Chalarodon steinkampi, Miralles, Aurelien, Glaw, Frank, Ratsoavina, Fanomezana M. & Vences, Miguel, 2015

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), pp. 201-220 : 208-214

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 Nomenclature1999)) 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<2<4<3, relative toe length 1<5<2<3<4, with toe 5 deeply separated from toes 1–4.

Rostral scale wider (2.7 mm) than tall (1.0 mm), wider as mental (1.7 mm). Nostrils directed laterally, not in contact with rostral. Supralabials smooth, with numerous pits; scale rows above supralabials with distinct keels and pits. Dorsal scales on head very heterogeneous in size, mostly with multiple, long, ornament-like keels. Parietal eye round, lense-like with shiny blackish surface, 0.5 mm in diameter, surrounded by a single ring-like scales, 1.5 mm in diameter, and a further ring consisting of 10 scales. Eyelids with numerous small conical tubercles in the periphery, with one row of enlarged flat scales around eye opening which is surrounded by a row of elevated enlarged tubercles. Eye opening 2.6 mm in diameter.

One heterogeneous row of distinctly enlarged vertebral scales each with one very distinct keel from the neck along the back, fading towards the base of the tail. Scales on the back generally small, not imbricated, with one distinct keel or a conical tubercle, with keels generally getting less distinct towards the flanks. Scales on dorsal surfaces of forelimbs, hindlimbs and tail distinctly keeled, partly with slight imbrications. Two strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca. No precloacal pores. Tail base laterally enlarged anteriorly, nearly round in cross section posteriorly, with sharply pointed tip, without distinct tubercles or spines.

Ventrally, a distinct skin fold between throat and chest. Ventral scales of chest and abdomen largely flat and mostly hexagonal without keels, gulars mostly small and rounded, but on the belly larger than scales on the flanks. Ventral scales on thighs without keels, on shanks ventrally with distinct keels, on feet and tail strongly keeled, except in the very anterior part of the tail, where scales are only slightly keeled. Subdigitals in single rows. Claws curving downwards, claws of fourth toe 1.7 mm, claws of third finger 1.0 mm.

Sublabial scales and ca. 3 adjacent scale rows towards the throat with distinct pit-like holes.

Mental triangular rounded, bordered by four scales (two infralabials and posteriorly by a pair of non-elongate, irregular hexagonal postmentals). Postmentals contact mental, other postmental, first infralabial, two enlarged lateral gulars, one smaller posterolateral gular and one tiny central gular. First infralabial taller than others. Gulars small, granular. Hemipenes with distinct long horn-like lobe.

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 (16S 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 Münchenberg 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 16S Accession c-mos number

madagascariensis

Antsalova (TM64) UADBA/FGZC 954 Antsalova -18.6667, 44.6167 - EU099705 EU099654

Betioky 1 (TM75) UADBA/FGZC 335 Betioky -23.7167, 44.3833 - EU099713 EU099662

Betioky 2 (TM76) UADBA/FGZC 336 Betioky -23.7167, 44.3833 - EU099714 EU099663 (TM85) UADBA 21059 (FGMV 2002.1465) Analalava (Isalo) -21.6917, 44.7833 - EU099706 EU099655

Toliara 1 (TM86) ZSM 946/2003 (FGMV 2002.1551) Toliara -23.35, 43.67 Yes EU099707 EU099656

Toliara 2 (TM87) UADBA (FGMV 2002.2017) Toliara -23.35, 43.67 - EU099708 EU099657

Toliara 3 (TM88) 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 ZSM422/2002 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 TM71)

Esomony 2 UADBA (FGZC 2363) (PT) Esomony -24.51762, 46.60405 - EU099711 EU099660 TM72)

Esomony 3 UADBA (FGZC 2549) (PT) Esomony -24.51762, 46.60405 - EU099704 EU099653 TM73)

Esomony 4 ZSM 140/2005 (FGZC 2550) (PT) Esomony -24.51762, 46.60405 Yes EU099712 KJ 170300 * TM74)

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)

TM100 UADBA (FGMV 2002.2093) Ifaty -23.15, 43.62 - EU099749 EU099697

Figures

FIGURE 3. Photographic comparisons of specimens of Chalarodon steinkampi sp. nov. (A: probably ZSM 836 / 2010, C: ZSM 835 / 2010, E: ZSM 834 / 2010, 835 / 2010, 836 / 2010) collected 30 km north of Amboasary with specimens of C. madagascariensis (B, D, F) collected at the closest known neighboring locality, 5 km north of Amboasary. (A – D) Specimens in life, photographed in the field and (E, F) ventral views of freshly dead specimens; (E) ZSM 834 – 836 / 2010, two paratypes and holotype of C. steinkampi; (F) ZSM 832 / 2010, ZSM 833 / 2010, and UADBA (ZCMV 12907).

FIGURE 5. Detailed views on mental region (A – B), belly scales (C – D) and hemipenes (E – F) of Chalarodon steinkampi sp. nov. (ZSM 835 / 2010) in comparison with C. madagascariensis (ZSM 832 / 2010). The ulcer-like tubercles on the belly in (C) were not studied but might be due to unidentified parasites. Note the long distal lobe of the hemipenis of C. steinkampi (F) and the pair of distinctly enlarged scales posterior to the cloaca of the males (E – F). Photographs by M. Franzen.

FIGURE 6. Holotype of Chalarodon steinkampi sp. nov. (ZSM 835 / 2010): (A) head with parietal eye in dorsal view, (B) dorsal view, (C) ventral view. Photographs by M. Franzen.

Tables

TABLE 1. (Continued)

Sample lab number Amboasary 3	Voucher number UADBA (ZCMV 12907)	Locality 5 km N Amboasary	GPS coordinates -24.9998, 46.3969	Morphology -	Accession 16S KJ170261	Accession c-mos KJ170288
Petriky 1	UADBA (MIR 223)	Petriky forest	-25.0484, 46.8629	-	KJ170262	KJ170289
Petriky 2	UADBA (MIR 230)	Petriky forest	-25.0484, 46.8629	-	KJ170263	KJ170290
Petriky 3	UADBA (MIR 234)	Petriky forest	-25.0484, 46.8629	-	KJ170264	KJ170291
Petriky 4	UADBA (MIR 233)	Petriky forest	-25.0484, 46.8629	-	KJ170265	KJ170292
Petriky 5	UADBA (MIR 239)	Petriky forest	-25.0484, 46.8629	-	KJ170266	KJ170293
Petriky 6	UADBA (ZCMV 12911)	Petriky forest	-25.0484, 46.8629	-	KJ170267	KJ170294
Petriky 7	ZSM 831/2010 (ZCMV 12912)	Petriky forest	-25.0484, 46.8629	Yes	KJ170268	KJ170295
Petriky 8	UADBA (MIR 227)	Petriky forest	-25.0484, 46.8629	-	KJ170269	KJ170296