Calumma ratnasariae, Prötzel, David, Scherz, Mark D., Ratsoavina, Fanomezana M., Vences, Miguel & Glaw, Frank, 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 : 52-54

Description of Calumma ratnasariae sp. nov.

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Remark: This new species refers to clade I of Fig. 2 and GEHRING et al. (2012).

Holotype: ZSM 35 /2016 ( MSZC 0066 ), adult male, collected in the Ampotsidy mountains (14.4146°S, 48.7115°E, 1400 m a.s.l.), Sofia Region, northern Madagascar, on 22 December 2015 by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, T. S.E. Razafimandimby, D.H. Nomenjanahary, J. Rabearivony.

Paratypes: ZSM 1724 /2010 ( ZCMV 12483 ), adult male and ZSM 2884 /2010 ( ZCMV 12273 ), adult female, both collected in Analabe Forest , near Antambato village (Ambodimanga mountain, 14.5048°S, 48.8760°E, 1361 m a.s.l.) on 24 June 2010 , ZSM 517 / 2014 ( DRV 6283 ), adult male collected in Andrevorevo (14.3464°S, 49.1028°E, 1717 m a.s.l.), Sofia Region, northern Madagascar on 21 June 2010 , all three by M. Vences, D. R. Vieites , R. D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison , T. Rajoafiarison ; ZSM 36 /2016 ( MSZC 0140 ), adult female, collected on the Ampotsidy mountains (14.4099°S, 48.7155°E, 1647 m a.s.l.) on 3 January 2016 , and ZSM 37 /2016 ( MSZC 0169 ), adult female, collected on the Ampotsidy mountains (14.4193°S, 48.7193°E, 1337 m a.s.l.), on 8 January 2016 , both by M.D. Scherz, J. Borrell, L. Ball, T. S.E. Razafimandimby, D.H. Nomenjanahary, J. Rabearivony .

Diagnosis (based on the type series; osteology based on micro-CT scans of ZSM 35/2016 and ZSM 517/2014, both males): Calumma ratnasariae sp. nov. is characterised by (1) a large size (male SVL 43.9– 52.0 mm, female SVL 48.7 – 51.5 mm; male TL 97.1 – 110.7, female TL 95.3 – 101.0); (2) a short (1.8 – 2.3 mm in males, 2.1 – 2.2 mm in females) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4 – 7) rostral, lateral, temporal (one tubercle on either side), and cranial crests present, (8) parietal crest distinct and running the length of the parietal bone, (9) a distinctly raised casque in males with a height of 1.3– 1.5 mm, (10) a dorsal crest of 7 – 12 cones in males, generally present in females (6 – 8 cones), (11) 10 – 13 supralabial scales with a straight upper margin, (12) absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 1.2 – 1.6 mm, (14) a frontoparietal fenestra in the skull, (15) parietal and squamosal in contact (n = 2), (16) parietal bone width at midpoint 17.8 – 18.5% of skull length, (17) a generally yellowish body colouration in males, greyish body colouration in females, (18) rostral appendage not accentuated from the body colouration, (19) a blue and yellow cheek colouration, (20) yellow in males and beige in females, and (21) brown stripe crossing the eye. Calumma ratnasariae sp. nov. is unique among the C. nasutum complex in having an elevated bony knob on the anterodorsal edge of the maxillary facial process (this character is similar to that seen in C. uetzi ).

Calumma ratnasariae sp. nov. can 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 all other species of the C. nasutum group without occipital lobes except C. fallax by the presence of a frontoparietal fenestra. It is also quite unusual in having an overall yellowish body colouration in males. In addition, it can be distinguished from C. vatosoa easily by the presence of a rostral appendage (vs absence); from C. vohibola by longer rostral appendage (RRS 3.8 – 4.8% vs 0.2 – 3.1%), parietal crest present (vs absent), fewer supralabials (10 – 13 vs 13 – 16) with a straight upper margin (vs serrated), larger temporal scale (1.2 – 1.6 mm vs 1.0 mm), broader parietal bone with a continuous parietal crest (vs smooth parietal); from C. nasutum as here redefined by a larger maximum total length in males (110.7 mm vs 89.0 – 100.8 mm), a distinct parietal crest (vs absent or indistinct), dorsal crest generally present in both sexes (vs generally absent and absent in females); from C. radamanus by larger total length (95.3– 110.7 mm vs 77.0 – 93.5 mm), tail length in males longer than SVL (vs shorter), rostral scale not integrated into the rostral appendage (vs generally integrated), parietal crest present (vs absent), supralabials with a straight upper margin (vs serrated), larger temporal scale (1.2 – 1.6 mm vs 0.6 – 0.9 mm), and parietal and squamosal in contact (vs widely separated); from C. emelinae sp. nov. by parietal crest distinct (vs general absence), higher casque in males (1.3 – 1.5 mm vs 0.5 – 1.1 mm), dorsal crest consisting of cones (vs spines) in males; larger temporal scale (1.2 – 1.6 mm vs 0.6 – 1.0 mm), and broad postparietal process (vs strongly tapering); from C. tjiasmantoi sp. nov. by larger body length of females (SVL 48.7 – 51.5 mm vs 43.9 – 46.1 mm), dorsal crest generally present in females (vs absent), fewer supralabials (11 – 12 vs 15 – 17), and larger diameter of temporal scale (1.2 – 1.6 mm vs 0.6 – 0.8 mm); and from C. fallax by generally shorter relative rostral appendage length in females (RRS 4.1 – 4.5% vs 4.2 – 7.6%), cranial crest present (vs generally absent), parietal crest longer and more distinct, dorsal crest generally present in females (vs generally absent), and a wider mid-parietal width (17.8 – 18.5% of skull length vs 6.7 – 15.7%).

Description of the holotype ( Figs. 4F , 16 A,B ): Adult male, with mouth closed, in good state of preservation, with everted hemipenes, one completely (on the left) and one incompletely (on the right); SVL 48.2 mm, tail length 54.2 mm, for other measurements, see suppl. Table 1 ; distinct rostral ridges that give the snout a right angle; laterally compressed dermal rostral appendage of oval tubercle scales that projects slightly downwards over a length of 2.3 mm with a diameter of 1.9 mm; 13 infralabial and 12 supralabial scales, both relatively large; supralabials with a smooth dorsal margin; distinct lateral crest running horizontally; temporal crest consisting of one tubercle per side; short cranial crest; distinct and long parietal crest ending in the tip of the casque with a height of 1.5 mm; no occipital lobes; dorsal crest present, consisting of 7 broad cones; no traces of gular or ventral crest. Body laterally compressed with fine homogeneous scalation and distinctly larger scales on extremities and head region, largest scale in temporal region with diameter of 1.3 mm and in cheek region of 1.1 mm; no axillary or inguinal pits.

Skull osteology of the holotype ( Fig. 14D ): Skull length 11.9 mm; snout-casque length 14.4 mm; narrow paired nasals anterior slightly connected; anterior tip of frontal exceeding the middle of the prefrontal fontanelle, which is fused with the naris; prominent and broad prefrontal; frontal and parietal smooth without any tubercles; frontal with a width of 2.6 mm (21.8% of skull length) at border to prefrontal extending to 4.4 mm (37.0%) at border to postorbitofrontal; large frontoparietal fenestra with a width of 2.5 mm (21.0%); broad parietal with distinct parietal crest tapering slightly from a width of 4.0 mm (33.6%) at the border to postorbitofrontal to a width at midpoint of 2.2 mm (18.6%); the posterodorsally broadened end is in weak contact with the squamosals; squamosals thin with a few tubercles. For further measurements, see Table 2 .

Hemipenial morphology, based on ZSM 1724/2010 ( Fig. 5E ), ZSM 517/2014, and ZSM 35/2016: small calyces (hemipenial character A); two pairs of finely denticulated rotulae of different size, on sulcal side large with about 12 – 15 tips, on asulcal side small with about 5 – 7 tips (B); papillary field of small, unpaired papillae (C); pair of short cornucula gemina (D), only visible when hemipenis fully everted.

Variation: For variation in measurements, see Table 1 . Sexual dimorphism: Males and females do not seem to differ in body size. Tail length is longer in males than in females (RTaSV 112 – 121% vs 94 – 98%). Relative rostral appendage length does not differ. Dorsal crest does not differ between males and females.

Colouration in life ( Fig. 16 ): Strong sexual dichromatism with males of yellow body colouration and turquoise stripes and females generally uniformly beige. Males with turquoise annulated tail and extremities and two brown blotches on the body side that is crossed by a diffuse white lateral stripe; no pattern in females; throat and ventral region slightly brighter than the flank; indistinct rostral appendage not accentuated from the body/head, can be spotted with blue or yellow dots; in males cheek region and eyelids with turquoise dots, in females blue dots can occur on rostral ridges and eyelids when stressed; a diffuse brown stripe crosses the eye in both sexes.

Etymology: The specific epithet is named after Yulia Ratnasari, in recognition of her support for taxonomic research and nature conservation projects in Madagascar through the BIOPAT initiative (http://biopat.de/en/).

Distribution ( Fig. 9 ): Calumma ratnasariae sp. nov. is only known from the Bealanana District of northern Madagascar. It is distributed from Analabe Forest in the south to Andrevorevo, about 20 km further north (for coordinates, see above), from an elevation of 1337– 1717 m a.s.l.

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. 16. Calumma ratnasariae sp. nov. from Ampotsidy mountains. (A, B) Adult male holotype (ZSM 35/2016), slightly displaying, and (C) adult female (MSZC 0130, UADBA uncatalogued) in relaxed state.

Fig. 14. Micro-computed tomography scans of the skulls of species with a frontoparietal fenestra in dorsal and lateral view. (A) Calumma fallax (MNHN 1899.317), male lectotype; (B) C. fallax (MNHN 1840.430), male; (C) C. fallax (ZSM 286/2016), male of genetic clade H, assigned to C. fallax; (D) C. ratnasariae sp. nov. (ZSM 35/2014), male holotype. Diagnostic characters are encircled in red. Scale bars: 2.0 mm.

Fig. 5. Micro-CT scans of hemipenes of representative males of the different new or redescribed Calumma species (as far as available), each in sulcal (left) and asulcal or lateral view (right). (A) C. fallax (ZSM 694/2003); (B) C. radamanus (ZSM 443/2005); (C) C. emelinae sp. nov. (ZSM 618/2009); (D) C. nasutum (ZSM 924/2003); (E) C. ratnasariae sp. nov. (ZSM 1724/2010). Scale bar=1 mm.

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.