Energy expenditure and food intake of territorial male Pachydiplax longipennis (Odonata: Libellulidae)

ABSTRACT.

• 1 Field observations of males of the dragonfly, Pachydiplax longipennis (Burmeister), were used to determine how individuals of this species allocate energy to different activities during territory occupancy.
• 2 The effects of biological and physical factors on the species’daily activity pattern were examined. The proportion of time spent in flight was independent of temperature but increased asymptotically with increasing population density.
• 3 Measurements of assimilation efficiency and the quantity of faeces produced per day were used to calculate daily intake of food. An independent estimate of food consumption was derived from data on gut contents and clearance rate.
• 4 Food intake appears to exceed only slightly the energy required to maintain a territory, with little available for other activities. The activity pattern may be determined in part by the amount of energy available to individuals.

     

Factors influencing age related dispersal of the nymphal OdonatesIschnura barberi (Currie),Pachydiplax longipennis (Burm.) andErythemis simplicicollis (Say)

Size related spatial patterns were observed in naiads of the three dominant odonate taxa in a man-made marsh. These patterns were correlated with various physical and chemical parameters, the vegetation gradient, and prey availability using linear and curvilinear regressions.Ischnura barberi andPachydiplax longipennis naiads increased linearly in head capsule widths from the marsh center to the marsh edge. Increasing naiad size correlated with changes in vegetation. Additionally, the spatial patterning ofPachydiplax longipennis was correlated with decreasing submerged plant surface area as naiad size increased. The head capsule width ofErythemis simplicicollis increased from the marsh center to the marsh edge and then continued to increase towards the marsh center. This complex dispersal pattern did not correlate with any of the measured physical or chemical parameters or the vegetation gradient. However, the dispersal pattern ofErythemis simplicicollis correlated with increased in consumable prey densities along the migratory path. No correlation existed between consumable prey and the dispersal pattern ofIschnura barberi orPachydiplax longipennis.     

Foraging behavior ofPachydiplax longipennis (Odonata: Libellulidae)

Food intake, prey availability, and prey capture behavior at feeding areas were quantified in the dragonflyPachydiplax longipennis by observing focal individuals on artificial perches, where they exhibited marked short-term site fidelity. Prey capture success was high and relatively constant, but the frequency and duration of feeding flights depended on the time of day and season (at least in part because of associated variation in temperature and other physical factors), study site, sex, density of other dragonflies, and prey density. Individuals rapidly responded temporally and spatially to changes in prey availability, particularly to localized prey concentrations.     

Microhabitat selection as an antipredator strategy in the aquatic insect Pachydiplax longipennis Burmeister (Odonata: Libellulidae)

Summary An investigation of the larval dragonfly fauna associated with the plant, Sagittaria platyphylla, was conducted in a small pond. Despite the presence of several larval anisopteran species in the pond, only Pachydiplax longipennis larvae were found on Sagittaria plants. A study of the microspatial distribution of P. longipennis larvae on S. platyphylla indicated that larvae use the various regions of a plant in a highly non-random fashion. Larvae show a strong preference for the leaf axil area. A generalized predator, the bluegill sunfish (Lepomis macrochirus), was allowed to selectively eat either of two larvae placed in various plant regions. This experiment indicated that larvae in a leaf axil area were significantly less susceptible to bluegill predation than larvae positioned in other plant regions. The microspatial distribution of starved larvae revealed that larvae with high hunger levels occupied the leaf axil area significantly less than well fed larvae, suggesting 1) larvae do not use these regions as feeding sites, and 2) high hunger levels may induce a behavioral shift in habitat use, with starved larvae forced into areas of high predation risk by the need to fulfill nutritional requirements.     

Fights at the Dinner Table: Agonistic Behavior in Pachydiplax longipennis (Odonata: Libellulidae) at Feeding Sites

Aggressive behavior of Pachydiplax longipennis during foraging was quantified by observing focal individuals on arrays of artificial perches. Pachydiplax apparently aggressively defend, for up to several hours at a time, one or a few feeding perches. Seventeen percent of all behaviors included agonistic actions, e.g., chasing or physical contact. The frequency of interactions was correlated positively with ambient temperature, solar radiation, prey density and density of other dragonflies. Both sexes initiated and responded to intra- and interspecific aggression; intraspecific interactions were more intense, however. Males had significantly higher interaction rates and fighting success than females, and intraspecific male–male contests were particularly intense. When prey were visibly localized, contest winners commonly gained perches closer to the prey swarm, and aggressive behavior was apparently correlated with feeding opportunity. Despite the frequency of aggression, these dragonflies allocated only about 19 s, on average, to agonistic behavior during 30-min observation periods. This and other costs appear small compared to foraging benefits of occupying a favorable perch, although at a very high interaction intensity high energy costs and lower intake reduce the net energy gain.     

Limited male dispersal in a social spider with extreme inbreeding

Cooperatively breeding animals commonly avoid incestuous mating through pre-mating dispersal. However, a few group-living organisms, including the social spiders, have low pre-mating dispersal, intra-colony mating, and inbreeding. This results in limited gene flow among colonies and sub-structured populations. The social spiders also exhibit female-biased sex ratios because survival benefits to large colonies favour high group productivity, which selects against 1 : 1 sex ratios. Although propagule dispersal of mated females may occasionally bring about limited gene flow, little is known about the role of male dispersal. We assessed the extent of male movement between colonies in natural populations both experimentally and by studying colony sex ratios over the mating season. We show that males frequently move to neighbouring colonies, whereas only 4% of incipient nests were visited by dispersing males. Neighbouring colonies are genetically similar and movement within colony clusters does not contribute to gene flow. Post-mating sex ratio bias was high early in the mating season due to protandry, and also in colonies at the end of the season, suggesting that males remain in the colony when mated females have dispersed. Thus, male dispersal is unlikely to facilitate gene flow between different matrilineages. This is consistent with models of non-Fisherian group-level selection for the maintenance of female biased sex ratios, which predict the elimination of male dispersal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 227–234.     

Sperm ejection as a possible cryptic female choice mechanism in Odonata (Insecta)

Abstract.  The few odonate studies of sperm use suggest that females spend apparently more sperm than appears necessary during oviposition (sometimes females may have their sperm stores reduced to 50% after a single oviposition episode). Furthermore, some studies document that females eject sperm during and after copulation. This raises the question of whether sperm reduction may be interpreted as a cryptic female choice mechanism. Using two zygopterans, Ischnura denticollis Burmeister and Enallagma praevarum Hagen, and one anisopteran, Pantala flavescens Fabricius, it is shown that females mate more than once, show a marked reduction in stored sperm, and that this is by ejection of sperm before to oviposition. The extent of sperm reduction is inversely related to the number of eggs laid. When mated to the same male, females show similar reductions in sperm stores and egg load and only rarely does the vaginal duct contain sperm. This suggests that marked sperm reduction is common in this insect order and is not explained by an excess of sperm obstructing the egg passage. It is suggested that female's sperm shortage is better explained as a cryptic female choice mechanism aimed at favouring the sperm of some males. This study provides exciting research avenues for future studies of female choice in an animal taxa whose sexual biology is otherwise regarded as controlled by males.     

Body size and its effect on male-male competition inHylaeus alcyoneus (Hymenoptera: Colletidae)

Body size largely determines the outcome of male-male competition in the banksia bee,Hylaeus alcyoneus. Large males invariably perch on and defend banksia flower spikes, whereas smaller males often nonaggressively patrol circuits that take them repeatedly to several flower spikes. Within the population of males perching on banksia spikes, larger individuals tend to monopolize inflorescences that are higher in banksia shrubs, whereas smaller males often occupy spikes closer to the ground. Perches defended by larger males are more quickly occupied by replacements when the original residents are experimentally removed and held in temporary captivity. When released, the original residents invariably return to and displace the smaller replacements that have taken their territories. When territory takeovers do occur, the winner is almost always larger than the previous resident, showing that residency effects are secondary to body size in determining territorial ownership.     

Observations on the breeding biology of the Writhed-billed Hornbill (Aceros waldeni) in the Philippines

Summary We report on some aspects of the breeding biology of the critically endangered Writhed-billed Hornbill (Aceros waldeni) on the island of Panay, Philippines. Observations were made at three nests during 1995–1997. Walling-in of the females commenced in the first week of March. One female remained incarcerated for 77 days, two of three broods completed fledging around May 20 (1995, 1997). Details on fledging of the female and her brood and postfledging care by both parents are reported.The food of the males at two nests was ca. 98% fruits and 2% invertebrates. The plants exploited comprised at least 14 species. Over a third of the fruits delivered were figs of a small number of species.Two males had average feeding rates of 0.56 and 0.88 times per hour respectively, and fed 1 to 66 (median 8) items per feeding visit at the nest. The hourly feeding rate increased after hatching, but the composition of the diet did not change noticeably. As a rule, food items were delivered singly and, during one visit, in runs of one, or rarely up to 3, species.In the three weeks following vacation of the nest, the male appeared to be the sole food provider while the female stayed continually with the 3 young (as sentinel?) in the vicinity of the nest.The nest environs were defended by the male against Tarictic Hornbills (Penelopides panini panini). Six vocalisations of the parents are mentioned. One was used in territorial skirmishes with Tarictic Hornbills.With perhaps less than 30 pairs of the Writhed-bill surviving, the future for the species looks bleak. Only drastic conservation measures can prevent the species' demise. Some have been started by the PESCP.This paper is publication No. 11 of the Philippine Endemic Species Conservation Project (PESCP) of the Frankfurt Zoological Society.     

Effects of experimental warming on survival,phenology, and morphology of an aquatic insect (Odonata)

1. Organisms can respond to changing climatic conditions in multiple ways including changes in phenology, body size or morphology, and range shifts. Understanding how developmental temperatures affect insect life‐history timing and morphology is crucial because body size and morphology affect multiple aspects of life history, including dispersal ability, whereas phenology can shape population performance and community interactions. 2. It was experimentally assessed how developmental temperatures experienced by aquatic larvae affected survival, phenology, and adult morphology of dragonflies [Pachydiplax longipennis (Burmeister)]. Larvae were reared under three environmental temperatures: ambient, +2.5, and +5 °C, corresponding to temperature projections for our study area 50 and 100 years in the future, respectively. Experimental temperature treatments tracked naturally‐occurring variation. 3. Clear effects of temperature were found in the rearing environment on survival and phenology: dragonflies reared at the highest temperatures had the lowest survival rates and emerged from the larval stage approximately 3 weeks earlier than animals reared at ambient temperatures. There was no effect of rearing temperature on overall body size. Although neither the relative wing nor thorax size was affected by warming, a non‐significant trend towards an interaction between sex and warming in relative thorax size suggests that males may be more sensitive to warming than females, a pattern that should be investigated further. 4. Warming strongly affected survival in the larval stage and the phenology of adult emergence. Understanding how warming in the developmental environment affects later life‐history stages is critical to interpreting the consequences of warming for organismal performance.     

Patch size and breeding status influence movement patterns in the threatened Malleefowl (Leipoa ocellata)

Information on the movement ecology of species can assist with identifying barriers to dispersal and appropriate management actions. We focus on the threatened Malleefowl (Leipoa ocellata) whose ability to move and disperse within fragmented landscapes is critical for their survival. We also investigate the possible effects of climate change on Malleefowl movement. We used solar-powered GPS telemetry to collect movement data and determine the influence of breeding status, remnant vegetation patches and environmental variables. Seven Malleefowl were tracked between 1 and 50 months, resulting in 20 932 fixes. While breeding, Malleefowl had significantly smaller home ranges (92 ± 43 ha breeding; 609 ± 708 ha non-breeding), moved shorter daily distances (1283 ± 605 breeding; 1567 ± 841 non-breeding) and stayed closer to the incubation mound (349 ± 324 m breeding; 3293 ± 2715 m non-breeding). Most Malleefowl effectively disassociated from the mound once breeding stopped, with two birds dispersing up to 10.2 km. Movement patterns were significantly correlated with the size of the remnant native vegetation patch, with smaller home ranges being utilized in small patches than in large patches. One male almost exclusively remained within a 107-ha patch for over 4 years, but a female crossed between closely spaced uncleared patches. Long-range movements of nearly 10 km daily displacement were recorded in large remnants almost exclusively when not breeding. Temperature and rain had a significant effect on movement: modelling suggests daily distances decline from 1.3 km at 25°C to 0.9 km at 45°C, with steeper declines over 30°C. The influence of patch size on movement patterns suggests that Malleefowl movement may be governed by the size of remnant patches and that habitat continuity may be important for facilitating recolonization after catastrophic events and maintaining genetic diversity. Climate change may reduce Malleefowl movement during hot, dry periods possibly affecting breeding success.     

Body size and alternative mating tactics in the beewolf Philanthus zebratus (Hymenoptera; Sphecidae)

A study of the mating behaviour of males of the beewolf Philanthus zebratus revealed that in one population males display variability in mating tactics and that this variability is related to male body size. There was a tendency for large males to patrol the airspace above the nesting area while smaller males were territorial adjacent to it. The mean sizes of the two groups of males were significantly different, although the size ranges of the two groups overlapped. Only 2.5% of the males were observed to undertake both mating tactics, at different times. Observations are presented on daily and seasonal activity patterns and on the relative location of nests, territories, and patrolling males. A second population, with lower nest density, was observed for several days, revealing only territorial males. It is suggested that the presence of patrolling males is related to the higher nest density of the one population. The fact that patrolling males tend to be relatively large is possibly related to flight energetics or simply to the ability of large males to seize females, which are usually larger than males, in mid-air.     

A comparative analysis of senescence in adult damselflies and dragonflies (Odonata)

Any population whose members are subject to extrinsic mortality should exhibit an increase in mortality with age. Nevertheless, the prevailing opinion is that populations of adult damselflies and dragonflies do not exhibit such senescence. Here, we challenge this contention by fitting a range of demographic models to the data on which these earlier conclusions were based. We show that a model with an exponential increase in age-related mortality (Gompertz) generally provides a more parsimonious fit than alternative models including age-independent mortality, indicating that many odonates do indeed senesce. Controlling for phylogeny, a comparison of the daily mortality of 35 odonate species indicates that although male and female mortalities are positively correlated, mortality tends to be higher in males of those species that exhibit territoriality. Hence, we show for the first time that territoriality may impose a survivorship cost on males, once the underlying phylogenetic relationships are accounted for.     

Sexual selection, sexual size dimorphism and Rensch's rule in Odonata

Odonata (dragonflies and damselflies) exhibit a range of sexual size dimorphism (SSD) that includes species with male-biased (males > females) or female-biased SSD (males < females) and species exhibiting nonterritorial or territorial mating strategies. Here, we use phylogenetic comparative analyses to investigate the influence of sexual selection on SSD in both suborders: dragonflies (Anisoptera) and damselflies (Zygoptera). First, we show that damselflies have male-biased SSD, and exhibit an allometric relationship between body size and SSD, that is consistent with Rensch's rule. Second, SSD of dragonflies is not different from unit, and this suborder does not exhibit Rensch's rule. Third, we test the influence of sexual selection on SSD using proxy variables of territorial mating strategy and male agility. Using generalized least squares to account for phylogenetic relationships between species, we show that male-biased SSD increases with territoriality in damselflies, but not in dragonflies. Finally, we show that nonagile territorial odonates exhibit male-biased SSD, whereas male agility is not related to SSD in nonterritorial odonates. These results suggest that sexual selection acting on male sizes influences SSD in Odonata. Taken together, our results, along with avian studies (bustards and shorebirds), suggest that male agility influences SSD, although this influence is modulated by territorial mating strategy and thus the likely advantage of being large. Other evolutionary processes, such as fecundity selection and viability selection, however, need further investigation.     

Fine-scale spatial genetic structure and dispersal among spotted salamander (Ambystoma maculatum) breeding populations

We examined fine-scale genetic variation among breeding aggregations of the spotted salamander (Ambystoma maculatum) to quantify dispersal, interpopulation connectivity and population genetic structure. Spotted salamanders rely on temporary ponds or wetlands for aggregate breeding. Adequate breeding sites are relatively isolated from one another and field studies suggest considerable adult site fidelity; therefore, we expected to find population structure and differentiation at small spatial scales. We used microsatellites to estimate population structure and dispersal among 29 breeding aggregations in Tompkins County, New York, USA, an area encompassing 1272 km(2). Bayesian and frequency-based analyses revealed fine-scale genetic structure with two genetically defined demes: the North deme included seven breeding ponds, and the South deme included 13 ponds. Nine ponds showed evidence of admixture between these two genetic pools. Bayesian assignment tests for detection of interpopulation dispersal indicate that immigration among ponds is common within demes, and that certain populations serve as sources of immigrants to neighbouring ponds. Likewise, spatial genetic correlation analyses showed that populations < or = 4.8 km distant from each other show significant genetic correlation that is not evident at higher scales. Within-population levels of relatedness are consistently larger than expected if mating were completely random across ponds, and in the case of a few ponds, within-population processes such as inbreeding or reproductive skew contribute significantly to differentiation from neighbouring ponds. Our data underscore the importance of these within-population processes as a source of genetic diversity across the landscape, despite considerable population connectivity. Our data further suggest that spotted salamander breeding groups behave as metapopulations, with population clusters as functional units, but sufficient migration among demes to allow for potential rescue and recolonization. Amphibian habitats are becoming increasingly fragmented and a clear understanding of dispersal and patterns of population connectivity for taxa with different ecologies and life histories is crucial for their conservation.