent), and hemipenis with two apical processes each with a distinct spine (vs. narrow hemipenis without apical ornaments); from B. dentata by presence of well-developed lateral spines on the tail (vs. absence); from B. exarmata by a larger adult body size (female SVL 27.3– 30.0 mm vs. 25.7–26.5 mm), and presence of well-developed lateral spines on the tail (vs. absence); from B. micra by a larger adult body size (male SVL 25.0–26.7 vs. 15.3–15.8 mm), presence of well-developed lateral spines on the tail (vs. absence), and hemipenis with two apical processes each with a distinct spine (vs. comb-like arranged papillae on apex); from B. minima by a larger adult body size (male SVL 25.0–26.7 vs. 15.0– 20.6 mm), presence of well-developed lateral spines on the tail (vs. indistinct), pelvic spine present (vs. absent or indistinct), and hemipenis with two apical processes each with a distinct spine (vs. balloon-like hemipenis without ornaments); from B. peyrierasi by a larger adult body size (male SVL 25.0–26.7 vs. 19.7–22.4 mm), presence of well-developed lateral spines on the tail (vs. indistinct), and hemipenis with two apical processes each with a distinct spine (vs. four spines on each lobe); from B. ramanantsoai by presence of welldeveloped lateral spines on the tail (vs. absence), and hemipenis with two apical processes each with a distinct spine (vs. balloon-like hemipenis without ornaments); from B. tristis by a larger adult body size (male SVL 25.0–26.7 vs. 18.0– 18.2 mm), and hemipenis with two apical processes each with a distinct spine (vs. small spinelike papillae on apex); and from B. tuberculata by a larger adult body size (male SVL 25.0–26.7 vs. 14.4–18.8 mm), presence of welldeveloped lateral spines on the tail (vs. absence), and hemipenis with two apical processes with a distinct spine (vs. a single crownlike structure on apex). B. desperata is most similar to B. karchei in body size, number of dorsolateral pointed tubercles (12–13) and distinct lateral tubercles on tail (Fig. 9 B). However, B. karchei differs from the new species by more pronounced supraocular and supranasal cones, more prominent and spiny posterior crest and only one enlarged tubercle at lateral side of head (three in B. desperata). Referencing a fragment of the 16S rRNA gene, B. desperata shows an uncorrected pairwise divergence of 6.4% to its sister species B. tristis, and divergences>6.6% to all other species of the B. minima group. Description of holotype.— Adult male in good state of preservation (Fig. 7; Supporting Information S 1). Both hemipenes everted. Measurements in Table 2. Head with low lateral crest starting at median level at the posterior edge of eye; prominent orbital crests with distinctly developed supraocular cone directed anteriorly, and a crest at the posterior edge of the head, that form a weakly developed dorsal helmet; a pair of slightly curved parasagittal crests that start above the eyes and begin to converge slightly before terminating at the posterior crest; between the parasagittal crests there is a second pair of short parallel longitudinal crests; three pointed tubercles on each side of posterior helmet crest, one at termination point of lateral crest, one at termination point of parasagittal crest, and one between parasagittal and lateral crests; three pointed tubercles on lateral surface of head, one below lateral crest in temporal region, one at posterior edge of orbital crest below lateral crest, and one between lower edge of eye slightly anterior to angle of jaws; orbital crest with few enlarged conical tubercles; no supraocular cone recognizable; supranasal cone does not project beyond snout tip; head longer (6.4 mm) than wide; chin and throat without longitudinal rows of slightly enlarged tubercles. Dorsal surface of body without a vertebral ridge or keel; 13 dorsolateral pointed tubercles form a complete longitudinal line on the body; most posterior (13th) pointed dorsolateral tubercle being distinctly largest, slightly posterior to insertion point of hindlimb, projecting backwards; pointed dorsolateral tubercles almost equally spaced, 8th to 12th slightly smaller than others, pointing out almost perpendicularly from body; slightly enlarged, rounded tubercles form weakly elevated curved transversal crests on either side of the vertebral line between 1 st and 12th dorsolateral pointed tubercles; slightly enlarged tubercles on dorsal surface of tail, forming two weakly recognizable longitudinal parallel lines; barely defined dorsal pelvic shield in sacral area; lateral surface of body with densely scattered enlarged conical tubercles; venter without enlarged tubercles; scattered, conical tubercles on limbs; no pointed tubercles around cloaca; longitudinal row of distinctly enlarged tubercles lateroventrally on tail, forming a longitudinal row from tail base to three fourth of tail length; no enlarged tubercles on ventral surfaces of tail. After almost four years in ethanol, flanks, dorsal surfaces of limbs and tail uniformly dark grey-brown; neck with distinct light grey spot; vertebral region mottled with brown and grey; throat grey, venter and ventral surfaces of tail grey-brown. Hemipenes whitish. Variation.— For morphological measurements and proportions see Table 2 and Supporting Information S1. In life, dorsal colouration of head, dorsum and tail light grey. Lateral parts of the body beige, brown or dark brown, with few dark brown spots; limbs almost uniformly dark brown. This pattern is likely to refer to a stress colouration (Fig. 9). When unstressed, most parts of the body beige, with a slightly lighter area on the head anterior to the eyes and in the vertebral region of the dorsum. In preservative, ZSM 2172/2007 is greyish with brown blotches and flecks on dorsum and flanks whereas the other paratypes are more or less uniformly dark brown. The tail base of the males is only slightly more thickened than in the females. Genital morphology.— Everted hemipenes were available from the holotype (ZSM 2170/2007) and from four paratypes (ZSM 2171/ 2007, 2176–2177 /2007, 2179/2007). Structures were characteristic and concordant among all five specimens and are described based on the holotype. Hemipenis length is 3.2 mm. Hemipenis width in the fully turgid state is about 2.3 mm. The retractor muscle is visible through the transparent hemipenis integument and bifurcates after about 1/2 of the hemipenis length. From the wide apex, two narrow tubular processes (0.5 mm in width) extend about 1.1 mm distally, with each end of the retractor muscle ending at the tip of these processes. The end of each of the tubes bears a distinct spine. The apex processes with their large spines are easily recognizable also in incompletely everted hemipenes (Fig. 6). When the hemipenis is everted but not turgid (in preserved specimens), the processes are very distinct and project much more distinctly from the apex. Etymology.— The species epithet is an adjective derived from the Latin ‘‘desperatus’’ meaning ‘‘desperate’’. Although the known range of the species is within a nature reserve established decades ago, its habitat is in truth barely protected and subject to numerous human-induced environmental problems resulting in severe habitat destruction [41], thus threatening the survival of the species. Distribution.— Only known from the southern edge of Forêt d’Ambre Special Reserve, northern Madagascar. Natural History.— All individuals were found roosting at night on small branches or leaves about 5–100 cm above the ground in disturbed rainforest. Individuals occurred in apparently high abundance and were also found at the border of forest clearings on banana plants. One female laid two large eggs when kept in a plastic bag. When stressed, individuals can quickly change colour and display a broad pale vertebral stripe contrasting with the darker flanks (Fig. 9 A).Published as part of Frank Glaw, Jörn Köhler, Ted M. Townsend & Miguel Vences, 2012, Rivaling the World's Smallest Reptiles: Discovery of Miniaturized and Microendemic New Species of Leaf Chameleons (Brookesia) from Northern Madagascar, pp. 1-24 in PLoS ONE 7 (2) on pages 17-18, DOI: 10.1371/journal.pone.0031314, http://zenodo.org/record/89404
When can chromosomes drive speciation? The peculiar case of the Malagasy tomato frogs (genus Dyscophus)
No full textExcept for polyploidy, parthenogenesis and hybridogenesis, the role of chromosome rearrangements in
speciation is still highly debated. This paper highlights an unusual karyotype diversification and a putative
case of chromosomal speciation in the endemic Malagasy tomato frogs.
The tomato frogs include Dyscophus insularis as well as the sister species D. antongilii and D. guineti,
which are characterized by a disjunct distribution and a low genetic divergence, probably due to a recent
hybridization. Our results show that the three Dyscophus species present 2n = 26 biarmed chromosomes,
differing in chromatin pattern. Interestingly, both sexes of D. guineti show a heteromorphic chromosome
pair 11, with one element larger (11a) than the other (11b) due to the presence of a large terminal tract
on the short arm with an euchromatic band enclosed between two heterochromatic bands. This fixed
condition reminds that of some newts of the genus Triturus, but in Dyscophus it probably characterizes the
beginning of a cladogenetic process. We hypothesize that in D. guineti the raising of the heteromorphic
11th chromosome pair followed the hybridization with D. antongilii, originating a post-zygotic barrier
and representing a case of sympatric or parapatric chromosomal speciation. This speciation mechanism
may also have had important biogeographic consequences, shaping the current disjunct distribution of
D. antongilii and D. guineti, so far not sufficiently explaine
Calumma juliae David Prötzel & Miguel Vences & Oliver Hawlitschek & Mark D. Scherz & Fanomezana M. Ratsoavina & Frank Glaw 2018, SP. NOV.
No full textCALUMMA JULIAE SP.NOV. urn:lsid:zoobank.org:act:6 A 92 C 95 D- 6383-45DC-9 FFC- 0943 B 08 F 064 F Remark: Despite intensive research at the type locality in January (by L. Randriamanana, F. G. and D. P.), August (by N. Raharinoro, K. Glaw, T. Glaw, J. Forster, F. G. and D. P.) and November 2016 (by A. Rakotoarison and M.D. S.) in the rainy and dry seasons, only female specimens of this new species were found. In total, eight adult females, two subadult females and two juveniles were encountered (but not all of them collected). Holotype: ZSM 143/ 2016 (FGZC 5235), adult female, collected in small forest fragment 5 km east of Moramanga, just south of the Route Nationale 2 (18.9520° S, 48.2707° E, at 950 m a.s.l.), Toamasina Province, eastern Madagascar on 6 January 2016 by F. Glaw, D. Prötzel and L. Randriamanana. Paratypes: ZSM 142 / 2016 (FGZC 5233), FGZC 5232 and FGZC 5234 (two uncatalogued specimens in UADBA), all three adult females, collected from the same location as the holotype (18.9519° S, 48.2705° E, within a radius of 50 m, at 950 m a.s.l.) on 6 January 2016 by F. Glaw, D. Prötzel, and L. Randriamanana. ZSM 254 / 2016 (FGZC 5274), adult female, and ZSM 255/ 2016 (FGZC 5275) and FGZC 5276 (uncatalogued specimen in UADBA), both juveniles, all three collected on 30 July 2016; FGZC 5277 (uncatalogued specimen in UADBA), subadult, collected on 31 July 2016 at the same location as above by F. Glaw, D. Prötzel, and N. Raharinoro. Diagnosis: Male specimens are unknown so far, hence the diagnosis refers only to females of this species. Calumma juliae sp. nov. is a member of the phenetic C. nasutum species group (Prötzel, Ruthensteiner & Glaw, 2016), on the basis of the presence of a soft, dermal unpaired rostral appendage, absence of gular or ventral crests, and heterogeneous scalation on the lower arm, consisting mostly of tubercles of a diameter of 0.7–0.8 mm. Within the group, it is a large (TL 105.3–111.6 mm), grey–beige chameleon that is characterised by a long and distally rounded rostral appendage, a dorsal crest of 11–14 tubercles, occipital lobes that are clearly notched but not completely separated, and absence of axillary pits. Calumma juliae sp. nov. differs from C. fallax, C. gallus, C. nasutum, C. peyrierasi, C. vatosoa, and C. vohibola of the C. nasutum group by the presence of occipital lobes; from C. gehringi, C. guibei, and C. lefona sp. nov. in the completely closed brain case (vs. frontoparietal fenestra); additionally, from female C. gehringi and C. guibei in body size of 53.3–59.4 mm SVL (vs. 47.5–52.3 mm); from female C. gehringi in the shorter rostral appendage of 2.3–2.7 mm (vs. 3.2–4.4 mm); from C. guibei (both sexes) in the notch between the occipital lobes of 0.2–0.8 mm (vs. completely separated with notch of 1.2–1.9 mm; see Brygoo, 1971); from C. lefona sp. nov. (one male) in the shorter (2.3–2.7 mm) and rounded rostral appendage (vs. 5.6 mm, pointed), the absence of a temporal and parietal crest (vs. presence) and the number of dorsal cones of 11–14 (vs. 23); from C. boettgeri (both sexes) by the higher number of large (0.7–0.8 mm diameter) juxtaposed tubercle scales on the extremities (17–19 in line vs. 7–14, diameter of 0.2–0.5 mm and isolated from each other); from female C. uetzi sp. nov. in the larger body size of 53.3–59.4 mm SVL (vs. 42.0 mm SVL in females), the absence of a temporal and parietal crest (vs. presence of both) and the higher number of infralabial scales of 14–15 (vs. 11–12); from its most similar taxon C. linotum by the clearly notched occipital lobes with a depth of 0.2–0.8 mm (vs. not or slightly notched with depth 0–0.2 mm), presence of a dorsal crest in females consisting of 11–14 conical scales in C. juliae sp. nov. (vs. zero in C. linotum and six in C. cf. linotum; one specimen from Andampy), higher number of infralabial scales in females of 14–15 (vs. 12–13), absence of temporal and parietal crest in females (vs. both present), and in generally larger body size in females of 53.3–59.4 mm SVL (vs. 42.7–54.5 mm). In skull morphology, the squamosal and parietal do not meet in female C. juliae sp. nov. (vs. broad in contact in male and female C. linotum (Prötzel et al., 2015); frontal of triangular shape and narrower, e.g. 16.8–18.2% of SkL at border to prefrontal (vs. 22.1–24.4%), 30.7– 31.1% of SkL at border to postorbitofrontal (vs. 34.4– 35.8%) and 25.0–26.3% of SkL bordering the parietal (vs. 32.5–33.6%); also, parietal is narrower at border to frontal with 28.5–29.5% of SkL (vs. 34.1–35.1%). Description of the holotype (Fig. 3 C): Adult female, in a good state of preservation; SVL 55.8 mm, tail length 55.4 mm; for other measurements, see Table 1; distinct rostral crest, laterally compressed dermal rostral appendage that projects forward of the snout tip over a length of 2.7 mm with a diameter of 2.2 mm, rounded distally; 14 infralabial and 13 supralabial scales; supralabials dorsally serrated; no supra-orbital crest; distinct lateral crest running horizontally; no temporal crest; occipital lobes clearly developed and separated but still slightly connected by a notch of 0.5 mm; casque raised; dorsal crest present, starting 4.3 mm from the base of the notch between the occipital lobes, consisting of a row of 12 separated, small conical scales spaced at regular intervals of ~ 2 mm, continuing on the tail with smaller and narrower spaced cones; no traces of gular or ventral crest. Body laterally compressed, with fine homogeneous scalation with the exception of the extremities and head region; limbs with rounded tubercle scales of maximal diameter 0.7 mm; heterogeneous scalation on the head, with largest scale on temporal region having a diameter of 1.3 mm; no axillary or inguinal pits. Skull osteology of the holotype (Fig. 4 C, Supporting information, Video S4): Skull length 13.2 mm; snout– casque length 16.1 mm; broad paired nasals meeting each other; anterior tip of frontal exceeding less than half of the naris and separated from premaxilla; prefrontal fontanelle and naris fused; frontal smooth, and parietal with only a few tubercles; frontal with width of 2.4 mm (18.2% of SkL) at border to prefrontal extending to 4.1 mm (31.1% of SkL) at border to postorbitofrontal; broad parietal tapering strongly from a width of 3.9 mm (29.5% of SkL) at the border to frontal to 0.8 mm (6.1% of SkL) at midpoint, then extending mostly straight beyond the posterodorsal extension of the squamosals, and finally tapering again at its tip; posterodorsally directed parietal does not meet the squamosal; squamosal thin without tubercles. For further measurements, see Table 2. Coloration of the holotype in preservative (Fig. 3C): The body of the holotype in preservative is of dark grey colour without any distinct pattern; inner side of extremities and tail beige; extremities and temporal and postocular region of the head speckled with greyish–blue tubercle scales. Variation: All adult female C. juliae sp. nov. that have been found so far show a consistent morphology and consequently agree well with the holotype. ZSM 254/2016 and FGZC 5234 have a slightly shorter rostral appendage than the holotype, at 2.4 and 2.3 mm (vs. 2.7 mm); the notch between the occipital lobes is deeper in FGZC 5232 at 0.8 mm and less deep in ZSM 142/ 2016 at 0.2 mm (vs. 0.5 mm); the dorsal crest of ZSM 255/ 2016 consists of only nine tubercles (vs. 12); this might be referred to its juvenile developmental stage. In osteology, no significant variations were found. Coloration in life (Fig. 7): Females are indistinctly grey–beige coloured; a netlike dark brown pattern on the skin between the scales and two dorsoventrally compressed blue blotches may occur on the body; tubercle scales on extremities and body may be bright green; rostral appendage not highlighted and of same colour as the body; a dark lateral stripe may stretch from the rostral appendage across the eyes to the occipital lobes. If stressed, only the head coloration changes to dark brown or green pattern at the dorsal head region, eyelids with radially aligned blue/violet spots, and rostral crest and appendage of turquoise/ green colour. Available names: Apart from C. boettgeri and C. linotum, there is no other valid species or synonym in the C. nasutum group with slightly notched occipital lobes. Etymology: D. P. dedicates the first new species he discovered himself to Julia Forster, in recognition of her generous support and understanding of our research on Madagascan chameleons and her help in collecting specimens of this species. Distribution: Calumma juliae sp. nov. has so far been recorded only from a fragment of degraded primary rainforest, covering an area of just 15 ha, east of Moramanga (Toamasina Province, eastern Madagascar; Figs 6, 8 A); for geographical coordinates, see the descriptions in the subsections ‘Holotype’ and ‘Paratypes’ above. The forest fragment spreads over a hill that rises to a maximum elevation of 1010 m a.s.l.; specimens were found only along the trail at 950 m a.s.l., but we expect them to occur on the hill as well. A further specimen was observed just north of the Route Nationale 2 (18.9513° S, 48.2721° E, 950 m a.s.l.) in secondary bushes. Natural history and ecology: Calumma juliae sp. nov. is an arboreal, diurnal species found in bushes and trees in a small forest fragment of degraded primary rainforest. Roosting sites at night were thin branches or, rarely, leaves that were not exposed, but hidden inside tree/bush cover 0.3–2 m above the ground. In contrast to the syntopically occurring population of C. cf. nasutum, the present species preferred a horizontal sleeping position (vs. head pointing downwards). When disturbed, some specimens dropped immediately and stayed curled and motionless on the ground. In appropriate habitat, specimens occurred a few metres from one another. In January 2016, only adult females were collected; in August 2016, additionally subadult and juvenile specimens; in November 2016, only one adult female was found. None of the collected females showed signs of being gravid. The size of the juveniles suggests that they hatched during the rainy season from approximately January to March. Under the occipital lobes of two specimens (ZSM 143/ 2016 and 142/2016) we found three and two mites, respectively; the lobes might function as mite pockets comparable to axillary pits in other chameleon species to limit and locate the damage caused by these ectoparasites (Arnold, 1986). Recommended IUCN status: As no attempts have yet been made to estimate the population size or status of C. juliae sp. nov. directly, we suggest that it should be assessed under the IUCN Red List criterion B (IUCN, 2012). The distribution area of the single known forest in which the species occurs has an area of ~ 0.15 km 2, which we interpret as the area of occupancy (AOO) of the species as defined by the IUCN (2012), but which at present is also equivalent to the EOO of the species (criterion B 1). This constitutes a single threat-defined location (criterion B, subcriterion a). The forest in which it occurs is under heavy active anthropogenic pressure and is, in our opinion, in imminent danger of disappearance [criterion B, subcriterion b(iii)]. As the AOO of the species is considerably 1500 m a.s.l. have one. Additionally, FF also occur in C. peyrierasi and C. tsaratananense (Prötzel et al., 2018; A. van’t Padje et al., unpublished observations); both are montane species that occur> 1900 m a.s.l. (Brygoo, 1971, 1978), the latter of which is certainly not closely related to the C. nasutum group (Tolley et al., 2013). All other investigated Calumma species have a closed skull roof (Prötzel et al., 2018; van’t Padje et al., unpubl.).Published as part of David Prötzel, Miguel Vences, Oliver Hawlitschek, Mark D. Scherz, Fanomezana M. Ratsoavina & Frank Glaw, 2018, Endangered beauties: micro-CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae), pp. 1-28 in Zoological Journal of the Linnean Society 2018 (20) on pages 15-20, DOI: 10.1093/zoolinnean/zlx112, http://zenodo.org/record/122648
Going Beyond Counting First Authors in Author Co-citation Analysis
Get PDFThe present study examines one of the fundamental aspects of author co-citation analysis (ACA) - the way co-citation
counts are defined. Co-citation counting provides the data on which all subsequent statistical analyses and mappings
are based, and we compare ACA results based on two different types of co-citation counting - the traditional type that
only counts the first one among a cited work's authors on the one hand and a non-traditional type that takes into
account the first 5 authors of a cited work on the other hand. Results indicate that the picture produced through this non-traditional author co-citation counting contains more coherent author groups and is therefore considerably clearer. However, this picture represents fewer specialties in the research field being studied than that produced through the traditional first-author co-citation counting when the same number of top-ranked authors is selected and analyzed. Reasons for these effects are discussed
Figure 10 in Endangered beauties: micro-CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae)
No full textFigure 10. DiceCT scan integrated into micro-CT scan of the skull of Calumma gehringi (ZSM 2840/2010), showing the position of the brain below the frontoparietal fenestra. A, lateral view on midsagittal section of three-dimensional model. B, dorsal view on three-dimensional model. C, lateral view on midsagittal section (diceCT only). D, correlation of elevation and presence/width of the frontoparietal fenestra in the Calumma nasutum group, based on data in Table 4. Scale bars: 2.0 mm.Published as part of David Prötzel, Miguel Vences, Oliver Hawlitschek, Mark D. Scherz, Fanomezana M. Ratsoavina & Frank Glaw, 2018, Endangered beauties: micro-CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae), pp. 1-28 in Zoological Journal of the Linnean Society 2018 (20) on page 21, DOI: 10.1093/zoolinnean/zlx112, http://zenodo.org/record/122648
Variations on the Author
Get PDF“Variations on the Author” discusses two of Eduardo Coutinho’s recent films (Um Dia na Vida, from 2010, and Últimas Conversas, posthumously released in 2015) and their contribution to the general question of documentary authorship. The director’s filmography is characterized by a consistent yet self-effacing form of authorial self-inscription: Coutinho often features as an interviewer that rather than express opinions propels discourses; an interviewer that is good at listening. This mode of self-inscription characterizes him as an author who is not expressive but who is nonetheless markedly present on the screen. In Um Dia na Vida, however, Coutinho is completely absent form the image, while Últimas Conversas, on the contrary, includes a confessional prologue that moves the director from the margins to the center of his films. This article examines the ways in which these works stand out in the filmography of a director who offers new insights into the notion of cinematic authorship
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