Zoosyst. Evol. 94 (2) 2018, 247-261 | DOI 10.3897/zse.94.21037 > PENSUFT. ifit=.. ‘2? NATURKUNDE BERLIN A distinctive new frog species (Anura, Mantellidae) supports the biogeographic linkage of two montane rainforest massifs in northern Madagascar Mark D. Scherz!*, Oliver Hawlitschek*, Jary H. Razafindraibe?, Steven Megson*, Fanomezana Mihaja Ratsoavina*, Andolalao Rakotoarison**, Molly C. Bletz*°, Frank Glaw', Miguel Vences* oO fF WN FP Zoologische Staatssammlung Miinchen (ZSM-SNSB), Miinchhausenstr. 21, 81247 Munich, Germany Zoologisches Institut, Technische Universitat Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo 101, Madagascar School of Science and the Environment, Manchester Metropolitan University, Manchester, M1 5GD, UK Department of Biology, University of Massachusetts Boston, 100 Morrissey Boulevard, Boston, MA 02125, USA http://zoobank. org/8A83 DE58-A2EE-494F-A03C-820DC836CDDF Corresponding author: Mark D. Scherz (mark.scherz@gmail.com) Received 16 September 2017 Accepted 26 February 2018 Published 15 March 2018 Academic editor: Johannes Penner Key Words Bioacoustics Biogeography Marojejy Montane Endemism Sorata Taxonomy Introduction Abstract We describe a new species of the genus Gephyromantis, subgenus Vatomantis (Mantelli- dae, Mantellinae), from moderately high elevation (1164-1394 m a.s.1.) on the Marojejy, Sorata, and Andravory Massifs in northern Madagascar. The new species, Gephyromantis (Vatomantis) lomorina sp. n. is highly distinct from all other species, and was immedi- ately recognisable as an undescribed taxon upon its discovery. It is characterised by a granular, mottled black and green skin, reddish eyes, paired subgular vocal sacs of partly white colour, bulbous femoral glands present only in males and consisting of three large granules, white ventral spotting, and a unique, amplitude-modulated advertisement call consisting of a series of 24—29 rapid, quiet notes at a dominant frequency of 5124—5512 Hz. Genetically the species is also strongly distinct from its congeners, with uncorrected pairwise distances >10 % in a fragment of the mitochondrial 16S rRNA gene to all other nominal Gephyromantis species. A molecular phylogeny based on 16S sequences places it in a clade with species of the subgenera Laurentomantis and Vatomantis, and we assign it to the latter subgenus based on its morphological resemblance to members of Vatoman- tis. We discuss the biogeography of reptiles and amphibians across the massifs of northern Madagascar, the evidence for a strong link between Marojeyy and Sorata, and the role of elevation in determining community sharing across this landscape. be confirmed whether or not they constitute new species (e.g. Vieites et al. 2012). Differing from this general pat- In recent decades, the number of frog species that have been discovered in Madagascar, while steadily increasing (Kohler et al. 2005), often included species that were not immediately recognizable as new to science, though with occasional exceptions, e.g. Boophis lichenoides (Vallan et al. 1998), Scaphiophryne boribory (Vences et al. 2003), and Tsingymantis antitra (Glaw et al. 2006). The major- ity of newly discovered taxa are assignable to existing complexes and must be investigated closely before it can tern, on a 2012 expedition to the Sorata massif in north- ern Madagascar, we discovered a small green frog of the genus Gephyromantis that was immediately recognisable as a new species. It was not given a candidate species number at the time, and no sequences of this species were included in the barcoding assessment of Perl et al. (2014). In a 2016 survey in Andravory, near Sorata, and a 2016 survey of Marojeyy National Park in northeastern Mada- gascar, we encountered the same species. Copyright Mark D. Scherz et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 248 At present, 44 species of Gephyromantis are rec- ognized and assigned to six subgenera (Asperomaniis, Duboimantis, Gephyromantis, Laurentomantis, Phylac- omantis, and Vatomantis) based on molecular and mor- phological criteria (Glaw and Vences 2006, Vences et al. 2017). This classification is largely in agreement with the molecular multi-gene phylogeny of Kaffenberger et al. (2012). However, this phylogenetic study revealed that the subgenera Laurentomantis and Vatomantis are closely related, and that Gephyromantis klemmeri Guibé, 1974, morphologically similar to other species of the subgenus Gephyromantis, is sister to the Laurentomantis clade, suggesting the need for an improved classification. We here provide a description of the new species, which has potential implications for the supraspecific taxonomy of Gephyromantis, and the biogeographical linkage of the rainforest massifs of northern Madagascar. Materials and methods Specimen collection and morphological measurement Specimens were collected at night using head torches along montane streams, euthanized using MS222 anaes- thesia and subsequent overdose, fixed in 96 % ethanol, and deposited in 75 % ethanol for long-term storage. Tissue samples were stored in 96 % ethanol. Field num- bers refer to the zoological collections of Miguel Venc- es (ZCMV), Frank Glaw (FGZC), and Steven Megson (SM). Specimens were deposited in the amphibian collec- tions of the Mention Zoologie et Biodiversité Animale, Université d’ Antananarivo (UADBA-A) and the Zoolo- gische Staatssammlung Munchen (ZSM). Morphological measurements were taken to the nearest 0.1 mm using a digital calliper. Measurement schemes followed generally previous work on the genus (e.g. Vences et al. 2017) with modifications to decrease the risk of damaging the fragile limbs of the specimens when ascertaining limb lengths: snout—vent length (SVL), maximum head width (HW), head length from posterior edge of tympanum to snout tip (HL), horizontal eye diam- eter (ED), horizontal tympanum diameter (TD), distance from eye to nostril (END), distance from nostril to snout tip (NSD), distance between nostrils (NND), upper arm length from the articulation of the arm with the trunk to the elbow (UAL), lower arm length from the elbow to the base of the hand (LAL), hand length from the base of the hand to the tip of the longest finger (HAL), fore- limb length (FORL*, given by the sum of UAL, LAL, and HAL), forearm length (FARL, given by the sum of LAL and HAL), thigh length from cloaca to knee (THIL), tibia length from knee to heel (TIBL), tarsus length from heel to base of foot (TARL), foot length from base of foot to tip of longest toe (FOL), hindlimb length (HIL*, given by the sum of THIL, TIBL, TARL and FOL), and length and width of femoral gland (FGL, FGW). Asterisks in this list indicate measurements that have the same abbreviation as the analogous single-measurement of previous studies zse.pensoft.net Scherz, M.D. et al.: New green Gephyromantis from Madagascar (e.g. Vences et al. 2017) but are cumulative here and there- fore not necessarily equivalent; comparison of such values must be done cautiously. Sequencing and analysis of DNA sequences DNA was extracted from tissue samples using a Qiagen DNeasy blood & tissue kit (Qiagen, Hilden, Germany), or standard salt extraction protocols. For two samples from Sorata and one sample from Marojejy (ZCMV 15269), we amplified a fragment of the mitochondrial 16S rRNA gene (hereafter 16S) in 25 ul polymerase chain reactions with the primers 16Sra-L and 16Sb-H (Palumbi et al. 1991), 1 ul of template DNA, and the following steps: initial de- naturation for 3 min at 94 °C, followed by denaturation with 35 cycles of 30 sec each at 94 °C, 30 sec of annealing at 55°C and 60 sec of elongation at 72 °C, and a final elon- gation step of 10 min at 72 °C. Sequencing was conduct- ed using the BigDye Terminator v1.1 Cycle Sequencing Kit on ABI 3730 and ABI 3130xI capillary sequencers. Newly determined sequences were deposited in GenBank (accession numbers MG926811—MG926823). For an ad- ditional nine specimens from Marojeyy, we sequenced a shorter, highly variable stretch of 250 bp of the same 16S region by an I/lumina amplicon approach (Vences et al. 2016) to confirm their identification (data not shown). For an exploratory analysis, we aligned the new se- quences with 16S sequences used by Kaffenberger et al. (2012) for all nominal species of Gephyromantis. Because the obtained tree (not shown) confirmed the new species to be related to the Laurentomantis/Vatomantis clade as also strongly suggested by morphology, we focused our analysis on this subgroup, 1.e., all nominal species of the subgenera Laurentomantis and Vatomantis, and G. klem- meri which is known to be related to these subgenera (Kaffenberger et al. 2012), as well as G. granulatus (sub- genus Duboimantis) as outgroup. We aligned sequences in MEGA 7 (Kumar et al. 2016), yielding an alignment of 532 positions of the sequenced stretch of the 16S rRNA gene. As only a few indels were found in this alignment, we did not exclude any positions for further analysis. We used the Bayesian Information Criterion in jModelTest 2.1.4 (Darriba et al. 2012) to determine a SYM+G substitution model as best-fitting our data. We implemented this model in MrBayes 3.2 (Ronquist et al. 2012) and computed a Bayesian infer- ence phylogenetic analysis, with two independent runs of 20 million generations, each comprising four Markov Chains (three heated and one cold), sampling every 1000 generations. Chain mixing and stationarity were assessed by examining the standard deviation of split frequencies and by plotting the -InL per generation using Tracer 1.5 software (Rambaut and Drummond 2007). Results were combined to obtain a 50 %-mayority rule consensus tree and the respective posterior probabilities of nodes, after discarding 25 % of the generations as burn-in (all compat- ible nodes with probabilities 50 % (bootstrap) or >90 % (posterior probabilities). Each specimen/species is followed by the corresponding GenBank accession number used in the alignment. Schematic drawings of femoral glands of all species in the subgenera Laurentomantis and Vatomantis as well as of G. klemmeri are shown to the right of the phylogeny, and coloured according to the subgenus to which they are assigned. by O. Hawlitschek, F. Glaw, A. Rakotoarison, F. M. Rat- soavina, T. Rajoafiarison, and A. Razafimanantsoa; ZSM 1545—1547/2012 (FGZC 3716, 3734, and 3664), adult males, and ZSM 1548/2012 (FGZC 3721), adult female, collected between 28 and 30 November 2012 from a creek below a bamboo forest on the Sorata massif (13.6772°S, 49.4413°E, 1394 m above sea level), Sava Region, north- eastern Madagascar, by O. Hawlitschek, F. Glaw, A. Ra- kotoarison, F. M. Ratsoavina, T. Rajoafiarison, and A. Razafimanantsoa; ZSM 318/2016 (SM AEA 063), adult female, and UADBA-A uncatalogued (SM AEA 062), un- sexed adult, collected between 18h45 and 18h50 on 30 May 2016 in Andravory (13.7385—13.7388°S, 49.53 10°E, 1164-1179 m a.s.l.), Sava Region, Antsiranana Province, northeastern Madagascar, by S. Megson, R. Walker, W.-Y. Crawley, and T. H. Rafeliarisoa (Figs 3-4). Diagnosis. A species assigned to the genus Gephyro- mantis on the basis of its granular skin, moderately en- zse.pensoft.net larged finger tips, small femoral glands consisting of a small number of large granules and present in males only (thus of type 2 as defined by Glaw et al. 2000), and bi- fid tongue. Within the genus Gephyromantis, assigned to the subgenus Vatomantis on the basis of its small size, connected lateral metatarsalia, absence of an outer meta- tarsal tubercle, paired subgular vocal sacs of partly whit- ish colour, greenish skin colouration, and riparian ecol- ogy. Gephyromantis lomorina sp. n. is characterized by the possession of the following suite of morphological characters: (1) granular skin, (2) reddish eyes, (3) mot- tled green and black skin, (4) males with paired subgular vocal sacs of partly white colour, (5) males with bulbous type 2 femoral glands consisting of a small number (2-3) of large granules, (6) white spots on the venter, (7) SVL 20.2—25.5 mm, and (8) fourth finger much longer than second. Furthermore, the species is characterised by dis- tinctive, 1681-1827 ms advertisement calls of relatively low intensity, consisting of 24—30 individual pulsed notes, Zoosyst. Evol. 94 (2) 2018, 247-261 25 Figure 2. The holotype of Gephyromantis lomorina sp. n., ZSM 419/2016 (ZCMV 15221) in life. (a) Dorsal; (b) ventral; and (c) dorsolateral view. Scale bars indicate 5 mm. with 2—4 pulses per note, an inter-note interval of 41—75 ms, and a dominant frequency of 5124-5555 Hz. DNA sequence data from the 16S gene fragment supports the high divergence of this taxon to all other Gephyromaniis, and is in agreement with its subgeneric assignment, albeit without statistical support (Fig. 1). Within the genus Gephyromantis, G. lomorina sp. n. can be distinguished from all subgenera except Laurentomantis and Vatomantis on the basis of the combination of femoral glands composed of few large granules (vs. composed of many, small granules; note that G. Alemmeri is here treated separately from all other subgenera, below, due to its unclear assignment), SVL < 26 mm (vs. > 27 mm in all other subgenera except Gephyromantis), absence of a white stripe along the upper lip (vs. general presence in subgenus Gephyromantis), and absence of distinctly enlarged supraocular spines (vs. presence in Asperomantis and some Duboimantis). It may be distinguished from all members of the subgenus Laurentomantis (G. ventrimaculatus (Angel), G. malagasius (Methuen & Hewitt), G. striatus (Vences, Glaw, Andreone, Jesu & Schimmenti), G. horridus (Boettger), and G. ranjomavo Glaw & Vences) by paired subgular vocal sacs (vs. single), absence of outer metatarsal tubercles (vs. presence), and at least partly greenish dorsal skin (vs. mostly yellowish to brown to reddish), and from several of these by the zse.pensoft.net 232 Scherz, M.D. et al.: New green Gephyromantis from Madagascar Table 1. Morphological data on specimens of Gephyromantis lomorina sp. n. Abbreviations: m = male, f = female, sa = subadult; for measurement abbreviations, see the Materials and methods. The holotype is bolded. Additive measurements (FARL, FORL, and HIL) are not explicitly shown but can be deduced from these data. m Catalogue (field number) ZSM 419/2016 (ZCMV 15221) /eND | NSD | ND] UAL LAL | HAL | THIL| TIBL| TARL| FOL | FGL | FGW 7.3 | 8.2 |13.4)13.9| 7.4 .L| 2.8 | 2.0 ZSM 421/2016 (ZCMV 15271) G0) 20a 12.2) hay 70 Gje22° | P18 ZSM 420/2016 (ZCMV 15222) B25 | Se | Ie Ss P1334) es |e: 22 | 1S ZSM 418/2016 (ZCMV 15220) UADBA-A 60294 (ZCMV 15270) UADBA-A 60298 (ZCMV 15273) UADBA-A 60296 (ZCMV 15223) UADBA-A 60297 (ZCMV 15272) 2.4) 1.4] 2.1 : 1.5 | 1.4 19/49] 6 7.7 | 84 |13.6} 14.8] 6.7 n/a oily | eZ UADBA-A 60295 (ZCMV 15219) 1.4 : é 8.1 ; | 7G 11.6) n/a UADBA-A 60299 (ZCMV 15247) 1.5 ; ; aes) A} 7.0 11.6) n/a ZSM 1549/2012 (FGZC 3714) ZSM 1545/2012 (FGZC 3716) ZSM 1546/2012 (FGZC 3734) ZSM 1547/2012 (FGZC 3664) ZSM 1548/2012 nee) ; 7 112.6 1.5 On 7 (MB. e279") 25h 13.4} 3.4 | 2.5 PSIG: | 6225.) e2Al PEP PPEEEEREREEE : ‘2 ZSM 318/2016 (SM AEA 063) Lee ores| VAD. | ie (2st) Gal ae0 13.6 |14:2| 7.2 n/a absence of tibial glands in males (vs. typical presence). notes in all species except G. ventrimaculatus); Within the subgenus Vatomantis, G. lomorina sp. 0. may be distinguished from all species by its more granular dorsal skin (vs. granular but not rough) and venter spotted with white (vs. generally without whitish spotting except on the chin and over the sternum); from G. rivicola (Vences, Glaw & Andreone) and G. webbi (Grandison) by its reddish iris colouration (vs. copper and greenish, respectively); from G. silvanus (Vences, Glaw & Andreone) by its smaller size (SVL 20.5—25.5 mm vs. 31 mm) and partly whitish vocal sacs (vs. yellowish); from G. webbi by femoral glands composed of few large granules (vs. composed of many, small granules) and large inner metatarsal tubercle (vs. small). Gephyromantis lomorina sp. n. may be distinguished from G. klemmeri by its roughly granular dorsal skin (vs. smooth to shagreened), greenish skin colour (vs. brownish), reddish iris (vs. gold), and strongly protruding inner metatarsal tubercle (vs. small and not protruding). The call of G. Jomorina sp. n. may be distinguished from all Vatomantis and Laurentomantis species in having notes that are clearly pulsed (vs. unpulsed zse.pensoft.net Gephyromantis ventrimaculatus has a higher number of pulses per note notes than G. /Jomorina sp. n. (ca. 6 pulses per note vs. 2-4 in G. Jomorina sp. n.). The call of G. lomorina sp. n. is somewhat similar to that of G. klemmeri, especially in having pulsed notes, but the call duration is much longer (1681-1827 ms vs. 626— 982 ms), the call has a more distinct amplitude decay (vs. complex amplitude modulation, see Vences et al. 1997), the notes of the call are more homogeneous (vs. distinct components of the call), and it lacks frequency modulation (vs. frequency modulated toward the end of the call). Description of the holotype. A specimen in a good state of preservation, a piece of tissue taken from the left thigh. SVL 23.3 mm; for other body measurements see Table 1. Body slender. Widest part of head marginally wider than widest part of body. Snout rounded in dorsal and lateral view, protruding slightly over upper jaw in lat- eral view. Nostrils not distinctly protruding, with lateral openings. Canthus rostralis distinct, concave. Loreal re- Figure 3. Morphological and chromatic variation among paratypes of Gephyromantis (Vatomantis) lomorina sp. n. from Marojejy in life. (a—b) ZSM 420/2016; (e-d) UADBA-A 60296; (e-f) UADBA-A 60295; and (g—h) ZSM 418/2016. Scale bars indicate 2 mm. gion concave, vertical. Tympanum distinct, fairly small, limbs and hindlimbs slender. Inner and outer metacarpal 53% of eye diameter. Supraocular spines absent. Weakly tubercle present, both indistinct. Finger discs enlarged, distinct supratympanic fold running from the eye over — round. Subarticular tubercles distinct, dark in colour. No the tympanum to above the insertion of the arm. Fore- webbing between fingers. Comparative finger lengths zse.pensoft.net 254 Scherz, M.D. et al.: New green Gephyromantis from Madagascar Figure 4. Photographs of Gephyromantis (Vatomantis) lomorina sp. n. and its habitat in Sorata. (a,d) ZSM 1545/2012; (bye) ZSM 1547/2012; and (¢,f) ZSM 1549/2012, not to scale; (g) habitat where several specimens were found in Sorata, showing (h,i) the appearance of the species in situ whilst calling at night. 1 <2 <4<3, fourth finger much longer than second fin- 1 <2<3=5 <4. Inner metatarsal tubercle rather large ger. Toe discs slightly enlarged, smaller than finger discs. (length about 1.3 mm), protruding strongly distally to Traces of webbing between toes. Comparative toe length resemble a toe. Outer metatarsal tubercle absent. Lateral zse.pensoft.net Zoosyst. Evol. 94 (2) 2018, 247-261 Frequency (kHz) mm & @ Ow Relative amplitude © 0 1000 Time (ms) 2000 Frequency (kHz 255 es 122) oO ep) Relative amplitude © 0 125 Time (ms) 250 Figure 5. Spectrogram (above) and waveform (below) of a call of the holotype of Gephyromantis (Vatomantis) lomorina sp. n., ZSM 419/2016, from Marojejy. (a) A full call (spectrogram shown using FFT of 512 points to visualise call structure); and (b) a 250 ms section from the middle of a call, showing the degree of pulsation of each note (spectrogram shown using FFT of 128 points to visual- ise note structure). metatarsalia connected. Dorsal skin granular, with nu- merous small tubercles arranged in mostly parallel lines running posteriorly over the dorsum, with convergent lines of tubercles on the posterior head, and weak rows of tubercles on the hindlimbs and forelimbs. Femoral glands round, consisting of three large granules with an inden- tation in their middle (similar to type 2 sensu Glaw et al. 2000). Vomerine teeth absent. Maxillary teeth present. Choanae small and lateral. Subgular vocal sacs whitish in distensible portion, blackish on the jaw, fairly small. Tongue bifid, free posteriorly. Colouration in life (Fig. 2) dorsally mottled with greens, browns, blacks, and yellows. Particularly green over the eyes. Raised ridges on the back were mostly yel- lowish, but some also with an orange hint. Flanks and lat- eral head as dorsum. Legs dark brown with yellow-green cross-bands, three on the thigh, three on the shank, and two on the tarsus. The tarsus and dorsal foot were a more ruddy brown than the rest of the body, mottled with a tan orange on the toes and on the heel. A few tubercles on the legs were red. A whitish annulus was present before the terminal disc of each toe and finger. The forelimbs were as the shanks and foot, ruddy brown mottled with yellow-green and dark brown, with a few red tubercles. Whitish spots were present in the inguinal region and the ventral portion of the flank, and also two cream stripes were present below the eye that continued on the bottom lip. The tympanum was distinctly brownish. The venter was umber in base colour with more reddish portions of translucent skin on the ventral side of the arms. The chin had white portions along the lip and especially on the vo- cal sacs, but the jaw itself was blackish. The venter had distinct white spots. The ventral hindlimbs were umber with irregular pale olive and yellow patches on the ven- tral thigh and shank. The ventral tarsus, foot, and hand were umber. The femoral glands were fleshy in colour, and the area ventral to the cloaca was pinkish. The iris was copper above and below, and rusty anteriorly and posteriorly, with blackish reticulations and a blackish line above and below the centre of the pupil. After six months in preservative, the colouration of the holotype has faded to become more uniformly brownish, and areas that were greenish in life have become cream. White areas of the venter are still immaculately white. Variation. All paratypes resemble the holotype in gross morphology; see Table 1 for morphological variation. Tympanum diameter ranges from 47—79 % of eye, without strong sexual dimorphism in tympanum size. Females are marginally but not significantly larger than males (‘-test, t = -].9215, df= 13, p = 0.07687). Several paratypes have smaller femoral glands than the holotype. Femoral glands are composed of 2 or 3 large granules (mean 2.875 + 0.35, n = 8; all but one of eight examined specimens with 3 granules). Females have miniscule raised bumps in the femoral area. There is considerable variation in coloura- tion of the specimens, with some individuals being much darker, and others being more green (Figs 3-4). The chin of females is more solidly dark than that of males, and they lack most white spots. A pair of cream stripes below the eye that continue on the lower lip is present in all specimens. Two specimens (UADBA-A 60299, and ZSM 1545/2012, Fig. 4) have a bright vertebral stripe. Bioacoustics. Call recordings were made in Marojejy from the holotype ZSM 419/2016 at its collection locali- ty at a distance of 0.5 m during light rain (Suppl. material 1, DOI: 10.7479/nmx8-aq7v). The call is interpreted as an advertisement call as it resembles the advertisement calls of the subgenus Laurentomantis, and was emitted without close proximity to other individuals, and while the frog was otherwise inactive (Kohler et al. 2017). Air temperature was not recorded. A strict FFT bandwidth filter was applied to the dataset to remove all sound be- low 400 Hz in order to remove wind artefacts. Two calls were recorded from the holotype, but numerous calls zse.pensoft.net 256 were heard whilst searching for this species along the river where it was found. Calls consisted of a rapid series of 24—29 extremely short notes (note duration 6.3 + 1.9 ms, range 2-10 ms, n = 53; Fig. 5a), each of which had 2.6 + 0.6 pulses (24 pulses, n= 50), the peak amplitudes of which were separated by 2.7 + 0.6 ms (1-4 ms, n= 53; Fig. 5b). Notes were separated by silent inter-note inter- vals of 64.6 + 5.5 ms (47—75 ms, n = 51). The call was amplitude modulated, increasing in amplitude quick- ly and slowly decaying toward the end of the call. Call duration was 1769-1827 ms (n = 2), with one inter-call interval recorded of 2399 ms. Generally, however, the calls appeared to be emitted rather irregularly. Dominant frequency was 5124-5512 Hz, and the 90 % bandwidth was from 2723-2759 to 6391-6462 Hz. Similar calls were recorded in Sorata from ZSM 1549/2012 at its collection locality (Suppl. material 2, DOI: 10.7479/nmx8-aq7v). Air temperature was not re- corded. The calls strongly resembled those recorded from the holotype. Three calls were recorded, but one was cut off and another had loud calls of Gephyromantis (Dubo- imantis) sp. in the background, so only one was analysed. The call consisted of a rapid series of 31 extremely short notes (note duration 6.9 + 0.8 ms, range 6—10 ms, n = 27 analysed), each of which had 2.0 + 0.2 pulses (2-3 puls- es, n = 27), the peak amplitudes of which were separated by 3.0 + 0.4 ms (2-4 ms, n = 27). Notes were separated by silent inter-note intervals of 46.3 + 3.8 ms (41—55 ms, n = 27). The call was amplitude modulated in the same way as that of ZSM 419/2016. Call duration was 1681 ms, and one inter-call interval was ca. 1900 ms. In gener- al however calling was irregular. The dominant frequen- cy was 5555 Hz, and the 90 % bandwidth was from 4979 to 6003 Hz. The call with a loud Gephyromantis (Dubo- imantis) sp. in the background was considerably shorter, and consisted of just 11 notes over a duration of 515 ms, but we suppose this call may have been disturbed as it lacked amplitude reduction toward its end. Distribution. The new species is known from three local- ities in northeastern Madagascar: (1) Marojejy National Park (type locality), (2) Sorata massif, and (3) Andravory massif (Fig. 6). All specimens were collected between 1164 and 1394 ma.s.l. Natural history. Specimens were collected near mountain streams in pristine montane riparian rainforest (Fig. 4g). In Marojeyy National Park they were encountered during and after light rain, sitting in inconspicuous locations, especial- ly on the fronds of tree ferns, but also on other low vege- tation, between a few centimetres and up to 2 m above the ground. Specimens in Sorata were found in similar posi- tions during dry weather, in the days just before the begin- ning of the rainy season. Males called irregularly and soft- ly (see the call description above). Population density in Marojeyy was remarkably high, with around three or four individuals being found along a 10 m stretch of stream. The observed density in Sorata was lower, possibly due zse.pensoft.net Scherz, M.D. et al.: New green Gephyromantis from Madagascar to the absence of rain during the observation period. The Species occurred in close sympatry with a number of other mantellids, but only few of these (especially Mantidactylus aff. femoralis) were found in the same microhabitat. Sev- eral specimens from Marojejy had pinkish mites (probably of the genus Endotrombicula, see Wohltmann et al. 2007) embedded within translucent whitish pustules on the skin of their fingers, toes, and bodies. Nothing is known about the reproduction of this species, but the calling sites sug- gest an association with lotic water. Available names. There are no other, earlier names cur- rently available (e.g., junior synonyms) that are assign- able to the subgenera Vatomantis or Laurentomantis and that could apply to the new species. Etymology. The specific epithet is the Malagasy word lomorina, meaning ‘covered in moss’, in reference to the green, mossy appearance of the species in life. It 1s used as an invariable noun in apposition to the genus name. Conservation. The species occurs in two regions with very different conservation situations: the highly protect- ed forests of Marojeyy National Park, and the unprotect- ed, isolated, and highly threatened forests of Sorata and Andravory. Mamunirina et al. (2008) report a study site in the rainforest of Sorata at 970 m a.s.l., but in our surveys in 2012, we detected larger patches of forest only at el- evations of ca. 1270 m and above. The new species was collected at lower elevation in Andravory (1164-1179 m a.s.l.), where forest persists. Higher elevation levels of Sorata are covered by high-elevation forests different to those where G. /omorina sp. n. was found, and these therefore may not support this species. In this area, the species 1s therefore directly threatened by the loss of the only forests in which it has been detected. By contrast in Marojeyy, forest extends down to rough- ly 200 maz.s.1., is highly protected, and the high elevation forest where this species occurs does not seem to be fac- ing any immediate threats. Although the tourist load to Marojejy is relatively high, and the area upslope from the collection locality of the holotype and several paratypes is somewhat polluted with refuse from the nearby tourist camp, the species was abundant around this stream during our survey there in 2016, and presumably inhabits other streams around the same elevation across the massif. Accommodating this spread of risk is a challenge for the IUCN Red List status. However, G. (V) lomorina sp. n. is not the first species to have almost exactly this distribution. Rhombophryne vaventy Scherz, Ruthensteiner, Vences & Glaw was recently recovered from Sorata (Peloso et al. 2016, Scherz et al. 2016, Lambert et al. 2017) after initially having been described from the same type locality as G. Jomorina sp. n. (Scherz et al. 2014). In the case of this species, Scherz et al. (2017a) argued for a classification of Endangered under IUCN criterion Blab(iii), 1.e. an extent of occurrence under 5000 km? (B1), known from fewer than five Zoosyst. Evol. 94 (2) 2018, 247-261 10 40 km 2o7 * ae Crs i) aes i) . h J Sarraka+ wey LEI ODE 3 a Figure 6. Distribution of Gephyromantis (Vatomantis) lomorina sp. n. in northern Madagascar. Areas with diagonal lines are official protected areas. The dotted outline indicates the proposed area with the scope of the WWF protection plan for this part of Madagas- car (Biodev Madagascar Consulting 2014, WWF Madagascar 2015). Three arc second SRTM data from Jarvis et al. (2008). threat-defined locations (a), and an observed, estimated, inferred, or projected decline (b) in the area, extent, and/ or quality of habitat (111). Given the similar situation in G. lomorina sp. n., 1.¢., very similar, limited distribution and ongoing reduction and threat to a substantial part of its habitat (1.e., the forests of Sorata and Andravory), we propose that the same threat status and justification be given for this species. Discussion Gephyromantis (Vatomantis) lomorina sp. n. 1s a distinc- tive species, mostly due to its granular, greenish skin, which is rougher than in all other members of the sub- genus Vatomantis, but not as rugose as in many species of the subgenus Laurentomantis. Indeed, it is 1n several aspects intermediate between these subgenera, having a zse.pensoft.net 258 call that sounds similar to both (Vences et al. 2006). Its phylogenetic position is at present basically unresolved between these two subgenera. However, its morphology is clearly more similar to Vatomantis than to Laurento- mantis, as it lacks an outer metatarsal tubercle (present in Laurentomantis), has a distinct brown tympanum (less distinct in Laurentomantis), lacks a broadened head (usu- ally distinctly broadened in Laurentomantis) and has paired subgular vocal sacs (single in Laurentomantis) (Glaw and Vences 2006). Gephyromantis (Vatomantis) rivicola, G. (V.) silvanus, G. (KV) lomorina sp. n., and most Laurentomantis species share a unique femoral gland morphology with glands be- ing composed of a small number of large, round granules (each granule representing a single gland within the fem- oral macrogland; Vences et al. 2007; Fig. 1). Glaw et al. (2000) interpreted these unusual glands as possible inter- mediate steps between Type 2 glands (sharply delimited groups of numerous granules of up to 0.9 mm diameter) toward Type 3 and 4 glands (a rounded structure composed of few, large granules and an external central depression). The position of G. (Vv) Jomorina sp. n. appears to make this situation more complicated; formerly, it seemed that granule size had increased and number decreased in G. (V.) rivicola and G. (V-) silvanus while G. (V.) webbi had re- tained Type 2 glands typical of most other Gephyromantis species (Glaw et al. 2000, Vences et al. 2007). However, given the split of G. (V.) lomorina from a more basal node in that clade (Fig. 1), and given the ubiquity of these un- usual glands in the sister subgenus Laurentomantis (Glaw et al. 2000, Kaffenberger et al. 2012), it seems that Type 2 femoral glands may have independently originated one or more times in this clade. A better resolved phylogeny of the clade will be necessary to better understand the evolu- tion of their femoral gland morphology. The apparently highly divergent G. (V.) Jomorina sp. n. sheds some light on questions regarding the relationships of G. klemmeri. Formerly, G. klemmeri was considered a member of the subgenus Gephyromantis, but Kaffen- berger et al. (2012) showed that it has affinities between Laurentomantis and Vatomantis. They forestalled action on transferring it to one of these subgenera until more data become available, as single genes disagreed as to its position. Gephyromantis klemmeri shares femoral gland morphology with both Laurentomantis and Vatomantis, having large glands with a small number of large gran- ules. This lends credence both to its phylogenetic position being close to these subgenera, and also to the hypothe- sis that smaller numbers of larger granules in the femoral glands may be ancestral in this clade. Kaffenberger et al. (2012) suggested three possible alternatives to dealing with the phylogenetic affinities of G. klemmeri: (a) including G. klemmeri in Laurentoman- tis (its position sister to Laurentomantis was supported with 94 % bootstrap support from maximum likelihood and >0.99 posterior probability, but was not supported in maximum parsimony analysis), (b) erecting a new mono- typic subgenus, or (c) redefining a more inclusive subge- zse.pensoft.net Scherz, M.D. et al.: New green Gephyromantis from Madagascar nus Laurentomantis that besides G. klemmeri would also include Vatomantis as a junior synonym (the clade con- taining Laurentomantis, Vatomantis, and G. klemmeri was supported with 100 % bootstrap support from maximum likelihood, >0.99 posterior probability, and 86 % bootstrap support from maximum parsimony). Determining the best course of taxonomic action will in part depend on the res- olution of the phylogenetic relationships of G. klemmeri and of G. (V.) lomorina sp. n., in the framework of a more comprehensive revision of Laurentomantis and Vatoman- tis, as these subgenera still contain further candidate spe- cles requiring in-depth analysis (Vieites et al. 2009). Gephyromantis (Vatomantis) lomorina sp. n. also sheds light on the biogeography of northern Madagascar, pro- viding yet more evidence for a strong link between Sorata and Marojejy. The environmental conditions of these two regions are similar (Brown et al. 2016), and various spe- cies originally described from one of the two areas have subsequently been discovered in the other, e.g. Rhombo- phryne vaventy (Peloso et al. 2016, Scherz et al. 2016, 2017a, Lambert et al. 2017), Gephyromantis (Asperoman- tis) tahotra (Glaw et al. 2011, Vences et al. 2017), and G. (D.) schilfi (Glaw and Vences 2000, Scherz et al. 2017b). These similarities are generally limited to species found above 1200 m, probably because forest below 1200 m in Sorata has been mostly eradicated. We predict that similarities between faunal composi- tions of the mountainous massifs of northern Madagas- car are limited by elevational connectivity. For instance, there is continued connectivity between regions of ele- vation up to 1400 m from Sorata to Marojeyy and indeed roughly to the Manongarivo massif as well. There is no connectivity above this elevation however; areas of over 1400 m across the different massifs are separated by low- er elevations, leading to island-like isolation of peak ar- eas. Therefore, we predict that species occurring above 1400 m will show a greater degree of microendemism, and those below this elevation will have a greater prob- ability of occurring more widely; the higher a species’ centre of elevational distribution is located, the greater its chance of being microendemic. No absolute threshold of turnover is expected, because major climate fluctuations in the past will likely have blurred elevational boundaries over time. So far, evidence appears to support this hypothesis; as already stated, several species from around 1300 m are shared between Marojeyy and Sorata (and Andravory, though at present only limited and generally unpublished data are available from this forest), and some species known from higher elevations are so far thought to be microendemic to either region, e.g. Rhombophryne lon- gicrus (Scherz et al. 2015), Gephyromantis (Duboiman- tis) tohatra (Scherz et al. 2017b), Calumma jejy, and C. peyrierasi. Assuming this hypothesis is correct, it raises questions about species that are microendemic at low- er elevations, but opportunities to study and understand these taxa are increasingly limited by the fact that forest at lower elevations is disappearing outside of protected Zoosyst. Evol. 94 (2) 2018, 247-261 areas. Conservation efforts must be redoubled to ensure that these study systems may remain long enough to be investigated and understood. Acknowledgements As always, we are grateful to the Malagasy authorities of the Ministry of Environments and Forests for providing us with permits. Field research was conducted under permit N° 215/16/MEEF/SG/DGE/DSAP/SCB.Re (dated 5 September 2016) and N° 265/12/MEF/SG/DGF/DCB.SAP/SCB (dat- ed 18 October 2012). Specimens were exported under per- mits N° O01ON-EA01/MG17 (dated 4 January 2017) and N° 163N-EA12/MG12 (dated 17 December 2012). This work was carried out in collaboration with the Mention Zoologie et Biodiversité Animale, Universite d’ Antananarivo, to whom we are also grateful for the loan of the paratype series. We are also grateful to R. Walker, W.-Y. Crawley, T. H. Rafeliarisoa, and the Andravory team for their help in Andravory, and A. Razafimanantsoa and T. Rajoafiarison for their help in Maro- jeyy and Sorata. AR and MCB were supported by fellowships of the Deutscher Akademischer Austauschdienst. MV and MDS were supported by grants of the Deutsche Forschungs- gemeinschaft (VE247/13-1 and 15-1). The fieldwork of OH, FG, AR, and FR was supported by the Mohamed bin Zayed Species Conservation Fund (project 11253064). The publica- tion of this article in Zoosystematics and Evolution was made possible by the Museum ftir Naturkunde Berlin. References Biodev Madagascar Consulting (2014) Document de référence sur la NAP complexe Ambohimirahavavy Marivorahona en vue de |’ eval- uation environnementale par le CTE. Unpublished report. Accessed 14 June 2017. Available from: https://goo.gl/4KgD9W [Archived by WebCite® at http://www. webcitation.org/6sa Y QDhEv] Brown JL, Sillero N, Glaw F, Bora P, Vieites DR, Vences M (2016) Spatial biodiversity patterns of Madagascar’s amphibians and reptiles. PLoS One 11(1): e0144076. https://doi.org/10.137 1/journal.pone.0144076 Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. 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Wohltmann A, du Preez L, R6del M-O, Kohler J, Vences M (2007) En- doparasitic mites of the genus Endotrombicula Ewing, 1931 (Acari: Prostigmata: Parasitengona: Trombiculidae) from African and Mad- agascan anurans, with description of a new species. 54: 225-235. https://doi.org/10.14411/fp.2007.031 WWE Madagascar (2015) Plan d’Aménagement et de gestion intégré du complexe d’aires protegées Ambohimirahavavy Marivorahona, Report Draft. Accessed 14 June 2017. Available from goo.gl/GGXP2D Zoosyst. Evol. 94 (2) 2018, 247-261 Supplementary material | Advertisement call of Gephyromantis lomorina sp. n. Authors: Mark D. Scherz, Oliver Hawlitschek, Jary H. Razafindraibe, Steven Megson, Fanomezana Mihaja Ratsoavina, Andolalao Rakotoarison, Molly C. Bletz, Frank Glaw, Miguel Vences Data type: WAV File (.wav) Explanation note: Call recording of Gephyromantis (Va- tomantis) lomorina sp. n. ZSM 419/2016 (ZCMV 15221). Calls recorded at 21h20 on 18 November 2016 near Camp Simpona (ca. 14.4366°S, ca. 49.7434°E, ca. 1325 ma.s.l.) in Marojeyy National Park, Sava Re- gion, Antsiranana Province, northeastern Madagascar, by M. D. Scherz. Frog was ca. 1 m above the ground on a fern near a small river, calling occasionally during light rain. Air temperature was not taken. Recording distance was 0.5 m. Animal Sound Archive: https:// doi.org/10.7479/nmx8-aq7V. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://do1.org/10.3897/zse.94.21037.suppl1 261 Supplementary material 2 Advertisement call of Gephyromantis lomorina sp. n. Authors: Mark D. Scherz, Oliver Hawlitschek, Jary H. Razafindraibe, Steven Megson, Fanomezana Mihaja Ratsoavina, Andolalao Rakotoarison, Molly C. Bletz, Frank Glaw, Miguel Vences Data type: WAV File (.wav) Explanation note: Call recording of Gephyromantis (Va- tomantis) lomorina sp. n. ZSM_ 1549/2012 (FGZC 3714). Calls recorded at night on 30 November 2012 on the Sorata massif (creek near campsite, 13.6829°S, 49.4403°E, 1325 m as.l.), Sava Region, Antsir- anana Province, northeastern Madagascar, by O. Hawlitschek. Ecological data not available. Air tem- perature and recording distance were not noted. An- imal Sound Archive: https://doi.org/10.7479/nmx8- aq7v. Copyright notice: This dataset is made available under the Open Database License (http://opendatacom- mons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Data- set while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://do1.org/10.3897/zse.94.21037.suppl2 zse.pensoft.net