Corresponding author: Shea M. Lambert (
Academic editor: Johannes Penner
We describe a new species from the cophyline microhylid genus
Over the past several decades, integrative approaches to taxonomy have shown that Madagascar’s anuran fauna is one of the most spectacular on earth, with current estimates approaching 600 species; 99.9% of which are endemic to the island (reviewed in
Phylogenetic relationships between
We herein describe
We collected specimens during the day through targeted searching, using the advertisement call to locate males. We euthanized specimens using 20% benzocaine, fixed them in ~10% formalin solution buffered with sodium phosphate to pH 7.0, and transferred them to 70% ethanol for long-term storage after approximately two weeks. We deposited the holotype in the Biodiversity Institute of the
Immediately following euthanasia, we removed the tongue and placed it in 95%
GenBank Accession numbers for all sequences used in phylogenetic analysis. Asterisks indicate newly generated sequences.
Species | Locality | Voucher number | Accession |
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Ambatomandondona |
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Torotorofotsy |
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Vohidrazana | CRH 286 |
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Vohidrazana |
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Andasibe | ZSM 3/2002 |
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Torotorofotsy | ZCMV 968 |
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Tsararano | MRSN A 2620 |
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Vohidrazana |
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Mandraka | ZSM 694/2001 |
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Betampona | FAZC 13887 |
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Betampona | MRSN A 6271 |
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Betampona | MRSN A 6347 |
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Tsaratanana | ZCMV 12401 |
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Montagne d’Ambre | ZSM 218/2004 |
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Montagne d’Ambre | FGZC 1052 |
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Sorata forest | FGZC 3651 |
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Nosy Mangabe | ZCMV 886 |
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Foret’d Ambre | FGZC 1888 |
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Foret’d Ambre | FGZC 1890 |
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Antsiranana, Andapa | AMNH 181903 |
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Marojejy | FGZC 2897 |
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Marojejy | FGZC 2899 |
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Tsaratanana Camp Matsaborimaika | DRV 6456 |
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Tsaratanana Camp Matsaborimaika | ZCMV 12382 |
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Tsaratanana Camp Matsaborimaika | ZCMV 12384 |
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Marojejy | ZCMV 2065 |
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Ambolokopatrika | FAZC 7292 |
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Ilampy | FAZC 10314 |
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Tsaratanana | MRSN A 2631 |
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Tsaratanana | ZSM 667/2001 |
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Ambolokopatrika | MRSN A 2640 |
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Marojejy | FGZC 2866 |
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Marojejy | ZCMV 2065 |
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Tsaratanana | 2001 G46 |
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Masoala | MRSN A 2115 |
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Ilampy | MRSN A 2610 |
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Andemaka |
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Makira | ZCMV 11473 |
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Tsaratanana Camp Matsaborimaika | ZCMV 12359 |
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Nosy Be | ZSM 474/2000 |
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Nosy Be | ZSM 475/2000 |
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Antsiranana | AMNH A167315 |
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Marojejy | FGZC 2842 |
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Torotorofotsy |
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Raw genetic distances at the 16s rRNA gene fragment between analysed taxa and
Taxon | Distance |
---|---|
3.80% | |
4.89–5.98% | |
5.98% | |
7.61% | |
9.78–10.32% | |
10.87% | |
11.41% | |
11.41% | |
12.50% | |
11.96–12.50% | |
11.96% | |
11.96% | |
12.50% | |
13.04% | |
13.04% | |
11.96% | |
12.50% | |
13.59% | |
13.59% | |
14.13% | |
13.59% | |
15.76% | |
15.22% | |
16.30% | |
19.02% | |
18.48% | |
23.37% |
We took morphological measurements using a digital caliper to 0.01 mm, rounded to 0.1 mm. We note that only the holotype was measured, as the paratype was unavailable for study. Measurements follow the standard for this genus and are repeated here verbatim from
We performed
We exported the volume as an “Analyze Volume” under standard settings in VG Studio Max 2.2, and imported the resulting .hdr file into Amira 6.1 (FEI Visualization Sciences Group, Burlington MA, USA), where a surface model was produced essentially following
We note that skeletal comparisons to other cophylines are based on largely unpublished
We recorded calls attributed to the holotype on two occasions using an Olympus LS-10 Linear PCM Field Recorder and a Sennheiser K6-ME66 super-cardioid shotgun microphone. The calls were recorded at a sampling rate of 44.1 kHz and 16 bits resolution in WAV format. Recordings were made at mid-day in overcast weather conditions. No precise temperature recordings are available, but we estimate that the ambient temperature was approximately 20° C at the time of recording. We note that the individual was not visible during the recordings, as it was calling from a burrow. We therefor e cannot be completely certain that the recordings are of the same individual, however, only a single individual at a time was heard calling from this location, and the collected individual was found with distended vocal sac shortly after the second recordings. Additionally, the measured call parameters from the two occasions are nearly completely overlapping (Fig.
We follow
Following
The advertisement call recorded for
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Species |
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Specimen number | NA | |
Locality | Ranomafana | Nosy Be |
N – calls | 7 | 4 |
Inter-call interval duration (s) | 42.5–99.5 (68.77 ± 24.0) | 5.98–10.1 |
Call duration (ms) | 505–544 (536 ± 1.7) | 828–896 |
Call envelope | 0.601–0.787 (0.663 ± 0.073) | - |
Number of amplitude peaks | 3–5 (3.4 ± 0.5) | 1 |
Fundamental frequency (Hz) | 236.9–279.9 |
258.4–279.9 |
Dominant frequency throughout call (Hz) | 528.3–538.8 (537.9 ± 9.2) | 538.3–555.9 |
Dominant frequency at peak amplitude (Hz) | 528.3–538.8 (537.9 ± 9.2) | 581.4–602.9 (586.8 ± 10.8) |
First Harmonic (Hz) | 775.2–818.3 |
775.2–796.7 (791.3 ± 10.8) |
The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (
We discovered a large-bodied cophyline microhylid frog near Andemaka within Ranomafana National Park in eastern Madagascar. Several obvious differences in morphology exist between the collected specimens and all known described and undescribed cophyline microhylids. Analysis of a fragment of its mitochondrial 16S rRNA gene recovered it with a close relationship to an undescribed population of
Map of Ranomafana National Park and the type locality of
A frog assigned to the cophyline genus
Although the bioacoustic repertoires of cophylines is far from completely known, bioacoustically, this species’ call is strongly distinct from the other known calls by being strongly amplitude modulated (Fig.
Body rotund; dorsal and ventral skin smooth, with subtle bumps on the dorsal skin (more rugose in life). Head considerably wider than long (
Photos in life of
Arms strongly built, relatively short; fingers without webbing, short, with distinct, rounded subarticular tubercles, relative lengths 1<2<4<3, the second finger marginally shorter than the fourth (and marginally longer than the first), without enlarged terminal discs; inner metacarpal tubercle strong, oblong, 28.1% of hand length; outer metacarpal tubercle indistinct, round. Legs relatively long and thick (
Anterior braincase laterally closed by the sphenethmoid. Interior braincase containing calcified material. Nasal in medial contact with contralalteral and posterior contact with frontoparietal. Frontoparietal broadening anteriorly from narrow waist anterior to lateral flanges, possessing a strong, posteriorly elongated dorsal process. Prechoanal vomer simple, triradiate. Neopalatine and postchoanal vomer distinguishable. Vomerine teeth not medially fused, without diastemata, oriented oblique to antero-posterior body axis, curved. Maxillary teeth minute. Otic capsule dorsally poorly ossified.
Sternum not ossified. Clavicle robust, curved. Humerus proximally broad, distally rather narrow; possessing a well-developed crista ventralis along roughly 50% of its length; crista lateralis weak. Terminal phalanges of fingers and toes with small distal knobs. Phalangeal formula of fingers 2-2-3-3; of toes 2-2-3-4-3. Femur without cristae. Prepollex strong, blade-like, half length of first metacarpal. Prehallux strong, approximately half length of first metatarsal.
Neural spines decrease in size posteriorly, the sixth and seventh lacking spines altogether. Neural arches of atlas fused. Dorsal crest of urostyle running roughly 80% along its shaft. Iliosacral articulation type IIA sensu
The paratype
We analysed a total of seven calls from
Each call is rapidly pulsed, with 3–5 (3.5 ± 0.534) amplitude modulated peaks occurring throughout the call, and peak amplitude occuring in the last 50% of the call. The call duration is 505–544 (536 ± 1.7) ms with an inter-call interval duration of 42.5–99.5 (68.8 ± 24.0) s. The fundamental frequency is 236.9–279.9 (261.5 ± 22.9) Hz. The mean dominant frequency throughout the call was 528.3–555.9 (537.9 ± 9.2) Hz and the first harmonic frequency is 775.2–818.3 (796.8 ± 17.6) Hz (Fig.
The osteology of
The specific epithet “
Due to morphological and size similarities, as well as geographic distribution, two existing names must be considered for this species:
Comparative spectrograms (top), oscillograms (center) and power spectra (bottom) between the calls of (
Although the type locality of
The discovery of
In addition to morphological distinctiveness,
Our limited genetic data suggests that
We thank the Malagasy authorities for issuing permits; field research was conducted under permit number 303/14/MEF/SG/DGF/DCB.SAP/SCB; specimens were exported under 017N-EV01/MG14. We also thank MICET and Centre ValBio for facilitating fieldwork. Finally, SML would like to thank Ralaivao Jean Fulgence and Emile Rajeriarison for their exceptional work in the field during the Andemaka expedition. If not for their dedication and ability,
File S1
Adobe PDF file
This file contains a PDF-embedded interactive 3D model of the skeleton of the holotype of