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The Diet of the Spiny-Cheek Crayfish Orconectes

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The Diet of the Spiny-Cheek Crayfish Orconectes Cent. Eur. J. Biol. • 9(1) • 2014 • 58-69 DOI: 10.2478/s11535-013-0189-y Central European Journal of Biology The diet of the spiny-cheek crayfishOrconectes limosus in the Czech Republic Research Article Renata Vojkovská, Ivona Horká*, Zdeněk Ďuriš University of Ostrava, Faculty of Science, Department of Biology and Ecology, 710 00 Ostrava, Czech Republic Received 30 June 2012; Accepted 25 March 2013 Abstract: Thecompositionofthedietoftheinvasivespiny-cheekcrayfishOrconectes limosus was studied using qualitative and quantitative analyses of stomach contents. A total of 368 specimens collected in 2003–2005 and 2008 in Czech localities were examined, predominantly fromtheLabe(Elbe)andVltavaRiverbasins.Foodcomponentswerecomparedforthreesizeclassesofcrayfishandbothsexes.The followingconclusionswerereached:(1)thespiny-cheekcrayfishisanomnivorousspeciesconsumingplants,animalsanddetritus; (2) quantitatively, the main food component of O. limosusisdetritus,whiletheplantcomponentwassecond;(3)O. limosus may swallow whole food particles up to 4 mm in size, and the bodies of small animals may sometimes be found undamaged in their stomachs. Keywords: Crayfish diet • Food particles • Invasive crayfish • Stomach content • Trophic index © Versita Sp. z o.o. 1. Introduction mussels [10], or small and middle-sized mussels and aquatic gastropods to large ones [11-13]. The spiny-cheek crayfish, Orconectes limosus Being omnivorous throughout their life cycle, crayfish (Rafinesque, 1817), is the most common alien crayfish in may prefer different foods in different stages of their life, European freshwater ecosystems [1-3]. Characteristics with freshly hatched juveniles feeding mainly on animal determining its competitive advantage in comparison plankton and later on benthic invertebrates, while adults with native species include: an r-strategist life cycle with consuming mostly plants and detritus [14,15]. The faster reproduction and earlier maturation; a greater selection of plant foods often depends on mechanical tolerance to polluted waters; and the ability to spread structure, nutritional values, or plant chemical defenses the crayfish plague pathogen, together with its own [7,16-18]. According to Nyström and Strand [19] and resistance to that illness [4]. Cronin et al. [18], crayfish prefer newly budding or finely Crayfish forage on a wide range of foods, including branching plants to those that are well grown and rigid. water macrophytes, algae, detritus and macro- When foraging on submerged and emerged macrophytes, invertebrates [5-7]. Despite being generally omnivorous crayfish may cause changes to deep water environments. animals, different food components may be preferred Invasive species, in particular, may negatively affect in accordance with availability and energetic value [8]. aquatic plant density and diversity [19-21]. Söderbäck et al. [9] also showed that, in experimental Alien species compete with native species over conditions, crayfish feeding on animal-originated food many biological resource aspects, including food grow faster and show higher foraging activity than consumption. Despite a fairly wide literature on feeding those kept on a detrital diet. Feeding selectivity has also of different crayfish species, from which only a limited been experimentally documented for food of animal review is given above, little data has been reported origin, with crayfish foraging on thin-shelled rather than on the food and feeding impacts of the invasive spiny- thick-shelled mollusks [7], preferring fish eggs to zebra cheek crayfish. Staszak and Szaniawska [22], as well * E-mail: [email protected] 58 R. Vojkovská et al. as Anwand and Valentin [23], described O. limosus as 2. Experimental Procedures an omnivore. Staszak and Szaniawska [22] noted that, at higher densities, cannibalistic behavior of spiny- Qualitative and quantitative analyses of the main and cheek crayfish frequently occurs. Chiesa et al. [24] also particular food items were performed for the stomach characterized O. limosus as omnivorous. Based on contents of 368 spiny-cheek crayfish specimens of qualitative analyses of stomach contents, they divided different sizes and sexes, collected in the Labe (Elbe) these contents into four categories – animal, plant, detrital and Vltava River basins. Crayfish were collected in 26 and others (the latter containing abiotic components); localities of the Czech Republic, usually from April to the main food item found was of plant origin, followed October, 2003–2005 (Table 1; Figure 1). The material by detritus and animal food. Chybowski [25] studied was originally collected for distributional and biometric the foraging of O. limosus in Polish lakes in relation to purposes [27,28]. Specimens were caught by hand diurnal and annual activity. He identified plant material during the morning hours and preserved in 70% as the primary food item found in crayfish stomachs, ethanol within 2–4 hours. Some additional specimens and stated that annual consumption amounted to only were collected from the Prudník Brook, Odra river, NW 0.27% of the available aquatic vegetation. Similar results Moravian-Silesian region in 2008 [29]. Stomach contents are presented also by Anwand and Valentin [23] for the were analyzed in the laboratory for 368 O. limosus species in Kleiner Döllnsee Lake in Germany. Chucholl specimens (212 males and 156 females, including 17 [26] demonstrated the omnivorous feeding habits of the ovigerous females) (Table 2). calico crayfish, Orconectes immunis (Hagen, 1870), a Because of low numbers of specimens collected competitor of O. limosus in the Rhine River, Germany. from particular localities, all the crayfish used in this The aim of this study is elucidate the diet study were pooled into one sample, then sexed and composition of the non-indigenous spiny-cheek crayfish divided into three size classes (S, M, and L) according O. limosus in the Czech Republic using qualitative and to the post-orbital carapace length (POCL), measured quantitative analysis of stomach contents, contributing in the dorsal body midline from the level of the posterior to knowledge on the ecology of invasive animals and orbital margins to the posterior margin of the carapace to the conservation of native species and natural water (see [27]: S – up to 13 mm, M – 13–28 mm, L – over ecosystems. 28 mm). Figure 1. Localities where samples of Orconectes limosus were taken for stomach content analyses. 59 Diet of Orconectes limosus Watercourse / body Nearest settlement Latitude (N) Longitude (E) Running waters Labe Hněvice 50°27’ 14°22’ Labe Kolín 50°02’ 15°13’ Labe Ostrá 50°10’ 14°54’ (confluence with Farský potok) Labe Litoměřice 50°32’ 50°32’ Labe Malé Březno 50°40’ 14°10’ Labe Nebočady 50°43’ 14°11’ Labe Obříství 50°18’ 14°29’ Labe Poděbrady 50°09’ 15°06’ Labe Štětí 50°27’ 14°22’ Labe Těchlovice 50°42’ 14°12’ Labe Ústí nad Labem 50°39’ 14°03’ Labe Třeboutice 50°31’ 14°12’ (confluence with Luční potok) Cidlina Libice nad Cidlinou 50°07’ 15°11’ Doubrava Záboří nad Labem 50°01’ 15°21’ Jickovický potok Jickovice 49°27’ 14°13’ Jizera Nový Vestec 50°11’ 14°44’ Ohře Doksany 50°27’ 14°09’ Ohře Bohušovice nad Ohří 50°30’ 14°09’ Prudník Osoblaha 50°17’ 17°44’ Vltava Vrbno u Mělníka 50°19’ 14°27’ Standing waters Barbora (quarry) Oldřichov u Teplic 50°38’ 13°45’ Cítov (sand pit) Vliněves 50°22’ 14°27’ Kojetice (quarry) Kojetice u Neratovic 50°14’ 14°30’ Lhota (sand pit) Lhota 50°15’ 14°40’ sand pit Stará Boleslav 50°12’ 14°40’ near the airport Borek Orlík (reservoir) Temešvár 49°21’ 14°16’ Table 1. Geographical details of sampling localities. Stomach contents were observed using an Arsenal particles and digested by crayfish immediately from the MBS-10-100 stereomicroscope and Arsenal LS 1001 substratum), together with more or less recognizable standard light microscope. The frequency of occurrence animal remains (e.g., sclerotized insect heads or legs, of a particular food item was calculated as a percentage mollusk shells), or plant parts (tree roots). of the number of analyzed stomachs containing an Stomach fullness was estimated visually in relation actual food item, i.e., excluding empty stomachs. Where to the potential total volume of a stomach and divided possible, animal remains in stomachs were determined into five classes: A – empty stomach; B – distinctly using identification keys. When not empty, stomachs less than half of the volume (e.g., one insect larva or a mostly contained any of three poorly identifiable low quantity of plant tissues); C and D – partially filled, food items – soft animal tissues (e.g., fresh muscles, around half or distinctly more than half, respectively, of fat bodies), plant tissues (fresh, with recognizable the stomach volume; E – full stomach. lignified vascular tissues), and detritus (particulate Based on the presence or absence of distinct food organic material consisting of minute fragments of items, the trophic index „D“ showing the wideness of the dead organisms exposed for a time among bottom food spectrum [30] was calculated: 60 R. Vojkovská et al. Classes Parameters Qualitative analyzes Quantitative analyzes Size Small (POCL < 13 mm) 38 2 Medium (POCL 13 – 28 mm) 271 83 Large (POCL > 28 mm) 59 27 Σ 368 112 Sex Males 212 62 Females 156 50 (included ovigerous females) (17) 0 Σ 368 112 spring 34 40 Seasons summer 237 33 autumn 54 39 Σ 368 112 Table 2. Number of specimens used for qualitative and quantitative analyses of stomach contents. non-parametric method to test for the equality of = log quantitative contents of main food items among crayfish groups (based on size classes and sexes). where pi is a frequency of occurrence of a distinct food item, and s is a sum of all food items. According to 3. Results Herrera [30] the limit for this index is 0<D<s log N where N is the size of a sample (here, the number of crayfish According to the level of stomach fullness, specimens stomachs analysed). The trophic index D is high in with mostly empty stomachs (i.e., up to half of its volume) generalist species (the theoretical maximum value is dominated in most cases, as shown here for crayfish for species feeding on all food items), and decreases in sexes (Figure 2) and size classes (Figure 3).
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