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Miralles, Aurelien, Glaw, Frank, Ratsoavina, Fanomezana M., Vences, Miguel (2015): A likely microendemic new species of terrestrial iguana, genus Chalarodon, from Madagascar. Zootaxa 3946 (2): 201-220, DOI: http://dx.doi.org/10.11646/zootaxa.3946.2.3

Identity of Chalarodon madagascariensis Peters, 1854 and designation of a lectotype The original description of Chalarodon madagascariensis by Peters (1854: 616) is extremely short, but in a later publication the same author provided further details and a figure (Peters 1882: 32) and stated that the original collection consisted of ten specimens from the Bay of St. Augustin. All of these were of small size although the species was expected to grow significantly larger. Of the ten syntypes, a minimum of six have been traced by Bauer et al. (1995: 58): ��� ZMB 4360 (2 specimens), 5617, 9214 (2 specimens), MNHN 270 (fide Brygoo [1989]); BMNH ?? (fide Boulenger [1885]).��� In the meantime, one of the two specimens with the number ZMB 4360 was renumbered to ZMB 69147, one of the two specimens with the number ZMB 9214 was renumbered to ZMB 69144, and the formerly overlooked skeleton ZMB 13569 was identified as an additional syntype, whereas ZMB 5617 is currently missing from the collection (F. Tillack, pers. comm. 27. Jan. 2015). As indicated by Bauer et al. (1995) one syntype has been exchanged with the Mus��um nationale d'Histoire naturelle at Paris and now bears the number MNHN 270 (see Brygoo 1989). According to the VertNet database (accessed on 27 Jan. 2015), MNHN- RA- 0.270 was collected by Peters, but the locality of this specimen is given as ���Sans localit�� pr��cise��� and its status as syntype is not indicated. It is apparently the only Chalarodon specimen present in the MNHN collected by Peters and therefore should be considered as syntype. One of the syntypes was exchanged with the collection of the Natural History Museum in London (Boulenger 1885: 128). Altogether, the list of syntypes according to current knowledge includes eight specimens: (1) ZMB 4360; (2) ZMB 5617 (missing from collection); (3) ZMB 9214; (4) ZMB 13569 (skeleton); (5) ZMB 69144 (ex 9214); (6) ZMB 69147 (ex ZMB 4360); (7) MNHN 0.270; (8) one syntype in BMNH (number unknown). Four of the syntypes (ZMB 4360, ZMB 9214, ZMB 69147, ZMB 69144) were available for our study, and as was expected from the work of Peters (1882) are all juveniles. The following account refers exclusively to these four ZMB syntypes examined by us. All are in relatively poor state of preservation, thus only basic data (not included in table 3) are given. The four specimens are characterized by distinctly keeled ventral and gular scales and in all of them six scales are in contact with the mental, thereby confirming the differences to the new species described below. The four specimens are characterized as follows: (1) ZMB 4360: SVL 42.9 mm, tail 68.3 mm (almost complete), two strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca, therefore probably a juvenile male. (2) ZMB 9214: SVL 36.0 mm, tail 18.2 (mutilated), no strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca, therefore probably a juvenile female. (3) ZMB 69144: SVL 37.0 mm, tail 34.8 (tail tip mutilated and remaining tail broken into two pieces), no strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca, therefore probably a juvenile female. (4) ZMB 69147: SVL 39.0 mm, tail ca. 68.7 mm (tail complete but almost broken at one point), belly ventrally opened, two strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca, therefore probably a juvenile male. Although the four syntypes examined are morphologically uniform, and the entire series was reported to originate from the same locality, we cannot exclude that any of the six syntypes not examined by us might turn out to belong to the new species below, or to yet another, still undiscovered taxon. Hence, considering the existence of at least two Chalarodon species and the unknown fate of several syntypes it appears important for taxonomic stability to define a single name-bearing type for C. madagascariensis. Following this rationale, we hereby designate ZMB 4360 as lectotype (Fig. 1). This is the largest of the four studied syntypes, has a nearly complete tail, a darkened throat, and is in the relatively best state of preservation. It therefore is taxonomically the most informative specimen. See Figure 1 for a photograph of the lectotype., Published as part of Miralles, Aurelien, Glaw, Frank, Ratsoavina, Fanomezana M. & Vences, Miguel, 2015, A likely microendemic new species of terrestrial iguana, genus Chalarodon, from Madagascar, pp. 201-220 in Zootaxa 3946 (2) on pages 203-204, DOI: 10.11646/zootaxa.3946.2.3, http://zenodo.org/record/232560, {"references":["Peters, W. (1854) Diagnosen neuer Batrachier, welche zusammen mit der fruher (24. Juli und 17. August) gegebenen Ubersicht der Schlangen und Eidechsen mitgetheilt werden. Vol. 1854. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglich preussischen Akademie der Wissenschaften zu Berlin, 15 pp. [pp. 614 - 628]","Peters, W. (1882) Naturwissenschaftliche Reise nach Mossambique. Zoologie. III. Amphibien. G. Reimer, Berlin, 191 pp.","Bauer, A. M., Gunther R. & Klipfel, R. (1995) The herpetological contributions of Wilhelm Peters (1815 - 1883). SSAR, Ithaca, New York, 714 pp.","Brygoo, E. R. (1989) Les types d'Iguanides (Reptiles, Saunens) du Museum national d'Histoire naturelle. Catalogue critique. Bulletin Museum National d'Histone Naturelle (Paris), Series 4 (11), Sec. A (3), Supplement, 1 - 112.","Boulenger, G. A. (1885) Catalogue of the lizards in the British Museum (Natural History). Second Edition. Vol. II. Iguanidae, Xenosauridae, Zonuridae, Anguidae, Anniellidae, Helodermatidae, Varanidae, Xantusiidae, Teiidae, Amphisbaenidae. Printed by order of the Trustees of the British Museum (Natural History), London, xiii + 497 pp. + 24 plates."]}

Chalarodon steinkampi sp. nov. (Figs. 3, 5, 6) Holotype. ZSM 835 / 2010 (field number ZCMV 12909), adult male, collected at a locality 30 km north of Amboasary Sud along the road to Esomony (- 24.7721, 46.4207, 160 m above sea level), on 17 December 2010, by A. Miralles and F. M. Ratsoavina. Paratypes. ZSM 836 / 2010 (ZCMV 12910), adult male, and ZSM 834 / 2010 (ZCMV 12908), female (possibly subadult), same locality and collection data as holotype; ZSM 139 / 2005 (FGZC 2330), adult male, from a site north of Tranomaro along the road to Esomony (- 24.51792, 46.59895, 420 m above sea level), collected by F. Glaw, M. Vences and P. Bora on 25 January 2005; ZSM 140 / 2005 (FGZC 2550), adult female, and two specimens not examined morphologically and deposited in UADBA collection (FGZC 2549 and 2363), all from the surroundings of Esomony village near crossing of Manambolo river (- 24.51762, 46.60405, 420 m above sea level), collected by F. Glaw, M. Vences and P. Bora on 1 February 2005. Note. Although the two specimens in the UADBA collection (FGZC 2549 and 2363) have not yet been catalogued and were not available for morphological examination, we still decided to assign them to the species (and thereby define them as part of the type series, as paratypes, in agreement with article 72.4. 1 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999)) because their identity was unambiguously determined through DNA sequences, and because their paratype status is in line with general taxonomic practice of distributing types among several collections. Diagnosis. The new species is a member of the genus Chalarodon, easily distinguished from Oplurus, the only other genus of iguanids present in Madagascar, by (i) a rather small size, (ii) absence of enlarged or spiny scales encircling the tail and (iii) the presence of a longitudinal dorsal crest (especially marked in sexually mature males) running from the occipital region to the mid-length of the tail. The new species is morphologically differentiated from Chalarodon madagascariensis, the only nominal species in the genus, by (1) smooth (unkeeled) gular and ventral scales (vs. strongly keeled with 1���3 keels per scale), (2) mental scale in contact with four scales (vs. with 5���8, usually 6 scales), (3) a spotted pattern extending from flanks onto belly (vs. this pattern restricted to flanks), (4) an unpigmented throat (vs. mostly pigmented throat), (5) a distinct, dark dorsal pattern (vs. absent or poorly delimited), (6) on average relatively shorter limbs (FOL/SVL ratio 0.40���0.43 vs. 0.40���0.49; HIL/SVL ratio 0.68���0.79 vs. 0.74���0.87), (7) on average a lower number of spines in dorsal crest (81���97 vs. 90���109), and (8) on average a lower number of scale rows around midbody (144���170 vs. 165���204). Furthermore, C. steinkampi is distinguished from C. madagascariensis by a relevant differentiation in mtDNA and nDNA, without indications for admixture despite occurrence in close geographical proximity, without obvious geographical barriers between populations. Description of the holotype. Adult male in good condition with apparently complete original tail and largely everted hemipenes. See Table 2 for morphometric measurements and numerical coloration characters, Table 3 for meristic characters, and Fig. 3 for coloration. Snout-vent length (56.3 mm) distinctly shorter than tail length (87.4 mm). Head slightly wider than neck. Snout pointed in dorsal view, rounded in lateral view. Canthus rostralis distinct. Ear opening (auditory meatus) vertical (height 3.1 mm; width 1.2 mm). Relative finger length: 1 ��� 5 Variation. See Table 2 for morphometric measurements and numerical chromatic characters. See Table 3 for meristic characters of the paratypes and Fig. 3 for ventral coloration. The general morphology and coloration of the paratypes are similar to the holotype, especially the scalation of the throat and the almost complete absence of keeled scales on throat and belly. Etymology. The specific name is dedicated to Martin Steinkamp, in recognition of his support of biodiversity research and nature conservation through the BIOPAT initiative. Distribution and natural history. The species is only known from a small area adjacent to the western slopes of the Andohahela Massif. Three sites are known, all along the road Amboasary-Tranomaro-Esomony, including the surroundings of Esomony village. All known specimens were active during the day on sandy soils among a vegetation of xerophytic bushes. TABLE 1. List of samples and specimens included in the present study, with their respective localities, voucher field numbers, institutional catalogue number (when available) and GenBank accession numbers (16 S and c-mos). Accessions of newly determined sequences are in bold. Where two voucher numbers are given, the second one (in parentheses) refers to the field number. For collection acronyms, see Materials and Methods. Additional abbreviations: HT, holotype and PT, paratype of C. steinkampi. Specimens analyzed for morphology are marked "Yes" in column "Morphology". NA, not applicable. An asterisk marks c-mos sequence of one C. steinkampi (ZSM 140 / 2005) which in the work of M��nchenberg et al. (2008) was determined as heterozygote (hybrid), formerly deposited in GenBank under the accession number EU099661, but was retrieved as homozygote in study. Sample lab Voucher number Locality GPS coordinates Morphology Accession 16 S Accession c-mos number madagascariensis Antsalova (TM 64) UADBA / FGZC 954 Antsalova - 18.6667, 44.6167 - EU099705 EU099654 Betioky 1 (TM 75) UADBA / FGZC 335 Betioky - 23.7167, 44.3833 - EU099713 EU099662 Betioky 2 (TM 76) UADBA / FGZC 336 Betioky - 23.7167, 44.3833 - EU099714 EU099663 (TM 85) UADBA 21059 (FGMV 2002.1465) Analalava (Isalo) - 21.6917, 44.7833 - EU099706 EU099655 Toliara 1 (TM 86) ZSM 946 / 2003 (FGMV 2002.1551) Toliara - 23.35, 43.67 Yes EU099707 EU099656 Toliara 2 (TM 87) UADBA (FGMV 2002.2017) Toliara - 23.35, 43.67 - EU099708 EU099657 Toliara 3 (TM 88) UADBA 21038 (FGMV 2002.2018) Toliara - 23.35, 43.67 - EU099709 EU099658 1 UADBA (ZCMV 12797) Ifaty site 1 -23.1266, 43.6340 - KJ 170247 KJ 170273 2 UADBA (ZCMV 12798) Ifaty site 1 -23.1266, 43.6340 - KJ 170248 KJ 170274 6 UADBA (MIR 160) Ifaty site 1 -23.1266, 43.6340 - KJ 170252 KJ 170278 7 UADBA (MIR 187) Ifaty site 2 -23.1241, 43.6496 - NA KJ 170279 3 UADBA (ZCMV 12840) Ifaty site 2 -23.1241, 43.6496 - KJ 170249 KJ 170275 4 UADBA (ZCMV 12839) Ifaty site 2 -23.1241, 43.6496 - KJ 170250 KJ 170276 5 UADBA (ZCMV 12838) Ifaty site 2 -23.1241, 43.6496 - KJ 170251 KJ 170277 Tombohina ZSM 828 / 2010 (ZCMV 12861) Tombohina - 23.8673, 44.0877 Yes KJ 170253 KJ 170280 Beloha ZSM 829 / 2010 (ZCMV 12865) Road Beloha ��� - 25.1375, 45.1550 Yes KJ 170254 KJ 170281 Tsiombe Beloha UADBA (ZCMV 12866) Road Beloha ��� - 25.1375, 45.1550 - KJ 170255 KJ 170282 Tsiombe Beloha UADBA (ZCMV 12867) Road Beloha ��� - 25.1375, 45.1550 - KJ 170256 KJ 170283 Tsiombe Beloha UADBA (ZCMV 12868) Road Beloha ��� - 25.1375, 45.1550 - KJ 170257 KJ 170284 Tsiombe Cap ZSM 830 / 2010 (ZCMV 12898) 2 km N Faux Cap - 25.55, 45.52 Yes KJ 170258 KJ 170285 Amboasary 1 ZSM 832 / 2010 (ZCMV 12905) 5 km N Amboasary - 24.9998, 46.3969 Yes KJ 170259 KJ 170286 Amboasary 2 ZSM 833 / 2010 (ZCMV 12906) 5 km N Amboasary - 24.9998, 46.3969 Yes KJ 170260 KJ 170287 ������continued on the next page ZSM 60 / 1907 Tsimanampetsotsa (N - 24.11, 43.84 Yes NA NA Mahafaly) ZSM 616 / 2000 (FGMV 2000.505) Ifaty - 23.15, 43.62 Yes NA NA ZSM 422 / 2002 Toliara -23.35,43.67 Yes NA NA . steinkampi sp. nov. Esomony 1 ZSM 139 / 2005 (FGZC 2330) (PT) N Tranomaro - 24.51792, 46.59895 Yes EU099710 EU099659 TM 71) Esomony 2 UADBA (FGZC 2363) (PT) Esomony - 24.51762, 46.60405 - EU099711 EU099660 TM 72) Esomony 3 UADBA (FGZC 2549) (PT) Esomony - 24.51762, 46.60405 - EU099704 EU099653 TM 73) Esomony 4 ZSM 140 / 2005 (FGZC 2550) (PT) Esomony - 24.51762, 46.60405 Yes EU099712 KJ 170300 * TM 74) Amboasary 4 ZSM 834 / 2010 (ZCMV 12908) (PT) 30 km N Amboasary - 24.7721, 46.4207 Yes KJ 170270 KJ 170297 Amboasary 5 ZSM 835 / 2010 (ZCMV 12909) (HT) 30 km N Amboasary - 24.7721, 46.4207 Yes KJ 170271 KJ 170298 Amboasary 6 ZSM 836 / 2010 (ZCMV 12910) (PT) 30 km N Amboasary - 24.7721, 46.4207 Yes KJ 170272 KJ 170299 Oplurus cyclurus outgroup) TM 100 UADBA (FGMV 2002.2093) Ifaty - 23.15, 43.62 - EU099749 EU099697, Published as part of Miralles, Aurelien, Glaw, Frank, Ratsoavina, Fanomezana M. & Vences, Miguel, 2015, A likely microendemic new species of terrestrial iguana, genus Chalarodon, from Madagascar, pp. 201-220 in Zootaxa 3946 (2) on pages 208-214, DOI: 10.11646/zootaxa.3946.2.3, http://zenodo.org/record/232560, {"references":["Munchenberg, T., Wollenberg, K. C., Glaw, F. & Vences, M. (2008) Molecular phylogeny and geographic variation of Malagasy iguanas (Oplurus and Chalarodon). Amphibia-Reptilia, 29, 319 - 327. http: // dx. doi. org / 10.1163 / 156853808785112101"]}

Fimbrios smithi sp. n. (Figs. 2���8) Holotype. IEBR 3157, an adult male, from Phong Nha���Ke Bang National Park, Quang Binh province, central Vietnam, collected by Doan Van Kien and Nguyen Van Hoan on 15 May 2005 in the Cha Noi region at an altitude of ca. 350 m above sea level. Etymology. We name this species in honour of Malcolm A. Smith, who described the monotypic genus Fimbrios with the species F. klossi, and in recognition of his outstanding herpetological contributions for Southeast Asia. Diagnosis. A species of the genus Fimbrios, characterized by a combination of the following characters: 1) rostral, mental and first three to four labials with raised, everted edges; 2) rostral separated from the internasals by a horizontal ridge of tissue; 3) suture between the internasals longer than that between the prefrontals; 4) a single pair of enlarged chin shields; 5) one large loreal, that extends from the nasal to the eye; 6) one preocular, one supraocular, two postoculars, and two suboculars; 7) 31 rows of keeled scales at midbody (those of the outer row enlarged); 8) 193 laterally rounded ventrals; 9) 72 unpaired subcaudals; 10) a total length of at least 440 mm in males (with a tail length of 94 mm, and a tail / total length ratio of 0.214); 11) dorsum greyish brown; with a paler flank area, and pale blotches and stripes at the neck region. Description of holotype. 440 mm total length (346 mm snout vent length, and 94 mm tail length); tail / total length ratio 0.214; body slender, cylindrical; head length 10.2 mm (from tip of snout to hind margin of parietal); head not distinct from neck, dorsally covered with large shields; eye small, with vertically subelliptic pupil; right maxilla with ca. 34 equal sized and curved teeth. Rostral partly destroyed, triangular, not visible from above, separated from the internasals by a horizontal ridge of tissue; suture between the internasals (1.4 mm) longer than that between the prefrontals (0.9 mm); nostril in the anterior part of a large, concave nasal; frontal slightly broader than long, broadly truncate in front, much shorter than the parietals; loreal large, pentagonal, extending from the nasal to the eye; one small preocular, higher than wide; one small supraocular, wider than high; two small postoculars; two suboculars, the anteriormost being distinctly larger; four anterior and five posterior temporals each; eight supralabials, the first four very small, with strongly raised edges, the last one much elongated; fifth and sixth supralabials in contact with the larger subocular; one mental, followed by ten infralabials; mental and anterior three infralabials with their edges raised like the supralabials; first pair of infralabials in contact with each other; first five infralabials in contact with the single pair of enlarged chin shields; chin shields covering nearly the whole of the chin in front, in contact with the first ventral; no posterior chin-shields. Dorsal scales elliptical, keeled from the neck region onwards; especially at the cloacal region, the middle keel is distinctly pointed posteriorly; 31 scale rows at midbody, those of the outer row enlarged; 36 scales round the anterior part of the body (one head length behind head), and 29 dorsal scale rows at posterior body (one head length before vent); 193 distinct, laterally rounded ventrals; 72 single subcaudals; anal entire; tail moderate. The invertedly dissected right hemipenis proved to be deeply forked, with the region proximal to the bifurcation being smooth, and the area distal to the bifurcation being spinose. The formaldehyde-fixed and subsequently ethanol-preserved holotype is dark greyish brown above, the lower flanks are pale brown; there are few pale blotches discernible in the dorsal and lateral neck region; these blotches comprise one to five scales; in addition, there exists each a thin, one to three scales wide longitudinal pale stripe in the lateral neck region; these stripes begin about 7 mm behind the eyes, obliquely descend for about 3 mm and then horizontally stretch for further 7���13 mm; the chin region of the preserved holotype is brownish grey; the ventral and subcaudal scales are yellowish grey, edged with dark grey (most obvious in the posterior body and below the tail). Distribution. The discovery of the new species, which is known only from the type specimen from Phong Nha���Ke Bang National Park (see Fig. 9), took place at the type locality of the skink species Tropidophorus noggei and Lygosoma boehmei (see Ziegler et al. 2005, 2007 b). Natural history. The male holotype of Fimbrios smithi sp. n. was discovered on the forest ground in steep primary karst forest, near A Bo Doi limestone cave. We did not find any water courses in the immediate vicinity, but the snake was found within the dry season. The snake was found killed by local people on the forest path nearby karst rock outcrops., Published as part of Ziegler, Thomas, David, Patrick, Miralles, Aurelien, Kien, Doan Van & Truong, Nguyen Quang, 2008, A new species of the snake genus Fimbrios from Phong Nha ��� Ke Bang National Park, Truong Son, central Vietnam (Squamata: Xenodermatidae), pp. 37-48 in Zootaxa 1729 on pages 39-41, DOI: 10.5281/zenodo.181276

In a previous study, Miralles & Vences (2013) compared seven different methods of species delimitation applied to the genus Madascincus. While focusing on methodological aspects their study involved an extensive data set of multilocus DNA sequences and of comparative morphology. On this basis they emphasized the need of revising the taxonomy of Madascincus, and revealed the existence of at least two well-supported candidate species. The present paper provides formal descriptions of these two taxa: (1) M. miafina sp. n., a species from dry areas of northern Madagascar, morphologically very similar to M. polleni (although both species are not retrieved as sister taxa), and (2) M. pyrurus sp. n., a montane species occurring >1500 m above sea level, endemic to the central highlands of Madagascar (Ibity and Itremo Massifs). Phylogenetically, M. pyrurus is the sister species of M. igneocaudatus, a taxon restricted to the dry littoral regions of the south and south-west of Madagascar in lowlands

In a previous study, Miralles & Vences (2013) compared seven different methods of species delimitation applied to the genus Madascincus. While focusing on methodological aspects their study involved an extensive data set of multilocus DNA sequences and of comparative morphology. On this basis they emphasized the need of revising the taxonomy of Madascincus, and revealed the existence of at least two well-supported candidate species. The present paper provides formal descriptions of these two taxa: (1) M. miafina sp. n., a species from dry areas of northern Madagascar, morphologically very similar to M. polleni (although both species are not retrieved as sister taxa), and (2) M. pyrurus sp. n., a montane species occurring >1500 m above sea level, endemic to the central highlands of Madagascar (Ibity and Itremo Massifs). Phylogenetically, M. pyrurus is the sister species of M. igneocaudatus, a taxon restricted to the dry littoral regions of the south and south-west of Madagascar in lowlands

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