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Mesoveliidae), Microvelia Reticulata and Velia Caprai (Veliidae) (Heteroptera: Gerromorpha

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Mesoveliidae), Microvelia Reticulata and Velia Caprai (Veliidae) (Heteroptera: Gerromorpha Eur. J. Entomol. 107: 579–587, 2010 http://www.eje.cz/scripts/viewabstract.php?abstract=1569 ISSN 1210-5759 (print), 1802-8829 (online) Effect of population density on the development of Mesovelia furcata (Mesoveliidae), Microvelia reticulata and Velia caprai (Veliidae) (Heteroptera: Gerromorpha) TOMÁŠ DITRICH1, 2 and MIROSLAV PAPÁýEK1 1Department of Biology, Faculty of Education, University of South Bohemia, Jeronýmova 10, 371 15 ýeské BudČjovice, Czech Republic; e-mails: [email protected]; [email protected] 2Department of Ecosystem Biology, Faculty of Science, University of South Bohemia, Branišovská 31, 370 05 ýeské BudČjovice, Czech Republic Key words. Population density, developmental rate, wing dimorphism, body size, oogenesis, Mesoveliidae, Veliidae, Heteroptera Abstract. Population density during nymphal development affects body size, developmental rate and wing polymorphism in semia- quatic bugs. Nymphs from crowded habitats grow faster and thus gain an advantage in the later stadia. Rapid development results in smaller body size in several gerrids. Macropterous adults develop more frequently at high population densities in most species, which enables the bugs to leave crowded habitats. Three European widespread species Mesovelia furcata Mulsant & Rey, 1852 (Mesoveliidae), Microvelia reticulata Burmeister, 1835 and Velia caprai Tamanini, 1947 (Veliidae) were reared individually and simultaneously either at a low or high population density. Duration of postembryonic development, wing morph, body size and length of distal oocyte in females were recorded. High population density accelerates development in Mesovelia furcata and Micro- velia reticulata. However, there was no trade-off between developmental rate and body size. Accelerated development without a decrease in body size was probably because maturation was delayed. Individually reared nymphs developed faster than nymphs from communal cohorts. No long-winged Microvelia reticulata specimen developed in any treatment. However, more macropterous indi- viduals developed in high-density treatments in Mesovelia furcata (significant) and Velia caprai (not significant). All the nymphs of the species that were reared individually developed into apterous adults. The results suggest that population density strongly influ- ences the life history of semiaquatic bugs. However, the only commonly shared response seems to be an increase in developmental rate when reared at high population densities. Other traits such as wing dimorphism, body size and rate of oogenesis differ at the spe- cies level. INTRODUCTION 1982). Incidence of the different wing morphs depends Crowding during nymphal development affects the life on genetic background and/or environmental conditions history of many insect species. A traditional object of during nymphal development (Spence & Andersen, research is the “gregaria phase” that develops in crowded 1994). Vepsäläinen (1973) has proposed a general model populations of Orthoptera and several other herbivores for the family Gerridae, which predicts a higher propor- including blissid bugs, noctuid caterpillars, aphids and tion of macropterous individuals in ephemeral and tempo- planthoppers (see Applebaum & Heifetz, 1999 for rary habitats. review). Population density strongly influences not only One of the main factors affecting the rate of develop- the life of plant feeding species, but also the traits of pre- ment and flight capability is probably population density. daceous semiaquatic bugs (Heteroptera: Gerro- Nymphs that are crowded should have a faster rate of morpha). Most gerromorphan bugs live on the surface of development and proportionally more of them should water, which is a well-restricted 2D zone, so their popula- develop into macropterous adults, which are capable of tion density is well defined. Non-migratory individuals dispersing. However, the responses of various species (e.g. nymphs and flightless adults) usually cannot easily reared at high vs. low population densities differ. As in escape from areas where population density is high. The most insect species (Applebaum & Heifetz, 1999) more habitat can thus become crowded when the area of water long winged (LW) individuals develop at high population surface decreases during droughts or when the eggs hatch densities in Aquarius paludum (Gerridae) (Harada et al., simultaneously. Population density of gerrid nymphs can 1997), Microvelia douglasi, Microvelia horvathi and reach up to several hundreds per m2 in the field (Harada Microvelia kyushuensis (Veliidae) (Muraji et al., 1989; & Spence, 2000). Muraji & Nakasuji, 1988). On the other hand, proportion- Semiaquatic bugs are predators and scavengers, feeding ally more micropterous specimens develop at high popu- on different arthropods trapped in the surface film of lation densities in species such as Gerris buenoi (Harada water. Almost all temperate species exhibit wing poly- & Spence, 2000). Gerris pingreenensis exhibits no or morphism. Most species have macropterous, brachyp- little effect of population density on wing morph develop- terous / micropterous and apterous morphs (Andersen, ment and nymphal duration (Harada & Spence, 2000). At high nymphal population densities the duration of nym- 579 phal development decreases in all gerrids tested (Harada, Culture conditions 1996; Harada & Spence, 2000; Harada et al., 1997). More Three different series of aquaria, all 10 cm high and filled to a rapid postembryonic development is advantageous when depth of 2 cm with water were used for the communal rearing of intraspecific competition is intense. nymphs: 10 × 15 cm for Mes. furcata, 10 × 12 cm for Micr. Nymphal population density during development also reticulata and 15 × 20 cm for V. caprai. influences body size and the physiological characteristics High and low population density groups were established of the adults. This effect can be sex dependent. When G. with 60 and 10 nymphs, respectively. The high-density groups thus corresponded to 50, 40 and 20 individuals/100 cm2, respec- buenoi and G. pingreenensis were reared at a low popula- tively. There were ten replicates of every low population density tion density, females were larger than those reared at a treatment and two of the high population density treatments. high population density, but not in the case of males For the individual treatments, 1st instar individuals of Micr. where there was no significant effect (Harada & Spence, reticulata and Mes. furcata were placed singly into 6 × 5 cm 2000). Females of A. paludum reared at a high population plastic containers and V. caprai into 8 × 6 cm containers. All density have a longer preoviposition period than those containers were filled to a depth of 1 cm with tap water. There reared at low population density, but other gerrids do not were forty-eight replicates of this treatment for each of the three show this response (Harada & Spence, 2000; Harada et species. All experiments were carried out at a constant tempera- al., 1997). Apparently, the response to different popula- ture 19 ± 1°C and under a long-day photoperiod of 16L: 8D. Several pieces of polystyrene were placed in every aquarium as tion densities experienced during development, expressed resting sites. in terms of wing polymorphism, adult body size and pre- Specimens in all groups were fed frozen Drosophila mela- disposition to reproduce, varies among Gerridae. nogaster daily as follows: Micr. reticulata: 0.5 flies per 1st instar This research is focused on the effect of nymphal popu- nymph; 1 fly per 2nd and 3rd instar and 2 flies per 4th and 5th lation density (treatments: isolated nymphs and nymphs instar; Mes. furcata: 1 fly per 1st and 2nd instar nymph, 2 flies reared at low and high population densities) on wing per 3rd instar and 3 flies per 4th instar and V. caprai: 1 fly per 1st polymorphism, body size and rate of gametogenesis in instar nymph and up to 5 flies per 5th instar nymph. three common and widely distributed temperate Old The aquaria were checked every day between 11 a.m. and 1 World non-gerrid species of semiaquatic bugs, Mesovelia p.m., flies were replaced with fresh ones, exuviae and dead nymphs removed. Water was changed every second day. Adults furcata, Microvelia reticulata and Velia caprai. These were removed immediately they developed and placed into species differ in their habitat preference, body size, life Bouin’s fixative. history traits (Saulich & Musolin, 2007) and phyloge- netical positions (relatedness) (Damgaard, 2008a, b). Source of nymphs Overwintered adults of Micr. reticulata (60 WL specimens) MATERIAL AND METHODS were collected during May 2008 at three ponds in South Terminology, abbreviations and species studied Bohemia (Motovidlo pond, GPS 48°59´58˝N, 14°22´52˝E; Jarval pond, 48°59´30˝N, 14°32´51˝E and Jizba pond, Following abbreviations are used in the text: LW – long- 48°52´50˝N, 14°26´22˝E) and reared at 24 ± 2°C under a natural winged, macroptery, macropterous; SW – short-winged, bra- photoperiod. All adults were kept in an aquarium (30 × 20 cm) chyptery, brachypterous; WL – wingless, aptery, apterous. Mes. filled to a depth of 2 cm with water, with several pieces of poly- – Mesovelia, Micr. – Microvelia. Last-mentioned abbreviations styrene as oviposition sites. Any eggs laid were removed daily are used instead of the more commonly used generic name and placed in an empty aquarium. Only nymphs that hatched abbreviation “M.” for both genera, in order to more clearly dif- within 12 h of one another were placed as a group
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