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Calumma radamanus (Mertens, 1933)
in Prötzel, Scherz, Ratsoavina, Vences & Glaw, 2020 |
Based on a large number of specimens and genetic samples, we revise the chameleons of the phenetic Calumma nasutum species group using an integrative taxonomic approach including external and hemipenial morphology, osteology, and sequences of a mitochondrial (ND2) and a nuclear gene (c-mos). After more than 180 years of taxonomic uncertainty, the eponymous species of the group, C. nasutum, is re-described, a lectotype is designated, and the species is assigned to a genetic clade that occurs in eastern (Anosibe An’Ala, Andasibe) and northern Madagascar (Sorata) based on morphology and osteology. The identity of C. fallax is also clarified and a lectotype is designated; it occurs at high elevation along the east coast from Andohahela (south) to Mandraka (central east). Calumma radamanus is resurrected from synonymy of C. nasutum; it lives at low elevations in eastern Madagascar from Tampolo (south) to its type locality Ambatondradama (north). However, up to five deep mitochondrial lineages and high morphological variation are identified within C. radamanus, which we consider a species complex still in need of further taxonomic revision. Furthermore, three new species of the C. nasutum group are described: C. emelinae sp. nov. is distributed in eastern Madagascar (Anosibe An’Ala in the south to Makira in the north), C. tjiasmantoi sp. nov. in southeastern Madagascar (from Andohahela in the south to Ranomafana NP in the north), and C. ratnasariae sp. nov. is known from the Bealanana District in northern Madagascar. There is only little variation in hemipenial morphology in this group; the cornucula gemina are present in all taxa except the C. radamanus complex. Due to this taxonomic revision the protection status of the treated six chameleon species needs to be newly assessed; at least two of the species appear to warrant threatened statuses.
Key words: Calumma nasutum species group; hemipenis morphology; integrative taxonomy; Madagascar; new species; osteology.
the Calumma nasutum group
Calumma nasutum (Duméril & Bibron, 1836)
Etymology: the Latin adjective nasutum meaning ‘big-nosed,’ in the neuter nominative singular; obviously in reference to the characteristic rostral appendage.
Distribution (Fig. 9): Calumma nasutum, as redefned here, is known from eastern Madagascar between Anosibe An’ala (...) and Andasibe in the central east to Sorata, more than 600 km further north (see coordinates above), from an elevation of 880 –1400 m a.s.l.
Calumma radamanus (Mertens, 1933)
Etymology: Radamanus means ‘from Radama’ in Latin (-anus declension meaning ‘of’ or ‘pertaining to’), and clearly refers to the origin of specimens from Col Pierre Radama. However, it is not clear from the word ‘radamanus’ nor from the original description whether this is to be treated as a substantive noun or an adjective. Thus article 31.2.2 of The International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999) has to be applied: ‘Where the author of a species-group name did not indicate whether he or she regarded it as a noun or as an adjective, and where it may be regarded as either and the evidence of usage is not decisive, it is to be treated as a noun in apposition to the name of its genus.’ Given that the taxon Chamaeleo radamanus disappeared soon after its description into the synonymy of C. nasutum, there was no decisive usage of this name (see above). Therefore, the name is to be considered an invariable noun in apposition, and its declension is not changed.
Distribution (Fig. 9): Calumma radamanus sensu stricto as redefned here, is known from eastern Madagascar between Tampolo and the type locality Ambatond’Radama about 250 km further north (for coordinates, see above), from an elevation of about 7–500 m a.s.l. (see Fig. 9). For the distribution of the C. radamanus complex, see Fig. 9. Specimens from Montagne d’Ambre were not analysed here, but probably also belong to the C. radamanus complex, based on overall appearance.
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Calumma emelinae sp. nov.
adult male (not collected) from the type locality in Makira in stress colouration. |
Calumma emelinae sp. nov.
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.
Etymology: The specifc 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.
Calumma tjiasmantoi sp. nov.
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) fve characteristic dorsoventral stripes of blue or brown colour, and (21) a diffuse brown stripe crossing the eye.
Etymology: The specifc 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).
Calumma fallax (Mocquard, 1900)
Etymology: A Latin adjective meaning ‘deceptive’ or ‘fallacious’ in the neutral nominative, with unclear justifcation.
Distribution (Fig. 9): Calumma fallax as redefned here, occurs in eastern Madagascar from Andohahela in the south to Mandraka about 650 km further north (coordinates, see above), from an elevation of 922–1781 m a.s.l.
Calumma ratnasariae sp. nov.
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).
Etymology: The specifc 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.
David Prötzel, Mark D. Scherz, Fanomezana M. Ratsoavina, Miguel Vences and Frank Glaw. 2020. Untangling the Trees: Revision of the
Calumma nasutum complex (Squamata: Chamaeleonidae).
Vertebrate Zoology. 70(1): 23-59. DOI:
10.26049/VZ70-1-2020-3
