The effects of temperature and prey density on the development rates and growth of damselfly larvae (Odonata: Zygoptera)

Abstract. 1. The effects of prey density and temperature on the feeding and development rates of several late instars of the larvae of three common European damselflies ( Lestes sponsa (Hansemann), Coenagrion puella (L.) and Ischnura elegans van der Linden) were investigated in a laboratory experiment.
2. Functional responses were used to estimate maximum feeding rates. Maximum feeding rates were compared between species, instars and temperatures by expressing prey consumption in terms of prey biomass consumed per unit predator biomass. Lestes was capable of feeding at almost twice the rate of either Coenagrion or Ischnura.
3. Higher feeding rates led to faster development rates and there was an interaction between species and temperature. With the exception of those with very low feeding rates, larvae maintained at higher temperatures, but similar feeding rates, developed faster. Under similar conditions of temperature and feeding rate, Lestes larvae developed faster than larvae of either Coenagrion or Ischnura.
4. Faster development rates at similar rates of food intake were achieved at the cost of reduced size-increases between instars.
5. The differences in the responses of the three species are discussed in the light of their respective life histories, and with reference to a recent model of population regulation in damselflies.     

Nitrogen deposition and herbivory affect biomass production and allocation in an annual plant

If environmental conditions vary, plasticity in life-history traits is predicted. A recent model indicates that males and females should differ in life-history traits, because sexes differ in optimal attributes depending on species ecology. In this study we test the impact of two biotic factors in combination (presence/absence of predators and low/high food level) on gender specific life-history traits in the damselfly Coenagrion puella (Odonata). Results show that predator presence and low food density decreased activity in both sexes. Additionally, individuals with less food grew more slowly, emerged later, remained smaller and had a higher mortality. At low food densities, however, and in contrast to former investigations, individuals from treatments with predator presence were the same size or larger than individuals without predators. Gender had a strong impact on larval activity and life-history traits and sexes differed in development. Females were less active and took longer to complete development, but emerged at a larger size, weight and fat content. This study highlights the importance of gender specific approaches in life-history research.     

Asymmetric intraspecific competition among larvae of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Abstract. 1. A laboratory competition experiment is described in which the growth and development rates of larvae of the damselfly Ischnura elegans (Lind.) were measured over an entire instar.
2. Two larval instars which commonly occur together in the field were used in the experiment; they were maintained with a superabundance of prey and either larvae from the same or the larger/smaller instar.
3. Small larvae suffered increased development times and decreased size increases at the moult in the presence of large larvae but similar interference effects were not evident when these smaller larvae were in the presence of other small larvae.
4. Development time and size increases of large larvae were not significantly affected by the presence of small larvae.
5. Irrespective of the instar combinations investigated, interference effects were reduced when there were more perches available, although in only a few cases was this reduction significant.
6. The consequences of the asymmetric competition reported in the experiment for the study of lifetime reproductive success in damselflies are discussed. Late emerging adults may incur reduced reproductive success.     

Embryonic development, larval growth and life cycle of Coenagrion puella (Odonata: Zygoptera) from an Austrian pond

SUMMARY. 1. Newly-laid eggs of Coenagrion puella (L.) from a pond near Herzogenburg (Lower Austria) were kept at constant water temperatures (range c .3.5°C to c .28°C)in the laboratory. Hatching success varied with temperature; no eggs hatched below 12°C and nearly all hatched at c .l6°C. Hatching time decreased with increasing temperature and the relationship between the two variables within the range 12–28 °C was well described by a power law. The length of the hatching period was less than 12 days. Hatching times estimated from the power-law equations and those obtained in the field experiments were similar. Therefore both the hatching time and the length of the hatching period in the field could be estimated from the laboratory data for the range 12–28°C.
2. The maximum number of instars from egg to imago was 11; the average body length increment (mm) per moult was proportionately constant at c .26% and Dyar's rule was applicable. The interval between moults decreased with increasing temperature up to the seventh instar and the relationship between the two variables within the range 12–28°C was well described by a power law. The moulting interval for instars 8–11 ranged from 23 to 48 days and was relatively independent of temperature. No moulting occurred at temperatures below 12°C.
3. Larval growth was logistic in the laboratory and variations in mean logistic growth rate (range 0–2.5% length day⁻¹) were related to mean temperature with no growth at temperatures <12°C. Larval growth rates in pond experiments were similar to those estimated from laboratory data, and therefore the regression equations obtained from the laboratory experiments are probably applicable to larval growth in the field.
4. Information on the life cycle of C. puella is briefly reviewed and it is concluded that C. puella from the pond near Herzogenburg has an univoltine life cycle.     

Regulation of damseifly populations: the effects of larval density on larval urvival, development rate and size in the field

SUMMARY. 1. The popuhttion density of Coenagrion pttella larvae was monitored in five populations, and of Ischntira elegans in two populations, between October 1982 and May 1983.
2. There was no measurable mortality of larvae over winter and no larval growth until April. Larvae in high density populations were smaller than those in low density populations and were more likely to have a semi- voltine life history.
3. The population density of C. ptiella was also monitored (more frequently) in two populations with differenl initial densities between July and November 1983. In the high density population there was a constant rate of larval mortality, while in the low density population there was no detectable larval mortality, indicating that larval mortality may be density dependent. Larvae in the high density population were again smaller, and more likely to be semi-voitine, than those in the low density population.
4. The role of density dependent larval growth, development and mortality in the regulation of damseifly populations is discussed.     

Effects of temperature and perch diameter on arboreal locomotion in the snake Elaphe guttata

Arboreality is widespread in multiple lineages of snakes and these habitats are important for foraging, escaping predators, and thermoregulation for many species. However, very little is known about factors influencing the arboreal locomotor abilities of snakes. Arboreal performance was assessed in a semi-arboreal snake (Elaphe guttata) using an artificial perch apparatus. Locomotor velocity, body posture, and balance was measured during movement on three perch diameters (3, 6, 10-cm) at three temperatures (10, 20, 30 degrees C). Velocities attained by E. guttata on perches are much slower than those of terrestrial lateral undulation and swimming and somewhat slower than concertina velocities recorded in other species across the same experimental temperatures. At higher temperatures, faster speeds were associated with a more elongated posture. At lower temperatures, snakes displayed a more looped body posture, but still fell more often than at higher temperatures. Our results suggest that temperature has a large influence on both balance and movement by snakes on perches. Although there were no differences in velocities resulting from perch diameter, snakes fell more often from thicker perches. This differs from arboreal velocities attained by limbed vertebrates, which decrease with decreasing perch diameter, suggesting that snakes have a size-relative advantage over limbed animals, such as lizards, when traversing a network of narrow branches. Future studies investigating arboreal locomotion among snakes that vary both phylogenetically and morphologically are needed to assess the potential benefits of limblessness in complex, three-dimensional environments.     

Better Hosts Dive: Detachment of Ectoparasitic Water Mites (Hydrachnellae: Arrenuridae) from Damselflies (Odonata: Coenagrionidae)

Water mite larvae parasitizing damselflies must detach while the host is in a suitable reproduction habitat for both parasites and itself. They should do so during the host's oviposition. In this paper I present experimental data for the detachment rate of water mite larvae (Arrenurus cuspidator) from different host species, Coenagrion hastulatum and C. puella, in relation to the host's oviposition behavior. C. hastulatum oviposits submerged, whereas C. puella oviposits at the water surface and aggregates with conspecifics. It was found that mite larvae detach at a significantly higher ratio from hosts with submerged oviposition. Experimental tests showed that this is not a species-specific effect. It is caused mainly by the oviposition behavior. The results are discussed in the light of different oviposition systems in damselflies.     

Effects of previous diet and frequency of feeding on development of larval damselflies

SUMMARY. 1. Food availability during instar F-3 affected duration of instar F-3 of larval Ischnura vertiealis (Say) maintained in the laboratory but did not affect the increase in head width at the moult to instar F-2.
2. Food availability during instar F-2 altered both duration in instar F-2 and weight per unit head width of newly moulted F-l larvae, but not the increase in head width at the moult to instar F-1.
3. Food availability during instar F-3 had no effect on instar duration, weight per unit head width, or increase in head width of F-2 larvae at the moult to instar F-l.
4. Survival of larval Coenagrion resolution (Hagen) reared in the laboratory was dependent on mean number of Daphnia offered per day. In addition, larvae fed less frequently, but on the same mean number of Daphnia per day. had lower survival rates than larvae fed more frequently.     

Does substrate quality influence take-off decisions in Common Starlings?

1. An instrumented perch was used to measure the reaction forces during take-off of starlings from perches of three thicknesses. There was no significant decrease in reaction force with decreasing perch thickness; however, over the range of perch sizes there was a 10-fold decrease in safety factor and a 400-fold increase in the energy of deflection of the thinnest perch.
2. The implications of these results are that birds do not change their take-off strategy when faced with perches of variable thickness and that, as a result, leaps from thin perches are very much less efficient than those from thicker perches. Birds appear not to adjust their leaping behaviour to minimize the risk of substrate failure during take-offs.     

Can bivalve veligers escape feeding currents of adult bivalves?

While the stock of introduced Pacific oysters (Crassostrea gigas) increased in the Oosterschelde estuary (SW Netherlands), so did the filtration pressure of all bivalve species together. In the same period, stocks of native bivalves declined slightly. The expansion of Pacific oysters in Dutch estuaries might be partially due to better abilities of their larvae to avoid or escape filtration, compared to larvae of native bivalves. In this context, escape and swimming abilities of Pacific oyster larvae and the larvae of the native blue mussel (Mytilus edulis) were compared.Swimming behaviour of C. gigas larvae and larvae of M. edulis was recorded in still water and in a suction current mimicking a bivalve feeding current, in a horizontal and in a vertical plane. Larval swimming behaviour in a suction flow field was reconstructed by subtracting local water movement vectors from the total movement of larvae, yielding movement paths due to larval swimming alone.Swimming speeds and the rate of displacement in vertical direction of C. gigas and M. edulis larvae were related to larval shell length, and to the pitch of up- or downward swimming.Larvae of both species did not show escape reactions in a suction flow field. With increasing shell length, larval swimming speeds of both species increased significantly. Swimming speeds of C. gigas larvae were significantly higher than swimming speeds of M. edulis larvae, resulting in a faster vertical displacement. The ability to migrate to more favourable water layers faster may offer C. gigas an advantage over native bivalves with slower swimming larvae.     

Pathways to fitness: carry‐over effects of late hatching and urbanisation on lifetime mating success

Life history theory and most empirical studies assume carry‐over effects of larval ­conditions to shape adult fitness through their impact on metamorphic traits (age and mass at metamorphosis). Yet, very few formal tests of this connection across metamorphosis exist, because this entails longitudinal studies from the egg stage and requires measuring fitness in (semi)natural conditions. In a longitudinal one‐year common‐garden rearing experiment consisting of an outdoor microcosm part for the larval stage and a large outdoor insectary part for the adult stage, we studied the effects of two factors related to time constraints in the larval stage (egg hatching period and urbanisation) on life history traits and lifetime mating success in the males of the damselfly Coenagrion puella. We reared early‐ and late‐hatched larvae from each of three rural and three urban populations from the egg stage throughout their adult life. Key findings were that both the hatching period and urbanisation shaped adult fitness, yet through different pathways. As expected, the more time‐constrained late‐hatched individuals accelerated their larval life history and this was associated with a lower lifetime mating success. A path analysis revealed this carry‐over effect was mediated by the changes in the two metamorphic traits (reduced age and lower mass at emergence). Notably, urban males had a 50% lower lifetime mating success, which was not mediated by age and mass at emergence, and possibly driven by their shorter lifespan. Our results point to long‐term carry‐over effects of the usually ignored natural variation in egg hatching dates, and further contribute to the limited evidence showing fitness costs of adjusting to an urban lifestyle.     

Characteristics of dispersing Ischnura elegans and Coenagrion puella (Odonata): age, sex, size, morph and ectoparasitism

In this study we assessed whether individuals of the damselfly species Ischnura elegans and Coenagrion puella that moved between ponds differed in their mean characteristics from individuals that did not move. Overall, the sex (female) and species ( C. puella ) that spent the most time away from the breeding site was more likely to move between ponds. Ischnura elegans males that dispersed had significantly longer forewings than males that did not, while male C. puella parasitised by water mites were more likely to disperse than unparasitised males. There was no evidence for differences in dispersal rates among the female colour forms of either I. elegans or C. puella . In general, the differences in dispersal characteristics between sexes and species could be explained by underlying variation in activity and mobility. The majority of dispersal between breeding sites by C. puella and I. elegans did not appear to be directed, but probably arose from chance movements occasionally taking individuals to a different pond from which they emerged.     

The responses of carrot fly larvae, Psila rosae, to components of their physical enrivonment

Abstract. 1. The responses of third instar Psila rosae (F.) larvae to light, temperature, humidity and soil moisture were investigated in the laboratory.
2. Larvae were photonegative and preferred a temperature of about 15°C. Temperatures between 30 and 40°C adversely affected movement and over 40° C were lethal.
3. In choice chambers, larvae preferred humidities of 70–100% r.h. and larvae in sand avoided dry conditions (2.5% field capacity). The latter response became more marked as larvae approached the pre-pupal stage when moistures of 40% field capacity and lower were avoided.
4. Most larvae were found at a depth of 8 cm in sand of uniform moisture content and temperature, but variation in moisture content could alter this preference.
5. In August, most larval damage in the field occurred near the tip of the carrot tap root but was more evenly distributed over the roots in November. It is uncertain whether this was due to soil near the surface being drier in August or whether it was caused by behavioural differences between the two generations of carrot fly larvae.
6. During the summer of 1975, low soil moisture levels resulted in the total absence of larval mines on the carrot roots even though pupae were found at depths of 20–30 cm in the soil. Temperature had no effect on the distribution of mines on carrot roots except at the top 2 cm of the soil profile.     

Effects of predator-induced thinning and activity changes on life history in a damselfly

We investigated how the lethal and non-lethal presence and absence of a fish predator, perch (Perca fluviatils), influenced behaviour, numbers emerging, size at emergence, and development rate of the damselfly Lestes sponsa. The experiment was carried out in outdoor artificial ponds and spanned from the egg stage to emergence of the damselflies. During the experiment food resources for the damselflies were continuously monitored. Damselflies exposed to a lethal predator showed a significantly lower activity level than those in the absence of predators or subjected to a non-lethal predator. Half-way through the larval stage the reduction in activity level was correlated with the presence of lethal predators, and at the end of the larval stage with higher zooplankton densities. Though larvae decreased activity level, size at emergence was larger and development time faster for individuals in the lethal predator treatment. Since fewer larvae emerged from that treatment we interpret the larger size at emergence to be an effect of a combination of thinning and higher zooplankton densities.     

Distribution of perch (Perca fluviatilis L.) during their first year of life in Lake Constance

The distribution and behaviour of larval and juvenile perch (Perca fluviatilis L.) were studied for two years in large, deep Lake Constance. After hatching larvae were transported by water currents to the open water. The majority of larvae remained in the pelagic zone for about one month. In both years, their return to the littoral zone coincided with the decline of pelagic zooplankton abundance. After returning to the littoral zone, juveniles stayed among submerged macrophytes within 5 m depth and lived apart from larger perch which lived at depths of 6–20 m. By late summer, juveniles changed their distribution pattern: during the day they stayed intensively close to piers and ports, but increased their swimming activity at dusk, cruising among shallow and deep waters and feeding on zooplankton, and rested on the bottom at night. This behaviour appears to be related to the decrease of inshore food resources and to the presence of predators in deeper water. 0⁺ perch left the littoral zone and moved into deep waters when autumnal mixing began in late October. They overwintered near the bottom at depths of more than 30 m. During most of the year, juvenile and adult perch were separated from each other. But as soon as they occupied the same habitat, the occurrence of cannibalism increased.     

Parasitism increases offspring size in a damselfly: experimental evidence for parasite-mediated maternal effects

The effects of parasites on host fitness and the fitness effects of maternal effects are widely discussed. In this study, I conducted an experiment linking both aspects. I manipulated the ectoparasite load (Acari: Arrenurus cuspidator) of damselflies, Coenagrion puella, and found that larvae from mothers with high parasite loads were larger (assessed by head width) than larvae from mothers with low parasite loads. Furthermore, there was a negative correlation between the number of eggs laid and parasite load. Parasitized mothers thus seemed to have fewer, but probably better, offspring. The ecological significance of these parasite-mediated maternal effects remains to be tested. However, size-dependent cannibalism almost certainly has important consequences for population dynamics. Copyright 1999 The Association for the Study of Animal Behaviour.     

Population biology and behavior of larval Eustrongylides tubifex (Nematoda: Dioctophymatida) in poikilothermous hosts

A total of 178 channel catfish (Ictalurus punctatus), 162 freshwater drum (Aplodinotus grunniens), 60 smallmouth bass (Micropterus dolomieui) and 2,488 yellow perch (Perca flavescens) were examined for larval Eustrongylides tubifex (Nitzsch 1819) J?gerskiold 1909 during 1973 and 1974. Older, large fish exhibited higher prevalence, larger mean numbers of larvae in infected fish, larger mean numbers of larvae in all fish examined, and wider range of intensity of infection. Infections were most extensive in yellow perch. The majority of larvae were removed from mesenteric cysts. Under experimental conditions, larvae migrated from cysts through the body wall when environmental temperatures reached or exceeded 40 C. Fourth-stage larvae removed from yellow perch successfully reinfected a variety of poikilothermous experimental hosts. Both young-of-the-year and I+ age/size class yellow perch exhibited increasing monthly larval worm burdens during portions of the sampling period. Older (II+) age/size class yellow perch did not exhibit significantly increasing worm burdens. No significant differences were noted in numbers of male and female larvae or in worm burdens in male and female yellow perch.     

Hybridization between redstart Phoenicurus phoenicurus and black redstart P. ochruros, and the effect on habitat exploitation

Although redstarts Phoenicurus phoenicurus and black redstarts P. ochruros breed in different habitats they have been found to interbreed and to produce viable and fertile offspring, which backcross to the parental species. In a dual choice experiment, I investigated the preference of black redstarts and redstarts, and F₁-hybrids for different habitat structures in an aviary. Black redstarts strongly preferred a perch type on which they could stand to a perch they had to cling to. Redstarts were less selective and used both perch types equally as long as food was offered between both perches on the ground. If food was given at one of the perches redstarts changed their preference depending on food location. Black redstarts, however, showed only a slight response. They always preferred the same perch type. Hybrids behaved like redstarts. They did not show a preference as long as food was placed on the ground. When food was offered at one of the perch sites they more often chose the one with food. Due to high phenotypic variability among individuals perch type preference overlapped between the three experimental groups.     

Ectoparasites do not affect ejaculate volume in the dragonfly Coenagrion puella

Abstract. Imagoes of the dragonfly Coenagrion puella are parasitized frequently by ectoparasitic water mites. In an experimental study of the parasite load we examined the influence of parasite burden on host sperm volume. Infection with ectoparasitic water mites did not affect sperm volume in the seminal vesicle (ejaculate volume). It is concluded that water mite parasitism does not affect male fitness in C. puella by reducing sperm production.     

Food competition among Scotophugu stercoraria Iarvae with emphasis on its effects on reproductive success

Abstract. 1. Laboratory experiments with yellow dung fly larvae, Scatophaga sfercoraria L. were carried out in the hope of clarifying the causal relationship underlying the extreme variation in body size in wild populations.
2. First, parents were collected in the field and newly hatched larvae exposed to different density levels under two temperature regimes (15°C and 20°C). Second, heritability experiments were done.
3. The results showed that density during the larval stage has a very significant effect on adult size. Male size is more affected than female size and the male larvae spend more time feeding than the female larvae with the result that the males are on average larger. Also, the additive genetic component is of little or no significance, supporting the notion that body size is positively related to fitness.
4. Finally, the results are discussed in relation to field data on the effects of size on both male and female reproductive success.