Calumma tjiasmantoi, Proetzel, Scherz, Ratsoavina, Vences & Glaw, 2020

Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 2020, Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae), Vertebrate Zoology 70 (1), pp. 23-59 : 45-48

Description of Calumma tjiasmantoi sp. nov.

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Remark: This new species refers to clade J of Fig. 2 and GEHRING et al. (2012). Due to uncertainties about the collection data of the only male specimen we designate a well-documented female as the holotype.

Holotype: ZSM 735 /2003 ( FG / MV 2002-497 ), adult female collected in Ranomafana National Park (21.2639°S, 47.4194°E, 983 m a.s.l.), Vatovavy-Fitovinany Region, eastern Madagascar, on 23 January 2003 by F. Glaw, M. Puente, L. Raharivololoniaina, M. Thomas, D. R. Vieites.

Paratypes: ZSM 312 /2006 ( ZCMV 2896 ), adult male, and UADBA uncatalogued ( ZCMV 2895 ), female, both collected in Ranomafana , probably Ambatolahy (21.2439°S, 47.4262°E, 919 m a.s.l.) on 21 February 2006 by M. Vences ; ZSM 723 /2003 ( FG / MV 2002- 0396 ) and ZSM 736 /2003 ( FG / MV 2002-0498 ), both adult females, same collection data as holotype ; ZSM 380 /2016 ( ZCMV 14835 ), adult female, collected in Sampanandrano (24.1399°S, 47.0742°E, 539 m a.s.l.), Atsimo-Antsinanana Region, southeastern Madagascar, on 16 December 2016 by A. Rakotoarison, E. Rajeriarison, J.W. Ranaivosolo .

Diagnosis (based on the type series, osteology based on micro-CT scan of ZSM 735/2003, female): Calumma tjiasmantoi sp. nov. is characterised by (1) a small size (male SVL 43.9 – 46.8 mm, female TL 84.1 – 94.8 mm); (2) a medium sized (1.1 – 2.1 mm) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4 – 8) rostral, lateral, temporal, cranial, and parietal crests present, (9) casque medium sized in males (1.3 mm), (10) a dorsal crest of 7 – 9 spines can be present in males (based also on photographs), absent in females, (11) 15 – 17 supralabial scales with a mostly straight upper margin, (12) general absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 0.6 – 0.8 mm, (14) frontoparietal fenestra absent, (15) parietal and squamosal in contact, (16) parietal bone width at midpoint 16.1% of skull length (n = 1) with a characteristic broad tip to the postparietal process, (17) bright green or yellowish body colouration in males, females generally browner and less conspicuous, (18) rostral appendage colour generally inconspicuous, (19) cheek colouration greenish to turquoise, (20) five characteristic dorsoventral stripes of blue or brown colour, and (21) a diffuse brown stripe crossing the eye.

Calumma tjiasmantoi sp. nov. can easily be distinguished from all species of the C. boettgeri complex (see above) by the absence of occipital lobes; from C. gallus by different length, shape and colour of its rostral appendage (see above); from C. vatosoa by presence of a rostral appendage (vs absence); from C. vohibola by longer rostral appendage in females (RRS 2.4 – 4.6% vs 0.2 – 0.7%), parietal crest present (vs absent), smaller temporal scale (0.6 – 0.8 mm vs 1.0 mm); from C. nasutum as redefined herein by the higher number of supralabials (15 – 17 vs 12 – 15), a shorter rostral appendage (4.3% of SVL vs 4.5 – 5.3% of SVL), a shorter parietal (36.3% of skull length vs 41.0 – 44.3%), broad postparietal process (vs narrow); from C. radamanus by a relatively longer tail in females (RTaSV 92 – 95% vs 79 – 89%), rostral scale not integrated in rostral appendage (vs generally integrated), parietal crest present (vs absent), more supralabials (15 – 17 vs 11 – 15) with a generally straight upper margin (vs serrated), parietal and squamosal in contact (vs widely separated); from C. emelinae sp. nov. by presence of parietal crest (vs usually absent), absence of lateral white stripe, and broader postparietal process; for diagnosis against C. fallax , see below. For diagnosis against the other species described herein, see its description below.

Description of the holotype ( Fig. 4D ): Adult female, with mouth closed, in good state of preservation except for a ventrally sliced body, with four eggs still fixed in the oviduct; SVL 45.5 mm, tail length 42.5 mm, for other measurements, see suppl. Table 1 ; bulging rostral ridges; laterally flattened and distally rounded rostral appendage of tubercle scales that projects straight forward over a length of 1.7 mm with a diameter of 1.4 mm not including the rostral scale; 15 infralabial and 15 supralabial scales, all rather small; most of the supralabials with a straight upper margin, only anterior scales serrated; distinct lateral crest running horizontally; temporal crest consisting of two tubercles; short cranial crest; short parietal crest; no occipital lobes; medium sized casque of 1.1 mm height; no dorsal, gular or ventral crest. Body laterally compressed with fine homogeneous scalation and only slightly larger scales on extremities and head region, largest scale in temporal region with diameter of 0.6 mm and in cheek region of 0.6 mm; no axillary or inguinal pits.

Skull osteology of the holotype ( Fig. 11B ): Skull length 12.4 mm; snout-casque length 14.4 mm; narrow paired nasals anteriorly still in contact; anterior tip of frontal not exceeding the middle of the prefrontal fontanelle, which is fused with the naris; prominent and broad prefrontal with laterally raised tubercles; frontal and parietal smooth with only a few tubercles; frontal with a width of 2.4 mm (19.4% of skull length) at border to prefrontal extending to 4.3 mm (34.7%) at border to postorbitofrontal; no frontoparietal fenestra; broad parietal with distinct parietal crest tapering slightly from a width of 4.3 mm (34.7%) at the border to postorbitofrontal to a width at midpoint of 2.0 mm (16.1%); the posterodorsally broadened end is in strong contact with the squamosals; squamosals broad with several tubercles. For further measurements, see Table 2 .

Hemipenial morphology, based on ZSM 312/2006 (no micro-CT scan available): medium sized calyces (hemipenial character A); two pairs of small rotulae on apex of about the same size (B), finely denticulated with about 7 – 9 tips each; papillary field of small, unpaired papillae (C); pair of medium sized cornucula gemina (D), only visible when hemipenis fully everted.

Variation: For variation in measurements, see Table 1 . For variation in colouration in life, see Fig. 13 . The female ZSM 134/2005 (FGZC 2508) from Andohahela at high elevation (1548 m a.s.l.) belongs genetically to clade J but shows substantial morphological differences so that we do not designate it as a paratype. Aside from a larger total length (97.0 mm), a longer rostral appendage (2.6 mm), and a low number of supralabials (12) it also has a unique skull morphology with a FF. This correlates well with a previous study where only high elevation species tend to have a FF ( PRÖTZEL et al., 2018b) but does not fit in the general pattern of C. tjiasmantoi sp. nov. Further studies are necessary to clarify this apparent contradiction in our dataset and potential taxonomic conclusions. Sexual dimorphism: Body size (SVL and TL) is slightly larger in males than females, tail length is longer in males than in females (RTaSV 103% vs 92 – 95%), and a dorsal crest is only present in males.

Colouration in life: Strong sexual dichromatism with males of bright green or yellowish body colouration and turquoise extremities and females generally browner and less conspicuous. In both sexes five diffuse dorsoventral stripes on the body – in males of blue colour with light spots, in females of brown colour; no lateral stripe across the body; tail annulated, continuing the blotches from the body; throat and ventral region can be beige or of same colour as the body; indistinct rostral appendage not accentuated from the head; cheek region turquoise in males; a diffuse brown stripe may cross the eye.

Etymology: The specific epithet is a patronym honouring Wewin Tjiasmanto in recognition of his support for taxonomic research and nature conservation projects in Madagascar through the BIOPAT initiative (http://biopat. de/en).

Distribution ( Fig. 9 ): Calumma tjiasmantoi sp. nov. is known from eastern Madagascar from Andohahela in the south to Ranomafana NP about 400 km further north (for coordinates, see above), from an elevation of 267 to 983 m a.s.l. (see Fig. 9 ).

Figures

Fig. 2. Maximum likelihood tree based on DNA sequences of the mitochondrial ND2 gene (582 bp) of species of the Calumma nasutum group. Numbers at nodes are bootstrap proportions expressed as a percentage (500 replicates). Tip labels are field numbers followed by collection locality. Note: Calumma lefona was listed with a wrong field number (DRV 6284) in Fig.1 of Prötzel et al. (2018b) and in Fig.2 of Prötzel et al. (2018a).

Fig. 4. Preserved type specimens of (A) Calumma nasutum (lectotype, MNHN 6643C), adult male; (B) C. radamanus (holotype, SMF 22132), adult male; (C) C. emelinae sp. nov.(holotype, ZSM 618/2009), adult male; (D) C. tjiasmantoi sp. nov.(holotype, ZSM 735/2003), adult female; (E) C. fallax (lectotype, MNHN 1899.317), adult male; (F) C. ratnasariae sp. nov. (ZSM 35/2016), adult male.

Fig. 11. Micro-computed tomography scans of the skulls of holotypes of species with a closed skull roof in dorsal and lateral view. (A) C. emelinae sp. nov.(ZSM 618/2009), male holotype; (B) C. tjiasmantoi sp. nov. (ZSM 735/2003), female holotype.Diagnostic characters are encircled in red. Scale bars: 2.0 mm.

Fig. 13. Calumma tjiasmantoi sp. nov. in life. (A, B) adult male (not collected) from Ranomafana NP in relaxed state, photos: M. Knauf; (C) adult male (probably in UADBA) from Andohahela (Camp 2 low elevation) in slightly stressed colouration; (D) adult male and (E) adult female, both from Andreoky, referring to tissue samples PSG_2766, 2768, 2786 in GEHRING et al. (2012) in relaxed state, photos: P-S. Gehring.

Fig. 9. Distribution map of six species of the Calumma nasutum group in Madagascar. Localities based on collected specimens and/or DNA sequences of GEH- RING et al. (2012) and new sequences shown in Fig. 2. Black stars within the symbols indicate type localities. Colours correspond to those used in Fig. 2.