Effects of resource level and habitat type on behavioral and morphological plasticity in Eurasian perch

Spatial and temporal heterogeneity in the environment is a common feature affecting many natural populations. For example, both the resource levels and optimal habitat choices of individuals likely change over time. One way for organisms to cope with environmental variation is to display adaptive plasticity in traits such as behavior and morphology. Since trait plasticity is hypothesized to be a prerequisite for character divergence, studies of mechanisms behind such plasticity are warranted. In this study, we looked at the interaction of two potentially important environmental variables on behavioral and morphological plasticity in Eurasian perch (Perca fluviatilis L.). More specifically, the plastic responses in activity and morphology of perch exposed to different resource levels and simulated habitat types were studied in an aquarium experiment. The resource level experienced had a large influence on plasticity in both activity and morphology. Behavioral adaptations have been thought to mediate morphological transitions, and we suggest that the morphological response to the resource level was mediated by differences in activity and growth rates. The habitat type also affected morphological plasticity but to a lesser extent, and there was no effect on activity from habitat type. Based on these results, we suggest that it is essential to include several environmental factors acting in concert when studying mechanisms behind trait plasticity. We also propose that variation in resource levels might play a key role in fostering trait plasticity in at least fish populations, while other environmental variables such as divergent habitat complexities and prey types might be less influential. Dynamics in resource levels and optimal habitat choices might thus be important factors influencing character divergence in natural populations.     

Inter‐ and intraspecific effects of body size on habitat use among sexually‐dimorphic macropodids

Body size affects key life‐history parameters including dietary requirements and predation risk. We examined these effects on diel habitat use in a community of three sexually‐dimorphic macropodid marsupial species: western grey kangaroo Macropus fuliginosus, red‐necked wallaby M. rufogriseus and swamp wallaby Wallabia bicolor. In particular, our study seeks evidence of these effects operating concurrently at the intra‐ and interspecific levels. We used radio‐tracking to quantify habitat use and characterised each used location by recording the cover of plant functional groups and the presence of plant species. During nocturnal foraging periods we predicted that smaller animals (between and within species) should use habitats with higher‐quality forage, which is often less abundant, than larger animals, as metabolic demand scales with body size. During diurnal resting periods we predicted that smaller animals (between and within species), being more vulnerable to predation, should use greater concealment cover than larger animals. Western grey kangaroos and swamp wallabies behaved as predicted during foraging periods, but red‐necked wallabies did not, using more open, poorer‐quality habitats than expected. Only western grey kangaroos showed a within‐species effect on habitat use: the relatively smaller females foraged in higher‐quality patches. Habitats used by animals during the resting period generally offered greater concealment cover than those used during the foraging period, but there were no clear body size effects on the density of vegetation used. In our system, body size alone could not explain all of the observed patterns, suggesting that there may also be individual differences in habitat requirements influenced by factors such as reproductive costs, predation risk and social facilitation.     

Macroevolutionary pattern of sexual size dimorphism in geckos corresponds to intraspecific temperature‐induced variation

Many animal lineages exhibit allometry in sexual size dimorphism (SSD), known as ‘Rensch’s rule’. When applied to the interspecific level, this rule states that males are more evolutionary plastic in body size than females and that male‐biased SSD increases with body size. One of the explanations for the occurrence of Rensch’s rule is the differential‐plasticity hypothesis assuming that higher evolutionary plasticity in males is a consequence of larger sensitivity of male growth to environmental cues. We have confirmed the pattern consistent with Rensch’s rule among species of the gecko genus Paroedura and followed the ontogeny of SSD at three constant temperatures in a male‐larger species (Paroedura picta). In this species, males exhibited larger temperature‐induced phenotypic plasticity in final body size than females, and body size and SSD correlated across temperatures. This result supports the differential‐plasticity hypothesis and points to the role phenotypic plasticity plays in generating of evolutionary novelties.     

Development of intraspecific size variation in black coucals,white-browed coucals and ruffs from hatching to fledging

Most studies on sexual size dimorphism address proximate and functional questions related to adults, but sexual size dimorphism usually develops during ontogeny and developmental trajectories of sexual size dimorphism are poorly understood. We studied three bird species with variation in adult sexual size dimorphism: black coucals (females 69% heavier than males), white-browed coucals (females 13% heavier than males) and ruffs (males 70% heavier than females). Using a flexible Bayesian generalized additive model framework (GAMM), we examined when and how sexual size dimorphism developed in body mass, tarsus length and bill length from hatching until fledging. In ruffs, we additionally examined the development of intrasexual size variation among three morphs (Independents, Satellites and Faeders), which creates another level of variation in adult size of males and females. We found that 27–100% of the adult inter- and intrasexual size variation developed until fledging although none of the species completed growth during the observational period. In general, the larger sex/morph grew more quickly and reached its maximal absolute growth rate later than the smaller sex/morph. However, when the daily increase in body mass was modelled as a proportion, growth patterns were synchronized between and within sexes. Growth broadly followed sigmoidal asymptotic models, however only with the flexible GAMM approach, residual distributions were homogeneous over the entire observation periods. These results provide a platform for future studies to relate variation in growth to selective pressures and proximate mechanisms in these three species, and they highlight the advantage of using a flexible model approach for examining growth variation during ontogeny.     

Environmental and organismal predictors of intraspecific variation in the stoichiometry of a neotropical freshwater fish

The elemental composition of animals, or their organismal stoichiometry, is thought to constrain their contribution to nutrient recycling, their interactions with other animals, and their demographic rates. Factors that affect organismal stoichiometry are generally poorly understood, but likely reflect elemental investments in morphological features and life history traits, acting in concert with the environmental availability of elements. We assessed the relative contribution of organismal traits and environmental variability to the stoichiometry of an insectivorous Neotropical stream fish, Rivulus hartii. We characterized the influence of body size, life history phenotype, stage of maturity, and environmental variability on organismal stoichiometry in 6 streams that differ in a broad suite of environmental variables. The elemental composition of R. hartii was variable, and overlapped with the wide range of elemental composition documented across freshwater fish taxa. Average %P composition was ～3.2%(±0.6), average %N～10.7%(±0.9), and average %C～41.7%(±3.1). Streams were the strongest predictor of organismal stoichiometry, and explained up to 18% of the overall variance. This effect appeared to be largely explained by variability in quality of basal resources such as epilithon N:P and benthic organic matter C:N, along with variability in invertebrate standing stocks, an important food source for R. hartii. Organismal traits were weak predictors of organismal stoichiometry in this species, explaining when combined up to 7% of the overall variance in stoichiometry. Body size was significantly and positively correlated with %P, and negatively with N:P, and C:P, and life history phenotype was significantly correlated with %C, %P, C:P and C:N. Our study suggests that spatial variability in elemental availability is more strongly correlated with organismal stoichiometry than organismal traits, and suggests that the stoichiometry of carnivores may not be completely buffered from environmental variability. We discuss the relevance of these findings to ecological stoichiometry theory.     

The production of hemoglobin by small pond Daphnia pulex: intraspecific variation and its relation to habitat

SUMMARY.

• 1 Hemoglobin production was measured for two populations of Daphnia pulex, one from a temporary pond (GR Pond) and one from a permanent pond (Rash Pond). Surface water in Rash Pond remains high in oxygen, while by late summer very little oxygen remains in GR Pond. Hemoglobin synthesis was induced in the laboratory by artificial oxygen deficiency.
• 2 Reproductive state influenced the level of hemoglobin in Daphnia. Females with parthenogenesis eggs had higher total body hemoglobin than ephippial females. Clutches laid by pale Daphnia immediately before exposure to low oxygen conditions were aborted. Offspring produced by hemoglobin-rich females later in the treatment survived.
• 3 Hemoglobin production was higher for Rash Pond Daphnia. Visual predators exist in GR Pond, but are absent in Rash Pond. Also, Daphnia are smaller in GR Pond. Visual predation may have selected against high levels of hemoglobin in the temporary pond. Thus oxygen availability in nature is not necessarily a good predictor of hemoglobin production by Daphnia.
• 4 Feeding behavior was altered in low oxygen conditions. Browsing was dominant in low oxygen treatments while Daphnia in control treatments filtered algae normally. Browsing may increase access to sedimented iron necessary for hemoglobin synthesis.

     

Effects of body size, trophic mode and larval habitat on Diptera stoichiometry: a regional comparison

Ecological stoichiometry has emerged as a tool for exploring nutrient demand and evolutionary responses to nutrient limitation. Previous studies of insects have found predictable variability in stoichiometry, both in relation to body size and trophic mode, at ordinal levels or higher. Our study further examines the evolutionary and ecological lability in these traits by comparing the effects of body size, trophic mode (larval and adult) and larval habitat on the stoichiometry of insects within one order (Diptera). The study also expands on previous work by analyzing trophic mode both at coarse (detritivore, herbivore, predator) and fine (high- vs low- nutrient quality resources within trophic categories) scales and by comparing nutrient stoichiometry in two geographical regions, Sweden and Arizona. As predicted, adults feeding on nectar or pollen had the lowest body N content in the dataset. Additionally, for Diptera with predatory larvae, species low N diets had lower body N content than those with high N diets. However, body N content was not consistently lower for all species with low N resources, as species feeding on plant material were indistinguishable in stoichiometry from predators with high N diets. We suggest that these results emerge because larval resource exploitation is poorly understood in herbivorous Diptera species. Body P content for Swedish Diptera decreased with body size for all trophic modes, and the only difference among trophic modes was that blood feeders had higher P content than other groups. The regional comparison further showed that the allometry of body P content is a labile trait that may vary at regional scales, as there was no allometric scaling of body P content in the Arizona data set, in contrast to the Swedish data set. These results are not easily explained by existing theoretical frameworks, but instead point to a general context-dependence of P stoichiometry, which should now be a focus for future work.     

Effects of size structure and habitat complexity on predator–prey interactions

1. A predator's ability to suppress its prey depends on the level of interference among predators. While interference typically decreases with increasing habitat complexity, it often increases with increasing size differences among individuals. However, little is known about how variation in intrinsic factors such as population size structure alters predator–prey interactions and how this intrinsic variation interacts with extrinsic variation. 2. By experimentally varying the level of vegetation cover and the size structure of the predatory damselfly Ischnura posita Hagen, we examined the individual and interactive effects of variation in habitat complexity and predator size structure on prey mortality. 3. Copepod prey survival linearly increased as the I. posita size ratio decreased and differed by up to 31% among different predator size structures. Size classes had an additive effect on prey survival, most likely because intraspecific aggression appeared size‐independent and size classes differed in microhabitat preference: large I. posita spent 14% more time foraging on the floor than small larvae and spent more time in the vegetation with increasing habitat complexity. Despite this difference in microhabitat use among size classes, habitat structure did not influence predation rates or interference among size classes. 4. In general, results suggest that seasonal and spatial variation in the size structure of populations could drive some of the discrepancies in predator‐mediated prey suppression observed in nature, and this variation could exceed the effects of variation in habitat structure.     

Increased propensity for aerial dispersal in disturbed habitats due to intraspecific variation and species turnover

Animal dispersal depends on multiple factors, such as habitat features and life‐history traits of the species. We studied the propensity for ballooning dispersal in spiders under standardized laboratory conditions. The 1269 tested individuals belonged to 124 species and originated from 16 sites with wide variation in habitat type. Spiders from disturbed habitats ballooned 5.5 times more than spiders from stable habitats. In Meioneta rurestris , for which we had enough data for a single‐species analysis, individuals were most dispersive if they originated from highly disturbed habitats. While the data for the other species were not sufficient for single‐species analyses, a hierarchical model that included the data simultaneously on all species suggested that dispersal propensity generally increases within species with the level of habitat disturbance. Dispersal probability showed a trend to increase with niche width, but the higher commonness of species with wide niches provides an alternative explanation for this pattern. As the prevalence of especially dispersive species was highest in disturbed habitats, variation in dispersal propensity was influenced by both inter‐ and intraspecific factors. We conclude that the positive correlation between niche width and dispersal propensity enables generalist species to utilize highly disturbed habitats, whereas the persistence of specialist species with restricted dispersal ability requires the conservation of stable habitats.     

Predicting the non‐linear collapse of plant–frugivore networks due to habitat loss

Habitat loss can trigger cascades of secondary extinctions, changing the organization of interacting assemblages. Until recently, most extinction models in interaction systems had limited ecological realism. Here, we estimate a realistic sequence of species extinctions resulting from habitat loss to assess its impacts on the structure of frugivory networks from the Brazilian Atlantic Forest. We show that realistic and random extinctions led to similar patterns. We also identified a threshold in the response of network structure to habitat loss. When forest cover was reduced to less than 40% of the landscape, network organization changed dramatically. Hence, the number of species being lost, rather than the order of species extinctions, is the key determinant of its impacts on the organization of frugivory networks. We highlight the need to conserve around 40% of forest cover to keep the basic organization of frugivory networks, a threshold already reached at the best‐preserved Brazilian Atlantic Forest bioregion.     

Life cycle, growth and diet of Balston's pygmy perch in its natural habitat of acidic pools in south-western Australia

Nannatherina balstoni is found in a few acidic pools (pH 3.9–6.0) in the extreme south-western corner of Australia. Although many of these pools become dry during summer and early autumn, they are recolonized by fish from nearby pools that overflow during winter floods. N. balstoni spawns at the end of its first year of life, when, on average, the males and females have reached 60 and 63 mm t.l ., respectively, and then usually die within the next few months. The largest fish, which was one of only three in its third year of life, measured 90 mm and weighed 7.3 g. The von Bertalanffy growth curve parameters for L₀, K and t₀ were 71.2 mm, 1.69 and – 0.078 for males and 82.6 mm, 1.31 and – 0.095 for females. Fecundity ranged from 550 to 1600. N. balstoni. spawns during the middle of winter, after heavy flooding and when water temperatures are at, or close to, their annual minima. This enables the larvae and young juveniles to capitalize on those aquatic organisms, especially Cladocera, which are very abundant amongst the flooded riparian vegetation that surrounds the pools in winter and spring. Hence, the fish grow rapidly and attain an appreciable size before summer, when the increases that occur in the densities of the larger carnivorous fish species, as a result of marked declines in water levels, increases the chances of predation. By spring, when most N. balstoni exceed 25 mm t .l ., the diet changes markedly to one that consists almost exclusively of terrestrial fauna. In contrast, three of the six co-occurring native species of teleost feed on aquatic and terrestrial fauna throughout the year, while a further two feed only within the water column and the sixth feeds on benthic invertebrates. The pronounced shift in diet exhibited by N. balstoni as it increases in size, allied to dietary differences amongst the other six co-occurring species, reduces any potential for interspecific competition for food resources during the summer and autumn, when such resources are declining.     

The nature of intraspecific and interspecific genome size variation in taxonomically complex eyebrights

Background and aimsGenome size varies considerably across the diversity of plant life. Although genome size is, by definition, affected by genetic presence/absence variants, which are ubiquitous in population sequencing studies, genome size is often treated as an intrinsic property of a species. Here, we studied intra- and interspecific genome size variation in taxonomically complex British eyebrights (Euphrasia, Orobanchaceae). Our aim is to document genome size diversity and investigate underlying evolutionary processes shaping variation between individuals, populations and species.MethodsWe generated genome size data for 192 individuals of diploid and tetraploid Euphrasia and analysed genome size variation in relation to ploidy, taxonomy, population affiliation and geography. We further compared the genomic repeat content of 30 samples.Key resultsWe found considerable intraspecific genome size variation, and observed isolation-by-distance for genome size in outcrossing diploids. Tetraploid Euphrasia showed contrasting patterns, with genome size increasing with latitude in outcrossing Euphrasia arctica, but with little genome size variation in the highly selfing Euphrasia micrantha. Interspecific differences in genome size and the genomic proportions of repeat sequences were small.ConclusionsWe show the utility of treating genome size as the outcome of polygenic variation. Like other types of genetic variation, such as single nucleotide polymorphisms, genome size variation may be affected by ongoing hybridization and the extent of population subdivision. In addition to selection on associated traits, genome size is predicted to be affected indirectly by selection due to pleiotropy of the underlying presence/absence variants.     

Territory occupancy by common loons in response to disturbance,habitat, and intraspecific relationships

Structure and distribution of animal territories are driven by a variety of environmental and demographic factors. A peninsular population of common loons (Gavia immer) nests on lakes in northwestern Montana, but does not occupy all apparently suitable breeding territories, suggesting unexplained limitations on population growth. To evaluate territorial dynamics of breeding loons in Montana, we created and tested occupancy models that evaluated the hypothesized effects of disturbance, habitat, and intraspecific relationships on territory occupancy by common loons in Montana from 2003 to 2007. Model-averaged results indicated that the abundance of feeding lakes within 10 km (i.e., forage quality) and the number of territorial pairs within 10 km (i.e., density of loons) were equally supported and related to probabilities of occupancy. We found substantial support that the population was in a state of equilibrium, with the numbers of occupied territories stable in time, but not space. We also found that density of territorial pairs was related to the likelihood that an existing territory would be abandoned, but did not influence the establishment of new territories, suggesting the presence of territorial pairs could be a stronger indicator of territory quality to loons than physical lake characteristics. Our index of human disturbance was not well-supported compared to other factors. Our results suggest management for stable or growing loon populations could be achieved using long-term monitoring and protection of occupied territorial lakes and nearby feeding lakes, because these factors most influenced the probability of occupancy of surrounding lakes. © 2011 The Wildlife Society.     

Effects of sample size and intraspecific variation in phylogenetic comparative studies: a meta‐analytic review

Comparative analyses aim to explain interspecific variation in phenotype among taxa. In this context, phylogenetic approaches are generally applied to control for similarity due to common descent, because such phylogenetic relationships can produce spurious similarity in phenotypes (known as phylogenetic inertia or bias). On the other hand, these analyses largely ignore potential biases due to within‐species variation. Phylogenetic comparative studies inherently assume that species‐specific means from intraspecific samples of modest sample size are biologically meaningful. However, within‐species variation is often significant, because measurement errors, within‐ and between‐individual variation, seasonal fluctuations, and differences among populations can all reduce the repeatability of a trait. Although simulations revealed that low repeatability can increase the type I error in a phylogenetic study, researchers only exercise great care in accounting for similarity in phenotype due to common phylogenetic descent, while problems posed by intraspecific variation are usually neglected. A meta‐analysis of 194 comparative analyses all adjusting for similarity due to common phylogenetic descent revealed that only a few studies reported intraspecific repeatabilities, and hardly any considered or partially dealt with errors arising from intraspecific variation. This is intriguing, because the meta‐analytic data suggest that the effect of heterogeneous sampling can be as important as phylogenetic bias, and thus they should be equally controlled in comparative studies. We provide recommendations about how to handle such effects of heterogeneous sampling.     

Morphological variation in pumpkinseed Lepomis gibbosus introduced into Iberian lakes and reservoirs; adaptations to habitat type and diet?

The morphology of pumpkinseed Lepomis gibbosus populations from five Catalonian waterbodies (north-eastern Spain) that vary in hydromorphometry was examined and compared to a native North American reference site that contained two morphological variants of this species. Populations exhibited significant differences in fin location, body depth and caudal peduncle length, which are known to have functional significance in the hydrodynamics of swimming and hence the foraging mode. Differences were also noted in internal morphological traits functionally related to prey selection. Pumpkinseed populations that fed extensively on zooplankton showed narrow gill raker spacing, and mollusc-feeding populations had longer and wider pharyngeal bones. Discriminant function analysis (DFA) provided significant separation of all populations on the basis of both external and internal morphology, with the main axis of separation being geographical rather than environmental. The secondary DFA axis, however, did separate populations that fed primarily on zooplankton from those that were primarily benthic invertebrate feeders. In this regard, a population that occupied an Iberian steep-sided reservoir with an unstable littoral zone showed similar morphological adaptations to the limnetic morphological variant native in North America, supporting previous studies showing that fish morphology is strongly affected by prey type and feeding mode. The results suggest that pumpkinseeds are able to adapt, morphologically, to the types of habitats and prey present in Catalonian waterbodies, and this may partially explain why they are so successful in areas where they have been introduced.     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.     

Population size,habitat fragmentation,and the nature of adaptive variation in a stream fish

Whether and how habitat fragmentation and population size jointly affect adaptive genetic variation and adaptive population differentiation are largely unexplored. Owing to pronounced genetic drift, small, fragmented populations are thought to exhibit reduced adaptive genetic variation relative to large populations. Yet fragmentation is known to increase variability within and among habitats as population size decreases. Such variability might instead favour the maintenance of adaptive polymorphisms and/or generate more variability in adaptive differentiation at smaller population size. We investigated these alternative hypotheses by analysing coding-gene, single-nucleotide polymorphisms associated with different biological functions in fragmented brook trout populations of variable sizes. Putative adaptive differentiation was greater between small and large populations or among small populations than among large populations. These trends were stronger for genetic population size measures than demographic ones and were present despite pronounced drift in small populations. Our results suggest that fragmentation affects natural selection and that the changes elicited in the adaptive genetic composition and differentiation of fragmented populations vary with population size. By generating more variable evolutionary responses, the alteration of selective pressures during habitat fragmentation may affect future population persistence independently of, and perhaps long before, the effects of demographic and genetic stochasticity are manifest.