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Composition and Scaling of Male and Female Alpine Newt (Mesotriton Alpestris) Prey, with Related Site and Seasonal Effects

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Composition and Scaling of Male and Female Alpine Newt (Mesotriton Alpestris) Prey, with Related Site and Seasonal Effects Ann. Zool. Fennici 49: 231–239 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 31 August 2012 © Finnish Zoological and Botanical Publishing Board 2012 Composition and scaling of male and female Alpine newt (Mesotriton alpestris) prey, with related site and seasonal effects Oldřich Kopecký1,*, Jiří Vojar2, František Šusta3 & Ivan Rehák3 1) Department of Zoology and Fish Farming, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences in Prague, Kamýcká 957, CZ-165 21 Prague, Czech Republic (*correspondig author’s e-mail: [email protected]) 2) Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences in Prague, Kamýcká 129, CZ-165 21 Prague, Czech Republic 3) Prague Zoological Garden, U Trojského zámku 3/120, CZ-171 00 Prague, Czech Republic Received 30 Sep. 2011, final version received 18 Apr. 2012, accepted 19 Apr. 2012 Kopecký, O., Vojar, J., Šusta, F. & Rehák, I. 2012: Composition and scaling of male and female Alpine newt (Mesotriton alpestris) prey, with related site and seasonal effects. — Ann. Zool. Fen- nici 49: 231–239. In amphibians, most species are female-biased sexually dimorphic and such dimor- phism is often accompanied by intersexual differences in prey composition. While many aspects of foraging ecology have been studied in this group, intersexual dif- ferences have rarely been described. We examined dietary composition of male and female Alpine newts (Mesotriton alpestris) from two localities in the Czech Republic during the entire breeding period by stomach flushing. Isopoda, Cladocera, Rana eggs were the most important prey. About 16% of newts did not contain prey items. At stud- ied localities, females were about 15% larger than males. Snout–vent length (SVL) of newts was related to the mass of consumed prey, but not its number and diversity. Ana- lyzing the residuals from the relationships between SVL and prey characteristics, we found the effect of sex on all observed prey variables — the females consumed more prey items, more diverse prey and also in greater mass than did the males. Introduction should be accompanied by dietary differences between sexes, because varied resources would Sexual size dimorphism (SSD) is a common be used by individuals of different sizes (e.g., characteristic of many animal taxa (Darwin Shetty & Shine 2002, Zalewski 2007). 1871, Andersson 1994). Male-biased sexual In fishes, snakes, raptors, owls and waders, size dimorphism (MSSD), has been considered females are usually the larger sex. In other rep- a product of male–male competition (Trivers tiles, other birds and most mammals, males 1972, Blanckenhorn 2005). SSD, however, can tend to be larger (Shine 1989, Andersson 1994, be also a consequence of intersexual resource Krüger 2005, Vincent & Herrel 2007). FSSD partitioning (Shine 1989, Krüger 2005). In any has been reported in the majority of amphibians case, the occurrence of SSD in particular species (Shine 1979), including newts from the genera 232 Kopecký et al. • ANN. ZOOL. FeNNICI Vol. 49 Lissotriton, Mesotriton and Triturus (Griffiths the Czech Republic near the town of Ledeč nad 1996, Malmgren & Thollesson 1999). While Sázavou. The first, hereafter referred to as local- resource partitioning between morphs (Denoël & ity A (49°42´45´´N, 15°16´50´´E; 452 m a.s.l.) is Andreone 2003, Denoël & Schabetsberger 2003, a fishless pond with a surface area of 36.0 m2 and Denoël et al. 2004), interspecific competition maximum depth of 0.8 m. The pond’s banks are (Fasola & Canova 1992, Joly & Giacoma 1992, planted with willows (Salix sp.), and the pond Jehle et al. 2000), and feeding relationships of is surrounded by pasture. The bottom is muddy larval newts (Braz & Joly 1994, Griffiths et al. and vegetation in the water is scant, consisting 1994, Babik 1998) are well known, the dietary mainly of common duckweed (Lemna minor), differences between males and females have starwort (Callitriche verna) and compact rush rarely been described, and even then generally (Juncus conglomeratus). The pond is artificial, only as a by-product of studies having a different its original purpose being to drain water from focus (Joly & Giacoma 1992, Rulík 1993, Denoël surrounding wet plots. During the study, the & Demars 2008). water’s pH ranged from 6.5 to 7.0. Because of newts’ well-hidden lifestyle The second locality, hereafter B (49°44´24´´N, during their terrestrial phase (Verrell 1985, Scha- 15°16´59´´E; 444 m a.s.l.), is a now fishless betsberger et al. 2004), the majority of dietary pond that was historically used for fish rearing. studies were carried out during newts’ breeding Pond B is situated in a spruce forest. It has a (aquatic) phase. Newts are prolonged breeders, surface area of 27.5 m2 and maximum depth of remaining in the water for several months (Grif- 0.3 m. The pond’s bottom is muddy. Water plants, fiths 1996, Arntzen 2002). During this time, envi- consisting mainly of the common waterweed ronmental conditions, rate of intraspecific com- (Elodea canadensis), starwort and reed man- petition, as well as prey patterns and availability nagrass (Glyceria aquatica), cover a large part can change considerably (Griffiths 1997, Denoël of the pond’s surface area. During the study, pH & Demars 2008, Kopecký et al. 2010). To inves- values were around 5.5. tigate potential intersexual dietary differences, A small number of smooth newts (Lissotriton one must examine the newts’ entire breeding vulgaris) and common frogs (Rana temporaria) period. However, this approach has not always inhabited both ponds. been taken (see Joly & Giacoma 1992, Jehle et al. 2000, Denoël & Schabetsberger 2003). In newts, as in other animals (Beck et al. Sampling and diet analysis 2007), the larger sex may consume larger prey items or greater numbers of smaller prey items. Adult Alpine newts were captured by net from Also, due to egg production, females may require the shore at three time points representing their more diverse food than males (Lewis et al. 2002). entire breeding season. Sampling was done Hence, the principal aim of this study was to during daylight hours in 1997: once in April, determine differences in prey scaling and prey once in May and once in June. The newts were composition (differences in prey mass, number kept in a container filled with water from the of prey items, and number of prey categories) pond and marked (Heyer et al. 1994) by toe- between sexes of the Alpine newt (Mesotriton clipping (Ferner 1979) different for each date of alpestris) while controlling for the different body capture. To avoid data dependence (Luiselli et al. length of the sexes and possible changes in newts’ 2007), only newts captured for the first time were foraging behavior across the entire aquatic period. used in the study. Each captured and unmarked newt was measured (snout–vent length, SVL) to the nearest 0.1 mm. The content of its stomach Materials and methods was extracted using a stomach flushing tech- nique described by Opatrný (1980): 1 ml syringe Study sites with 80 mm hypodermic flexible needle with last 30 mm before the tip covered by plastic tubing The study was conducted at two localities in was used. This terminal part of the needle was ANN. ZOOL. FeNNICI Vol. 49 • Prey differences between sexes of Alpine newts 233 gently inserted through mouth of the newt into Newts containing only non-prey items like plants, the stomach and 1 ml of water was injected. algae and mud or which had completely empty Usually two and maximally three flushes were stomachs were analyzed in a separate model. conducted. Measured and flushed newts were not We intended to compare sex ratio between anesthetized. Newts were released back into the localities across the entire sampling period water immediately after these procedures. (April–June). Since a non-significant result might The stomach contents were individually be obtained even when the sex ratio is biased as stored in vials and preserved in 4% formalde- long as sex ratios are the same across all months, hyde. Prey items were identified using a stere- we took into consideration the month of sampling oscopic microscope and sorted into 19 taxo- as an additional variable. Hence, we used a gener- nomic and four biomass categories — 0.001 g alized linear model with Poisson’s distribution of for Cladocera, Cyclopoida, Megaloptera larvae the response variable (numbers of newts) and sex, and Turbellaria; 0.01 g for Chironomidae larvae, locality, month and particularly the interactions Chironomidae pupae, Culicidae pupae, Bivalvia, between sex, month and locality as the explan- Gastropoda, and Dytiscidae larvae; 0.1 g for atory variables. To obtain a minimal adequate Ephemeroptera larvae, Ephemeroptera pupae, model, we simplified the maximal model using a Lepidoptera larvae, Plecoptera larvae, Trichop- chi-squared deletion test (Crawley 2007). tera larvae, Arachnida, Isopoda, and Rana eggs; We used the same statistical procedure for and 1 g for Lumbricus — due to the technical comparing newt numbers in relation to their impossibility of weighing all (1417) prey items replete status (replete vs. newts with empty thus obtained. Categorization of prey taxa into stomachs), sex, locality, month and the interac- prey categories was based on the previously pub- tions of these as the explanatory variables. lished works (Opatrný 1968). We wanted to distinguish intersexual diet Sex of the newts was determined using exter- differences from the pure effect of different nal secondary sexual characters. Sexual size body sizes between sexes. The first step was to dimorphism was determined using the sexual determine SVL differences between males and dimorphism index (SDI) proposed by Lovich females. To take into account possible vari- and Gibbons (1992): ability in SVL between localities and within the long breeding period (from April to June), we SDI = (size of larger sex/size of smaller sex) ± 1 analyzed the effects of sex, locality and month [+1 if males are larger, –1 if females are larger].
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