Comparative larval morphology in Madagascan frogs of the genus Guibemantis (Amphibia: Mantellidae) Author Vejarano, Silvia Author Thomas, Meike Author Vences, Miguel text Zootaxa 2006 1329 39 57 journal article 10.5281/zenodo.174179 d0606cb6-b406-40d1-8bfc-f023cebeca32 1175-5326 174179 Guibemantis liber (Peracca) , central-eastern Madagascar Series examined: ZSM 466–467/2004 (field number 2002.2014; Genbank accession number DQ054499 ), series of 14 specimens , ZSM 468/2004 (fieldnumber 2002.2016; DQ054500 ) one specimen, ZSM 469/2004 (fieldnumber 2002.2017; DQ054501 ), one specimen, ZSM 470/2004 (fieldnumber 2002.2019; DQ054502 ), series of two specimens and ZSM 659/2004 (fieldnumber 2002.1610; DQ054503 ) series of 34 specimens , all from An'Ala forest ( 18°56'S , 48°28'E , 840 m above sea level), collected in different shallow ponds not deeper than 50 cm with swampy and leaf-covered grounds on 1 March 2003 . Description (based on the DNA voucher of series ZSM 466-467/2004). Tadpole in stage 34. Coloration in preservative light brown without any distinct pattern. The intestinal wall also pigmented and intestinal spiral visible. Ventrolateral and ventral parts of the body lack pigmentation. Anterior part of the tail with the same coloration of the body. Most of the tail and caudal fin missing because of tissue sampling. Tadpole type IV ( Orton 1953 ). Body shape oval. Snout in dorsal view almost rounded. Eyes small (about 0.06 of body length) dorsally positioned (fig. 1a and b). Spiracle sinistral. Oral disc laterally emarginated, and ventrally positioned (fig. 1c). Dorsal gap of the marginal papillae 1.03 mm . Oral disc bordered by 46 short marginal papillae more or less equal in length (˜ 0.05 mm ). No submarginal papillae. LTRF 1: 3+3/3 ( Dubois 1995 ) or 4(2–4)/3(–) ( Altig & McDiarmid 1999 ). The lower lip and tooth rows possibly damaged by fixation or preservation; LTR1 continues on the left side until it meets LTR3 and appears as one single continuous row; LTR2 incomplete on the left side. UTR ca. 52–80 teeth per mm and LTR ca. 13–16 teeth per mm. From UTR2 on, they are separated by the upper jaw sheath. Jaw sheaths distinct and black, with serrated edges (ca. 55 on each jaw). Sheath of upper jaw not curved in the middle, lower jaw sheath V-shaped. Variation within the series. Series ZSM 466–467/2004 with 13 additional tadpoles in stages 27–40. Number of marginal papillae variable (31–64); submarginal papillae 0–3 on each side of the upper lip and 1–4 on the lower lip, except one specimen with seven on the right side of the lower lip. LTRF 3(2–3)/3(1) or 4(2–3)/3(1), independent from the stage. In the remaining specimens, general and oral morphology very similar but certainly different from the DNA voucher specimen in the oral disc (differences might be artefacts). Series ZSM 468/2004 and 469/2004 only consisting of their respective DNA voucher specimen. Series ZSM 470/2004 with two specimens, the DNA voucher in stage 27 and a second specimen in stage 31 that has LTRF 5(2–5)/3(1), 53 marginal and three submarginal papillae on the left side of the upper and lower lips and one on the right side. Series ZSM 659/2004 with 34 tadpoles in stages 28–41. LTRF 4(2–4)/3(1) in most specimens except five with LTRF 3(2–3)/3(1) and one with 5(2–5)/3(1) in which UTR5 has only four teeth on each side. In two specimens the oral disc is not in a good state of preservation, and misses parts of the lips or tooth rows. Coloration varies between individuals from light to dark brown, but the pattern is the same as in the DNA voucher of series ZSM 466–467/2004. Marginal papillae 32–65, but in most specimens 51–59. Number of submarginal papillae on each side of the upper lip usually 2–3, lacking in some specimens, and four in other individuals; number in the lower lip varies more, 1–8 on each side. This series is very homogeneous, in spite of the large number of specimens and their different developmental stages. Morphometric data of all DNA vouchers of G. l i b e r are shown in tables 1 and 2. TABLE 1. Morphometric data (all in mm) of DNA voucher specimens of Guibemantis liber . * DNA vouchers used for descriptions and drawings. ** No oral disc morphology data because of advanced developmental stage. Central-Eastern Madagascar LTR 1 2.5 0.69 0.40 0.68

* ZSM 466-467/2004	ZSM 659/2004	ZSM 470/2004	ZSM 468/2004	** ZSM 469/2004
Stage	34	36	27	34	43
BL	8.1	10.00	6.50	9.00	10.00
BW	5.0	5.50	4.00	5.00	4.00
ODW	1.9	2.58	1.43	1.98	2.65

DGMP	1.1	1.35	1.08	1.33	1.38
IOD	2.9	3.69	2.44	2.50	4.00
ED	0.6	1.38	0.75	1.06	1.75
TH	2.7	1.56	2.25	2.56	3.00
TMW	2.1	2.63	1.75	2.03	2.00
MP	0.05	0.13	0.10	0.13	0.05
LTRF	4(2–4)/3(?)	3(2–3)/3(1)	3(2–3)/3(1)	4(2–4)/3(1)
UTR1 UTR2 UTR3 UTR4	1.3 0.4 0.2 0.08	1.69 0.81 0.25 -	0.90 0.30 0.18 -	1.78 0.78 0.38 0.18

LTR 2 2.6 1.63 1.05 1.18 LTR 3 4.0 1.38 0.75 0.80 LLTR3 0.125 0.050 0.050 LUTR1 0.075 0.063 0.075 to be continued. TABLE 1 (continued).

South-Eastern Madagascar
* ZSM 433/2004 ZSM 452/2004	ZSM 451/2004	ZSM 449/2004
Stage	33 27	27	27
BL	8.3 5.50	5.50	6.00
BW	6.1 3.50	3.50	3.00
ODW	1.9 1.45	1.50	1.58
DGMP	1.2 0.88	0.75	1.00
IOD	1.7 1.69	1.93	1.90
ED	1.0 0.69	0.73	0.68
TH	2.2 0.90	0.73	0.73
TMW	2.2 0.65	0.48	0.48
MP	0.13 0.10	0.08	0.10
LTRF	4(2–4)/3(1) 2(2)/3(1)	2(2)/3(1)	2(2)/3(1)
UTR1	1.6 1.05	0.95	0.93
UTR2	0.6 0.39	0.30	0.30
UTR3	0.3 -	-	-
UTR4	0.2 -	-	-
LTR 1	0.7 0.41	0.40	0.40
LTR 2	1.3 0.85	0.88	0.98
LTR 3	1.2 0.70	0.65	0.80
LLTR3	0.025	0.025	0.075
LUTR1	0.050	0.050	0.075

Variation among the series. Complete morphometric data of all DNA voucher specimens are shown in table 1. DNA voucher of series ZSM 468/ 2004 in stage 34; LTRF 4 (2–4)/3(1); 45 marginal papillae; 1–2 submarginal papillae on each side of the upper lip and one on each side of the lower lip; in coloration and general morphology agreeing with the DNA voucher specimen of series ZSM 466–467/2004. DNA voucher of series ZSM 469/ 2004 in stage 43. Oral disc and mouth parts modified. Dorsal coloration of the body agreeing with the DNA voucher specimen of series ZSM 466–467/2004; tail and posterior legs with dark brown spots. DNA voucher of series ZSM 470/ 2004 in stage 27; LTRF 3(2–3)/3(1); 44 marginal papillae; two submarginal papillae on each side of the lower lip and one on the right side of the upper lip. Coloration of the body brown, but softer and lighter than in the DNA vouchers of all other series. DNA voucher of series ZSM 659/ 2004 in stage 36. LTRF 3(2–3)/3(1); 51 marginal papillae; two on each side of the upper lip and four on each side of the lower lip. Coloration of the body dark brown. In other characteristics, as far as recognizable, there are no relevant differences among the DNA voucher specimens of all series. TABLE 2. Number of marginal and submarginal papillae (MP, SMP) and number of teeth per row of all DNA voucher specimens of Guibemantis liber . For rows divided by a gap, the number of teeth presented on the table is the average of the numbers on each side of the tooth row. * DNA vouchers used for descriptions and drawings. ** No oral disc morphology data because of advanced developmental stage. Central-Eastern Madagascar UTR 2 26 29 25 42

* ZSM 466-467/2004	ZSM 659/2004	ZSM 470/2004	ZSM 468/2004	** ZSM 469/2004
MP	46	51	44	45	34
SMP	0	11	5	5	5
UTR1	68	99	62	104

UTR 3 16 18 11 21 UTR4 6 - - 9 LTR 1 33 39 28 28 LTR 2 41 89 66 66 LTR 3 54 68 48 48 continued. South-Eastern Madagascar * ZSM 433/2004 ZSM 452/2004 ZSM 451/2004 ZSM 449/2004 MP 52 36 35 40 SMP 39 7 6 8 UTR1 68 47 40 45 UTR 2 25 16 13 15 UTR3 12 - - - UTR4 6 - - - LTR 1 29 22 18 19 LTR 2 90 43 38 39 LTR 3 88 34 29 37 Previous descriptions. The tadpole of G. l i b e r has been described by Blommers- Schlösser (1975) under the name Gephyromantis liber : 68 tadpoles collected in Andasibe, the Mandraka valley, and Tampoketsa d’Ankazobe (all localities in central eastern Madagascar ) were examined at that time, and the only noticeable difference between the stages was the size (body length 3.3–13 mm ); general body shape was as a typical type IV tadpole ( Orton 1953 ), the oral disc emarginated, directed downwards and surrounded by small papillae except the dorsal margin. LTRF was variable between 4(2–4)/3(1), 3(2–3)/ 3(1) and 5(2–5)/3(1).