Calumma emelinae, 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 : 44-45

Description of Calumma emelinae sp. nov.

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

Holotype: ZSM 618 /2009 ( ZCMV 11292 ), adult male with completely everted hemipenes, right hemipenis cut off for micro-CT scanning, collected in the Makira plateau, Angozongahy or Ampofoko (about 15.44°S, 49.12°E, 1000 m a.s.l.), Analanjirofo Region, northeastern Madagascar, on 22– 25 June 2009 by M. Vences, D. R. Vieites, F.M. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajofiarison, J. Patton.

Paratypes: ZSM 660 /2014 ( DRV 5899 ) , ZSM 663 /2014 ( DRV 5898 ), both adult males, collected in Angozongahy , western side of Makira plateau camp 1 (15.4370°S, 49.1186°E, 1009 m a.s.l.), Analanjirofo Region, northeastern Madagascar, on 26 June 2009 by M. Vences, D . R. Vieites, F.M. Ratsoavina , R. D. Randrianiaina, E. Rajeriarison , T. Rajofiarison, J. Patton ; ZSM 553 /2001 ( MV 2001- 239 ), adult female, collected in Andasibe (about 18.93°S, 48.42°E, 900 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 16 – 18 February 2001 by M. Vences, D . R. Vieites ; ZSM 135 /2005 ( FGZC 2692 ) and ZSM 136 /2005 ( FGZC 2693 ), both adult females and collected in Vohidrazana (about 18.95°S, 48.50°E, 700– 800 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 09 February 2005 by F. Glaw , R.D. Randrianiaina , R. Dolch ; ZSM 661 /2014 ( DRV 5677 ) and ZSM 662 /2014 ( DRV 5708 ), both adult females, collected in Mahasoa campsite near Ambodisakoa village (NE Vohimena, NE Lake Alaotra, 17.2977°S, 48.7019°E, 1032 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 13– 15 February 2008 by D . R. Vieites , J. Patton, P. Bora, M. Vences ; ZSM 148 /2016 ( FGZC 5236 ), adult female, collected east of Moramanga in ‘ Julia Forest’ (18.9511°S, 48.2719°E, 941 m a.s.l.) on 6 January 2015 ; ZSM 147 /2016 ( FGZC 5175 ), adult female collected south of Moramanga (19.0192°S, 48.2341°E, 903 m a.s.l.), Alaotra-Mangoro Region, eastern Madagascar, on 4 January 2016, both by F. Glaw, D. Prötzel, L. Randriamanana .

Diagnosis (based on the type series; osteology based on micro-CT scan of ZSM 618/2009, male): Calumma emelinae sp. nov. is characterised by (1) a medium size (male SVL 46.6 – 48.7 mm, female SVL 40.1 – 49.1 mm; male TL 93.6 – 103.2 mm, female TL 82.7 – 95.8 mm), (2) a medium (2.3 – 2.9 mm in males 1.5 – 1.8 mm in females) and distally rounded rostral appendage, (3) rostral scale not integrated into the rostral appendage, (4) rostral crest present, (5) lateral crest present, (6) temporal crest present, (7) cranial crest variable, (8) parietal crest usually absent, (9) casque low in males with a height of 0.5 – 1.1 mm, (10) a dorsal crest of 7 – 10 spines in males, absent in females, (11) 12 – 16 supralabial scales with a mostly straight upper margin, serrated anteriorly, (12) absence of axillary pits, (13) diameter of the largest scale in the temporal region of the head 0.6 – 1.0 mm, (14) no frontoparietal fenestra, (15) parietal and squamosal in contact, (16) parietal bone width at midpoint 16.2% of skull length (n = 1), (17) a generally greyish to greenish body colouration, (18) rostral appendage colour generally unremarkable, (19) a green cheek colouration, (20) suggestions of two weak bluish lateral blotches, and (21) no strong eye colouration.

Calumma emelinae 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 generally longer relative rostral appendage length (RRS 3.1 – 6.1% vs 0.1 – 1.4%), dorsal crest always present in males (vs generally absent), and pointed tip of postparietal process (vs relatively broad), and crenate prefrontal (vs smooth); from C. nasutum as redefined herein by a lower casque in males (0.5 – 1.1 mm vs 1.5 – 2.0 mm), dorsal crest present in males consisting of spines (vs general absence or consisting of cones if present), and scales more homogeneous (largest temporal scale in males 0.7 mm vs 0.9 – 1.6 mm); from C. radamanus by relatively longer tail in males (longer than SVL vs shorter), longer rostral appendage in males (RRS 4.7 – 6.1% vs 2.9 – 3.6%), rostral scale not integrated into rostral appendage (vs integrated), supralabials with a largely straight upper margin (vs serrated), and parietal and squamosal in contact (vs widely separated); for diagnosis against C. fallax , see below. For diagnosis against the other species described herein, see their respective descriptions below.

Description of the holotype ( Fig. 4C ): Adult male, with mouth closed, in good state of preservation, both hemipenes fully everted; SVL 47.9 mm, tail length 50.7 mm, for other measurements, see suppl. Table 1 ; rostral ridges that give the snout a right angle; laterally flattened and distally rounded rostral appendage of small tubercle scales that projects straight forward over a length of 2.9 mm with a diameter of 2.4 mm not including the rostral scale; 14 infralabial and 15 supralabial scales, both rather small; supralabials with a straight upper margin; distinct lateral crest running horizontally; no temporal or cranial crest; low parietal crest; no occipital lobes; very low casque of 0.5 mm height; dorsal crest consisting of 10 spines; no gular or ventral crest. Body laterally compressed with fine homogeneous scalation and slightly larger scales on extremities and head region, largest scale in temporal region with diameter of 0.7 mm and in cheek region of 0.9 mm; no axillary or inguinal pits.

A B

Skull osteology of the holotype ( Fig. 11A ): Skull length 11.7 mm; snout-casque length 13.8 mm; narrow paired nasals still slightly connected at anterior end; anterior tip of frontal extending to about the middle of the prefrontal fontanelle, which is fused with the naris; prominent and broad prefrontal with laterally raised tubercles; frontal and parietal covered with a few tubercles; frontal with a width at border to prefrontal of 3.0 mm (25.6% of skull length) extending to 4.4 mm (37.6%) at border to postorbitofrontal; no frontoparietal fenestra; narrow parietal with a width at the border to postorbitofrontal of 4.2 mm (35.9%) and a width at midpoint of 1.9 mm (16.2%) tapering strongly posterodorsally; parietal laterally in strong contact with the squamosals; squamosals relatively thick and covered with several tubercles. For further measurements, see Table 2 .

Hemipenial morphology, based on ZSM 618/2009 ( Fig. 5C ), ZSM 660/2014 and ZSM 663/2014: medium sized calyces (hemipenial character A); two pairs of small rotulae on apex of about the same size (B), roughly denticulated with about 9 – 12 tips each; 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 are usually larger than females (mean TL of 98.5 mm vs 88.3 mm). Tail length is generally longer in males than in females (RTaSV> 100% vs <100%), as well as the length of the rostral appendage (> 2.0 mm vs <2.0 mm).

Colouration in life ( Fig. 12 ): Both sexes with an indistinct brown to beige body colouration, extremities and tail of same colour as the body; three diffuse/scattered brown dorsoventral blotches can occur on the body; males can show a beige lateral stripe; throat and ventral region beige; rostral appendage not accentuated; cheek region can be light green; eyelids crossed by a brown stripe and occasionally with radiating dark green stripes in both sexes.

Etymology: The specific epithet is named after Emelina Widjojo, the mother of Wewin Tjiasmanto, 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 emelinae sp. nov. is known in eastern Madagascar from Anosibe An’Ala to Angozongahy (Makira) about 500 km further north (for coordinates, see above), from an elevation of 750 1030 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. 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. 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. 12. Calumma emelinae sp. nov. in life. (A) adult male (not collected) from the type locality in Makira in stress colouration; (B) sub- adult male (not collected) from Mahasoa, in relaxed state; (C) adult female (not collected) from Makira, slightly stressed; (D) adult female (FGZC 5273, UADBA uncatalogued) from east of Moramanga (‘Julia Forest‘) in relaxed state; (E) adult female (uncollected) from same location as (D), relaxed.

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.