Gephyromantis granulatus (Boettger, 1881)

Randrianiaina, Roger Daniel, Wollenberg, Katharina C., Hiobiarilanto, Tahiry Rasolonjatovo, Strauss, Axel, Glos, Julian & Vences, Miguel, 2011, Nidicolous tadpoles rather than direct development in Malagasy frogs of the genus Gephyromantis, Journal of Natural History 45 (47 - 48), pp. 2871-2900: 2880-2894

publication ID

http://doi.org/ 10.1080/00222933.2011.596952

persistent identifier

http://treatment.plazi.org/id/274FF905-FFB3-CE0F-A0AA-FE2306E4FC67

treatment provided by

Felipe

scientific name

Gephyromantis granulatus (Boettger, 1881)
status

 

Gephyromantis granulatus (Boettger, 1881)  

( Figures 1A View Figure 1 , 2 View Figure 2 )

Material examined

The following description refers to one tadpole in developmental stage Gosner 40 (field number Tad 2004-75 – ZSM 298 View Materials /2008, BL 5.8 mm, TL 20.7 mm, accession number HQ188939 View Materials ) collected by R. D. Randrianiaina, M. Puente and F. Glaw on 19–23 February 2004 in Montagne d’Ambre National Park in a brook crossing the track “Voie des milles arbres” (coordinates at stream not taken, but not far from 12 ◦ 31.667’ S, 49 ◦ 10.667’ E, 1050 m a.s.l.). The 16S rRNA sequence of this specimen was 100% identical to the reference sequence of a G. granulatus   adult specimen (accession AJ 315926 View Materials ) in Genbank GoogleMaps   .

Description

In dorsal view, body ovoid, maximal body width attained between 2/5 and 3/5 of the body length (SBW 44% of BL), snout broadly rounded. In lateral view, body depressed (BW 132% of BH), maximal body height attained between 3/5 and 4/5 of body length (SBH 62% of BL), snout rounded. Eyes large (ED 17% of BL), visible from ventral view, positioned high (EH 61% of BH) laterally and directed laterally, situated between 2/10 and 3/10 of body length (SE 25% of BL). Distance between eyes wide (IOD 68% of BW). Nares small (ND 1.4% of BL), round, countersunk, positioned low (NH 33% of BH) laterally and oriented ventrally, situated nearer to snout than eye (RN 42% of NP) and below eye level (NH 55% of EH). Distance between nares moderately wide (IND 56% of IOD). Dark spot posterior to nares absent, other ornamentation absent. Spiracle sinistral, moderately large (SL 14% of BL), directed posteriorly, visible from ventral view and conspicuous from lateral view. Its inner wall absent and its aperture opens posteriorly. Opening elliptical, situated at 3/5 of body length (SS 60% of BL), located low on body (SH 39% of BH) and below height of contact of axis of tail myotomes with body (SH 61% of HAB). Vent tube medial, moderately long (VL 14% of BL), not attached to ventral fin. No dorsolateral glands visible. Tail very long (TAL 305% of BL). Maximal tail height higher than body height (MTH 115% of BH). Tail height at mid-tail higher than body height but lower than maximal tail height (THM 108% of BH and THM 94% of MTH). Tail height at beginning of tail lower than body height (TH 66% of BH). Caudal musculature well developed (TMW 49% of BW, TMH 66% of BH, TMH 100% of TH and 58% of MTH, TMHM 72% of THM, TMHM 67% of MTH). Tail muscle reaches tail tip. Fins very low (DF 18% of TMHM, VF 23% of TMHM), dorsal fin lower than ventral fin at mid-tail (DF 81% of VF). Dorsal fin originates on anterior 1/10 of tail musculature, rises progressively to attain its maximal height at maximal tail height, and then descends slightly towards tail tip. Ventral fin originates on caudal musculature just behind vent tube, increases gradually to attain maximal height at maximal tail height, and then declines towards tail tip. Maximal tail height located between 3/5 and 4/5 of tail length (DMTH 69% of TAL). Caudal vein conspicuous all along tail, myosepta perceptible on anterior 1/2 of tail musculature. Point of contact of axis of tail myotomes with body located on upper half of body height (HAB 65% of BH), axis of tail myotomes parallel with axis of trunk. Tail tip narrowly rounded. Oral disc very small (ODW 15% of BW), positioned and directed ventrally, not visible from dorsal view, not connected to snout. Oral disc opening triangular. Lower labium absent, upper labium folded to form a triangular opening. Two protuberances on each base of upper labium. Papillae, jaw sheaths and keratodonts absent.

Colouration in preservative

Predominantly pale brownish. Light, pale brown melanophoric pigment covers dorsum. Dark brown patches scattered irregularly on skin and condensed to form dark patches especially above neurocranium and whole dorsum. Laterally, jugal area and flank covered by light brown condensed reticulations, mainly between eye and spiracle, leaving obvious transparent spiracle on pale body wall. Tail musculature overlain by dense light brown reticulations leaving conspicuous lateral tail vein all along tail. Fins pale and unpigmented. Ventrally, oral disc, gular and branchial regions beige with a few blotches; venter covered by light brown condensed reticulations. Intestinal coils not visible.

Variation

All 19 non-DNA-voucher specimens of this series show the same external morphology as the voucher specimen.

Gephyromantis sculpturatus (Ahl, 1929)  

( Figures 1B View Figure 1 , 3A View Figure 3 , 4 View Figure 4 )

Material examined

The following description refers to one tadpole in developmental stage Gosner 39 (field number ZCMV 4833 ZSM 16 View Materials /2008, BL 6 mm, TL 18 mm, accession number HQ188940 View Materials ) collected by R. D. Randrianiaina, T. Rasolonjatovo H., S.H. Ndriantsoa, E. Reeve, A. Strauss and J. Glos on 11 February 2007 in Ranomafana National Park at Piste X 175 site (21 ◦ 15.846’ S, 47 ◦ 25.161’ E, 966 m a.s.l.). The 16S rRNA sequence of this specimen was 100% identical to a reference sequence of a G. sculpturatus   adult specimen (accession AY848432 View Materials ) from the same locality GoogleMaps   .

Description

In dorsal view, body elliptical. Maximal body width attained at mid-body (SBW 50% of BL), snout pointed. In lateral view, body depressed (BW 138% of BH), maximal body height attained between 3/5 and 4/5 of body length (SBH 63% of BL), snout broadly rounded. Eyes large (ED 19% of BL), visible from ventral view, positioned high (EH 64% of BH) laterally, directed laterally, situated between 2/10 and 3/10 of body length (SE 24% of BL). Distance between eyes wide (IOD 74% of BW). Nares small (ND 1.7% of BL), round, countersunk, positioned moderately high (NH 41% of BH) laterally, oriented ventrally, situated nearer to snout than eye (RN 34% of NP) and below eye level (NH 63% of EH). Distance between nares moderately wide (IND 47% of IOD). Dark spot posterior to nares absent, other ornamentation absent. Spiracle sinistral, small (SL 8% of BL), directed posterodorsally, visible neither from ventral nor from dorsal view, but perceptible laterally. Inner wall absent. Opening round, situated between 3/5 and 4/5 of body length (SS 63% of BL), located low on body (SH 36% of BH) and below height of contact of axis of tail myotomes with body (SH 56% of HAB). Vent tube medial, moderately long (VL 11% of BL), not attached to ventral fin. No dorsolateral glands visible. Tail very long (TAL 303% of BL). Maximal tail height higher than body height (MTH 106% of BH). Tail height at mid-tail almost equal to body height and as high as maximal tail height (THM 105% of BH and THM 100% of MTH). Tail height at beginning of the tail lower than body height (TH 80% of BH). Caudal musculature well developed (TMW 53% of BW, TMH 70% of BH, TMH 88% of TH and 66% of MTH, TMHM 66% of THM, TMHM 66% of MTH). Tail muscle reaches tail tip. Fins very low (DF 20% of TMHM, VF 32% of TMHM), dorsal fin lower than ventral fin at mid-tail (DF 61% of VF). Dorsal fin originates on the anterior 1/4 of tail, increases progressively to attain maximal height at maximal tail height, and then descends slightly towards tail tip. Ventral fin originates on caudal musculature just behind vent tube, increases gradually to attain maximal height at maximal tail height, and then declines towards tail tip. Maximal tail height located at 3/5 of tail length (DMTH 60% of TAL). Caudal vein and myosepta not visible. Point of contact of axis of the tail myotomes with body located on upper half of body (HAB 65% of BH), axis of tail myotomes parallel with axis of trunk. Tail tip narrowly rounded. Oral disc very small (ODW 8% of BW), elliptical, positioned and directed ventrally. Four small papillae present, two each ventrolaterally of oral disc opening. Inner papillae larger (0.05 mm) than outer papillae (0.03 mm). Jaw sheaths and keratodonts absent.

Colouration in life ( Figure 3A View Figure 3 )

Typically yellowish. Dorsal skin covered by brown variegated melanophores and some silver iridophoric spots or patches. Dorsolateral and lateral regions with same pattern as dorsal. Non-pigmented spiracle perceptible. Tail musculature yellowish with irregular brown blotches grouped to form irregular patches; their density increases towards tail tip. Sporadic silver iridophoric blotches present. Fins transparent; dorsal fin with many dark patches, ventral unpigmented. Ventral side covered by silver iridophoric blotches, mainly on the venter. Oral disc and gular region transparent; branchial area reddish and beating heart visible. Venter yellow with some brown and iridophoric blotches on skin. Intestinal coils not visible.

Colouration in preservative

Largely brownish. Light, pale brown melanophoric pigment covers dorsum. Dark brown patches scattered irregularly on skin and condensed to form dark patches especially above neurocranium and vertebral region. Laterally, jugal area and flank with sporadic light brown blotches leaving a noticeable pale transparent spiracle on pale body wall. Tail musculature overlain with scattered light brown reticulations. Fins pale and with reticulations, mainly close to tail tip. Ventrally, oral disc, gular and branchial regions pale; venter covered by light brown reticulations, no intestinal coils visible.

Gephyromantis tschenki (Glaw and Vences, 2001)  

( Figures 1C View Figure 1 , 3B View Figure 3 , 5 View Figure 5 )

Material examined

The following description refers to one tadpole in developmental stage Gosner 35, Del Pino and Escobar 23–25 (field number ZCMV 4335 ZSM 142 View Materials /2007, BL 4.6 mm, TL 16.7 mm, accession number GU975156 View Materials ) collected by R. D. Randrianiaina, T. Rasolonjatovo H., S.H. Ndriantsoa, E. Reeve, A. Strauss and J. Glos on 25 February 2007 in Ranomafana National Park at Bibiango site (21 ◦ 15.442’ S, 47 ◦ 25.096’ E, 962 m a.s.l.). The 16S rRNA sequence of this specimen was 98.5% identical to a reference sequence of a G. tschenki   adult specimen (accession AY848374 View Materials ) from the same locality GoogleMaps   .

Description

In dorsal view, body elliptical, maximal body width attained between 3/5 and 4/5 of body length ( SBW 64% of BL), snout broadly rounded. In lateral view, body depressed ( BW 117% of BH), maximal body height attained at 3/5 of body length ( SBH 60% of BL), snout pointed. Eyes large ( ED 17% of BL), visible from ventral view, positioned high ( EH 63% of BH) laterally, directed laterally, situated between 1/10 and 2/10 of body length ( SE 19% of BL). Distance between eyes wide ( IOD 62% of BW). Nares small (ND 2% of BL), round, countersunk, positioned moderately high (NH 41% of BH) laterally, oriented ventrally, situated nearer to snout than to eye ( RN 55% of NP) and below eye level (NH 65% of EH). Distance between nares wide ( IND 65% of IOD). Dark spot posterior to nares absent, other ornamentation absent. Spiracle sinistral, small (SL 9% of BL), directed posterodorsally, visible in ventral view and perceptible laterally. Its inner wall absent. Opening elliptical, situated at mid-body (SS 50% of BL), located low on body (SH 38% of BH) and below height of contact point of axis of tail myotomes with body (SH 64% of HAB). Vent tube medial, long ( VL 18% of BL), not attached to ventral fin. No dorsolateral glands visible. Tail very long ( TAL 303% of BL), maximal tail height as high as body height ( MTH 100% of BH), tail height at mid-tail lower than body height and maximal tail height ( THM 87% of BH and THM 88% of MTH), tail height at beginning of tail lower than body height (TH 73% of BH). Caudal musculature well developed (TMW 47% of BW, TMH 60% of BH, TMH 81% of TH and 60% of MTH, TMHM 68% of THM, TMHM 60% of MTH). Tail muscle reaches tail tip. Fins very low (DF 25% of TMHM, VF 22% of TMHM), dorsal fin higher than ventral fin at mid-tail ( DF 114% of VF). Dorsal fin originates after dorsal body–tail junction, increases progressively to attain its maximal height at maximal tail height, and then descends slightly towards tail tip. Ventral fin originates on caudal musculature just behind vent tube, increases gradually to attain its maximal height at maximal tail height, and then declines towards tail tip. Maximal tail height located between 3/5 and 4/5 of tail length ( DMTH 69% of TAL). Caudal vein visible on anterior 3/4 of tail, myosepta invisible. Point of contact of axis of tail myotomes with body located in upper half of body ( HAB 60% of BH), axis of tail myotomes not parallel with axis of trunk. Tail tip narrowly rounded. Oral disc very small ( ODW 15% of BW), positioned and directed ventrally, not visible from dorsal view and not connected to the snout. Oral disc opening triangular, lower labium absent and upper folded to form a triangular opening. Papillae, jaw sheaths and keratodonts absent.

Colouration in life ( Figure 3B View Figure 3 )

Typically yellowish. Dorsal skin covered by brown variegated melanophores. Dorsolateral and lateral regions with same pattern as dorsal part. Slightly unpigmented spiracle perceptible. Tail musculature yellowish with irregular brown blotches grouping to form irregular networks. Their density increases towards tail tip. Fins yellow, reticulated. Ventrally, oral disc and gular region whitish with brown blotches, branchial area reddish and beating heart visible, venter yellow. Intestinal coils not visible.

Colouration in preservative

Largely brownish. Brown melanophoric pigment covers dorsum. Dark brown patches scattered irregularly across the skin and condense to form larger dark patches especially above neurocranium, vertebral and abdominal regions. Laterally, jugal area and flank with sporadic light brown blotches condensing to form reticulations. Spiracle difficult to discern. Tail musculature overlain by scattered light brown reticulations leaving lateral tail vein perceptible all along tail. Fins pale and reticulated, mainly close to tail tip. Ventrally, oral disc, gular and branchial regions pale with sporadic brown reticulations; venter pale, no intestinal coils visible.

Gephyromantis ventrimaculatus (Angel, 1935)  

( Figures 1D View Figure 1 , 3C View Figure 3 , 6 View Figure 6 )

Material examined

The following description refers to one tadpole in developmental stage Gosner 39 (field number ZCMV 4927 ZSM 852 View Materials /2007, BL 6.4 mm, TL 20.4 mm, accession number GU975158 View Materials ) collected by R. D. Randrianiaina, T. Rasolonjatovo H., S.H. Ndriantsoa, E. Reeve, A. Strauss and J. Glos on 2 March 2007 in Ranomafana National Park at Sahateza site (21 ◦ 15.453’ S, 47 ◦ 21.609’ E, 1164 m a.s.l.). The 16S rRNA sequence of this specimen was 100% identical to a reference sequence of a G. ventrimaculatus   adult specimen (accession FJ559200 View Materials ) from Ranomafana (Ranomafanakely) GoogleMaps   .

Description

In dorsal view, body elliptical, maximal body width attained between proximal 2/5 and 3/5 of the body ( SBW 45% of BL), snout broadly rounded. In lateral view, body depressed ( BW 117% of BH), maximal body height attained between 3/5 and 4/5 of body length ( SBH 65% of BL), snout rounded. Eyes large ( ED 16% of BL), visible from ventral view, positioned moderately high ( EH 59% of BH) laterally and directed laterally, situated between 2/10 and 3/10 of body length ( SE 23% of BL). Distance between eyes wide ( IOD 76% of BW). Nares small (ND 1.7% of BL), round, countersunk, positioned moderately high (NH 45% of BH) laterally and oriented ventrally, situated nearer to snout than to eye ( RN 86% of NP) and below eye level (NH 75% of EH). Distance between nares moderately wide ( IND 48% of IOD). Dark spot posterior to nares absent, other ornamentations absent. Spiracle sinistral, small (SL 6% of BL), directed dorsally, visible from ventral and lateral views, inner wall free from body, aperture opens laterally. Opening ovoid, situated almost at mid-body length (SS 54% of BL), located low on body (SH 32% of BH) and below the height of the contact point of axis of tail myotomes with body (SH 54% of HAB). Vent tube medial, short ( VL 10% of BL), not attached to ventral fin. No dorsolateral glands visible. Tail very long ( TAL 316% of BL). Maximal tail height as high as body height ( MTH 97% of BH). Tail height at mid-tail almost equal to body height and maximal tail height ( THM 96% of BH and THM 98% of MTH). Tail height at beginning of tail lower than body height (TH 62% of BH). Caudal musculature well developed (TMW 54% of BW, TMH 59% of BH, TMH 94% of TH and 60% of MTH, TMHM 72% of THM, TMHM 71% of MTH). Tail muscle reaches tail tip. Fins very low (DF 18% of TMHM, VF 20% of TMHM), dorsal fin lower than ventral fin at mid-tail ( DF 86% of VF). Dorsal fin originates on dorsal body–tail junction, rises progressively to attain its maximal height at maximal tail height, and then descends slightly towards tail tip. Ventral fin originates on caudal musculature just behind ventral terminus of body, increases gradually to attain maximal height at maximal tail height, and then declines towards tail tip. Maximal tail height located between 3/5 and 4/5 of tail length ( DMTH 64% of TAL). Caudal vein and myosepta visible all along tail. Point of contact of axis of the tail myotomes with the body located on the upper half of the body ( HAB 59% of BH), axis of tail myotomes not parallel with axis of trunk. Tail tip rather rounded. Oral disc very small ( ODW 19% of BW), positioned and directed ventrally. Oral disc opening elliptical. Papillae, jaw sheaths and keratodonts absent.

Colouration in life ( Figure 3C View Figure 3 )

Typically black, covered by scattered silver iridophoric pigments.

Colouration in preservative

Largely black, area occupied by iridophoric pigment gives light pattern. Spiracle perceptible, intestine not visible.

Gephyromantis sp.   aff. blanci  

( Figures 7 View Figure 7 , 8 View Figure 8 )

Material examined

These data refer to a population of small terrestrial and diurnal frogs from Ranomafana National Park, considered as G. blanci   by Vieites et al. (2009). However, our own unpublished molecular and bioacoustic data indicate that in fact this population represents an undescribed candidate species that we here refer to as G. sp. aff. blanci   . A clutch of four eggs was collected by K.C. Wollenberg on 5 March 2007 in Ranomafana National Park, at a site locally known as Ranomafanakely (21 ◦ 14.921’ S, 47 ◦ 22.307’ E, 1134 m a.s.l.). Weather conditions were moist with constant rain at the time when the clutch was found. The site contained a forested slope, overgrown with lianae and moss. On the bottom of the slope, a ca. 10 cm thick layer of leaf litter covered the forest floor. Many males of G. enki   were calling from there. At the more elevated positions of the slope, dead wood overgrown with moss was under the leaf litter. It was forming a thick, porous layer with many cavities of up to 1 m depth. The clutch was found on such a cavity, overgrown with moss but partially exposed to daylight and attached to the surface of a dead leaf. Rainwater was dripping on the clutch from the moss layer above it. The clutch was found while searching for a male specimen of G. sp. aff. blanci   that was calling approximately 30 cm from the clutch from an elevated position. Other specimens of G. sp. aff. blanci   were heard calling from other sites further up the slope. We suspect that the eggs might have been guarded by the male, as this behaviour has been observed in other species of the G. boulengeri   group (K.C. Wollenberg personal observations). The clutch was taken to the lab, and one egg was immediately removed and preserved with field number ZCMV 5253 in 90% ethanol for species identification by DNA barcoding (accession number HQ188941 View Materials ). The remaining three eggs were left on the leaf, kept in a terrarium with leaf litter, and watered regularly. Their development was followed for 24 days until the last metamorph left the clutch. A single egg had a diameter of about 3 mm, the yolk being pale yellow in a transparent jelly.

Development of embryos and larvae ( Figure 7 View Figure 7 )

The observation started on day 1 after collection (6 March). On this day, the embryos were visible and were a pale yellowish colour. The outline of the tail was starting to develop. They were developmental stage 17–18 according to Gosner (1960) or 13 following Del Pino and Escobar (1981).

On day 3, the embryos were weakly pigmented and the outline of the eye was visible.

On day 4, TAL reached about 120% of BL, ED about 8% of BL, and the tail had transparent fins. The beating heart was visible and the cornea was transparent. There was clear tail elongation allowing first movements (hatchling stage according to Gosner 1960; staging after DelPino and Escobar 1981 not applicable). Arteries across the yolk were formed.

On day 7, TAL was about 170% of BL, ED about 15% of BL. The toes began to differentiate and develop (indentation 4–5). The vitelline volume decreased. The cornea was visible and the dorsal body was pigmented.

On day 10, TAL was 193% of BL, ED 17% of BL. There were indentations in toes 2–3, the vitelline reduced by about 1/3 compared to day 7. The nares and the blood vessel in the posterior part of the caudal musculature were visible. The body was more pigmented and the jelly of the two tadpoles was united.

On day 11, TAL was about 180% of BL, ED 16% of BL. The transverse muscular structure was visible. The pigmentation increased laterally to ventrally and there were indentations in toes 1–2.

On days 13 and 14 (18 and 19 March), TAL was 177% of BL, ED 9% of BL, TAL 211% of BL and ED 15% of BL. Toes 3–5 were separated.

On day 16, TAL was 200% of BL, ED 12% of BL. All toes were separated and the outline of the mouth was visible.

On day 17, TAL 177% of BL, ED 15% of BL.

On days 19 and 20 TAL 210% of BL, ED 16% of BL, TAL 180% of BL and ED 15% of BL. The feet tubercles were visible.

On day 24, the last hatchling left the jelly with four well-developed legs; the tail length was about 126% of BL.

Description of metamorphosed froglet

The following description refers to one metamorphosed froglet in Gosner stage 44 ( ZSM 649/2008, BL 5.3 mm TL 9 mm) from the batch described above. In dorsal view, body elliptical, maximal body width attained between 3/5 and 4/5 of body length ( SBW 64% of BL), broadly rounded snout. In lateral view, body depressed ( BW 112% of BH), maximal body height attained between 3/5 and 4/5 of the body length ( SBH 72% of BL), snout round. Eyes large ( ED 17% of BL), visible from ventral view, positioned moderately high ( EH 52% of BH) laterally and directed laterally, situated between 2/10 and 3/10 of body length ( SE 22% of BL). Distance between eyes wide ( IOD 87% of BW). Nares small (ND 1.9% of BL), round, countersunk, positioned low (NH 34% of BH) laterally and oriented ventrally, situated nearer to snout than eye ( RN 70% of NP). Distance between nares moderately wide ( IND 58% of IOD). Dark spot posterior to nares absent, other ornamentation absent. Spiracle, vent tube not visible. Tail largely resorbed, therefore very short ( TAL 126% of BL). Mouth opening moderately large ( ODW 42% of BW), not connected to snout, positioned and directed ventrally, already being transformed into a frog mouth structure. Yellowish structure, probably the tongue, visible inside the mouth. All typical structures of tadpole oral disc absent.

Colouration in preservative

General colouration yellowish. Body dorsum, laterally, and abdominal surface covered by light brown reticulations. Gular and branchial regions brown, intestinal coils not visible, tail musculature covered by some blotches.

R

Departamento de Geologia, Universidad de Chile

AJ

Central Research Laboratories

T

Tavera, Department of Geology and Geophysics

IND

Indiana University

TAL

Jardin botanique de Talence

TMH

Tasmanian Museum and Art Gallery

DMTH

Britannia Royal Naval College

ZSM

Bavarian State Collection of Zoology

Kingdom

Animalia

Phylum

Chordata

Class

Amphibia

Order

Anura

Family

Mantellidae

Genus

Gephyromantis