Morphological and behavioral defenses in dragonfly larvae: trait compensation and cospecialization

Many animals have two basic traits for avoiding being killedby a predator: behavioral modification and morphological defense.We examined the relationship between antipredator behavior andmorphological defense in larvae of three closely related dragonflyspecies within the genus Leucorrhinia. The three species differwith regard to their morphological defense as expressed in thelength of the larval abdominal spines. Results showed that longerabdominal spines provided protection against an attacking fishpredator (perch) because the probability of being rejected afteran attack was significantly higher in the species with the longestabdominal spines. In contrast to other studies, the specieswith the strongest morphological defense did not show the leastbehavioral predator avoidance. Instead, the species with intermediatemorphological defense showed the least predator behavioral avoidance.The results suggest that the Leucorrhinia system is a mixtureof trait cospecialization (a positive correlation between antipredatorbehavior and morphological defense) and trait compensation (anegative correlation between antipredator behavior and morphologicaldefense). Differences in the relationship between morphologicaland behavioral defense between species might be related to abundancepatterns of the three species in lakes with and without fishpredators.     

Behaviour and growth of dragonfly larvae along a permanent to temporary water habitat gradient

Abstract.  1. Freshwaters form a gradient from small temporary waters to large permanent waters. Identifying and examining traits that restrict the distribution of species along this gradient are crucial to the understanding of community structure in these habitats.
2. Using dragonfly larvae species, differences in traits important for growth and survival were studied.
3. The traits were studied in a series of laboratory experiments using two pairs of dragonfly species that coexist in the Namibian semi-desert. One species pair was from the most temporary part of the water permanence gradient and the other species pair from an intermediate part of the gradient.
4. As predicted, activity, capture rate, and growth rate were significantly greater in the two temporary water species. Contrary to the prediction made in the work reported here, species differences in microhabitat selection were not related to the species' habitat origin. Cannibalism did not differ between species.
5. The results lend support to the hypothesis that selection has favoured certain combinations of trait values and that these traits are important for a successful life in temporary and permanent waters.     

Relationship between morphology,dispersal and habitat distribution in three species of Libellula (Odonata: Anisoptera)

Morphology is an important determinant of flight performance and can shape species’ dispersal behaviour. This study contrasted the morphology of flight-related structures in dragonfly species with different dispersal behaviours to gain insights into the relationship between morphology and dispersal behaviour. Specifically, wing size, wing shape and thorax size were compared in three co-occurring species from different clades within the genus Libellula (Odonata: Anisoptera: Libellulidae) to assess how these morphological traits are related to differences in dispersal behaviour and to how broadly their larvae occur across a habitat gradient. Two species had broad larval habitat distributions as well as high rates and distances of dispersal. These two species had relatively larger wings and thoraces than the third species, which was found only in permanent lakes and had limited dispersal. The hind-wings of more dispersive species also had lower aspect ratios and a relatively wider basal portion of the wing than the less dispersive species. Broad hind-wings may facilitate the use of gliding flight and reduce the energetic costs of dispersal. Determining the morphological traits associated with alternative dispersal behaviours may be a useful tool to assess the differential dispersal capacities of species or populations.     

Polymorphic microsatellite loci to study population dynamics in a dragonfly,the libellulid Trithemis arteriosa (Burmeister, 1839)

One of the most widely distributed dragonfly species in Africa is the red‐veined‐dropwing Trithemis arteriosa. It is an indicator for permanent water bodies, which are freshwater ecosystems of high environmental value especially in arid regions. For studies to determine population structures, assess species viability and monitor environmental changes, a panel of 10 polymorphic microsatellite loci was developed. The number of alleles per locus ranged from four to 12, with an observed heterozygosity ranging from 0.149 to 0.843.     

The development of dragonfly larvae (Odonata: Anisoptera) from two streams in north-eastern Victoria, Australia

Developmental stages of larvae of eight species of Anisoptera from the Kiewa River and Middle Creek, Victoria, were determined and their phenology investigated. The last six to nine instars of each species were distinguished by size frequency and scatter plots, using labium width, metafemur and wing-pad measurements, and the early instars were estimated from Dyar's Law. This suggested between 11 to 14 instars. Four species appeared to be univoltine and four species semivoltine.     

The macrophyte, Stratiotes aloides, protects larvae of dragonfly Aeshna viridis against fish predation

Predation could be one force determining which contemporary species occupy a certain habitat. Aeshna viridis is an endangered dragonfly species with a larval distribution strongly associated with lakes where the water plant, water soldier, Stratiotes aloides, occurs. In this study, the larvae were almost exclusively found in patches of S. aloides. To study larval association with S. aloides further, we conducted a series of laboratory experiments. Behavioural experiments indicated that larvae preferred S. aloides. Aeshna viridis larvae were nocturnal and rather inactive. Larvae on S. aloides were less susceptible to predation by the perch, Perca fluviatilis, than larvae on another water plant, Myriophyllum alterniflorum. According to our study it seems that occurrence of S. aloides may limit the distribution of A. viridis, and the protection of refuges may be essential in planning the conservation of this endangered species.     

Density-dependent population dynamics in larvae of the dragonfly Pachydiplax longipennis: a field experiment

Summary Several features of dragonfly population biology suggest that population regulation occurs in the larval stage. This study was designed to determine if density-dependent interactions among larval odonates can affect survival, growth and emergence. First-instar larvae of the dragonfly Pachydiplax longipennis were raised in outdoor experimental ponds at initial densities of 38, 152, and 608 larvae · m^-2, under two levels of food availability. Food availability was supplemented in half the pools by volumetric addition of zooplankton every other day. Pools in the low food treatment did not receive the zooplankton supplement.There was a strong negative effect of density on the mean growth rate of survivors, which included both emerging tenerals and individuals overwintering in the larval stage. A higher proportion emerged from low density than high density pools. Metamorphs from high density populations were smaller and emerged slightly later than those from low density, but the absolute number of metamorphs did not differ significantly among density treatments. Food supplementation significantly increased the proportion of overwintering larvae. There were no significant food-by-density interactions, indicating that food and density acted independently on larval population dynamics. Density-dependent mechanisms can clearly contribute to odonate population regulation, especially by controlling the number of larvae which emerge and the average age at reproduction. Population-level responses to density may be a result of interference among larvae.     

Predicting habitat distribution and frequency from plant species co-occurrence data

Aim Species frequency data have been widely used in nature conservation to aid management decisions. To determine species frequencies, information on habitat occurrence is important: a species with a low frequency is not necessarily rare if it occupies all suitable habitats. Often, information on habitat distribution is available for small geographic areas only. We aim to predict grid‐based habitat occurrence from grid‐based plant species distribution data in a meso‐scale analysis. Location The study was carried out over two spatial extents: Germany and Bavaria. Methods Two simple models were set up to examine the number of characteristic plant species needed per grid cell to predict the occurrence of four selected habitats (species data from FlorKart, http://www.floraweb.de ). Both models were calibrated in Bavaria using available information on habitat distribution, validated for other federal states, and applied to Germany. First, a spatially explicit regression model (generalized linear model (GLM) with assumed binomial error distribution of response variable) was obtained. Second, a spatially independent optimization model was derived that estimated species numbers without using spatial information on habitat distribution. Finally, an additional uncalibrated model was derived that calculated the frequencies of 24 habitats. It was validated using NATURA2000 habitat maps. Results Using the Bavarian models it was possible to predict habitat distribution and frequency from the co‐occurrence of habitat‐specific species per grid cell. As the model validations for other German federal states were successful, the models were applied to all of Germany, and habitat distribution and frequencies could be retrieved for the national scale on the basis of habitat‐specific species co‐occurrences per grid cell. Using the third, uncalibrated model, which includes species distribution data only, it was possible to predict the frequencies of 24 habitats based on the co‐occurrence of 24% of formation‐specific species per grid cell. Predicted habitat frequencies deduced from this third model were strongly related to frequencies of NATURA2000 habitat maps. Main conclusions It was concluded that it is possible to deduce habitat distributions and frequencies from the co‐occurrence of habitat‐specific species. For areas partly covered by habitat mappings, calibrated models can be developed and extrapolated to larger areas. If information on habitat distribution is completely lacking, uncalibrated models can still be applied, providing coarse information on habitat frequencies. Predicted habitat distributions and frequencies can be used as a tool in nature conservation, for example as correction factors for species frequencies, as long as the species of interest is not included in the model set‐up.     

Micro-morphological adaptations of the wing nodus to flight behaviour in four dragonfly species from the family Libellulidae (Odonata: Anisoptera)

Adult dragonflies can be divided into two major groups, perchers and fliers, exhibiting notably different flight behaviour. Previous studies have yielded conflicting results regarding the link between the wing macro-morphology and flight style in these two groups. In this study, we present the first systematic investigation of the micro-morphological differences of wings of percher and flier dragonflies in four closely related species from the family Libellulidae. Our results suggest that the shape and material composition of wing microstructural components and, in particular, the nodus are adapted to facilitate the specific wing functioning in fliers and perchers. The findings further indicate a decreasing trend in the area proportion of the soft resilin-dominated cuticle in the nodus in the series of species from typical perchers to typical fliers. Such a reduction in the resilin proportion in the nodus of fliers is associated with an increase in the wing aspect ratio. The knot-shaped protrusion at the nodus of perchers, which becomes notably smaller in that of strong fliers, is likely to act as a mechanical stopper, avoiding large wing displacements. This study aims to develop a novel framework for future research on the relationship between wing morphology and flight behaviour in dragonflies.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

Correlation between population dynamics of mosquito larvae and their habitat qualities

Surveillance of anopheline and culicine larvae was conducted fortnightly to determine population composition and density at seven sites in the area of the Mathura Refinery, Mathura, India during 2005–2006. The correlation between population of the vectors and their habitat quality was established. Temperature (0.978 and 0.85°C), pH (0.99 and 0.95), conductivity (0.98 and 0.98 mho) and calcium ion hardness (0.978 and 0.85 p.p.m.) were positively correlated with the populations of both larval species. In contrast, the correlation coefficients between the parameters total hardness, magnesium ion hardness, biochemical oxygen demand and chemical oxygen demand varied with the larval species. The anopheline larval population was dominant over the culicine population in the surveillance area. The distribution of anopheline mosquito larvae during the study period was found to be constant, and the average density of anopheline and culicine larvae was 84.70 and 15.30%, respectively, across all seven sites. Thus, the study demonstrated spatial and temporal population differences of mosquito larvae with respect to environmental factors, including water quality.     

The distribution patterns of dragonflies (Insecta: Odonata) along the Kiewa River,Australia, and their relevance in conservation assessment

Sampling of larval and adult Odonata from 16 sites along the Kiewa River, Victoria, yielded 34 species: 10 Zygoptera, 24 Anisoptera. Patterns of larval and adult incidence were appraised, and showed that most species were restricted in incidence to several consecutive sites along the river, and that there is clear distinction also between the faunas of the potamon, rhithron and eucrenon regions. Different species of some genera of Anisoptera displayed different zonational distributions, and patterns of incidence and relative abundance of larvae and adults confirmed zonational occupancy. For larvae, these distribution patterns transcended the mode of collection, although many species were found most abundantly in one microhabitat or by one of several sampling methods employed at each site. Sampling of the two stages separately showed considerable concurrence of distributional patterns, so that either stage alone may provide data of value in faunal and conservation assessment. This revised version was published online in July 2006 with corrections to the Cover Date.     

Factors determining mammal species richness on habitat islands and isolates: habitat diversity, disturbance, species interactions and guild assembly rules

• 1 For over three decades the equilibrium theory of island biogeography has galvanized studies in ecological biogeography. Studies of oceanic islands and of natural habitat islands share some similarities to continental studies, particularly in developed regions where habitat fragmentation results from many land uses. Increasingly, remnant habitat is in the form of isolates created by the clearing and destruction of natural areas. Future evolution of a theory to predict patterns of species abundance may well come from the application of island biogeography to habitat fragments or isolates.
• 2 In this paper we consider four factors other than area and isolation that influence the number and type of mammal species coexisting in one place: habitat diversity, habitat disturbance, species interactions and guild assembly rules. In all examples our data derive from mainland habitat, fragmented to differing degrees, with different levels of isolation.
• 3 Habitat diversity is seen to be a good predictor of species richness. Increased levels of disturbance produce a relatively greater decrease in species richness on smaller than on larger isolates. Species interactions in the recolonization of highly disturbed sites, such as regenerating mined sites, is analogous to island colonization. Species replacement sequences in secondary successions indicate not just how many, but which species are included. Lastly, the complement of species established on islands, or in insular habitats, may be governed by guild assembly rules. These contributions may assist in taking a renewed theory into the new millennium.

     

The geometry of habitat fragmentation: Effects of species distribution patterns on extinction risk due to habitat conversion

Land‐use changes, which cause loss, degradation, and fragmentation of natural habitats, are important anthropogenic drivers of biodiversity change. However, there is an ongoing debate about how fragmentation per se affects biodiversity in a given amount of habitat. Here, we illustrate why it is important to distinguish two different aspects of fragmentation to resolve this debate: (a) geometric fragmentation effects, which exclusively arise from the spatial distributions of species and habitat fragments, and (b) demographic fragmentation effects due to reduced fragment sizes, and/or changes in fragment isolation, edge effects, or species interactions. While most empirical studies are primarily interested in quantifying demographic fragmentation effects, geometric effects are typically invoked as post hoc explanations of biodiversity responses to fragmentation per se. Here, we present an approach to quantify geometric fragmentation effects on species survival and extinction probabilities. We illustrate this approach using spatial simulations where we systematically varied the initial abundances and distribution patterns (i.e., random, aggregated, or regular) of species as well as habitat amount and fragmentation per se. As expected, we found no geometric fragmentation effects when species were randomly distributed. However, when species were aggregated, we found positive effects of fragmentation per se on survival probability for a large range of scenarios. For regular species distributions, we found weakly negative geometric effects. These findings are independent of the ecological mechanisms which generate nonrandom species distributions. Our study helps to reconcile seemingly contradictory results of previous fragmentation studies. Since intraspecific aggregation is a ubiquitous pattern in nature, our findings imply widespread positive geometric fragmentation effects. This expectation is supported by many studies that find positive effects of fragmentation per se on species occurrences and diversity after controlling for habitat amount. We outline how to disentangle geometric and demographic fragmentation effects, which is critical for predicting the response of biodiversity to landscape change.     

Activity budgets, habitat use and body size in two coexisting species of sand wasps (Microbembex: Sphecidae, Hymenoptera)

ABSTRACT. 1. Two superficially similar species of Microbembex, M.monodonta (Say) and M.cubana Bohart (Hymenoptera, Sphecidae, Nyssoninae), occur syntopically on Staniel Cay, a small Bahamian island (15 km²).
2. Nesting (female) and territorial (male) aggregations of M.cubana were smaller and more numerous than those of M.monodonta in the study site and on the island as a whole.
3. Both species had regular patterns of daily activities that required them to use a variety of microhabitats in the study area.
4. Males of both species defended territories from c. 09.00 to 12.00 hours EST (in May) and the left to feed on nectar until they retired to sleeping burrows, typically from 13.00 to 15.00 hours. Male M.monodonta were active longer each day and intermixed feeding and territorial defence more than did male M.cubana.
5. Females of both species were active for longer each day, from c. 08.30 to 15.30 hours, and intermixed their respective activities – hunting for provisions, feeding, and nest digging and maintenance – during each day more than did males. Female M.monodonta were active longer each day, rising earlier and retiring later, than female M.cubana.
6. In M.monodonta , females were larger (length, X =11.00 mm) than males (10.5 mm). In M.cubana , males were larger (11.1 mm) than females (10.4 mm). Although mean differences were small, they were highly significant ( F =22.42; P <10⁻⁴) for the species-sex interaction.
7. Differences in activity budgets, habitat use, and sexual size dimorphism appear to be related. A combination of factors, particularly thermal constraints, possible competition, and different mating systems, could explain these patterns.     

Genetic variation in three species in the Lampropholis delicata (Lacertilia: Scincidae) complex

The Lampropholis delicata complex consists of three closely related species, which have very different habitat requirements and geographic distributions. Percentage polymorphism and average heterozygosity per population are lowest in the species with the narrowest habitat requirements. Genetic differentiation between populations is also lowest in this species, which had a very limited and almost continuous distribution. For one species, which occurs in a very wide range of habitats, both natural and disturbed, genetic variability is compared between samples from disturbed and undisturbed habitats. Average heterozygosity is significantly lower in samples from disturbed than in those from undisturbed sites.     

A systematic relationship between phytochrome-controlled development and species habitat,for plants grown in simulated natural radiation

A survey of the responsiveness of plant species, typical of open and shade habitats, to simulated natural shade-light quality (i.e. white light plus supplementary far-red) has demonstrated a systematic relationship between habitat and certain developmental responses. Supplementary far-red light has a much greater effect on stem extension rate, petiole length, and leaf dry weight: stem dry weight ratio of the open habitat, shade-intolerant species. Far-red effects on leaf chlorophyll content show no such systematic grading. These results are discussed in relation to habitat adaptation. In most cases, the relationship between developmental response and the estimated phytochrome photoequilibrium, which is established by the radiation treatment, is linear. This is taken as an indication of phytochrome involvement in shade perception.Abbreviations P _fr far-red light absorbing form of phytochrome - P _total total phytochrome - PAR photosynthetically active radiation Paper 8 in the series The Function of Phytochrome in the Natural Environment; for paper 7 see Morgan and Smith (1978)