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Herpetological Journal FULL PAPER Volume 24 (October 2014), 245–253 FULL PAPER Herpetological Journal Published by the British Variation in dietary composition of granular spiny frogs Herpetological Society (Quasipaa verrucospinosa) in central Vietnam Binh Van Ngo1, Ya-Fu Lee1 & Chung Dac Ngo2 1Department of Life Sciences, National Cheng Kung University, Tainan 70101, Taiwan 2Faculty of Biology, College of Education, Hue University, Hue 47000, Vietnam The granular spiny frog, Quasipaa verrucospinosa (Bourret, 1937), is native to Vietnam and classified as Near Threatened due to environmental degradation, loss of forest and stream habitats and human exploitation. We collected stomach contents of Q. verrucospinosa using a nonlethal stomach-flushing technique from three stream sites in the rain forests of Thừa Thiên- Huế Province, central Vietnam, to investigate their food habits. Dietary analysis identified 2645 prey items of 27 orders and nine classes. Prey comprised mainly invertebrates, but also fishes, frogs and conspecific sub-adults. The major prey items as determined by frequency of occurrence, item count and percent volume were spiders, beetles, crabs, hymenopterans, grasshoppers, crickets and cicadas. Insects alone accounted for an importance value of 59.8%. The mean monthly prey volume consumed was positively and negatively correlated to temperature and rainfall, respectively. Consistent with the increased energetic needs prior to the main breeding season, the number of prey items and volume of prey consumed per frog were highest in the little rainy season. The volume of prey consumed was positively correlated with snout-vent length and mouth width of frogs, supporting the gape limitation hypothesis. Despite their larger size, however, females did not consume greater numbers of prey items or larger-sized prey than males. Adults consumed a higher diversity of prey and higher proportions of Araneae and Hemiptera than sub-adults, whereas females had a more even diet than males and consumed a higher proportion of Orthoptera. Key words: Anura, diet, feeding, frogs, stomach content, Quasipaa verrucospinosa, Vietnam INTRODUCTION or terrestrial species (Hothem et al., 2009; Rosa et al., 2011). It also generally helps to explain why anurans nimal food habits offer essential information for living in tropical-subtropical regions have more diverse Aconstructing food webs (Elton, 2001). Dietary diets than temperate species (e.g., Moseley et al., 2005; variation may provide additional insights for our Ugarte et al., 2007; Hothem et al., 2009; Brito et al., understanding of life history features, resource and 2013). Nevertheless, anuran diets that specialise on habitat use, population dynamics and interspecific specific prey types such as Acari, ants, beetles or termites interactions of species (e.g., Toft, 1980). In terrestrial are also reported from the tropics (Isacch & Barg, 2002; and freshwater habitats, anurans are predators of many Bonansea & Vaira, 2007; Valderrama-Vernaza et al., invertebrates and sometimes even vertebrates, and play a 2009; Brito et al., 2013; Rodrigues & Santos-Costa, 2014), crucial role in energy and nutrient cycling between these and may be associated with foraging mode, season and two systems (Duellman & Trueb, 1994; Wells, 2007). habitat (Toft, 1980). However, detailed information on ecological needs of The granular spiny frog, Quasipaa verrucospinosa many anurans is either insufficient or lacking. Anurans (Bourret, 1937) is a dicroglossid frog native to Vietnam are among the most threatened terrestrial vertebrates (Frost, 2014). All 12 species of the genus Quasipaa (Stuart et al., 2004; Alford, 2011), and population are narrowly distributed from northeastern Indochina declines are likely to result in a particularly high diversity to southern and southwestern China (Nguyen et al., loss in tropical regions, where vast numbers of species 2009; Frost, 2014), and are classified as either at least reside and often exhibit high endemism (e.g., Fauth et Near Threatened or as Data Deficient (IUCN, 2013). In al., 1989; Allmon, 1991; Vitt & Caldwell, 1994). Vietnam, Q. verrucospinosa resides only in headwaters Anurans are usually considered as opportunistically of a few montane streams in primary forests in the foraging generalist predators (Wells, 2007). This western and southern Thừa Thiên-Huế Province (Ngo intuitively predicts that more aquatic anurans consume & Ngo, 2011), where intact rainforest habitats remain larger numbers of aquatic prey than do semiaquatic and represent a recognised biodiversity hotspot rich in Correspondence: Ya-Fu Lee ([email protected]) 245 B.V. Ngo et al. Fig. 1. Map of Thừa Thiên-Huế Province showing the three localities: (1) Mang Stream, Bach Ma National Park; (2) Ba Rang Stream, Sao La Conservation Area; and (3) Dong Ngai Stream, Phong Dien Nature Reserve (A Luoi District), where granular spiny frogs were collected. endemic species (Myers et al., 2000). In many parts of this mean of 173±23 mm; Nguyen et al., 2004). The study region, however, frog populations have been declining area is dominated by montane rainforests at elevations over recent decades due to over-exploitation and other of 700~1400 m and cloud forests from above 1400 large-scale disturbances such as degradation of forested m up to the summit at 1774 m (Nguyen et al., 2004). habitats and stream sedimentation from unselective or destructive logging (Ngo & Ngo, 2009; Ngo et al., 2009). Sample collections This study presents the first comprehensive We captured frogs from three localities at roughly the investigation on the dietary composition and same elevations: Mang Stream (1455 m) in Bach Ma spatiotemporal variation of granular spiny frogs across National Park, Ba Rang Stream (1546 m) in the Sao La its range in central Vietnam. We also compared the diets Conservation Area and Dong Ngai Stream (1579 m) in the of frogs of different sexes and age groups to examine sex- A Luoi District (Fig. 1). During two nights in each month dependent dietary variation, and tested the hypothesis from January 2008 to December 2009, a team of six that the size of prey consumed and the diet breadth are people conducted nocturnal surveys from 2000 to 0200 correlated with morphometry. Our study is the first to hours in each of the three streams. We visually searched focus on dietary variation and its ecological correlates for for frogs using spotlights, aided by their calls, in water one of the least-studied dicroglossid frogs in southeast and up to 15 m away from the stream over a length of Asia, providing baseline information for understanding 3 km, collecting frogs by hand. Sex was determined by resource use patterns of the genus Quasipaa. the number of spinules on back warts, the presence or absence of pectoral spines, hypertrophied forearms, MATERIALS AND METHODS callosities on finger I of the forelegs and a distinct tympanum. Sub-adults were distinguished from adults Study sites by snout-vent length (SVL) and body colour, with reddish Field work took place in Thừa Thiên-Huế Province, brown characterising sub-adults and chrome yellow Vietnam (15o59’30”~16o44’30”N, 107o00’56”~108o characterising adults (Ngo & Ngo, 2011). We measured 12’57”E). This area is characterised by tropical climate, mouth width (MW) and SVL with a digital caliper and with annual temperature averaging 24.4±0.41oC (ranging body mass (BM) with an electronic balance. We used from 15.8±0.52oC in January to 29.7±0.71oC in June) visible implant elastomer tags (Nauwelaerts et al., 2000; and an annual mean precipitation of 4980±377 mm. A Hoffmann et al., 2008) to mark each frog. relatively dry period extends from January to April, with We placed frogs individually in labelled bags for monthly rainfall of around 120 mm (mean: 121±19 mm). stomach contents collection within 30 min of capture Most rainfall is concentrated in the monsoon season (Caldwell, 1996). We adopted stomach-flushing (Griffiths (September to December, monthly mean of 738±96 mm) 1986; Leclerc & Courtois, 1993) to obtain stomach and the little rainy season from May to August (monthly contents without sacrificing frogs. We used different 246 Quasipaa verucospinosa diet in Vietnam Table 1. Dietary composition (%) of granular spiny frogs (Quasipaa verrucospinosa) with regard to frequency of occurrence (F), number of items (N), and volume (V), and the overall importance value (I) of each prey (n=539 frogs considered). Prey Frequency (F) Number (N) Volume (V) Importance (I) Arachnida (Araneae) 13 21.5 15.2 16.57 Insecta Blattodea 2.8 2.1 3.4 2.76 Coleoptera 12.9 18.7 17.6 16.39 Collembola 0.3 0.3 0.1 0.21 Dermaptera 4.3 3.1 2.5 3.32 Diptera 4.4 3.4 3.7 3.81 Ephemeroptera 0.2 0.2 0.3 0.24 Hemiptera 5.4 4 7.7 5.71 Hymenoptera (Formicidae) 7.8 5.9 0.7 4.83 Hymenoptera (others) 9.9 7.2 1 6.01 Isoptera 0.3 0.2 0.1 0.18 Lepidoptera 2.5 1.9 4.7 3.04 Neuroptera 0.2 0.2 0.2 0.2 Odonata 0.4 0.3 0.7 0.46 Orthoptera 12.2 9.6 6.6 9.45 Phasmatodea 0.5 0.5 1.6 0.86 Insecta larvae 2.6 2.1 2.3 2.34 Diplopoda 1.8 1.4 2.2 1.81 Chilopoda 0.4 0.3 0.7 0.49 Crustacea Decapoda 13.3 13.6 20.7 15.86 Isopoda 1.7 1.3 0.7 1.24 Clitellata Lumbriculida 0.3 0.3 0.3 0.3 Megadrili 0.1 0.04 0.04 0.04 Gastropoda (Mesogastropoda) 0.9 0.7 1.7 1.09 Osteichthyes Cypriniformes 0.2 0.1 1.4 0.56 Perciformes 0.5 0.3 1.6 0.78 Amphibia (Anura) 0.3 0.2 1.6 0.73 Unidentified 0.8 0.6 0.7 0.73 sizes of soft catheter tubes (2, 3 or 5 mm inner diameter) and reference keys (Thai, 2001; Triplehorn & Johnson, with appropriately-sized syringes and different amounts 2005).
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