Life Cycle, Feeding and Production of Isoptena Serricornis(Pictet, 1841
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Internat. Rev. Hydrobiol. 89 2004 2 165–174 DOI: 10.1002/iroh.200310726 TOMÁSˇ DERKA1*, JOSÉ MANUEL TIERNO DE FIGUEROA2 and IL’JA KRNO1 1Department of Ecology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-2, SK-84215 Bratislava, Slovakia; e-mail: [email protected], [email protected] 2Departamento de Biología Animal y Ecología, Facultad de Ciencias, Universidad de Granada, 18071 Granada, Spain; e-mail: [email protected] Life Cycle, Feeding and Production of Isoptena serricornis (PICTET, 1841) (Plecoptera, Chloroperlidae) key words: Plecoptera, feeding, life cycle, production, Slovakia Abstract Some aspects of the biology and ecology (life cycle, feeding and production) of a population of Isoptena serricornis in the Rudava River (Slovakia) are studied, reported and discussed. The life cycle is annual, with slow growth in autumn-winter and fast growth in late summer and spring. The growth decreased two weeks before the Fall Equinox and increased two weeks after the Spring Equinox. The flight period spans from the end of May to the beginning of July. The presence of large sand particles in the gut of all studied nymphs is of note, and indicates that I. serricornis acts as a deposit- collector species. Nymphal food is principally composed of detritus, unicellular organisms and, in nymphs of intermediate or large size, Chironomidae larvae. Adult food is composed fundamentally of different types of pollen grains. Males usually have lower food content than females. Annual produc- tion of this species (~694–750 mg · m–2) is very high in relation to other previously studied Chloro- perlidae. This is probably largely responsible for I. serricornis being one of the most abundant com- ponents of the macroinvertebrate community in its habitat in the Rudava River. A negative correlation between production and temperature was observed. 1. Introduction Isoptena is a monospecific genus (one of the four Chloroperlidae genera present in Europe) with I. serricornis (PICTET, 1841) occurring in Northern, Central and Eastern Europe (ZWICK, 1973). ILLIES (1953) included this taxon among the species penetrating from the east to the west during the post-Pleistocene period. It is a rare and endangered species in Central Europe (KRNO, 1998b; SOLDÁN et al. 1998) and is affected by stream pollution (RAUSERˇ , 1971). The biology of I. serricornis is little known. According to WINKLER (1957), KITTEL (1976, 1980) and LILLEHAMMER (1988), nymphs inhabit rivers and slowly flowing lowland streams with sandy beds, and adults emerge in May-July. It is supposed to be a borrowing animal, probably with a one-year life cycle in Northern Europe (LILLEHAMMER, 1988). This species was reported to be one of the most abundant in the lowland Polish Pilica River, where it lives buried in the sandy bottom (KITTEL, 1976). A study in a Slovakian river (the Rudava River) showed that I. serricornis was strongly associated with the sandy substrate when, along with some members of the Chironomidae, it was the most important component of the macroinvertebrate community (DERKA et al., 2001). KAISER (1977) observed the presence of sand particles in the gut of this species. Recently, a study of I. serricornis eggs pointed out * Corresponding author © 2004 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1434-2944/04/205-0165 166 T. DERKA et al. that the maximum quantity of eggs found per dissected female is less than 50, which shows low fecundity for this species (TIERNO DE FIGUEROA and DERKA, 2003). Studies on the life cycles of some Plecoptera species have been carried out at different latitudes by several authors. Thus, considerable variation in life cycle characteristics has been described. This variation reflects the species studied (SÁNCHEZ-ORTEGA and ALBA-TER- CEDOR, 1991), and also the ecological conditions (particularly the climatic ones). Given this, it is generally accepted that Chloroperlidae have a one-year life cycle pattern (univoltine cycle) or two-year cycle pattern (semivoltine cycle) (HYNES, 1976), but periods of fast and slow growth and egg development are different among the studied species (HYNES, 1976; STEWART and STARK, 1988). In Central Europe, and particularly in Slovakia, life cycles of some Plecoptera species, including some Chloroperlidae species, have been studied in detail (KRNO, 1982, 1984, 1996, 1998a); but previous to now there had not been an annual study on the life cycle of the genus Isoptena. Feeding in adult stoneflies is a poorly known aspect of their biology (TIERNO DE FIGUEROA and FOCHETTI, 2001). Although traditionally it was thought that adult Perloidea species (at least the European ones) did not ingest food, subsequent work demonstrated that some Chloroperlidae [Siphonoperla torrentium (PICTET, 1841)] can metabolize food ingested in the adult stage (RUPPRECHT, 1990) and gain weight through imaginal feeding (ZWICK, 1990). ZWICK (1973) and SURDICK (1985) indicated that some adult Chloroperlidae feed on the pollen of coniferous plants. TIERNO DE FIGUEROA and SÁNCHEZ-ORTEGA (1999) and TIERNO DE FIGUEROA et al. (1998) showed that adults of one species of the family Chloroperlidae (Chloroperla nevada ZWICK, 1967), and three species of the family Perlodidae changed the proportions of dietary components over the flight period. This change reflected the availability of various components. In contrast, it was found that adult feeding was unim- portant in Perlidae and large-sized Perlodidae (TIERNO DE FIGUEROA and SÁNCHEZ-ORTEGA, 1999; TIERNO DE FIGUEROA and FOCHETTI, 2001). In Plecoptera, as in other orders of amphibious insects, nymphal feeding has been studied more extensively than adult feeding (STEWART, 1994). Studies have concentrated on the eco- logical role nymphs play in structuring aquatic communities, e.g., in the processing and cycling of nutrients (MERRIT et al., 1984), as primary consumers (LAMBERTI and MOORE, 1984) or as secondary consumers (PECKARSKY, 1984). According to MERRIT et al. (1984), nymphs of Chloroperlidae can be classified as engulfers according to their feeding mecha- nism. However, STEWART and STARK (1988) pointed out that the food habits of an unstudied species can not be inferred or deduced from the placement of a genus or a higher taxon in a generalized grouping based on studies of congeners. Moreover, the nymphal diet of a par- ticular species can change with individual size during the developmental cycle (BERTHÉLEMY and LAHOUD, 1981; LILLEHAMMER, 1988). The feeding of European Chloroperlidae species in particular is almost unknown. Knowledge of the secondary production of aquatic insects is of considerable ecological importance (applied as well as theoretical) from population and community perspectives. In terms of population dynamics, it combines two parameters that are considered to be of major ecological significance (individual growth and population survivorship) in a single measurement (BENKE, 1984). Nevertheless, study of the contribution of Plecoptera assemblages to overall benthic community production in various biotopes is in its infancy (STEWART and STARK, 1988). In contrast to this general affirmation, studies on stonefly pro- duction in Central Europe have increased in the last twenty years (KRNO, 1982, 1996, 1997, 1998a, 2000). More research is needed, however, to complete this knowledge, especially in lowland rivers which have not yet been studied. Although there have been studies on the production of some European and North American Chloroperlidae species (see BENKE, 1984; STEWART and STARK, 1988), there is no such information published for the genus Isoptena despite its importance in the macroinvertebrate community of its habitat (KRNO et al., 1994; DERKA et al., 2001). © 2004 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Life Cycle, Feeding and Production of Plecoptera 167 The aim of this study is to increase the overall knowledge of Isoptena serricornis biology through discussion of its life cycle, production, and nymphal and adult feeding. In addition to expanding the available information about a particular stonefly species, in this case a monospecific genus whose biology is almost unknown, it is hoped that this study will make an important contribution to the general knowledge of stonefly biology. 2. Study Area The study was carried out in the Rudava River, a tributary of the Morava River in south- western Slovakia. The sampling site (GPS coordinates 48° 30′ 41.1′′ N; 17° 07′ 26.8′′) is situated at 180 meters above sea level, close to Studienka Village. The river catchment area is 280.32 km2, and the average discharge is 0,73 m3 ·s–1. The discharge fluctuated from 0,35 to 2.61 m3 ·s–1. The river width is approximately 7 m, mean depth is 34 cm and maximum depth is 75 cm. The bottom consists of sand (55%), detritus (25.5%), woody debris (14.5%), submersed root mats of riparian tress (3.9%), and hard mud (2.1%). The riparian vegetation consists mainly of the alder Alnus glutinosa (L.) (DERKA et al., 2001). I. serricornis inhabits the study site along with the other stonefly species such as Perlodes dispar (RAMBUR, 1842), Isoperla tripartita ILLIES, 1954, Taeniopteryx nebulosa (LINNAEUS, 1758), Nemoura flexuosa AUBERT, 1949, and Leuctra hippopus KEMPNY, 1899 (KRNO et al., 1994; DERKA et al., 2001). The studied river is not affected by pollution or human activities and is includ- ed among the areas protected by the Ramsar Convention. 3. Material and Methods Quantitative and qualitative samples of I. serricornis nymphs were taken at approximately three-week intervals from July 2001 to June 2002. Three to five quantitative samples were collected using Kubícˇek’s benthic sampler (area 0.1 m2, mesh size 0.5 mm). Qualitative samples were taken using a kick net (mesh size 0.5 mm). Adults were collected during their flying period by sweeping the riparian vegetation (May to June, 2002). Specimens were preserved in 70% alcohol in plastic dram bottles and, in the laboratory, were labelled and preserved in glass vials. Water temperature data were obtained from the Slovak Hydrometeorological Institute. To study the production and life cycle, all collected nymphs were measured (total length) with an ocular micrometer of a binocular microscope.