This species was discovered and scientifically named by Glaw and Vences (1994) based on a holotype specimen collected at low elevations in the Marojejy Massif. No vouchered call recording referring to the name-bearing type (holotype) had been recorded, and no DNA sequence was so far available from this specimen. Calls heard from near the type locality consisted of short and long notes whereas populations of the B. marojezensis complex from higher elevations in Marojejy emitted calls with a greater number of very short notes only. We here provide a 16S sequence of the type obtained via archival DNA extraction and Illumina shotgun sequences which confirms that the name B. marojezensis is to be assigned to the lineage found in Marojejy at low elevation as well as in a few other sites in northern Madagascar (lineage F), characterized by advertisement calls containing two different note types (long and short).

Figure 6. 

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

Vieites et al. (2009) and Perl et al. (2014) used the specimen FGZC 2857 as their reference individual for B. marojezensis. That individual belongs to a different lineage, the one we refer to here as lineage C, which was referred to as B. sp. Ca53 by Randrianiaina et al. (2012) based on specimens from Tsaratanana Strict Nature Reserve. A second specimen, ZSM 326/2000 from Vohidrazana, included by Vieites et al. (2009) as a deep conspecific lineage of B. marojezensis, is shown by our trees to belong to lineage D. The true B. marojezensis was referred to as B. sp. 25 by Vieites et al. (2009) and Randrianiaina et al. (2012), herein corresponding to lineage F.

Figure 7. 

Individuals of Boophis marojezensis in life. A Male holotype (ZFMK 57401) from Marojejy (low elevation). B Male paratype (ZSM 567/1999, previously ZFMK 57402). C Individual from Masoala probably assignable to this species (not sequenced). D Male from Masoala (ZSM 250/2016, FGZC 5439).

ZFMK 57401, by original designation. Type locality: “the Marojezy massif at low altitude, NE-Madagascar”. A partial 16S sequence of the holotype is available from GenBank under accession number PQ278105.

One paratype: ZSM 567/1999 (previously ZFMK 57402), adult male, with same collection data as holotype.

In addition to the type material, we examined ZSM 208/2022 (FGZC 6510), probably an adult male, collected on 24 March 2022 by J.M. Rafanoharana, H. Raherinjatovo and F. Glaw at Analanjirofo (near Simpona Lodge), Makira Reserve (15.19917°S, 49.62083°E, 410 m a.s.l.); and ZSM 250/2016 (FGZC 5439), adult male, collected on 12 August 2016 by F. Glaw, D. Prötzel, J. Forster, K. Glaw, and T. Glaw at Masoala, around the “Eco-Lodge chez Arol” (ca. 15.7122°S, 49.9640°E, ca. 21 m a.s.l.).

A small treefrog assigned to the genus Boophis, subgenus Boophis, in the family Mantellidae based on its 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 20.0–25.7 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 the outer iris area, and advertisement calls at a dominant frequency of 4118–4441 Hz, consisting of 7–8 notes of different length (short notes 15–51 ms; long whistling notes 142–259 ms). Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16S 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 16S sequence of Mantella baroni): “C” in the site 107, “G” in the site 162, “C” in the site 251.

Within the B. blommersae group, distinguished from B. blommersae by calls containing multiple frequency-modulated whistles (vs. pulsed trills), and from B. vittatus by calls containing multiple frequency-modulated whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). For a distinction from other species of the B. marojezensis complex described herein, see accounts of these new species below.

The tadpole of B. marojezensis (under the name B. marojezensis [Ca25]) was described and illustrated by Randrianiaina et al. (2012), based on the DNA barcoded specimen ZSM 1611/2007 (FGZC 2929; GenBank accession number FJ559146). 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 (222 marginal and 315 submarginal; without dorsal gap). They are characterized by a pattern of several rounded patches formed by condensation of spots on the posterior half of the tail musculature.

An arboreal, nocturnal treefrog found in humid rainforests along fast flowing streams. Little is known of the ecology of the species. Many calling individuals of this species are sitting too high in trees to reach, or calling from perches where they are difficult to see and their calls are also often difficult to localize. Glaw et al. (2001) also reported calling activity in March, suggesting continuous reproductive activity throughout the rainy season.

Advertisement calls of Boophis marojezensis recorded at low elevation near a site known as “Camp 0”, Marojejy National Park, on 26 November 2016 (22.8°C air temperature) consist of two different note types, namely calls starting with a short series of short, fast repeated notes, followed by three distinctly longer notes, which are separated by longer intervals. All notes are tonal in character, with long notes exhibiting a distinct upward frequency modulation, with a frequency shift comprising approximately 300 Hz. Amplitude across the entire call is slightly increasing, with the first short notes being relatively soft. Long notes have the maximum call energy in the middle of the note, but amplitude modulation within notes is somewhat irregular. Numerical parameters of two analyzed calls of different individuals are as follows: call duration 1129–1222 ms; notes/call 7–8; short note duration 30–51 ms (34.7 ± 7.5 ms); long note duration 147–259 ms (199.7 ± 44.7 ms); inter-note interval 19–179 ms (62.5 ± 55.9 ms); dominant frequency 4118–4347 Hz (4258 ± 82 Hz); prevalent bandwidth 3800–4500 Hz.

These calls are in general agreement with those recorded at Marojejy on 20 March 1994, which, however, differ slightly in shorter inter-note intervals. Numerical parameters of two analyzed calls of the 1994 recording are as follows: call duration 764–1030 ms; notes/call 7–8; short note duration 15–49 ms (27.8 ± 10.6 ms); long note duration 142–231 ms (176.5 ± 38.1 ms); inter-note interval 22–61 ms (35.8 ± 14.1 ms); dominant frequency 4289–4441 Hz (4369 ± 64 Hz); prevalent bandwidth 4000–4700 Hz.

According to the molecular data summarized herein, the species is known from (1) the type locality, the Marojejy Massif at low elevation (close to “Camp Mantella”), (2) a second site at Marojejy (sample THC 302, sequence downloaded from GenBank; collected at 14.4467°S, 49.8251°E, 225 m a.s.l. by T.R. Fulgence), (3) the Masoala Peninsula near the Eco-Lodge “Chez Arol”, and (4) the Makira Reserve, at Analanjirofo near Simpona Lodge. Boophis marojezensis is a low-elevation species, known from 21–410 m a.s.l.

This species has previously been referred to as B. marojezensis [Ca51 JQ518198] = B. marojezensis [Ca51] by Randrianiaina et al. (2012) and B. sp. Ca51 in Hutter et al. (2018). It was included in B. marojezensis sensu lato by Glaw et al. (2001) and Glaw and Vences (2007) and not explicitly included or mentioned in the studies of Vieites et al. (2009) and Perl et al. (2014).

Figure 8. 

Individuals of Boophis kirki sp. nov. in life. A, D Male paratype KU 336967 (CRH 20) from Ranomafana in dorsolateral and ventral view. B, E Male paratype UADBA-CRH 15 (CRH 15) from Ranomafana in dorsolateral and ventral view. C Male paratype KU 336874 (CRH 17) from Ranomafana. F Male paratype KU 336875 (CRH 18) from Ranomafana. G Female (no type status) UADBA-CRH 430 (CRH 430) from Vohidrazana. H Male (no type status) KU 340727 (CRH 449) from Vohidrazana. I, J Male (no type status) KU 340730 (CRH 455) from Vohidrazana in dorsolateral and ventral view. Note the variation in iris color in both populations (Ranomafana and Vohidrazana), with some individual having red color in the outer iris area while others only have light orange color.

ZSM 699/2003 (FGMV 2002.331), adult male, collected by F. Glaw, M. Puente, L. Raharivololoniaina, M. Thomas, and D.R. Vieites on 20 January 2003 at Kidonavo bridge, near Vohiparara, Ranomafana National Park (21.2167°S, 47.3667°E), ca. 1000 m a.s.l., Southern Central East of Madagascar.

ZSM 700/2003 (FGMV 2002.332), adult male, with same collection data as holotype. KU 336874 (CRH 17), adult male collected by C.R. Hutter, Z.F. Andriampenomanana, and S. Justin on 13 January 2014 at Valohoaka, Ranomafana National Park (21.2987°S, 47.4385°E, 1085 m a.s.l.). KU 336875 (CRH 18) adult male collected by C.R. Hutter, Z.F. Andriampenomanana, and S. Justin on 13 January 2014 at Valohoaka, Ranomafana National Park (21.2987°S, 47.4386°E, 1086 m a.s.l.). UADBA-CRH 15 (CRH 15), adult male collected by C.R. Hutter, Z.F. Andriampenomanana, and S. Justin on 13 January 2014 at Valohoaka, Ranomafana National Park (21.2973°S, 47.4389°E, 1067 m a.s.l.). KU 336876 (CRH 19), adult male, collected by C.R. Hutter, Z.F. Andriampenomanana, and S. Justin on 14 January 2014 at Valohoaka, Ranomafana National Park (21.2975°S, 47.4390°E, 1065 m a.s.l.). KU 336967 (CRH 20), adult male, collected by C.R. Hutter, Z.F. Andriampenomanana, and S. Justin on 14 January 2014 at Valohoaka, Ranomafana National Park (21.2975°S, 47.4390°E, 1065 m a.s.l.). KU 336877 (CRH 105), adult male, collected by C.R. Hutter, Z.F. Andriampenomanana, E. Rajery, and S. Justin on 28 January 2014 at Maharira, Ranomafana National Park (21.3399°S, 47.4108°E, 1272 m a.s.l.). KU 336880 (CRH 185), adult male, collected by C.R. Hutter, Z.F. Andriampenomanana, and S. Justin on 26 December 2013 at Valohoaka, Ranomafana National Park (21.2978°S, 47.4389°E, 1066 m a.s.l.). MRSN-A 2245 (FAZC 11467), adult, collected by F. Andreone between 30 January and 3 February 2003 at Farihimazava (= Farimazava) Forest, near Antoetra (ca. 20.8350°S, 47.3325°E, 1380–1420 m a.s.l.).

The following specimens from Vohidrazana are not included in the paratype series due to their relatively high genetic divergence compared to specimens from the type locality: UADBA-CRH 430 (CRH 430), adult female, collected by C.R. Hutter, S.M. Lambert, and Z.F. Andriampenomanana on 3 January 2015 at Vohidrazana (18.9861°S, 48.5015°E, 1164 m a.s.l.). KU 340727 (CRH 449) and KU 340730 (CRH 455), two adult males, collected by C.R. Hutter, S.M. Lambert, and Z.F. Andriampenomanana on 4 January 2015 at Vohidrazana (18.9794°S, 48.5181°E, 1105 m a.s.l.).

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 20.0–23.4 mm), predominantly brownish dorsal coloration, suctorial stream-dwelling tadpoles, and molecular phylogenetic relationships. Within the B. blommersae group, defined by absence of dorsolateral bands, presence of red color in the outer iris area, especially at the dorsal and ventral edges, in many specimens, and advertisement calls at 3499–5604 Hz, consisting of a series of 9–19 whistling notes of successively increasing durations of 54–105 ms. Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16S 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 16S sequence of Mantella baroni): “A” in the site 17, “C” in the site 50, “T” in the site 159, “A” in the site 179.

Within the B. blommersae group, distinguished from B. blommersae by calls consisting of frequency-modulated whistles (vs. pulsed trills); and from B. vittatus by calls consisting of frequency-modulated whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from B. marojezensis by presence of red color in outer iris area in many specimens (vs. absence), and calls consisting of 9‒19 whistling notes of successively increasing durations of 54–105 ms (vs. 7–8 notes, with short and long notes distinguishable). For a distinction from other species of the B. marojezensis complex described herein, see accounts of these new species below.

Adult male, in good state of preservation, SVL 23.4 mm, tongue and right forelimb removed as tissue samples for molecular analysis. Body slender; head wider than long, wider than body; snout rounded in dorsal view, truncate in lateral view; nostrils directed laterally, nearer to tip of snout than to eye; canthus rostralis indistinct, slightly concave in dorsal view, loreal region slightly concave; tympanum indistinct but recognizable, round, TD 50% of ED; supratympanic fold not recognizable; vomerine odontophores weakly developed, well-separated in two very small rounded aggregations, positioned posteromedial to choanae; choanae medium-sized, rounded; maxillary teeth present. Tongue removed. Arms slender, subarticular tubercles single, round; metacarpal tubercles not recognizable; fingers weakly webbed and with lateral dermal fringes; webbing formula 1(traces), 2i(traces), 2e(traces), 3i(traces), 3e(1.75), 4(1); 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 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(0.5), 2i(1), 2e(0), 3i(1), 3e(0.25), 4i(1.5), 4e(1.5), 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, 20 years after collection (Fig. 6), dorsally reddish brown with a distinct and strongly contrasted dark brown hourglass-shaped marking on anterior part of the dorsum, and a dark brown transverse bar, somewhat chevron-shaped, on the posterior part of the dorsum. In addition, a few small dark brown and cream spots are scattered across the dorsum. A distinct light beige vertebral stripe runs from snout to cloaca and interrupts the dark markings. Limbs light brown with darker brown crossbands: 4–5 on forearm, 6–7 on shank, 6 on thigh. Ventrally cream, white on belly, with some dark pigment only on ventral side of feet. Color of holotype in life not recorded.

The paratype ZSM 700/2003 has a brown dorsal ground color with an irregular pattern of strongly contrasted small light markings, and neither vertebral stripe nor hourglass-patch. For variation of color in life in other paratypes, see Figure 8. A vertebral stripe as in the holotype is also observed in KU 336875, but not in the remaining specimens. Some individuals show a fine dotting with dark brown spots dorsally (e.g., KU 336967 and KU 336875, others have a dense spotting of small yellowish spots, mostly dorsolaterally (e.g., paratypes KU 336874 and KU 340730). Some specimens in life had relatively distinct red color ventrally on hindlimbs (Fig. 8D, E) while this was not apparent in others (e.g., Fig. 8J). The outer iris area was reddish in some specimens (KU 336874 from Ranomafana; KU 340730 from Vohidrazana) but only light orange in others (e.g., KU 336967). One female collected in January had light-colored oocytes visible through the ventral skin (UADBA-CRH 430; Fig. 8G). Iris periphery turquoise.

Named after the fictional character Captain James T. Kirk, first portrayed by William Shatner in Gene Roddenberry’s Star Trek (The Original Series), and also portrayed by Chris Pine in J.J. Abrams’ Star Trek films.

The tadpole of this species (under the name B. marojezensis [Ca51]) was described and illustrated by Randrianiaina et al. (2012), based on the DNA barcoded specimen ZSM 267/2008 (ZCMV 3691; GenBank accession number JQ518198) from Ranomafana. 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 (297 marginal and 309 submarginal; without dorsal gap). They are characterized by absence of a lateral transparent area of the integument, tail muscle covered by reticulations mainly on the anterior half, and eyes situated between the anterior 3/10 and 4/10 of the body.

An arboreal, nocturnal treefrog found in humid rainforests along fast flowing streams. Little is known of the ecology of the species. At Valohoaka in December 2013 and January 2014, male specimens were found calling sitting on leaves or branches, 1‒2 m above the ground along streams. Additionally, calling males generally were perched 1 meter to occasionally several meters above the ground, at places that often could not be reached for observation.

Advertisement calls of B. kirki sp. nov. recorded at Ranomafana on 1 March 1996 (ca. 22–23°C air temperature) consist of multiple tonal notes of medium duration, repeated at regular intervals. In each call, the first note is always lower in dominant frequency (4640–4895 Hz) compared to subsequent notes (5528–5668 Hz). Moreover, the first 3–5 notes of each call have a lower relative amplitude than subsequent notes. Each note exhibits a distinct upward frequency modulation, encompassing a shift in dominant frequency range of about 700 Hz from the beginning to the end of the note. Within calls, notes tend to become slightly longer in duration from the beginning to the end of the call. Call energy is distributed in a narrow frequency band. Numerical parameters of four analyzed calls are as follows: call duration 2036–2355 ms (2170.0 ± 165.8 ms); notes/call 16–18 (16.8 ± 0.9); note duration 60–105 ms (75.7 ± 10.0 ms); inter-note interval 45–65 ms (54.0 ± 5.2 ms); note repetition rate within calls varies between ca. 6.5–8.2 notes/second; dominant frequency 5538–5604 Hz (5567 ± 34 Hz); prevalent bandwidth 4600–6100 Hz.

Calls recorded at Valohoaka, Ranomafana, on 12–13 January 2014 (air temperature not recorded) and corresponding to the voucher specimens KU 336874–336876 agree with the calls described above in overall character, although some shorter calls are evident in the recordings. Numerical parameters of seven analyzed calls of three individuals are as follows: call duration 1180–2630 ms (1944.7 ± 564.5 ms); notes/call 10–19 (14.5 ± 3.2); note duration 54–104 ms (74.4 ± 15.9 ms); inter-note interval 33–71 ms (59.8 ± 9.9 ms); dominant frequency 4981–5604 Hz (5371 ± 205 Hz); prevalent bandwidth 4000–5900 Hz.

Calls recorded at Vohidrazana, Andasibe region, on 4 January 2015 (air temperature 19°C) and corresponding to voucher specimen CRH 455 also agree in character with those from the Ranomafana region described above, but are lower in dominant frequency. Numerical parameters of four analyzed calls are as follows: call duration 1370–1537 ms (1427.3 ± 85.5 ms); notes/call 9–12 (10.0 ± 1.4); note duration 59–105 ms (86.5 ± 13.5 ms); inter-note interval 60–104 ms (72.8 ± 16.3 ms); dominant frequency 3499–4242 Hz (3998 ± 300 Hz); prevalent bandwidth 3300–4500 Hz.

According to the molecular data summarized herein, the species is known from (1) the type locality, Vohiparara, and many other sites in Ranomafana National Park (Ambatolahy, Ambatovory, Fompohonina, Imaloka, Maharira, Mariavaratra, Ranomena, Sahateza, Samahalaotra, Valohoaka, Vatoharanana, Vohimanara), (2) from Antoetra (Farihimazava), based on barcoded specimen MRSN A2245 (Andreone et al. 2007); and (3) from Vohidrazana in the Andasibe region. The elevational range spans from mid elevations (e.g., 915 m a.s.l. at Ambatolahy) to relatively high elevations (1272 m a.s.l. at Maharira, up to 1420 m a.s.l. at Antoetra).

This species has previously been referred to as B. sp. Ca68 in Hutter et al. (2018). It was included in B. marojezensis sensu lato by Glaw et al. (2001), Glaw and Vences (2007), Vieites et al. (2009), and Rosa et al. (2012), and not explicitly included or mentioned in the studies of Randrianiaina et al. (2012), and Perl et al. (2014). Adult specimens from Mandraka considered to represent B. majori by Blommers-Schlösser (1979b) are probably to be referred to this species.

ZSM 264/2006 (ZCMV 1447), adult male, collected by D.R. Vieites, M. Vences, F. Rabemananjara, P. Bora, C. Weldon, and J. Patton on 7–8 February 2006 at An’Ala (18.9193°S, 48.4880°E, 889 m a.s.l.), Northern Central East of Madagascar.

ZSM 262/2006 (ZCMV 1436), ZSM 263/2006 (ZCMV 1437), ZSM 265/2006 (ZCMV 2364), three adult males with same collection data as holotype. ZSM 266/2006 (ZCMV 2403), ZSM 1969/2006 (ZCMV 1494), ZSM 1970/2006 (ZCMV 2417), three adult males with same collecting locality and collectors as holotype, but collected between 7–10 February 2006. ZSM 326/2000, adult male, collected by F. Glaw on 10 April 2000 at Vohidrazana (18.9658°S, 48.5103°E, 731 m a.s.l.). ZSM 189/2002 (FGMV 2001.1160), adult male, collected by M. Vences on 26–27 November 2001 at Vohidrazana (18.9658°S, 48.5103°E, 731 m a.s.l.). ZSM 1021/2003 (FGMV 2002.2359), adult male, collectors unknown, collected in 2003 at the type locality An’Ala (ca. 18.9193°S, 48.4880°E, ca. 880 m a.s.l.). KU 340631 (CRH 261), adult female, and KU 340641 (CRH 275), adult male, collected by C.R. Hutter, S.M. Lambert, Z.F. Andriampenomanana, and S. Justin on 10 December 2014 at Vohimana (18.9209°S, 48.5122°E, 787 m a.s.l.).

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 21.3–23.2 mm), predominantly brownish dorsal coloration, absence of red color on webbing or ventral side of limbs of many specimens, calling males along streams, and molecular phylogenetic relationships. Within the B. blommersae group, defined by absence of dorsolateral bands, presence of distinct red color in outer iris area, especially its dorsal and ventral edges, in most specimens, and advertisement calls with high dominant frequencies of 4903–5819 Hz consisting of 17–25 whistling notes comprising multiple short (19‒78 ms) and a few long notes (90–225 ms). Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16S 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 16S sequence of Mantella baroni): “G” in the site 177, “G” in the site 233, “C” in the site 314.

Within the B. blommersae group, distinguished from B. blommersae by calls mainly consisting of frequency-modulated whistles (vs. pulsed trills); and from B. vittatus by calls mainly consisting of frequency-modulated whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from B. marojezensis by presence of red color in outer iris area in most specimens (vs. absence), and advertisement calls consisting of 17–25 notes (vs. 7–8 notes), and from B. kirki sp. nov. by advertisement calls consisting of two types of (short and long) notes (vs. notes of successively increasing duration). For a distinction from other species of the B. marojezensis complex described herein, see accounts of these new species below.

Adult male, in good state of preservation, SVL 23.0 mm, tissue from right thigh removed as tissue samples for molecular analysis and posterior venter cut open for parasitological examination. Body slender; head wider than long, wider than body; snout rounded in dorsal view, truncate in lateral view; nostrils directed laterally, about equidistant to tip of snout and eye; canthus rostralis indistinct, slightly concave in dorsal view, loreal region slightly concave; tympanum indistinct but recognizable, round, TD about 44% of ED; supratympanic fold very indistinct, largely 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), 2i(traces), 2e(traces), 3i(traces), 3e(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 indistinct, recognizable as unpigmented weak swelling on first finger. 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(0.5), 2i(0.75), 2e(0.25), 3i(1), 3e(0), 4i(1.75), 4e(1.5), 5(0.5); 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, 17 years after collection (Fig. 6), dorsally reddish brown with a distinct and moderately contrasted dark brown hourglass marking on anterior part of the dorsum, and a dark brown transverse bar on the posterior part of the dorsum. Many dark spots of different sizes are scattered across the dorsum. Limbs light brown with a few rather poorly contrasted darker brown crossbands: 1–2 on forearm, 2–3 on shank, 2–3 on thigh. Ventrally cream, white on belly, with some dark pigment only on ventral side of feet. Color of holotype in life not recorded.

Several paratypes from An’Ala in preservative are characterized by a distinct dorsal hourglass pattern (plus additional patch posterior to it), particularly contrasted in ZSM 1970/2006 and 1969/2006. ZSM 268/2006 has a contrasted pinkish patch above the right eye, ZSM 326/2000 has a pinkish marking on the central dorsum and many small white-pinkish spots on the anterior dorsum, whereas ZSM 265/2006 has the dorsum covered with numerous larger pink patches (ca. 15 partly fused patches). ZSM 189/2002 from Vohidrazana features, in addition to the dorsal hourglass marking, a fine light vertebral line. In life, the dark dorsal pattern is often only weakly recognizable (Fig. 9). In one paratype (KU 340641), the hindlimbs have an orange tint in life but no deep red color is recognizable (Fig. 9B). The outer iris color can be deep red (Fig. 9C; see also Rosa et al. 2012) or light orange (Fig. 9A). Iris periphery light blue.

Figure 9. 

Individuals of Boophis picardi sp. nov. in life. A, B male paratype KU 340641 (CRH 275) from Vohimana in dorsolateral and ventral view. C, D male paratypes from An’Ala, not assignable to specific voucher specimens.

Named after the fictional character Captain Jean-Luc Picard, first portrayed by Sir Patrick Stewart in Gene Roddenberry’s Star Trek: The Next Generation, and later in Akiva Goldsman, Michael Chabon, Kirsten Beyer, and Alex Kurtzman’s Star Trek: Picard.

The tadpole of this species is unknown.

An arboreal, nocturnal treefrog found in humid rainforests along fast flowing streams. Little is known of the ecology of the species. At Vohidrazana, male specimens were collected calling along a stream, from 1 meter to occasionally several meters above the ground. Many other calling males could be heard calling higher but were unable to be reached.

Advertisement calls of B. picardi sp. nov. recorded at An’Ala on 12 February 1995 (21.5°C air temperature) consist of two different note types, namely a series of rather short, quickly repeated notes, followed by 2–3 distinctly longer notes separated by slightly longer intervals. All notes are tonal in character and exhibit a distinct upward frequency modulation, with a frequency shift comprising 400–500 Hz in long notes and about half that in short notes. Amplitude across the entire call is increasing, with the first short notes being relatively soft, reaching maximum call energy at about one third of the call’s duration. Within notes, no distinct amplitude modulation is recognizable. Numerical parameters of three analyzed calls of different individuals are as follows: call duration 1554–1832 ms (1673.3 ± 143.1 ms); notes/call 19–21 (20.3 ± 1.2); short note duration 24–53 ms (34.4 ± 6.6 ms); long note duration 112–200 ms (157.9 ± 26.8 ms); inter-note interval 25–96 ms (41.1 ± 17.5 ms); note repetition rate of short notes within the call vary around 16 calls/second; dominant frequency 4903–5444 Hz (5267 ± 152 Hz); prevalent bandwidth 4200–5600 Hz.

Calls of B. picardi sp. nov. recorded at Betampona on 30 October 2007, 22:30 h (air temperature 18°C) (from Rosa et al. 2011, 2012), are in general agreement with those described from An’Ala above. A slight difference is obvious for overall amplitude modulation across the call, with amplitude of notes slightly decreasing again after having reached maximum call energy at one third of the call’s duration. Also, inter-note intervals are slightly longer compared to calls from An’Ala. Numerical parameters of two analyzed calls are as follows: call duration 1991–2000 ms; notes/call 17; short note duration 28–43 ms (32.8 ± 5.1 ms); long note duration 90–205 ms (140.8 ± 39.4 ms); inter-note interval 42–119 ms (64.6 ± 24.3 ms); note repetition rate of short notes within the call vary around 13 calls/second; dominant frequency 5442–5819 Hz (5667 ± 111 Hz); prevalent bandwidth 4500–6200 Hz.

The character of calls of B. picardi sp. nov. from Vohidrazana, recorded on 17 February 2001 (air temperature not recorded), generally agrees with those from An’Ala and Betampona described above, but Vohidrazana calls contain more long notes (6) following the short notes. Numerical parameters of two analyzed calls are as follows: call duration 2362–2388 ms; notes/call 24; short note duration 27–62 ms (37.8 ± 10.9 ms); long note duration 112–162 ms (139.0 ± 17.4 ms); inter-note interval 28–145 ms (57.5 ± 33.7 ms); note repetition rate of short notes within the call vary around 17 calls/second; dominant frequency 5292–5560 Hz (5417 ± 89 Hz); prevalent bandwidth 4900–5800 Hz.

Calls recorded at Vohidrazana, in December 2015 and January 2016 (air temperatures 17.9 and 20.3°C) and corresponding to call vouchers KU 342939 (CRH 971) and KU 342967 (CRH 1044) have the following numerical parameters (six calls analyzed): call duration 2010–2376 ms (2182.3 ± 119.7 ms); notes/call 20–25 (22.2 ± 1.8); short note duration 28–78 ms (38.0 ± 12.6 ms); long note duration 124–209 ms (163.3 ± 29.6 ms); inter-note interval 27–122 ms (55.8 ± 28.0 ms); note repetition rate of short notes within the call vary around 15 calls/second; dominant frequency 4971–5122 Hz (5060 ± 64 Hz); prevalent bandwidth 4200–5500 Hz.

Calls recorded at Mantadia, on 14 January 2017 (air temperature 19.1°C) and corresponding to the voucher specimen KU 347246 (CRH 1932) are also in agreement with the calls described above. Numerical parameters of three analyzed calls are as follows: call duration 2012–2330 ms (2165.0 ± 159.3 ms); notes/call 21–24 (23.0 ± 1.7); short note duration 19–75 ms (32.6 ± 14.0 ms); long note duration 123–225 ms (169.3 ± 33.9 ms); inter-note interval 35–89 ms (53.4 ± 17.7 ms); note repetition rate of short notes within the call vary around 15 calls/second; dominant frequency 5033–5388 Hz (5178 ± 119 Hz); prevalent bandwidth 4300–5600 Hz.

According to the molecular data summarized herein, the species is known from several sites in the wider area around the village of Andasibe, i.e., (1) the type locality, An’Ala, (2) Vohidrazana, (3) Vohimana, and it also has been recorded from (4) Betampona (Sahambendrana and Sahabefoza sites, according to Rosa et al. 2012). The elevational range spans from 349 (Betampona, Sahabefoza) to 880 m a.s.l. (An’Ala).

This species has been previously referred to as B. marojezensis [Ca53 JQ518216] = B. marojezensis [Ca53] in Randrianiaina et al. (2012), and B. sp. Ca53 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).

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.

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.).

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.).

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 16S 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 16S sequence of Mantella baroni): “C” in the site 86, “A” in the site 108, “G” in the site 254.

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.

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), 2i(traces), 2e(traces), 3i(traces), 3e(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), 2i(1.5), 2e(0.5), 3i(1.75), 3e(0.5), 4i(1.75), 4e(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.

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. 10E). 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).

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.

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.

Tadpoles of this species were described and illustrated under the names B marojezensis and B. marojezensis [Ca53] by Randrianiaina et al. (2012), based on the DNA barcoded specimens ZSM 1528/2007 (FGZC 2277; GenBank accession number JQ518196) from Marojejy National Park, and ZSM 573/2010 (ZCMV 13200; GenBank accession number JQ518216) 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.

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.

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.

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.

This species has been previously referred to as B. marojezensis [Ca52 JQ518215] = B. marojezensis [Ca52] in Randrianiaina et al. (2012), and B. sp. Ca52 in Perl et al. (2014) and Hutter et al. (2018). It was included in B. marojezensis sensu lato in Glaw and Vences (2007) and not explicitly included or mentioned by Glaw et al. (2001) and Vieites et al. (2009).

ZSM 614/2001 (FGMV 2001.64), adult male (call voucher) collected by F. Andreone, F. Mattioli, J. Randrianirina, and M. Vences on 3 February 2001 at Antsahamanara campsite, Manarikoba forest, Tsaratanana Massif (14.045°S, 48.784°E, ca. 1000 m a.s.l.), Sambirano region, Madagascar.

ZSM 615/2001 (FGMV 2001.71) and ZSM 616/2001 (FGMV 2001.80), two adult males with same collecting data as holotype but collected 4‒9 February 2001 (ZSM 615/2001 not DNA barcoded). ZSM 10/2016 (MSZC 186), adult male, collected by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D.H. Nomenjanahary, and J. Rabearivony on 9 January 2016 at Ampotsidy, 15.7 km NNW of Bealanana (8.7 km NNW of Beandrarezona; 14.4276°S, 48.7223°E, 1320 m a.s.l.).

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 25.0–27.2 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 dominant frequencies of 4688–5332 Hz, consisting of 7–12 whistling notes of 44–220 ms duration, each with constant upward frequency modulation. Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16S 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 16S sequence of Mantella baroni): “G” in the site 109, “T” in the site 163, “A” in the site 233.

Within the B. blommersae group, distinguished from B. blommersae by calls consisting of a series of whistles (vs. pulsed trills); and from B. vittatus by calls consisting of a series of whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from B. marojezensis by somewhat larger body size (male SVL 25.0–27.2 vs. 20.0–25.7 mm), advertisement calls consisting of notes of rather similar duration (vs. clear distinction of short and long notes); from B. kirki sp. nov. by larger body size (male SVL 25.0–27.2 vs. 20.0–23.4 mm), advertisement calls with notes of maximum duration of 220 ms (vs. max. duration of 105 ms) and with regularly ascending frequency modulation (vs. very steep initial frequency ascent in the beginning of each note, slowing down towards end of note), and absence of red color in outer iris area (vs. presence in some specimens); from B. picardi sp. nov. by larger body size (male SVL 25.0–27.2 vs. 21.3–23.2 mm), advertisement calls consisting of notes of rather similar duration (vs. clear distinction of short and long notes), and absence of red color in outer iris area (vs. distinct in many specimens); and from B. pikei sp. nov. by larger body size (male SVL 25.0–27.2 vs. 21.4–25.0 mm), and advertisement calls consisting of 7–12 notes (vs. 25–33 notes) of maximum duration of 220 ms (vs. max. duration of 98 ms). For a distinction from other species of the B. marojezensis complex described herein, see accounts of these new species below.

Adult male, in excellent state of preservation, SVL 25.8 mm, muscle tissue removed from left thigh for molecular analysis. Body moderately slender; head slightly wider than long and slightly wider than body; snout rounded to truncate in dorsal view, moderately rounded to sloping in lateral view; nostrils directed laterally, nearer to tip of snout than to eye; canthus rostralis rather weakly expressed, concave in dorsal view, loreal region slightly concave; tympanum rather distinct, round, TD 39% of ED; supratympanic fold distinct, slightly curved in its anterior and straight in its posterior half; vomerine odontophores weakly developed, well-separated in two very small rounded aggregations, positioned posteromedial to choanae; choanae small to 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), 2i(traces), 2e(traces), 3i(traces), 3e(2), 4(1); 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(0.5), 2i(0.75), 2e(0.5), 3i(1.25), 3e(0.5), 4i(2), 4e(2), 5(0.5); 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, 22 years after collection (Fig. 6), dorsally red brown with a distinct and moderately contrasted dark brown hourglass marking on anterior part of the dorsum, and a dark brown curved transverse bar on the posterior part of the dorsum. A narrow dark transverse bar is visible between the eyes. Dorsum with many poorly delimited and indistinct small dark spots. Limbs light brown with distinct darker brown crossbands: about 3 on forearm, 3–5 on shank, 5 on thigh. Ventrally cream, white on belly and with dark pigment on ventral side of feet. In life (Fig. 11), similar but dorsal color light brown rather than reddish brown. Outer iris color yellowish, inner iris color light brown, iris periphery turquoise.

Figure 11. 

Individuals (adult males) of Boophis siskoi sp. nov. in life. A paratype ZSM 10/2016 (MSZC 186) from Ampotsidy. B holotype ZSM 614/2001 (FGMV 2001.64) from the Tsaratanana Massif (Antsahamanara campsite).

All paratypes in preservative have the typical pattern of a dark hourglass-patch on the anterior and a transverse bar or inverted U-patch on the posterior dorsum. ZSM 616/2001 has a small pink dot on the central dorsum. In life, the dark dorsal markings are sometimes poorly contrasted (Fig. 11A).

Named after the fictional character Captain Benjamin Sisko, first portrayed by Avery Brooks in Rick Berman and Michael Piller’s Star Trek: Deep Space Nine.

The tadpole of this species (under the name B. marojezensis [Ca52]) was described and illustrated by Randrianiaina et al. (2012), based on the DNA barcoded specimen ZSM 541/2010 (ZCMV 13168; GenBank accession number JQ518215) from Ambinanitelo. 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 (258 marginal and 522 submarginal; without dorsal gap). They are characterized by presence of a (poorly recognizable) lateral transparent area of the integument, and absence of dark (melanophoric) pigment on the tail.

An arboreal, nocturnal treefrog found in humid rainforests along fast flowing streams. Little is known of the ecology of the species. At Ampotsidy, this species was only encountered along a large river, and was not heard along narrower streams at higher elevations.

Advertisement calls of B. siskoi sp. nov. recorded at the type locality, Antsahamanara, Tsaratanana massif, on 3 February 2001 (24°C air temperature) consist of multiple notes (almost tonal in character, but with some irregular amplitude modulation) of variable duration, repeated at variable intervals. Within calls, note duration and inter-note intervals become longer from the beginning to the end of the call. Each note exhibits upward frequency modulation, which is most expressed in the last note (comprising a frequency shift of about 300 Hz) and much less in the first notes of the call. Overall frequency of the call exhibits a slight drop from the beginning to the end of about 200 Hz from the first notes compared to the last note (Fig. 5). However, in some calls the first 2–3 notes exhibit a lower frequency than subsequent ones. Each note is amplitude modulated with maximum call energy usually present somewhere in the first half of the note’s duration. Numerical parameters of four analyzed calls are as follows: call duration 1549–2168 ms (1877.0 ± 311.2 ms); notes/call 10–12 (10.7 ± 1.2); note duration 44–220 ms (89.8 ± 44.5 ms); inter-note interval 41–162 ms (72.6 ± 31.7 ms); dominant frequency 4890–5332 Hz (5125 ± 222 Hz); prevalent bandwidth 4000–5600 Hz.

A call of B. siskoi sp. nov. recorded at Ampotsidy, on 13 January 2016 at 18:43 (air temperature not recorded), differs from the calls described from Tsaratanana by less variation in note duration and lack of a distinct overall frequency drop from the beginning to the end of the call (Fig. 5). Numerical parameters of this analyzed call are as follows: call duration 1805 ms; notes/call 7; note duration 114–165 ms (144.3 ± 18.5 ms); inter-note interval 88–180 ms (125.7 ± 30.8 ms); dominant frequency 4688 Hz; prevalent bandwidth 3800–5200 Hz.

According to molecular data summarized herein, the species is known from (1) the type locality, Antsahamanara campsite on the western versant of the Tsaratanana Massif, (2) Ambinanitelo forest (14.2254°S, 48.9635°E, 1182 m a.s.l.; Randrianiaina et al. 2012), and (3) Ampotsidy forest. The elevational range of the species spans between ca. 1000–1320 m a.s.l.

This species has been previously referred to as B. sp. aff. marojezensis [CaHM364579] by Rosa et al. (2012), and as B. marojezensis by Perl et al. (2014). It was not explicitly included or mentioned in the studies of Glaw et al. (2001), Glaw and Vences (2007), Vieites et al. (2009), Randrianiaina et al. (2012), and Hutter et al. (2018).

ZSM 472/2009 (ZCMV 11468), adult male, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison, and J. Patton on 21 June 2009 at Angozongahy campsite, western side of Makira plateau (15.4370°S, 49.1186°E, 1009 m a.s.l.), North East of Madagascar.

ZSM 473/2009 (ZCMV 11486) and ZSM 474/2009 (ZCMV 11487), two adult males, with same collection data as holotype. ZSM 470/2009 (ZCMV 11270) and ZSM 471/2009 (ZCMV 11272), two adult males, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison, and J. Patton on 23–24 June 2009 at a campsite near the source of Fotsialanana river, western side of the Makira plateau (15.4668°S, 49.1289°E, 1067 m a.s.l.). ZSM 207/2022 (FGZC 6519), adult male, collected by J.M. Rafanoharana, H. Raherinjatovo, and F. Glaw on 24 March 2022 at Analanjirofo (near Simpona Lodge), Makira Reserve (15.1992°S, 49.6208°E, 410 m a.s.l.).

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 25.2–28.8 mm), predominantly brownish dorsal coloration, absence of red color on webbing or ventral side of limbs, calling males occurring along streams, 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 low dominant frequencies of 2687–3404 Hz, consisting of 3–5 whistling notes of 238–604 ms duration. Also characterized by numerous diagnostic nucleotide positions in the mitochondrial 16S 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 16S sequence of Mantella baroni): “G” in the site 149, “T” in the site 190, “G” in the site 191.

Within the B. blommersae group, distinguished from B. blommersae by calls consisting of a series of whistles (vs. pulsed trills); and from B. vittatus by calls consisting of a series of whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from B. marojezensis by advertisement calls with lower dominant frequency (2687–3404 vs. 4118–4441 Hz), with notes emitted at longer maximum inter-note intervals (639 vs. 179 ms), and larger body size (male SVL 25.2–28.8 vs. 20.0–25.7 mm); from B. kirki sp. nov. by advertisement calls with lower dominant frequency (2687–3404 Hz vs. 3499–5604 Hz), longer note duration (238–604 ms vs. 54–105 ms), larger male body size (SVL 25.2–28.8 vs. 20.0–23.4 mm), and absence of red color in outer iris area (vs. presence in some specimens); from B. picardi sp. nov. by advertisement calls with lower dominant frequency (2687–3404 Hz vs. 4903–5819 Hz), consisting of 3‒5 notes (vs. 17‒25 notes), longer note duration (238–604 ms vs. 19–225 ms), larger body size (male SVL 25.2–28.8 vs. 21.3–23.2 mm), and absence of red color in outer iris area (vs. distinct in many specimens); from B. pikei sp. nov. by advertisement calls consisting of 3–5 notes (vs. 25–33 notes) of 238–604 ms note duration (vs. max. duration of 98 ms), with lower dominant frequency (2687–3404 vs. 5174–5507 Hz), and larger body size (male SVL 25.2–28.8 vs. 21.4–25.0 mm); and from B. siskoi sp. nov. by advertisement calls consisting of 3–5 notes (vs. 7–12 notes) of 238–604 ms note duration (vs. max. duration of 220 ms) and with lower dominant frequency (2687–3404 vs. 4688–5332 Hz). For a distinction from other species of the B. marojezensis complex described herein, see accounts of these new species below.

Adult male, in excellent state of preservation, SVL 27.4 mm, muscle tissue removed from right thigh for molecular analysis. Body moderately slender; head slightly wider than long and slightly wider than body; snout rounded in dorsal and lateral views; nostrils directed laterally, about equidistant between tip of snout and eye; canthus rostralis distinct, concave in dorsal view, loreal region slightly concave; tympanum indistinct, round, TD about 54% of ED; supratympanic fold not recognizable (traces posterior to tympanum); 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), 2i(traces), 2e(traces), 3i(traces), 3e(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 between nostril and 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(0), 2i(0.5), 2e(0.25), 3i(1.25), 3e(0.25), 4i(1.75), 4e(1.75), 5(0.5); 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, 14 years after collection (Fig. 6), dorsally light brown with an indistinct and poorly contrasted brown hourglass marking on anterior part of the dorsum. No dark transverse bar is visible on the posterior part of the dorsum, but a comparatively wide dark transverse bar is present between the eyes. Dorsum with an irregular pattern of dark brown, black and whitish small spots, many of which are poorly contrasted. Limbs light brown with darker brown crossbands: 3–4 on forearm, about 5 on shank, about 5 on thigh. Ventrally cream, white on belly and with dark pigment on ventral side of feet. In life (Fig. 12), similar but the dorsal hourglass pattern almost not recognizable, crossbands on limbs poorly contrasted, but a few black and white dorsal spots well visible and contrasted. Outer iris color yellowish, inner iris color beige, iris periphery turquoise.

Figure 12. 

Individuals of Boophis janewayae sp. nov. in life. A male holotype ZSM 472/2009 (ZCMV 11468) from Angozongahy campsite, western side of Makira plateau. B calling male (with deflated vocal sac, during a pause between calls) photographed in the field at night, at the type locality.

Two paratypes from Makira West (ZSM 470/2009 and 471/2009) in preservative are characterized by a contrasted dark hourglass-marking on the anterior dorsum and an inverted U-shaped marking on the posterior dorsum. ZSM 473/2009 has only a poorly contrasted hourglass-marking but features a light vertebral line. ZSM 474/2009 has a pattern of irregular fine light spots.

Named after the fictional character Captain Kathryn Janeway, first portrayed by Kate Mulgrew in Rick Berman, Michael Piller, and Jeri Taylor’s Star Trek: Voyager.

The tadpole of this species is unknown.

An arboreal, nocturnal treefrog. Little is known of the ecology of the species. It has been found in humid rainforests along relatively slow-moving streams. Calling males were perched 1.5 to 2 m above the ground on leaves.

Advertisement calls of B. janewayae sp. nov. recorded from the holotype at Angozongahy, Makira area, on 21 June 2009 (air temperature not recorded) consist of a series of comparatively long tonal notes of variable duration, emitted at somewhat irregular intervals. Within calls, the first (and sometimes also the last) note is longest in duration. Amplitude is modulated within notes, with maximum call energy being present at each note’s end. Each note exhibits an upward frequency modulation comprising a shift in frequency of ~400 Hz at maximum from beginning to the end of the note. Numerical parameters of two analyzed calls are as follows: call duration 2358–2846 ms; notes/call 4–5; note duration 238–604 ms (347.6 ± 118.2 ms); inter-note interval 206–639 ms (304.3 ± 152.5 ms); dominant frequency 2687–2996 Hz (2865 ± 111 Hz); prevalent bandwidth 2100–3300 Hz; harmonic frequency bands are evident at around 6000, 9000, and 12000 Hz.

Calls of B. janewayae sp. nov. recorded at Betampona, on 15 November 2007, 23:43 h (21°C air temperature) (from Rosa et al. 2011, 2012), generally agree in character with those described above from Angozongahy, but differ slightly in showing more regular inter-note intervals and slightly higher dominant frequency. Numerical parameters of two analyzed calls are as follows: call duration 1571–2064 ms; notes/call 3–4; note duration 276–536 ms (365.4 ± 116.1 ms); inter-note interval 141–278 ms (220.2 ± 55.8 ms); dominant frequency 3036–3404 Hz (3231 ± 135 Hz); prevalent bandwidth 2500–3600 Hz; harmonic frequency bands are evident at around 6340, 9500, and 12630 Hz.

According to molecular data summarized herein, the species is known from (1) the type locality, western side of the Makira Reserve (Angozongahy and Fotsialanana), (2) the eastern side of the Makira Reserve (around “Simpona Lodge”), and (3) Betampona Reserve (sites: Sahambendrana, Sahabefoza, Sahaindrana, Vohitsivalana; Rosa et al. 2012). The elevational occurrence of the species spans from 349 m a.s.l. (Sahaindrana site in Betampona; Rosa et al. 2012) to 1067 m a.s.l. (Fotsialanana source, Makira).

This species has been newly discovered in this study and has not been included in any of the previous studies including B. marojezensis and 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).

ZSM 12/2016 (MSZC 213), adult male, collected by M.D. Scherz and M. Rakotondratisma on 15 January 2016 at Bevitagnono forest, 33.1 km SW of Bealanana on the road RN31 (14.7387°S, 48.5170°E, 1016 m a.s.l.), North West of Madagascar.

ZSM 11/2016 (MSZC 198), adult male, collected by M.D. Scherz and M. Rakotondratsima on 14 January 2016 at Andranonafindra forest, 30 km SW of Bealanana on the road RN31 (14.7360°S, 48.5481°E, 1169 m a.s.l.); ZSM 13/2016 (MSZC 243), adult female, collected by M.D. Scherz and M. Rakotondratsima on 17 January 2016 at Andranonafindra forest, 30 km SW of Bealanana on the road RN31 (14.7360°S, 48.5489°E, 1138 m a.s.l.).

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 25.0–25.3 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. blommersae group, 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 16S 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 16S sequence of Mantella baroni): “D” in the site 253, “T” in the site 277, “A” in the site 303.

Within the B. blommersae group, distinguished from B. blommersae by calls consisting of a series of whistles (vs. pulsed trills); and from B. vittatus by calls consisting of a series of whistles (vs. series of short clicks), and absence of dorsolateral stripes (vs. presence). Furthermore, distinguished from B. marojezensis by 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. marojezensis complex described herein, see accounts of these new species below.

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 TD about 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), 2i(traces), 2e(traces), 3i(traces), 3e(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), 2i(0.75), 2e(0), 3i(1.25), 3e(0), 4i(1.75), 4e(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, B male holotype ZSM 12/2016 (MSZC 213) from Bevitagnono forest in dorsal and dorsolateral view. C male paratype ZSM 11/2016 (MSZC 198) from Andranonafindra forest. D female paratype, ZSM 13/2016 (MSZC 243) from Andranonafindra forest.

The female ZSM 13/2016 in preservative 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.

Named after the fictional character Captain Jonathan Archer, first portrayed by Scott Bakula in Rick Berman and Brannon Braga’s Star Trek: Enterprise.

The tadpole of this species is unknown.

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.

Advertisement calls of B. archeri sp. nov., recorded at Bevitagnono and Andranonafindra forests from the holotype and the male paratype on 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 MSZC 198 and MSZC 213) 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.

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 m a.s.l.

This species has been previously referred to as B. sp. 26 by Vieites et al. (2009), B. marojezensis [Ca26] by Randrianiaina et al. (2012), and B. sp. Ca55 by Perl et al. (2014) and Hutter et al. (2018). Note that B. sp. Ca26 in Hutter et al. (2018) refers to another lineage. This species was not explicitly included or mentioned by Glaw et al. (2001) and Glaw and Vences (2007).

ZSM 492/2014 (DRV 6295), adult male, collected on 21 June 2010 by F.M. Ratsoavina at a site locally called Andrevorevo (campsite “A”) (14.3464°S, 49.1028°E, 1717 m a.s.l.), on the border of the North East and North West regions of Madagascar.

ZSM 491/2014 (DRV 6293), adult female with same collection data as holotype. ZSM 392/2016 (ZCMV 15171), probably a young or subadult male, collected by M.D. Scherz, A. Rakotoarison, M. Bletz, M. Vences and J. Razafindraibe close to Camp 2 “Marojejia”, Marojejy National Park (14.4348°S, 49.7660°E, 616 m a.s.l.).

A small-sized treefrog assigned to the genus Boophis, subgenus Boophis, in the family Mantellidae based on occurrence in 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 25.0–27.2 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 and absence of red color in outer iris area. As advertisement calls are unknown, this species can formally mainly be defined by its numerous diagnostic nucleotide positions in the mitochondrial 16S 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 16S sequence of Mantella baroni): “A” in the site 161, “C” in the site 179.

Within the B. blommersae group, distinguished from B. vittatus by absence of dorsolateral stripes (vs. presence). Because calls of this species are unknown, a bioacoustic pairwise diagnosis with other species of the group is not possible and therefore, the species can only be distinguished from some of its close relatives (B. archeri sp. nov., B. blommersae, B. janewayae sp. nov., B. siskoi sp. nov.) by molecular diagnostic sites (see Definition above). The species can be distinguished from B. kirki sp. nov. by absence of red color in outer iris area (vs. presence in some specimens), and presence of a lateral transparent area of the integument of tadpoles (vs. absence); from B. picardi sp. nov. by larger body size (male SVL 25.0–27.2 vs. 21.3–23.2 mm), and absence of red color in outer iris area (vs. distinct in many); from B. pikei sp. nov. by larger body size (male SVL 25.0–27.2 vs. 21.4–25.0 mm); and from B. marojezensis by absence of rounded patches on the posterior half of the tail musculature of tadpoles (vs. presence).

Adult male, in excellent state of preservation, SVL 25.5 mm, muscle tissue removed from left thigh for molecular analysis. Body moderately slender; head slightly wider than long, of similar width as body; snout rounded in dorsal view, moderately rounded to sloping in lateral view; nostrils directed laterally, about equidistant between tip of snout and eye; canthus rostralis distinct and concave in dorsal view, loreal region slightly concave; tympanum indistinct, difficult to recognize, somewhat ovoid (higher than wide), TD 43% of ED; supratympanic fold not recognizable anteriorly and dorsally of tympanum, weakly recognizable and regularly curved posterior of tympanum; vomerine odontophores weakly developed, well-separated in two 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 moderately webbed and with lateral dermal fringes; webbing formula 1(traces), 2i(traces), 2e(1), 3i(2.5), 3e(1.75), 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 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(0.25), 2i(1), 2e(0), 3i(1.25), 3e(0.25), 4i(1.75), 4e(1.75), 5(0.5); 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, 13 years after collection (Fig. 6), dorsally light reddish brown with a moderately contrasted but distinct and complete brown hourglass marking on anterior part of the dorsum, a discontinuous 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 which partly fuse to larger markings. Limbs light brown with darker brown crossbands: about 3–5 poorly marked crossbands on forearm, 5 on shank, 7–8 on thigh. Ventrally cream, white on belly and with some dark pigment on ventral side of feet. Color of holotype in life not recorded.

The female paratype ZSM 491/2014 in preservative has a rather uniform grayish dorsal color, with a few isolated dark spots and traces of an hourglass marking, and with three pink spots on the posterior dorsum close to the cloaca. In life, paratype ZSM 392/2016 (Fig. 14) had a reddish brown color with brown hourglass marking, a beige iris with orange color in the outer iris area, especially dorsally, a relatively distinct network of brown lines, and a turquoise iris periphery.

Figure 14. 

A paratype individual (probably a young male) of Boophis burnhamae sp. nov. in life: ZSM 392/2016 (ZCMV 15171) from Marojejia campsite in the Marojejy Massif.

Named after the fictional character Captain Michael Burnham, first portrayed by Sonequa Martin-Green in Bryan Fuller and Alex Kurtman’s Star Trek: Discovery.

The tadpole of this species (under the name B. marojezensis [Ca26]) was described and illustrated by Randrianiaina et al. (2012), based on the DNA barcoded specimen ZSM 1612/2007 (FGZC 2930; GenBank accession number JQ518197) from Marojejy. 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 (234 marginal and 430 submarginal; without dorsal gap). They are characterized by presence of a lateral transparent area of the integument surrounding the body.

An arboreal, nocturnal treefrog found in humid rainforests along streams. Little is known of the ecology of the species. The paratype ZSM 392/2016 was encountered at rest during the day on a leaf overhanging the path.

Unknown.

According to molecular data summarized herein, the species is reliably known from: (1) the type locality, Andrevorevo, and (2) Marojejy at mid-elevation near Camp Marojejia. The elevational range spans between 616–1717 m a.s.l.