Boophis marojezensis Glaw & Vences, 1994

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

Boophis marojezensis Glaw & Vences, 1994

Figures 6 , 7

Identity.

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 16 S 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).

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. Ca 53 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.

Holotype.

ZFMK 57401 , by original designation. Type locality: “ the Marojezy massif at low altitude, NE-Madagascar ”. A partial 16 S sequence of the holotype is available from GenBank under accession number PQ 278105.

Paratypes.

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

Material examined.

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.) .

Definition.

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 16 S rRNA gene: MolD identified the following robust diagnostic nucleotide combination compared to all other species in the B. marojezensis complex (sites given relative to the full-length 16 S sequence of Mantella baroni ): “ C ” in the site 107, “ G ” in the site 162, “ C ” in the site 251.

Diagnosis.

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.

Tadpole.

The tadpole of B. marojezensis (under the name B. marojezensis [Ca 25]) was described and illustrated by Randrianiaina et al. (2012), based on the DNA barcoded specimen ZSM 1611 / 2007 (FGZC 2929; GenBank accession number FJ 559146 ). 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.

Natural history.

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.

Calls.

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.

Distribution.

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.

Figures

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

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).