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Photographs of the tomato frog, Dyscophus antongilii and its habitat. (A) Study pond with partial view of the garden. (B) Adult male. (C) Adult female. (D) Study pond and surrounding vegetation. 

Photographs of the tomato frog, Dyscophus antongilii and its habitat. (A) Study pond with partial view of the garden. (B) Adult male. (C) Adult female. (D) Study pond and surrounding vegetation. 

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Based on daily monitoring around an urban pond in the coastal town of Maroantsetra, from 2003-2011, we provide an analysis of the yearly reproductive activity of the tomato frog (Dyscophus antongilii), a large-sized and prominent red-coloured microhylid frog from north-eastern Madagascar. Frogs were observed all year round but despite the limited c...

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... Saenz et al., 2006), or the availability of suitable sites for breeding (Sullivan, 1982). Temporally short breeding bouts were also suggested as an adaptation to reduce predation pressures (Woodward and Mitchell, 1990; Lucas et al., 1996), cannibalism (Petranka and Thomas, 1995), and the energetic costs of reproduction (McCauley et al., 2000). In temperate and subtropical regions that are char- acterized This study by distinct was carried dry and out wet in a seasons, pond within the reproduction the urban area of of pond-breeding Maroantsetra, anurans a small coastal occurs typically town located during in the north-eastern wet season while Madagascar in those that tropical is located and subtropical within an regions area of with very high little climatic yearly rainfall. variation The throughout town consists the year, mainly more of small species buildings demonstrate of one largely or two aseasonal storeys built breeding on sandy phenologies ground (Crump, 1974). with a network Duellman and Trueb (1994) of open ditches running throughout divided an the Amazonian town to drain anuran rainwater community and that partly reproduces sewage water. all year Most round houses into have continuous gardens breeders and numerous that potential- semi- abandoned ly breed every parcels night, are opportunistic scattered throughout breeders that the breed town. regularly D. antongilii after occurs heavy rains, at many and parts spo- radic within wet the and town, dry breeding breeders both that in breed ditches at irregular and in small intervals and after large heavy ponds. rains or throughout dry The spells, study respectively. pond (located They at suggest 15°25’47”S, that the 49°44’23”E) availability had of a breeding diameter sites of 6-7 dictates m and inter- was species surrounded variation by fences in breeding and garden phenology. areas, very close to several huts and houses, partial- The true tomato frog or northern tomato frog, Dyscophus antongilii is a representative of the Madagascar-endemic microhylid subfamily Dyscophinae that is related to Asian microhylids (Van Bocxlaer et al., 2006; Van der Meijden et al., 2007). Dyscophus is the sole genus in the subfamily, and it contains three species: D. insularis occurring in dry areas of Madagascar’s west coast, D. antongilii , living in a small area of the north-east and east coast, and D. guineti, the false tomato frog, living in eastern mid-altitude rainforest areas (Glaw and Vences, 2007). Dyscophus insularis is a medium sized (40-50 mm snout- vent length) and cryptically coloured species that breeds explosively in temporary as well as permanent ponds and has small tadpoles that complete their metamorphosis very fast (Glos, 2003; Grosjean et al., 2007; J. Glos, pers. comm.), the two closely related eastern species D. antongilii and D. guineti are larger (60-101 mm SVL), bright red-orange col- oured, breed in temporary as well as larger permanent ponds and have large-sized tad- poles that, as far as known, require a longer time to complete metamorphosis (Pintak, 1987; Glaw and Vences, 1994, 2007). While D. guineti is regularly exported from Mada- gascar for the exotic pat-trade and assessed within the Least Concern threat category, D. antongilii is considered to be Near Threatened and is listed in the Appendix I of the Con- vention on the International Trade in Endangered Species (CITES) (Raxworthy and Nuss- baum, 2000) which largely inhibits legal exports for the pet trade taking place from Mad- agascar (Andreone et al., 2005, 2008). Due to the prominence of D. antongilii which in the area of the town of Maroantsetra and the nearby Masoala National Park is considered as an important flagship species for conservation, several studies have addressed aspects of its genetic structure (Chiari et al., 2006) and ecology (Tessa et al., 2007, 2011). Basic information on its activity, activity cycles, reproduction, and habitat choice is however still missing. Here we contribute to closing this gap in knowledge and provide an analy- sis on the yearly activity of a population of D. antongilii located in an urban area within Maroantsetra. This study was carried out in a pond within the urban area of Maroantsetra, a small coastal town located in north-eastern Madagascar that is located within an area of very high yearly rainfall. The town consists mainly of small buildings of one or two storeys built on sandy ground with a network of open ditches running throughout the town to drain rainwater and partly sewage water. Most houses have gardens and numerous semi- abandoned parcels are scattered throughout the town. D. antongilii occurs at many parts within the town, breeding both in ditches and in small and large ponds. The study pond (located at 15°25’47”S, 49°44’23”E) had a diameter of 6-7 m and was surrounded by fences and garden areas, very close to several huts and houses, partial- ly surrounded by a fence, and by some vegetation at the edges and in the water (Fig. 1), and populated by domestic ducks. This anthropogenic habitat is in the property of a local nature guide in Maroantsetra living in an adjacent house. The Dyscophus popula- tion found in and around the study pond was relatively large. For instance, from 12-15 February 2003 we performed intensive searches and collected 78 individuals, and on 24 February 2004 we found six males (snout-vent length 62-67 mm, hereafter: SVL) and 10 females (84-101 mm SVL) of which 3-5 were toe-clipped recaptures from the previous year. During a total of nine years (2003-2011) the local nature guide in whose proper- ty the Dyscophus study pond was located regularly undertook a daily monitoring of the tomato frog population in and around the pond, noting (1) the number of adult frogs observed Our long active term on the survey ground based and on in the the data water, from (2) 2003-2011 presence or provides absence evidence of egg clutches that D. which antongilii float in on Maroantsetra the water surface, breed almost and (3) all emission year round of calls as reflected by the frogs. in the Because temporal Dysco- dis- phus tribution males of often adults, call egg from clutches, concealed and calls positions (Figs and 2-3). the The design mean of rank our of study the number was so to of obtain adults observed daily data differed by untrained between observers, months calling (Kruskal-Wallis; intensity was H = estimated 20.87; df in = four 11; P relatively > 0.05) rough and between categories: years 0 (Kruskal-Wallis; = no calls heard H in = the 16.26; respective df = 8; afternoon/evening, P > 0.05), with counts 1 = few being isolated par- calls ticularly heard, high 2 = through regular 2003 calling, and 3 particularly = large choruses. low through A distinction 2007 (Fig. 4). between Despite calls of the differ- lack ent of apparent individuals climatic was seasonality not possible, in but the region the calling the breeding in large of choruses D. antongilii certainly can referred be roughly to multiple divided according individuals to calling its activity simultaneously. level into a high Presence activity of period freshly between laid eggs January-May was scored as and 0 = a lower no eggs, activity 1 = few period eggs, between and 2 = June-December large quantity (Fig. 2). of eggs corresponding to multiple clutch- es. When Freshly egg-laying laid eggs were took found place on in all consecutive months except days, November, a distinction and of with newly highest laid inci- eggs from dence those between laid April previously and May, probably while was calling not was always heard possible in all months for the untrained (Fig. 3) during observers, both but mornings because and the evenings. eggs hatch The after number 36 hours of adults (Pintak, 1987) observed this was possible relatively error stable is limited over the to a years, maximum with no period trend of of two declining days. Due values to (Fig. 4). the limitations The maximum concerning number the call of adult and individu- egg data, most als observed of our statistical per day during analyses the are transect based on walks the around counts the of adults. pond were Although 12 in call, 2004, egg 20 and in adult 2005, count 17 in 2006, data is 20 available in 2007, for 35 in every 2008, year 27 in from 2009, 2003 30 to in 2011, 2010, and due 35 to in logistic 2011. reasons in several years there are gaps of several months in the dataset (e.g., no complete data for January-February 2003). For the period January 2003 to January 2004, data on daily minimum and maximum temperature (°C) as well as daily accumulated precipitation (mm) and daily mean rela- tive humidity (%) were obtained from the meteorological station of Maroantsetra, at a dis- tance of less than 1 km from the study site; unfortunately, these meteorological data were not available for subsequent years. We used a non-parametric test, Kruskal-Wallis, to test for difference in adult numbers between years and between months. For the correlation analysis of 2003 climatic data with the phenological data we used Pearson pairwise cor- relation for the adult counts and Spearman rank correlation for the categorical eggs and calls. Statistical analysis was carried out using the statistical software JMP IN version 5.0 by SAS Institute, Inc. and SigmaPlot version 10.0 by Systat software, Inc. for the exponen- tial curve-fitting of the non-linear regression analysis. A table with all original data has been deposited in the Dryad data repository ( Our long term survey based on the data from 2003-2011 provides evidence that D. antongilii in Maroantsetra breed almost all year round as reflected in the temporal dis- tribution of adults, egg clutches, and calls (Figs 2-3). The mean rank of the number of adults observed differed between months (Kruskal-Wallis; H = 20.87; df = 11; P > 0.05) and between years (Kruskal-Wallis; H = 16.26; df = 8; P > 0.05), with counts being par- ticularly high through 2003 ...

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