Boophis pikei, Vences & Köhler & Hutter & Preick & Petzold & Rakotoarison & Ratsoavina & Glaw & Scherz, 2024

Vences, Miguel, Köhler, Jörn, Hutter, Carl R., Preick, Michaela, Petzold, Alice, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Glaw, Frank & Scherz, Mark D., 2024, Communicator whistles: A Trek through the taxonomy of the Boophis marojezensis complex reveals seven new, morphologically cryptic treefrogs from Madagascar (Amphibia: Anura: Mantellidae), Vertebrate Zoology 74, pp. 643-681 : 643-681

Boophis pikei sp. nov.

Lineage C Figures 6 , 10

Identity.

This species has been previously referred to as B. marojezensis [Ca 53 JQ 518216 ] = B. marojezensis [Ca 53] in Randrianiaina et al. (2012), and B. sp. Ca 53 in Perl et al. (2014) and Hutter et al. (2018). This lineage also includes those specimens referred to B. marojezensis sensu stricto by Randrianiaina et al. (2012). It was included in B. marojezensis sensu lato by Glaw and Vences (2007) and Vieites et al. (2009), and not explicitly included or mentioned by Glaw et al. (2001).

Holotype.

ZSM 393 / 2016 (ZCMV 15251), adult male, collected by M. D. Scherz, A. Rakotoarison, M. Bletz, M. Vences, and J. Razafindraibe on 18 November 2016 at Camp 3 “ Simpona ” , Marojejy National Park (14.4366 ° S, 49.7434 ° E, 1325 m a. s. l.), North East of Madagascar.

Paratypes.

ZSM 394 / 2016 (ZCMV 15259), adult male, same collection data as holotype . ZSM 1514 / 2012 (FGZC 3732), adult male, collected by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina, and A. Razafimanantsoa on 30 November 2012 in bamboo forest above a campsite on the Sorata massif (ca. 13.6752 ° S, 49.4410 ° E, ca. 1485 m a. s. l.) . ZSM 107 / 2005 (FGZC 2853), ZSM 108 / 2005 (FGZC 2857), ZSM 109 / 2005 (FGZC 2871), ZSM 110 / 2005 (FGZC 2874) four adult males, collected by F. Glaw, M. Vences and R. D. Randrianiaina on 16 February 2005 at Camp 3 “ Simpona ” , Marojejy National Park (14.4367 ° S, 49.7434 ° E, 1326 m a. s. l.) .

Additional material.

ZSM 1515 / 2012 (FGZC 3657), adult male (not DNA barcoded) collected by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina, and A. Razafimanantsoa on 28 November 2012 in Sorata (above campsite, ca. 13.6811 ° S, 49.4455 ° E, ca. 1398 m a. s. l.) .

Definition.

A small treefrog assigned to the genus Boophis , subgenus Boophis , in the family Mantellidae based on occurrence on Madagascar, presence of intercalary element between ultimate and penultimate phalanx of fingers and toes (verified by external examination), presence of webbing between fingers, presence of nuptial pads in males, and absence of femoral glands in males. Assigned to the Boophis blommersae group based on small body size (male SVL 21.4–25.0 mm), predominantly brownish dorsal coloration, absence of red color on webbing or ventral side of limbs, suctorial stream-dwelling tadpoles, and molecular phylogenetic relationships. Within the B. blommersae group, defined by absence of dorsolateral bands, absence of red color in outer iris area, and advertisement calls with high dominant frequencies of 5174–5507 Hz, consisting of fast series of 25–33 short whistling notes of 12–98 ms duration. Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16 S rRNA gene: MolD identified the following robust diagnostic nucleotide combination compared to all other species in the B. marojezensis complex (sites given relative to the full-length 16 S sequence of Mantella baroni ): “ C ” in the site 86, “ A ” in the site 108, “ G ” in the site 254.

Diagnosis.

Within the B. blommersae group, distinguished from B. blommersae by calls consisting of a fast series of short whistles (vs. pulsed trills); and from B. vittatus by calls consisting of a fast series of short whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from B. marojezensis by advertisement calls consisting of 25–33 notes (vs. 7–8 notes); from B. kirki sp. nov. by advertisement calls consisting of 25–33 notes (vs. 9–19 notes) and absence of red color in outer iris area (vs. presence in some specimens); and from B. picardi sp. nov. by advertisement calls consisting of 25–33 notes (vs. 17–25 notes) of maximum duration of 98 ms (vs. max. duration of 225 ms), and absence of red color in outer iris area (vs. distinct in many specimens). For a distinction from other species of the B. marojezensis complex described herein, see accounts of these new species below.

Description of the holotype.

Adult male, in excellent state of preservation, SVL 22.3 mm, muscle tissue removed from right thigh for molecular analysis. Body slender; head slightly wider than long, much wider than body; snout rounded in dorsal view and rounded to slightly sloped in lateral view; nostrils directed laterally, equidistant to eye and tip of snout; canthus rostralis indistinct, slightly concave in dorsal view, loreal region slightly concave; tympanum indistinct but recognizable, round, TD 50 % of ED; supratympanic fold poorly recognizable, mostly straight; vomerine odontophores weakly developed, well-separated in two very small rounded aggregations, positioned posteromedial to choanae; choanae medium-sized, rounded; maxillary teeth present. Tongue ovoid, posteriorly bifid, free. Arms slender, forearms of slightly larger diameter, subarticular tubercles single, round; metacarpal tubercles not recognizable; fingers weakly webbed and with lateral dermal fringes; webbing formula 1 (traces), 2 i (traces), 2 e (traces), 3 i (traces), 3 e (2), 4 (1.25); relative length of fingers 1 <2 <4 <3 (finger 2 distinctly shorter than finger 4); finger discs enlarged, rounded; nuptial pads recognizable as unpigmented swelling on first finger. Hindlimbs slender; tibiotarsal articulation reaching beyond tip of snout 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 (1), 2 i (1.5), 2 e (0.5), 3 i (1.75), 3 e (0.5), 4 i (1.75), 4 e (1.75), 5 (1); relative length of toes 1 <2 <3 ≤ 5 <4; toe discs enlarged, rounded. Skin smooth on dorsal surfaces, throat, chest, and ventral surface of thighs, finely granular on belly; cloacal region surrounded by an area of distinct, large, white-colored granules.

In preservative, 7 years after collection (Fig. 6 ), dorsally brown with a distinct and moderately contrasted dark brown hourglass marking on anterior part of the dorsum, and a dark brown chevron-like transverse marking on the posterior part of the dorsum. An indistinct narrow dark transverse bar is visible between the eyes. Very few poorly contrasted light spots are scattered across the dorsum. Limbs light brown with distinct darker brown crossbands: about 3 on forearm, 3–5 on shank, 5 on thigh. Ventrally cream, white on belly, with dark pigment on ventral side of feet. In life (Fig. 10 ), similar but overall lighter and dorsal pattern less contrasted. Iris yellowish to beige, iris periphery turquoise.

Variation.

A dark dorsal hourglass-marking in preservative is apparent in most paratypes and very extended, merging into a blackish patch covering most of the head in ZSM 109 / 2005 . The female ZSM 107 / 2005 has only weakly contrasted traces of the hourglass-marking and is rather uniformly colored, with scattered light spots across the dorsum. ZSM 1514 / 2012 has a rather uniformly grayish dorsum and light flanks. In life, the dark dorsal markings are often only poorly contrasted or even absent (Fig. 10 ). Ventrally, in one photographed specimen, the belly is white, flanks unpigmented, and only traces of orange color are visible on hindlimbs (Fig. 10 E ). A sharply contrasted patch on the posterior-most dorsum is present in ZSM 1514 / 2012 . The iris coloration can be rather uniform beige (e. g., ZSM 1514 / 2012 ) or include a light orange outer and beige inner iris area ( ZSM 394 / 2016 ) (Fig. 10 ).

Etymology.

Named after the fictional character Captain Christopher Pike, first portrayed by Sean Kenney and Jeffrey Hunter in Gene Roddenberry’s Star Trek (The Original Series), and later portrayed by Anson Mount in Akiva Goldsman, Alex Kurtzman, and Jenny Lumet’s Star Trek: Strange New Worlds.

Tadpole.

Tadpoles of this species were described and illustrated under the names B marojezensis and B. marojezensis [Ca 53] by Randrianiaina et al. (2012), based on the DNA barcoded specimens ZSM 1528 / 2007 (FGZC 2277; GenBank accession number JQ 518196 ) from Marojejy National Park, and ZSM 573 / 2010 (ZCMV 13200; GenBank accession number JQ 518216 ) from Tsaratanana, respectively. As typical for all tadpoles of the group, the larvae belong to the “ suctorial ” ecomorphological guild. They have a large oral disk used to adhere to stones in fast-flowing water, a labial tooth row formula of 7 (5-7) / 3, and large numbers of oral papillae (243–290 marginal and 452–660 submarginal; without dorsal gap). They are characterized by presence of a lateral transparent area of the integument.

Natural History.

An arboreal, nocturnal treefrog found in humid rainforests along fast flowing streams. Little is known of the ecology of the species. In November 2016, a dense aggregation of calling males was found at night calling on low leaves and branches (ca. 1.2 m above the ground) between two narrow streams. At close range, the call is extremely loud.

Calls.

Advertisement calls of B. pikei sp. nov. recorded at Camp Simpona, Marojejy, on 18 November 2016, 21: 35 h (air temperature not recorded) consist of a high-pitched trill-like call composed of multiple very short tonal notes repeated at rather regular intervals and rapid succession. Within calls, usually the last 2–3 notes are slightly longer in duration when compared to leading notes (48–98 vs. 12–33 ms), and amplitude is modulated among notes, with call energy steadily increasing from the first to the last note of the call. Each note exhibits an upward frequency modulation, which is only very slightly expressed in the short notes, but more distinct in the last 2–3 notes of the call, comprising a shift in frequency ~ 100 Hz at maximum from beginning to the end of the note. Frequency modulation across the entire call may show some slight downward shift (Fig. 5 ). Numerical parameters of four analyzed calls from different individuals are as follows: call duration 921–1218 ms (1134.3 ± 142.6 ms); notes / call 25–33 (29.0 ± 3.4); note duration 12–98 ms (29.1 ± 22.6 ms); inter-note interval 17–31 ms (21.7 ± 4.1 ms); note repetition rate within calls varies between ca. 26–30 notes / second; dominant frequency 5174–5507 Hz (5367 ± 144 Hz); prevalent bandwidth 4800–5700 Hz.

Distribution.

According to molecular data summarized herein, the species is known from (1) the type locality, higher elevations at the Marojejy Massif (around Camp Simpona), and (2) the Sorata Massif. Based on tadpoles, it also has been recorded (3) on the Tsaratanana Massif at a site called Antevialambazaha (14.1743 ° S, 48.9452 ° E, 1699 m a. s. l.), by Randrianiaina et al. (2012). The elevational range of the species spans between 1325–1699 m a. s. l.

Figures

Figure 5. Comparative spectrograms and oscillograms illustrating representative calls of various species in the Boophis marojezensis complex, all at 4000 ms time scale.

Figure 6. Preserved holotypes in dorsal and ventral views of the eight nominal species in the Boophis marojezensis complex described in this study.

Figure 10. Individuals of Boophis pikei sp. nov. in life. A male paratype ZSM 394 / 2016 (ZCMV 15259) from Marojejy (Camp Simpona). B male holotype ZSM 393 / 2016 (ZCMV 15251) from Marojejy (Camp Simpona). C male from Marojejy (Camp Simpona), photo not assignable to a specific voucher specimen, photographed in 2016. D, E male paratype ZSM 1514 / 2012 (FGZC 3732) from Sorata.