Communicator whistles: A Trek through the taxonomy of the Boophis marojezensis complex reveals seven new, morphologically cryptic treefrogs from Madagascar (Amphibia: Anura: Mantellidae) Vences, Miguel Köhler, Jörn Hutter, Carl R. Preick, Michaela Petzold, Alice Rakotoarison, Andolalao Ratsoavina, Fanomezana M. Glaw, Frank Scherz, Mark D. Vertebrate Zoology 2024 2024-10-14 74 643 681 Vences & Köhler & Hutter & Preick & Petzold & Rakotoarison & Ratsoavina & Glaw & Scherz 2024 Amphibia Mantellidae Boophis Animalia Anura Chordata species archeri sp. nov.   Lineage E Figures 6, 13   Identity. This species has been newly discovered in this study and has not been included in any of the previous studies including  B. marojezensisand allied candidate species (e. g., Glaw et al. 2001; Glaw and Vences 2007; Vieites et al. 2009; Randrianiaina et al. 2012; Perl et al. 2014; Hutter et al. 2018).    Holotype.    ZSM12 / 2016( MSZC213), adult male, collected by M. D. Scherzand M. Rakotondratismaon  15 January 2016at Bevitagnono forest,  33.1 kmSW of Bealanana on the road RN 31(  14.7387 ° S, 48.5170 ° E,  1016 ma. s. l.), North Westof Madagascar.    Paratypes.    ZSM11 / 2016( MSZC198), adult male, collected by M. D. Scherzand M. Rakotondratsimaon  14 January 2016at Andranonafindra forest,  30 kmSW of Bealanana on the road RN 31(  14.7360 ° S, 48.5481 ° E,  1169 ma. s. l.);   ZSM13 / 2016( MSZC243), adult female, collected by M. D. Scherzand M. Rakotondratsimaon  17 January 2016at Andranonafindra forest,  30 kmSW of Bealanana on the road RN 31(  14.7360 ° S, 48.5489 ° E,  1138 ma. s. l.).   Definition. A small treefrog assigned to the genus  Boophis, subgenus Boophis, in the family Mantellidaebased 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 blommersaegroup based on small body size (male SVL25.0– 25.3 mm), predominantly brownish dorsal coloration, absence of red color on webbing or ventral side of limbs, males calling along streams, and molecular phylogenetic relationships. Within the  B. blommersaegroup, defined by absence of dorsolateral bands, absence of red color in outer iris area, and advertisement calls with dominant frequencies of 4130–4799 Hz, consisting of a series of 3–6 whistling notes of 158–308 ms duration and strong frequency modulation (frequency ascending and then descending in each note). Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16 SrRNA gene: MolD identified the following robust diagnostic nucleotide combination compared to all other species in the  B. marojezensiscomplex (sites given relative to the full-length 16 Ssequence of  Mantella baroni): “ D ” in the site 253, “ T ” in the site 277, “ A ” in the site 303.   Diagnosis. Within the  B. blommersaegroup, distinguished from  B. blommersaeby calls consisting of a series of whistles (vs. pulsed trills); and from  B. vittatusby calls consisting of a series of whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from  B. marojezensisby advertisement calls consisting of notes of rather similar duration (vs. clear distinction of short and long notes), and minimum note duration of 158 ms (vs. 15 ms); from  B. kirki  sp. nov.by advertisement calls consisting of 3–6 notes (vs. 9–19 notes) and longer note duration (158–308 ms vs. 54–105 ms), and absence of red color in outer iris area (vs. presence in some specimens); from  B. picardi  sp. nov.by advertisement calls consisting of notes of rather similar duration (vs. clear distinction of short and long notes), consisting of 3–6 notes (vs. 17–25 notes), minimum note duration 158 ms (vs. 19 ms), and absence of red color in outer iris area (vs. distinct in many specimens); from  B. pikei  sp. nov.by advertisement calls consisting of 3–6 notes (vs. 25–33 notes) of 158–308 ms note duration (vs. max. duration of 98 ms); from  B. siskoi  sp. nov.by advertisement calls consisting of 3–6 notes (vs. 7–12 notes), with each note strongly frequency-modulated with an initial ascent and final descent of frequency (vs. regularly ascending frequency); and from  B. janewayae  sp. nov.by advertisement calls with each note strongly frequency-modulated with an initial ascent and final descent of frequency (vs. regularly ascending frequency), and higher dominant frequency (4130–4799 vs. 2687–3404 Hz). For a distinction from other species of the  B. marojezensiscomplex described herein, see accounts of these new species below.   Description of the holotype. Adult male, in excellent state of preservation, SVL 25.3 mm, muscle tissue removed from left thigh for molecular analysis. Body moderately slender; head slightly wider than long, as wide as body; snout rounded in dorsal view, sloped to rounded in lateral view; nostrils directed laterally, nearer to eye than to tip of snout; canthus rostralis indistinct, slightly concave in dorsal view, loreal region slightly concave; tympanum very indistinct, estimated TDabout 43 % of ED; supratympanic fold poorly recognizable, slightly curved in its anterior and more straight in its posterior half; vomerine odontophores weakly developed, well-separated in two small rounded aggregations, positioned posteromedial to choanae; choanae medium-sized, rounded to ovoid; 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 (1.5); 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 anterior edge of eye 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 (0.25), 2 i (0.75), 2 e (0), 3 i (1.25), 3 e (0), 4 i (1.75), 4 e (1.75), 5 (0.25); 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 light reddish brown with a moderately contrasted but incomplete and somewhat discontinuous brown hourglass marking on anterior part of the dorsum, a broad brown transverse bar on the posterior part of the dorsum, and a narrow dark transverse bar between the eyes. Dorsum densely spotted with poorly contrasted small brown spots. Limbs light brown with darker brown crossbands: about 4 poorly marked crossbands on forearm, 5 on shank, 7–8 on thigh. Ventrally cream, white on belly and with dark pigment on ventral side of feet. In life (Fig. 13), dorsally cream and all dark dorsal elements poorly contrasted. Iris rather uniformly beige, iris periphery turquoise.   Figure 13. Individuals of  Boophis archeri  sp. nov.in life. A, Bmale holotype  ZSM12 / 2016( MSZC213) from Bevitagnono forest in dorsal and dorsolateral view. Cmale paratype  ZSM11 / 2016( MSZC198) from Andranonafindra forest. Dfemale paratype,  ZSM13 / 2016( MSZC243) from Andranonafindra forest.   Variation. The female   ZSM13 / 2016 inpreservative has a pattern of fine blackish spots across the dorsum, in addition to a moderately contrasted hourglass-patch on the anterior dorsum and second dark patch on the posterior dorsum. In life, the female had a more reddish brown dorsal color, and reddish brown color also in the iris; it contains a large number of mature oocytes, recognizable without dissection.   Etymology. Named after the fictional character Captain Jonathan Archer, first portrayed by Scott Bakula in Rick Berman and Brannon Braga’s Star Trek: Enterprise.   Tadpole. The tadpole of this species is unknown.   Natural history. An arboreal, nocturnal treefrog found in humid rainforests. Little is known of the ecology of the species. At the type locality, there was a remarkable density of these frogs in diminutive riparian forest fragments. Males frequently emitted calls whilst moving among the narrow twigs overhanging the stream.   Calls. Advertisement calls of  B. archeri  sp. nov., recorded at Bevitagnono and Andranonafindra forests from the holotypeand the male paratypeon 14–15 January 2016(air temperature not recorded), consist of multiple tonal notes, sounding like whistles. Within calls, inter-note intervals become longer from the beginning to the end of the call. All notes exhibit a distinct upward frequency modulation, with a frequency shift comprising approximately 400 Hz. The first note of each call is lower in relative amplitude when compared to subsequent notes. Amplitude modulation within notes is only slightly expressed (most distinct in first note), with increasing amplitude reaching its maximum at the second half of the note’s duration. Numerical parameters of nine analyzed calls of three different individuals (among them call vouchers MSZC198 and MSZC213) are as follows: call duration 889–1582 ms (1242.0 ± 231.8 ms); notes / call 3–6 (4.3 ± 1.0); note duration 158–308 ms (209.3 ± 33.6 ms); inter-note interval 77–162 ms (120.7 ± 28.8 ms); dominant frequency 4130–4799 Hz (4493 ± 191 Hz); harmonic frequency band present at around 9000 Hz; prevalent bandwidth 3400–5000 Hz.   Distribution. According to molecular data summarized herein, the species is known from (1) the type locality, Bevitagnono forest, and (2) Andranonafindra forest. The known elevational range of the species spans from 1016–1169 ma. s. l. 4952150323 2016-01-15 ZSM, MSZC M. D. Scherz & M. Rakotondratisma Madagascar 1016 -14.7387 Bevitagnono forest 7 48.517 North West ZSM 12 / 2016 1 1 holotype 4952150311 2016-01-14 ZSM M. D. Scherz & M. Rakotondratsima 1169 -14.736 Andranonafindra forest 7 48.5481 ZSM 11 / 2016 1 1 4952150328 2016-01-17 ZSM M. D. Scherz & M. Rakotondratsima 1138 -14.736 Andranonafindra forest 7 48.5489 ZSM 13 / 2016 1 1