Boophis spinophis Glaw & Köhler & Riva & Vieites & Vences 2010, sp. nov.
Creators: name: Glaw, Frank
name: Köhler, Jörn
name: Riva, Ignacio De La
name: Vieites, David R.
name: Vences, Miguel
Publisher: name: Zenodo
Year: 2010
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Mantellidae
Boophis
Boophis spinophis

description: Boophis spinophis sp. nov.   
(Fig. 13, Appendix 9)  
 Remark. This species has been referred to as Boophis sp. by Glaw & Vences (2007:164–165) and as Boophis sp. 15 in Vieites et al. (2009).   
 Holotype. ZSM 376 /2004 (ZCMV 691), adult male, from Ambatolahy forest, 21°14.632' S, 47°25.573' E, 915 m a.s.l., southeastern Madagascar, collected on 10 February 2004 by D. R. Vieites and C. Woodhead.   
 Etymology. The specific name is used as a noun in apposition and makes reference to the unique morphology of this species. In fact, when first examining the single specimen, we were unable to decide whether it belonged to the genus Boophis, or to the genus Spinomantis in the subfamily Mantellinae. The specific name is therefore a composite of these two generic names, and also makes reference to the dermal flaps and tubercles and tubercles and spines characterizing the species.   
 Diagnosis. Assigned to the genus Boophis based on the presence of an intercalary element between ultimate and penultimate phalanges of fingers and toes (verified by external examination), presence of nuptial pads and absence of femoral glands in the male, absence of gular glands in the male, enlarged terminal discs of fingers and toes, lateral metatarsalia separated by webbing, absence of outer metatarsal tubercle, molecular phylogenetic relationships (see Vieites et al. 2009 for a complete molecular analysis of Boophis), and overall similarity to other Boophis species. Assigned to the Boophis goudoti group based on the following combination of characters: relatively large size (male SVL 57 mm), brownish dorsal ground colour, non-transparent ventral skin, absence of red ventral colour; presence of webbing between fingers; apparently single subgular vocal sac; presence of vomerine teeth; and especially the molecular phylogenetic relationships of the species (see Vieites et al. 2009). Boophis spinophis differs distinctly from all described species in the B. goudoti group by strong genetic differentiation (see below), dorsal colouration and distinct dermal tubercles along the lateral parts of the tarsus and around the elbow (in life). It furthermore differs from B. boehmei, B. rufioculis, B. burgeri, B. reticulatus, and B. axelmeyeri by its larger size (SVL of the single known male 56.5 vs. 27–43 mm in the other mentioned species, see Glaw & Vences 2007) and from most of these species by iris colouration (absence of red colour in the outer iris area as is typical for B. axelmeyeri, B. boehmei, and B. rufioculis; presence of a strong pattern of dark stripes in iris which are absent in B. burgeri and usually less expressed in B. reticulatus). It also differs from B. goudoti and B. periegetes by the presence of a distinct dark pattern in the iris.   
 Description of the holotype. Adult male, SVL 56.5 mm. Body moderately slender; head approximately as long as wide, slightly wider than body; snout rounded in dorsal and lateral view, nostrils directed laterally, slightly nearer to tip of snout than to eye; canthus rostralis moderately sharp in cross section, but comparatively indistinct, slightly concave in dorsal view, loreal region slightly concave; tympanum distinct, rounded, TD 41% of ED; supratympanic fold thin, distinct; vomerine odontophores distinct, well separated in two elongated patches, positioned posteromedial to choanae; choanae medium-sized, elongated. Tongue posteriorly bifid, free.   
Arms slender; distal subarticular tubercles of fingers 1, 3 and 4 bifid, round; metacarpal tubercles not recognizable; fingers scarcely webbed, with lateral dermal fringes; webbing formula 1(1), 2i(2), 2e(1.25), 3i(2), 3e(1.5), 4(1.5); relative length of fingers 1<2<4<3 (finger 2 distinctly shorter than finger 4); finger discs enlarged. Hindlimbs slender; tibiotarsal articulation reaching nostril when hindlimb is adpressed along body; lateral metatarsalia separated by webbing; inner metatarsal tubercle small, distinct, elongated; no outer metatarsal tubercle; toes broadly webbed; webbing formula 1(0), 2i(0.25), 2e(0), 3i(0.5), 3e(0), 4i(0.75), 4e(0.75), 5(0); relative length of toes 1<2<5=3<4; toe discs enlarged. Dorsal skin smooth with small warts, smooth on throat and chest, granular on belly and ventral surfaces of thighs. Heel, lateral parts of tarsus, external finger and elbow region were covered with distinct dermal tubercles in life, which however are poorly recognizable in preservative. A tissue sample was removed from the right thigh.  
Measurements (in mm): SVL 56.5, HW 20.0, HL 20.1, ED 6.8, END 4.4, NSD 4.7, NND 6.8, TD 6.8, TL 29.9, HAL 18.7, FOL 26.5, FOTL 42.9.  
After almost four years in preservative, ground colour of upper surface of head and dorsum brown with irregular darker blotches and small cream spots. Ground colour of ventral surfaces uniformly creamy yellow.  
In life, dorsal surfaces of head, body and limbs pale brown with pale green dots and larger, darker brown circular blotches with pale green inside. A whitish subocular bar between upper jaw, tympanum and eye. Iris golden with brown markings, more densely arranged at the edges of iris; upper eye periphery blue (Fig. 13).  
 Natural history. Largely unknown. The single known specimen was collected at night along a stream next to Ambatolahy, in disturbed rainforest, perched on the vegetation and not emitting any call.   
 Molecular relationships. This species turned out to be phylogenetically highly divergent from all other species in the B. goudoti group, with pairwise distances>9% to all species. Phylogenies based on additional mitochondrial genes (not shown here) confirm, however, that B. spinophis belongs indeed into this group. The sequence of the holotype and single specimen from Ambatolahy clusters with high support together with a sequence that refers to a juvenile specimen from Ambohitantely (ZSM 118/2005, not studied here). The divergence between these two sequences is quite high (3.5%), indicating that these two geographically distant populations are possibly not conspecific.   
 Distribution. At present Boophis spinophis is only reliably known from its type locality, a forest patch just outside Ranomafana National Park, near the village of Ambatolahy (Appendix 10). A second locality, Ambohitantely, is supported by DNA barcoding analysis (low pairwise sequence divergence) of a juvenile specimen from this site, but the corresponding sequence had a substantial differentiation and the identity of this population requires further study (see section on molecular relationships above).
description: Published as part of Glaw, Frank, Köhler, Jörn, Riva, Ignacio De La, Vieites, David R. & Vences, Miguel, 2010, Integrative taxonomy of Malagasy treefrogs: combination of molecular genetics, bioacoustics and comparative morphology reveals twelve additional species of Boophis 2383, pp. 1-82 in Zootaxa 2383 (1) on pages 29-31, DOI: 10.11646/zootaxa.2383.1.1, http://zenodo.org/record/5310003