Acta Herpetologica 11(2): 119-125, 2016 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-18009 The unexpectedly dull tadpole of Madagascar’s largest frog, Mantidactylus guttulatus Arne Schulze1,*, Roger-Daniel Randrianiaina2,3, Bina Perl3, Frank Glaw4, Miguel Vences3 1 Hessisches Landesmuseum Darmstadt (HLMD), Friedensplatz 1, 64283 Darmstadt, Germany. *Corresponding author. E-mail: arne. schulze@hlmd.de 2 Département de Biologie Animale, Université d’Antananarivo, BP 906, Antananarivo 101, Madagascar 3 Division of Evolutionary Biology, Zoological Institute, Technical University of Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany 4 Zoologische Staatssammlung München (ZSM-SNSB), Münchhausenstr. 21, 81247 München, Germany Submitted on 2016, 11th February; revised on 2016, 13th April; accepted on 2016, 9th July Editor: Marco Sannolo Abstract. The Madagascar-endemic mantellid genus Mantidactylus contains one subclade with two described frog species characterized by very large body sizes. This subclade is classified as the subgenus Mantidactylus and is wide- spread in eastern and northern Madagascar, but their reproductive biology and larval stages are still unknown. We here provide a detailed description of the larvae of one species in this subgenus, M. guttulatus, on the basis of genetic assignment (16S DNA barcoding). The tadpoles were collected in the dry season from shallow waters near a stream in the Mahajanga Province in northwestern Madagascar. Their body and tail shape is remarkably generalized as typical for stream-adapted tadpoles, and the oral disc and labial keratodont row formula (4(2-4)/3(1)) are similar to those of other lotic mantellid frog larvae with generalized mouthparts like those in the subgenus Brygoomantis. The well-sep- arated positions of these subgenera in the mantellid phylogeny suggest extensive homoplasy in the evolution of larval mouthpart morphology within Mantidactylus. Keywords. Amphibia, Madagascar, Mantellidae, Mantidactylus, generalized oral disc, tadpole morphology. INTRODUCTION Among Madagascar’s native frogs, the family Man- tellidae is the most diverse clade with 212 named species (Amphibiaweb, 2016) and numerous undescribed spe- cies (Vieites et al., 2009; Perl et al., 2014). Mantellids are endemic to Madagascar and the Comoros and include a fascinating diversity in ecomorphology and reproduc- tive modes. The largest mantellids are classified in a well- supported subclade of the genus Mantidactylus (i.e., in the nominal subgenus Mantidactylus): Mantidactylus gut- tulatus, M. grandideri, and the candidate species M. sp. aff. grandideri “North”, although their alpha-taxonomy is in need of revision (see comments under Materials and Methods). With up to 120 mm snout-vent length M. gut- tulatus is the largest frog in Madagascar and is common in rainforest streams of the northern and eastern part of the island (Glaw and Vences, 2007). Despite their size and local abundance, informa- tion on the reproduction of this frog species is scarce and basically limited to one report of a calling specimen (Vences et al., 2004). Because for decades no tadpoles could be assigned to Mantidactylus guttulatus or its close relatives, it was assumed that these species lack a larval phase or that the pre-metamorphic tadpoles develop in a nesting burrow (Glaw and Vences, 1994, 2007). Altig and McDiarmid (2006) described a tadpole with reduced oral structures from the Ranomafana region and tentatively 120 Arne Schulze et alii assigned it as belonging to M. guttulatus. Randrianiaina et al. (2011) provided molecular evidence for an assign- ment of these tadpoles to M. majori whose juveniles are morphologically similar to those of M. guttulatus. During a survey in northwestern Madagascar, we obtained a small series of three Mantidactylus tadpoles that we initially identified by morphology as belonging into the subgenus Brygoomantis, despite being more elon- gated than other, syntopic Brygoomantis larvae. Molecu- lar evidence demonstrated that these tadpoles instead belonged to Mantidactylus guttulatus, and we provide a detailed description of their morphology. MATERIALS AND METHODS Three tadpoles (field numbers ZCMV 13332, 13333 and 13334) were collected by R.D. Randrianiaina, F.M. Ratsoavina, A.S. Rasamison, A. Rakotoarison, D.R. Vieites, and M. Vences in the dry season on 29 June 2010. They were found in an oppor- tunistic encounter survey near a large stream close to the Anal- amisondrotra mobile phone pylon, between 56-57 km along the national road N°31 from Bealanana to Antsohihy (14.72602°S, 048.55497°E; 1175 m a.s.l.) in the Mahajanga Province. Tadpoles were euthanized in a chlorobutanol solution shortly after collection. A tissue sample from the first third of the tail musculature of each tadpole was preserved in 99% etha- nol. After tissue sampling, all specimens were preserved in 5% formalin and two of them were deposited in the Zoologische Staatssammlung München, Germany (ZSM; collection numbers ZSM 704/2010, ZSM 705/2010). Tadpoles were identified by DNA barcoding based on a fragment of the mitochondrial 16S rRNA gene (Thomas et al., 2005). The fragment of about 550 bp was amplified with prim- ers 16Sar-L and 16Sbr-H from Palumbi et al. (1991) and stand- ard protocols resolved on automated sequencers were compared to a nearly complete database of sequences of adult Mala- gasy frog species. DNA sequences were deposited in GenBank (accession numbers KX023902, KX023903, KX023904). For species names in the subgenus Mantidactylus, we here follow the taxonomy suggested by Glaw and Vences (2007) who defined M. guttulatus as the species with a rather tuber- cular dorsum occurring mostly in northern Madagascar, and M. grandidieri as the species with smooth skin widespread in the southern and central east of the island. This differs from the definition of Altig and McDiarmid (2006) who applied the name M. guttulatus to populations from the southern central east. However, it is obvious that this taxonomy is in need of revision and it is likely that the available names (Rana guttula- ta Boulenger, 1881; Mantidactylus grandidieri Mocquard, 1895; and Rana pigra Mocquard, 1900, currently a synonym of M. guttulatus) will have to be applied in a different way to the bio- logical entities in the subgenus than in current practice. In fact, the tadpoles described herein might turn out to belong to a yet undescribed species. Independent from this taxonomic conun- drum, however, the subgenus Mantidactylus is well defined and the molecular data leave no doubt that the tadpoles described herein belong into this clade. A Canon DSLR with 100 mm 2.8L and MP-E 65 mm lenses mounted on an electronic-driven macro rail was used to obtain the digital images of the preserved specimens. A stack of 10-15 images was taken and merged with Helicon Pro software to achieve images with a wide depth of focus. Morphological descriptions and measurements were done on the basis of digital and scaled images of preserved tadpoles. Terminology of morphological characters follows Altig and McDiarmid (1999). Gosner’s (1960) classification was used to identify developmental stages. Structures of the oral apparatus were described according to Altig (1970), except for the term “keratodont,” which is used for the keratinized structures on the labia of the oral disc and presented as the labial keratodont row formula (LKRF). Marginal papillae are considered separately for the region of the upper labium, the lateral region, and the region of the lower labium and the “marginal papillae row for- mula” (MPRF) is provided according to Schulze et al. (2015). All morphological landmarks and distances considered for the description are described and specified in Table 1. Comparing measurements, we consider them as “almost equal” if ratios of the measured values are 95-96% or 104-105%, “equal” if they are in the range 97-103%, as almost “in the middle” if they are in the range 45-46% or 54-55% and “in the middle” if they are in the range 47-53% (Randrianiaina et al., 2011). RESULTS Three tadpoles identified by 16S DNA barcoding as Mantidactylus guttulatus were collected within rainforest and close (ca. 20 m) to a large stream of 25 m width. The tadpoles were in a seepage area of a small, very shallow puddle (1-2 cm deep) with a slow steady flow of water. The 16S rDNA sequences of these tadpoles were 99% identical to a reference sequence of an adult M. guttu- latus from the Tsaratanana Massif (GenBank accession no. FJ559237). The following description refers to one of these tadpoles in Gosner stage 26 (field number ZCMV 13332 / ZSM 704/2010, body length 9.5 mm, tail length 30.7 mm; Figs. 1, 2; Tab. 1). In dorsal view body ellipti- cal, maximal body width attained almost at mid-body length, snout narrowly rounded. In lateral view, body depressed, maximal body height attained between the 3/5 and 4/5 of the body length, snout rounded. Eyes moder- ately large, not visible from ventral view, positioned high laterally and directed anterolaterally, situated between the 2/10 and 3/10 of the body length. Distance between eyes wide. Nares rounded and small, marked with a mar- ginal rim, positioned moderately high dorsolaterally and directed anterolaterally, situated closer to snout than to eye and lower than eye. Distance between nares wide. Spiracle sinistrally positioned and short, directed pos- teriorly, visible from ventral view, invisible from dorsal 121Tadpole of Mantidactylus guttulatus view and perceptible from lateral view; posterior third of inner wall free from body and formed that aperture is lateroposteriorly directed, its opening rounded, narrower than tube, situated between the 2/5 and 3/5 of the body length, located low on the body at the height of the hind limb insertion. Long medial vent tube with dextral wall shorter than sinistral, causing a dextral directed open- ing, fully attached to ventral fin. Glands absent. Tail long, maximal tail height higher than body height, tail height at mid-tail higher than body height and as high as maxi- Table 1. Measurements of landmarks (in mm) and their ratios (in %) of the preserved tadpole specimen of Mantidactylus guttulatus (ZCMV 13332 / ZSM 704/2010) at Gosner stage 26: A1 = first upper keratodont row; BH = maximal body height; BL = body length; BW = maximal body width; DF = dorsal fin height at region of mid-tail; DG = size of the gap of marginal papillae in the region of the upper labium; DMTH = distance of maximal tail height from the tail-body junction; ED = eye diameter; HAB = height of the point where the axis of the tail myotomes contacts the body, measured from the lower curve of the belly; IND = inter-narial distance, measured from the centre of the eyes; IOD = inter-orbital distance; JW = maximal width of keratinized upper jaw sheath; MTH = maximal tail height; NH = naris height, measured from the lower curve of the belly to the centre of the naris; NP = naris-pupil distance; ODW = maximum width of opened oral disc; RN = rostro-narial distance, measured from the centre of the nares; SBH = distance between snout and the point of maximal body height; SBW = distance between snout and the point of maximal body width; SE = snout-eye distance, measured to the centre of the pupil; SH = spiracle height; SL = spiracle length, measured from its visible edges; SS = snout-spiracle distance, measured from the centre of the spiracle opening; TAL = tail length, measured from medium point of body-tail junction; TH = tail height at the body-tail junction; THM = tail height at mid-tail; TL = total length; TMH = tail muscle height at the body-tail junction; TMHM = tail muscle height at mid-tail; TMW = tail muscle width at the body-tail junction; VF = ventral fin height at mid-tail. Landmarks mm Ratio % BH 4.1 SBW - BL 57 BL 9.5 BW - BH 117 BW 4.8 SBW - BL 57 DF 0.9 ED - BL 11 DG 1.4 SE - BL 25 DMTH 7.5 IOD - BW 71 ED 1.0 ND - BL 3 EH 2.1 NH - BH 46 HAB 2.8 RN - NP 47 IND 2.3 IND - IOD 67 IOD 3.4 SL - BL 8 JW 1.2 SS - BL 53 MTH 4.8 SH - BH 34 NH 1.9 SH - HAB 50 NP 1.7 TAL - BL 213 ODW 2.5 MTH - BH 117 RN 0.8 THM - BH 117 SBH 6.5 THM - MTH 100 SBW 5.4 TH - BH 88 SE 2.4 TMW - BW 54 SH 1.4 TMH - BH 63 SL 0.8 - MTH 54 SS 5.0 TMHM - THM and MTH 60 TAL 20.3 DF - TMHM 31 TH 3.6 VF - TMHM 38 THM 4.8 DF - VF 82 TL 30.7 DMTH - TAL 37 TMH 2.6 HAB - BH 68 TMHM 2.9 ODW - BW 52 TMW 2.6 DG - ODW 56 VF 1.1 A1 - ODW 83 JW - ODW 48 122 Arne Schulze et alii mal tail height, tail height at body-tail junction lower than body height. Caudal musculature well developed. Tail muscle reaches tail tip. Tail fins very low, dorsal fin slightly lower than ventral fin at mid-tail, but slightly higher in posterior third. Dorsal fin originates slightly behind dorsal body-tail junction, with shallow, gradually rising until the anterior 1/3 of the tail where it increases brusquely to attain its maximal height behind mid-tail and then continues gradually until the posterior 3/4 of the tail where it descends abruptly towards the tail tip. Ventral fin originates at the ventral terminus of the body, rises meticulously until the anterior 1/4 of the tail, and then remains almost parallel to the ventral border of the tail muscle until close to the tail tip. Maximal tail height located behind mid-tail, lateral line vein and myosepta imperceptible, point where the axis of the tail myotomes contacts the body located in the upper half of the body height, axis of the tail myotomes parallel with the axis of body length. Tail tip narrowly rounded. Moderately wide generalized oral disc, positioned almost ventrally and directed anteroventrally, clearly laterally emargin- ated. Oral disc not visible from dorsal view, upper labium as a continuation of snout. Marginal papillae uniseri- ate and interrupted by a wide gap on the upper labium, gap on the lower labium absent, total number of mar- ginal papillae 48 (MPRF: (1)/1/1). Sixteen submarginal papillae present (8 on each side of the jaw sheaths folds). LKRF 4(2-4)/3(1), A1 keratodont row very long. Density of keratodonts varies from 20/mm to 71/mm, A1 59/mm (total 118). Gap in the A2 row narrow (>1% of A2 row) and distinctly wider in A3 and A4. Gap in the P1 row less than the width of three keratodonts. Alignment of anteri- or and posterior rows regular and nearly of same length. Distal keratodonts of same length as those in the centre; prominent space between marginal papillae and kera- todont rows. Jaw sheaths partially keratinized, only the half section close to the edge coloured black; with finely pointed serrations. Upper jaw sheath moderately wide and slightly arched, with a very shallow medial concav- ity. Lower jaw sheath V-shaped, partially keratinized and partially hidden by the upper jaw sheath when closed. Colouration in life uniformly dark brownish. Dor- sally, body covered by homogeneous dark brown mel- anophoric pigments. Laterally, area below eyes, flank, and abdominal region densely reticulated. Ventrally, oral disc and gular region reticulated, branchial regions red- dish and spotted, beating heart visible; venter transparent, regularly spiralled intestinal coils visible. Tail musculature yellowish coloured, and coarsely reticulated. Fins patched with dark small spots with fringy edges. Fig. 1. Images of the living tadpole specimen of Mantidactylus guttulatus (ZCMV 13332 / ZSM 704/2010) at Gosner stage 26; a-c), in dor- sal, lateral and ventral view (scale bar = 10 mm). 123Tadpole of Mantidactylus guttulatus Colouration in preservative uniformly brownish col- oured. Brown melanic pigment in layers deeper than the skin covered the dorsum and flank, leaving laterally a slightly transparent area. Some dark brown blotches scat- tered on the dorsum skin, condensed to form dark patch- es above the brain and the vertebral region. Laterally, area below eyes and flank covered by dark brown reticula- tions, leaving out a perceptible transparent spiracle on the body wall. Lower part of the flank spotted. Tail muscu- lature overlaid by dark brown spots which condensed in some area to form reticulations. Fins covered by brown spots. Ventrally, oral disc, gular and branchial regions reticulated; venter pale and spotted, intestinal coils visible with regular spiral shaped. In total, three tadpoles were captured, but due to a transportation problem, the second specimen (ZCMV 13333) was destroyed. The external morphology of the third voucher specimen (ZCMV 13334 / ZSM 705/2010; GS 25) from the same locality shows the same characters and an identical oral disc configuration as the described above. DISCUSSION For decades, searches for the tadpole of Mantidacty- lus guttulatus and its relatives in the subgenus Mantidac- tylus have been unsuccessful, and scientists eventually hypothesized a nidicolous developmental mode for this species with a nest hidden very deep in the soil or even direct development (e.g., Glaw and Vences, 2007). Dur- ing our tadpole surveys in many streams in Ranomafana National Park mainly during the rainy seasons between 2006 and 2009, no tadpole assignable to this subgenus was encountered (Strauß et al., 2013), even at sites where many adults were present. One possible explanation for the absence might be a shifted onset of their reproductive season. Contrary to many other species that start their reproductive efforts at the beginning of the warm-rainy season, the reproduction period of these frogs might peak at the end of each rainy season towards the begin- ning of the cool-dry season. The avoidance of reproduc- tive competition with co-occurring species would be one benefit of this shift. An indication for this hypothesis is the early developmental stage of these tadpoles which suggests that they hatched in May. On the other hand, the single report of a calling individual from February (Vences et al., 2004) indicates that some reproductive activity occurs during the peak of the warm-rainy season. The noticeable fact that the tadpoles were found in very shallow water and, moreover, in the seepage area of a small water body could be seen as an indication of fossorial habits. However, morphological adaptions for fossoriality like a prominent tubular spiracle or particu- larly small eyes present in other fossorial tadpoles e.g. Otophryne robusta (Wassersug and Pyburn, 1987), Lepto- brachella mjobergi (Haas et al., 2006) or Micrixalus her- rei (Senevirathne et al., 2016) are absent in Mantidac- tylus guttulatus. Due to small sample size and the close vicinity of a large stream from which the tadpoles could have been washed away during a heavy rainfall this enigma requires further studies. Also, because we did not hypothesize these tadpoles would belong to M. gut- tulatus when encountering them in the wild we under- took no special efforts to further investigate the seepage area in which they occurred. For instance, we cannot exclude that upstream the seepage would originate from some kind of cavity, more suitable for such a large frog to deposit its eggs. Fig. 2. Images of the preserved tadpole specimen of Mantidactylus guttulatus (ZCMV 13332 / ZSM 704/2010) at Gosner stage 26; a-c) in dorsal, lateral and ventral view (scale bar = 10 mm); d) wide open oral disc with anterior (A1-A4) and posterior (P1-P3) keratodont rows (white outline for better visibility, median gap in A1 row caused by preparation, scale bar = 1 mm); e) spiracle and f ) vent tube in closer view (white outline for better visibility, scale bar = 1 mm). 124 Arne Schulze et alii Mantidactylus guttulatus tadpoles show the typi- cal morphology of stream-adapted, Orton (1953) Type IV tadpoles with a large and muscular tail and low fins. According to Altig and Johnston (1989) they can be classified as lotic-benthic and thus assigned to the eco- morphological guild Section I, Guild 7. The tadpoles of M. guttulatus are similar to those of the subgenus Bry- goomantis (Schmidt et al., 2009) which are considered as rather generalized lotic tadpoles. They share an oral disc with a large dorsal gap of marginal papillae, and a LKRF of 3-5 keratodont rows on the anterior labium with only the first being continuous and the others are interrupted by medial gaps, and three keratodont rows on the pos- terior labium of which the first usually has a very small medial gap. Instead, the larvae of several other subgen- era have highly specialized mouthparts, such as funnel- shaped structures (Chonomantis), poorly developed and reduced keratinized parts (Ochthomantis, Hylobatra- chus) or a reduced number of keratodont rows in com- bination with unpigmented jaw sheaths (Maitsomantis) (Glaw and Vences, 1994; Vejarano et al., 2006; Grosjean et al., 2011; Randrianiaina et al., 2011). The well resolved phylogeny of Wollenberg et al. (2011) suggests that the subgenera with generalized mouthparts (Mantidactylus and Brygoomantis) are not sister clades. While the sub- genus Mantidacylus branches off from the basal node of the Mantidactylus clade, Brygoomants is a sister clade to Chonomantis (Wollenberg et al., 2011). If these relation- ships are confirmed, it suggests extensive homoplasy in the evolution of tadpole mouthparts — either mul- tiple independent evolution of specialized mouthparts, or reversal towards generalized mouthparts in the Bry- goomantis clade. It is surprising that a frog like Mantidactylus guttula- tus, whose reproductive mode has intrigued researchers for decades, has such a dull tadpole as described herein. The reproductive behaviour and the unusual microhabitat of the species still remains a mystery. Where the species deposits its eggs and whether it displays any kind of pre-hatching parental care requires being elucidated by future studies. ACKNOWLEDGEMENTS We are grateful to F. M. Ratsoavina, A. S. Rasami- son, A. Rakotoarison, and D. R. Vieites for assisting dur- ing fieldwork for this study. This study was carried out in the framework of a cooperation accord between the Département de Biologie Animale of the University of Antananarivo, Madagascar and the Technical University of Braunschweig. Permission for collection was granted by the Ministère des Eaux et Forêts, Service de la Gestion Faune et Flore; Autorisation de Recherche 064/070/10/ MEF/SG/DGF/DCB.SAP/SLRSE, delivered 25 March 2010;  including the export permit 135N-EAO07/MG10, delivered 8 July 2010. Financial support was granted by the Volkswagen Foundation to MV, RDR, and AS, and by the Deutscher Akademischer Austauschdienst to RDR. REFERENCES Altig, R. (1970): A key to the tadpoles of the continental United States and Canada. Herpetologica 26: 180-207. Altig, R., Johnston, G.F. (1989): Guilds of anuran larvae: relationships among developmental modes, morphol- ogies, and habitats. Herp. Monogr. 3: 81-109. Altig, R., McDiarmid, R.W. (1999): Body plan. Develop- ment and morphology. In: Tadpoles: the Biology of Anuran Larvae, pp. 24-51. McDiarmid, R.W., Altig, R. Eds, Chicago University Press, Chicago. Altig, R., McDiarmid, R.W. (2006): Descriptions and biological notes on three unusual mantellid tadpoles (Amphibia: Anura: Mantellidae) from south-eastern Madagascar. Proc. Biol. Soc. Wash. 119: 418-425. AmphibiaWeb: Information on amphibian biology and conservation. [web application] (2016). Berkeley, Cali- fornia: AmphibiaWeb. Available: http://amphibiaweb. org/. (Accessed: 05 February 2016). Glaw, F., Vences, M. (1994): A Fieldguide to the Amphib- ians and Reptiles of Madagascar. Second Edition. Vences & Glaw Verlag, Köln. Glaw, F., Vences, M. (2007): A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edi- tion, Vences & Glaw Verlag, Köln. Gosner, K.L. (1960): A simplified table for staging anuran embryos and larvae with notes on identification. Her- petologica 16: 183-190. Grosjean, S., Strauß, A., Glos, J., Randrianiaina, R.D., Ohler, A., Vences, M. (2011): Morphological and eco- logical uniformity in the funnel-mouthed tadpoles of Malagasy litter frogs, subgenus Chonomantis. Zool. J. Linn. Soc-Lond. 162: 149-183. Haas, A., Hertwig, S., Das, I. (2006): Extreme tadpoles: the morphology of the fossorial megophryid larva, Leptobrachella mjobergi. Zoology 109: 26-42. Orton, G.L. (1953): The systematics of vertebrate larvae. Syst. Biol. 2: 63-75. Palumbi, S.R., Martin, A., Romano, S., McMillian, W.O., Stine, L., Grabowski, G. (1991): The simple fool’s guide to PCR, v.2.0. Honolulu: Department Zoology, Kewalo Marine Laboratory, University of Hawaii. Perl, R.G.B., Nagy, Z.T., Sonet, G., Glaw, F., Wollenberg, K.C., Vences, M. (2014): DNA barcoding Madagas- 125Tadpole of Mantidactylus guttulatus car’s amphibian fauna. Amphibia-Reptilia 35: 197-206. Randrianiaina, R.D., Strauß, A., Glos, J., Glaw, F., Venc- es, M. (2011): Diversity, external morphology and “reverse taxonomy” in the specialized tadpoles of Mal- agasy river bank frogs of the subgenus Ochthomantis (genus Mantidactylus). Contrib. Zool. 80: 17-65. Schmidt, H., Strauß, A., Glaw, F., Teschke, M., Vences, M. (2009): Description of tadpoles of five frog species in the subgenus Brygoomantis from Madagascar (Man- tellidae: Mantidactylus). Zootaxa 1988: 48-60. Schulze, A., Jansen, M., Köhler, G. (2015): Tadpole diver- sity of Bolivia’s lowland anuran communities: molecu- lar identification, morphological characterisation, and ecological assignment. Zootaxa 4016: 1-111. Senevirathne, G., Garg, S., Kerney, R., Meegaskumbura, M., Biju, S.D. (2016): Unearthing the fossorial tad- poles of the Indian dancing frog family Micrixalidae. PLoS ONE 11: e0151781. Strauß, A., Randrianiaina, R.D., Vences, M., Glos, J. (2013): Species distribution and assembly patterns of frog larvae in rainforest streams of Madagascar. Hyd- robiologia 702: 27-43. Thomas, M., Raharivololoniaina, L., Glaw, F., Vences, M., Vieites, D.R. (2005): Montane tadpoles in Madagas- car: molecular identification and description of the larval stages of Mantidactylus elegans, Mantidactylus madecassus, and Boophis laurenti from the Andringi- tra Massif. Copeia 2005: 174-183. Vejarano, S., Thomas, M., Glaw, F., Vences, M. (2006): Advertisement call and tadpole morphology of the clutch-guarding frog Mantidactylus argenteus from eastern Madagascar. African Zoology 41: 164-169. Vences, M., Andreone, F., Glaw, F. (2004): Voice of a giant: bioacoustic data for Mantidactylus guttulatus (Amphibia: Mantellidae). Amphibia-Reptilia 25: 112- 115. Vieites, D.R., Wollenberg, K.C., Andreone, F., Köhler, J., Glaw, F., Vences, M. (2009): Vast underestimation of Madagascar’s biodiversity evidenced by an integrative amphibian inventory. PNAS 106: 8267-8272. Wassersug, R., Pyburn, W. (1987): The biology of the Pe- ret’ toad, Otophryne robusta (Microhylidae), with spe- cial consideration of its fossorial larva and systematic relationships. Zool. J. Linn. Soc-Lond. 91: 137-169. Wollenberg, K.C., Vieites, D.R., Glaw, F., Vences, M. (2011): Speciation in little: the role of range and body size in the diversification of Malagasy mantellid frogs. BMC Evol. Biol. 11: 217. Acta Herpetologica Vol. 11, n. 2 - December 2016 Firenze University Press Predator-prey interactions between a recent invader, the Chinese sleeper (Perccottus glenii) and the European pond turtle (Emys orbicularis): a case study from Lithuania Vytautas Rakauskas1,*, Rūta Masiulytė1, Alma Pikūnienė2 Effective thermoregulation in a newly established population of Podarcis siculus in Greece: a possible advantage for a successful invader Grigoris Kapsalas1, Ioanna Gavriilidi1, Chloe Adamopoulou2, Johannes Foufopoulos3, Panayiotis Pafilis1,* The unexpectedly dull tadpole of Madagascar’s largest frog, Mantidactylus guttulatus Arne Schulze1,*, Roger-Daniel Randrianiaina2,3, Bina Perl3, Frank Glaw4, Miguel Vences3 Thermal ecology of Podarcis siculus (Rafinesque-Schmalz, 1810) in Menorca (Balearic Islands, Spain) Zaida Ortega*, Abraham Mencía, Valentín Pérez-Mellado Growth, longevity and age at maturity in the European whip snakes, Hierophis viridiflavus and H. carbonarius Sara Fornasiero1, Xavier Bonnet2, Federica Dendi1, Marco A.L. Zuffi1,* Redescription of Cyrtodactylus fumosus (Müller, 1895) (Reptilia: Squamata: Gekkonidae), with a revised identification key to the bent-toed geckos of Sulawesi Sven Mecke1,*,§, Lukas Hartmann1,2,§, Felix Mader3, Max Kieckbusch1, Hinrich Kaiser4 The castaway: characteristic islet features affect the ecology of the most isolated European lizard Petros Lymberakis1, Efstratios D. Valakos2, Kostas Sagonas2, Panayiotis Pafilis3,* Sources of calcium for the agamid lizard Psammophilus blanfordanus during embryonic development Jyoti Jee1, Birendra Kumar Mohapatra2, Sushil Kumar Dutta1, Gunanidhi Sahoo1,3,* Mediodactylus kotschyi in the Peloponnese peninsula, Greece: distribution and habitat Rachel Schwarz1,*, Ioanna-Aikaterini Gavriilidi2, Yuval Itescu1, Simon Jamison1, Kostas Sagonas3, Shai Meiri1, Panayiotis Pafilis2 Swimming performance and thermal resistance of juvenile and adult newts acclimated to different temperatures Hong-Liang Lu, Qiong Wu, Jun Geng, Wei Dang* Olim palus, where once upon a time the marsh: distribution, demography, ecology and threats of amphibians in the Circeo National Park (Central Italy) Antonio Romano1,*, Riccardo Novaga2, Andrea Costa1 On the feeding ecology of Pelophylax saharicus (Boulenger 1913) from Morocco Zaida Ortega1,*, Valentín Pérez-Mellado1, Pilar Navarro2, Javier Lluch2 Notes on the reproductive ecology of the rough-footed mud turtle (Kinosternon hirtipes) in Texas, USA Steven G. Platt1, Dennis J. Miller2, Thomas R. Rainwater3,*, Jennifer L. Smith4 Heavy traffic, low mortality - tram tracks as terrestrial habitat of newts Mikołaj Kaczmarski*, Jan M. Kaczmarek Book Review: Sutherland, W.J., Dicks, L.V., Ockendon, N., Smith, R.K. (Eds). What works in conservation. Open Book Publishers, Cambridge, UK. http://dx.doi.org/10.11647/OBP.0060 Sebastiano Salvidio