Climate change and sexual size dimorphism in an Arctic spider

Climate change is advancing the onset of the growing season and this is happening at a particularly fast rate in the High Arctic. However, in most species the relative fitness implications for males and females remain elusive. Here, we present data on 10 successive cohorts of the wolf spider Pardosa glacialis from Zackenberg in High-Arctic, northeast Greenland. We found marked inter-annual variation in adult body size (carapace width) and this variation was greater in females than in males. Earlier snowmelt during both years of its biennial maturation resulted in larger adult body sizes and a skew towards positive sexual size dimorphism (females bigger than males). These results illustrate the pervasive influence of climate on key life-history traits and indicate that male and female responses to climate should be investigated separately whenever possible.     

Sex differences in phenotypic plasticity of a mechanism that controls body size: implications for sexual size dimorphism

The degree and/or direction of sexual size dimorphism (SSD) varies considerably among species and among populations within species. Although this variation is in part genetically based, much of it is probably due to the sexes exhibiting differences in body size plasticity. Here, we use the hawkmoth, Manduca sexta, to test the hypothesis that moths reared on different diet qualities and at different temperatures will exhibit sex-specific body size plasticity. In addition, we explore the proximate mechanisms that potentially create sex-specific plasticity by examining three physiological variables known to regulate body size in this insect: the growth rate, the critical weight (which measures the cessation of juvenile hormone secretion from the corpora allata) and the interval to cessation of growth (ICG; which measures the time interval between the critical weight and the secretion of the ecdysteroids that regulate pupation and metamorphosis). We found that peak larval mass of males and females did not exhibit sex-specific plasticity in response to diet or temperature. However, the sexes did exhibit sex-specific plasticity in the mechanism that controls size; males and females exhibited sex-specific plasticity in the growth rate and the critical weight in response to both diet and temperature, whereas the ICG only exhibited sex-specific plasticity in response to diet. Our results suggest it is important for the sexes to maintain the same degree of SSD across environments and that this is accomplished by the sexes exhibiting differential sensitivity of the physiological factors that determine body size to environmental variation.     

Intersexual trophic niche partitioning in an ant-eating spider (Araneae: Zodariidae)

Background

Divergence in trophic niche between the sexes may function to reduce competition between the sexes (“intersexual niche partitioning hypothesis”), or may be result from differential selection among the sexes on maximizing reproductive output (“sexual selection hypothesis”). The latter may lead to higher energy demands in females driven by fecundity selection, while males invest in mate searching. We tested predictions of the two hypotheses underlying intersexual trophic niche partitioning in a natural population of spiders. Zodarion jozefienae spiders specialize on Messor barbarus ants that are polymorphic in body size and hence comprise potential trophic niches for the spider, making this system well-suited to study intersexual trophic niche partitioning.

Methodology/Principal Findings

Comparative analysis of trophic morphology (the chelicerae) and body size of males, females and juveniles demonstrated highly female biased SSD (Sexual Size Dimorphism) in body size, body weight, and in the size of chelicerae, the latter arising from sex-specific growth patterns in trophic morphology. In the field, female spiders actively selected ant sub-castes that were larger than the average prey size, and larger than ants captured by juveniles and males. Female fecundity was highly positively correlated with female body mass, which reflects foraging success during the adult stage. Females in laboratory experiments preferred the large ant sub-castes and displayed higher capture efficiency. In contrast, males occupied a different trophic niche and showed reduced foraging effort and reduced prey capture and feeding efficiency compared with females and juveniles.

Conclusions/Significance

Our data indicate that female-biased dimorphism in trophic morphology and body size correlate with sex-specific reproductive strategies. We propose that intersexual trophic niche partitioning is shaped primarily by fecundity selection in females, and results from sex-differences in the route to successful reproduction where females are selected to maximize energy intake and fecundity, while males switch from foraging to invest in mating effort.     

Sex-specific trait architecture in a spider with sexual size dimorphism

Sexual dimorphism, or sex-specific trait expression, may evolve when selection favours different optima for the same trait between sexes, that is, under antagonistic selection. Intra-locus sexual conflict exists when the sexually dimorphic trait under antagonistic selection is based on genes shared between sexes. A common assumption is that the presence of sexual-size dimorphism (SSD) indicates that sexual conflict has been, at least partly, resolved via decoupling of the trait architecture between sexes. However, whether and how decoupling of the trait architecture between sexes has been realized often remains unknown. We tested for differences in architecture of adult body size between sexes in a species with extreme SSD, the African hermit spider (Nephilingis cruentata), where adult female body size greatly exceeds that of males. Specifically, we estimated the sex-specific importance of genetic and maternal effects on adult body size among individuals that we laboratory-reared for up to eight generations. Quantitative genetic model estimates indicated that size variation in females is to a larger extent explained by direct genetic effects than by maternal effects, but in males to a larger extent by maternal than by genetic effects. We conclude that this sex-specific body-size architecture enables body-size evolution to proceed much more independently than under a common architecture to both sexes.     

Effects of copulation temperature on female reproductive output and longevity in the wolf spider Pardosa astrigera (Araneae: Lycosidae)

The copulation duration of male wolf spider Pardosa astrigera, was significantly influenced by environmental temperature, as had been found in some insect species. Therefore, temperature during male courtship and copulation may influence the amount of sperm and seminal fluids transferred during copulation, which in turn could influence female fitness. In order to test this hypothesis, we subjected pairs of male and female P. astrigera to five temperature groups from 16 to 32 °C at an interval of 4 °C, and investigated whether and to what extent the various temperatures during male courtship and copulation influenced female reproductive output and female adult longevity under controlled laboratory conditions. With the increase of copulation temperature, females were more likely to lay egg sacs. The total egg sacs and lifetime fecundity of female were positively influenced by copulation temperature, whereas female lifetime spiderlings and adult longevity were independent of copulation temperature.     

Seasonal variations in body melanism and size of the wolf spider Pardosa astrigera (Araneae: Lycosidae)

Variations in species morphology and life‐history traits strongly correlate with geographic and climatic characteristics. Most studies on morphological variations in animals focus on ectotherms distributed on a large geographic scale across latitudinal and/or altitudinal gradient. However, the morphological variations of spiders living in the same habitats across different seasons have not been reported. In this study, we used the wolf spider, Pardosa astrigera, as a model to determine seasonal differences in adult body size, melanism, fecundity, and egg diameter both in the overwintering and the first generation for 2010 and 2016. The results showed that in 2010, both females and males of the overwintering generation were significantly darker than the first generation. Moreover, the overwintering females were markedly larger and produced more and bigger eggs than the first generation in both 2010 and 2016. Considering the overwintering P. astrigera experiencing low temperature and/or desiccation stress, these results suggest that substantially darker and larger body of the overwintering generation is adaptive to adverse conditions.     

High-Arctic butterflies become smaller with rising temperatures

The response of body size to increasing temperature constitutes a universal response to climate change that could strongly affect terrestrial ectotherms, but the magnitude and direction of such responses remain unknown in most species. The metabolic cost of increased temperature could reduce body size but long growing seasons could also increase body size as was recently shown in an Arctic spider species. Here, we present the longest known time series on body size variation in two High-Arctic butterfly species: Boloria chariclea and Colias hecla. We measured wing length of nearly 4500 individuals collected annually between 1996 and 2013 from Zackenberg, Greenland and found that wing length significantly decreased at a similar rate in both species in response to warmer summers. Body size is strongly related to dispersal capacity and fecundity and our results suggest that these Arctic species could face severe challenges in response to ongoing rapid climate change.     

农田狼蛛亚群落结构的研究

本文对南宁市郊农田九种生境狼蛛亚群落结构进行了讨论。群落有7属37种狼蛛。文中揭示了农田各生境区间狼蛛亚群落的主要成分及其主要属性差异的原因。以狼蛛类群为指示动物,用群落相似性比例和聚类分析法,能较好地区分幼豹蛛(Pardosa pusiola)与奇异獾蛛(Trochosa ruricola)和拟环纹豹蛛(Pardosa pseudoannulata)与拟水狼蛛(Pirata subpiraticus)是构成农田区旱地和水田生境型具有代表性的两个狼蛛亚群落主要成分。     

Sex-specific responses to fecundity selection in the broad-nosed pipefish

Fecundity selection, acting on traits enhancing reproductive output, is an important determinant of organismal body size. Due to a unique mode of reproduction, mating success and fecundity are positively correlated with body size in both sexes of male-pregnant Syngnathus pipefish. As male pipefish brood eggs on their tail and egg production in females occurs in their ovaries (located in the trunk region), fecundity selection is expected to affect both sexes in this species, and is predicted to act differently on body proportions of males and females during their development. Based on this hypothesis, we investigated sexual size dimorphism in body size allometry and vertebral numbers across populations of the widespread European pipefish Syngnathus typhle. Despite the absence of sex-specific differences in overall and region-specific vertebral counts, male and female pipefish differ significantly in the relative lengths of their trunk and tail regions, consistent with region-specific selection pressures in the two sexes. Male pipefish show significant growth allometry, with disproportionate growth in the brooding tail region relative to the trunk, resulting in increasingly skewed region-specific sexual size dimorphism with increasing body size, a pattern consistent across five study populations. Sex-specific differences in patterns of growth in S. typhle support the hypothesis that fecundity selection can contribute to the evolution of sexual size dimorphism.     

The impact of snow accumulation on a heath spider community in a sub-Arctic landscape

Patterns of snow cover across the Arctic are expected to change as a result of shrub encroachment and climate change. As snow cover impacts both the subnivean environment and the date of spring melt, these changes could impact Arctic food webs by altering the phenology and survival of overwintering arthropods, such as spiders (Araneae). In this field study, we used snow fences to increase snow cover across a series of large (375 m²) heath tundra plots and examined the effects on the local spider community during the following growing season. Fences increased snow cover and delayed melt on the treatment plots, paralleling the conditions of nearby shrub sites. Frequent sampling over the season revealed that increased snow cover did not affect spider abundance across different genera nor did it affect overall community composition. Further, our snow treatment did not affect the dates when plots achieved seasonal catch milestones (25, 50, 75 % of total seasonal catch). Increased winter snow cover did, however, produce higher body masses in adults and juveniles of the dominant species Pardosa lapponica (Lycosidae), beginning immediately after snow melt until midway through the growing season. In addition, ovary/oocyte mass of mature P. lapponica females was significantly higher on treatment plots during the peak reproductive period. This is the first experimental manipulation study to report a significant effect of landscape-level changes to winter snow cover on the biomass of an Arctic macroarthropod.     

Size dimorphism in Rankinia [Tympanocryptis] diemensis (Family Agamidae): sex-specific patterns and geographic variation

Sexual dimorphism has implications for a range of biological and ecological factors, and intersexual morphological differences within a species provide an ideal opportunity for investigating evolutionary influences on phenotypic variation. We investigated sexual size dimorphism (SSD) in an agamid species, Rankinia [Tympanocryptis] diemensis , to determine whether overall size and/or relative morphological trait size differences exist and whether geographic variation in size dimorphism occurs in this species. Relative morphological trait proportions included a range of head, limb, and inter-limb measurements. We found significant overall intersexual adult size differences; females were the larger sex across all sites but the degree of dimorphism between the sexes did not differ between sites. This female-biased size difference is atypical for agamid lizards, which are usually characterized by large male body size. In this species, large female-biased SSD appears to have evolved as a result of fecundity advantages. The size of relative morphological trait also differed significantly between the sexes, but in the opposite direction: relative head, tail, and limb sizes were significantly larger in males than females. This corresponds to patterns in trait size usually found in this taxonomic group, where male head and limb size is important in contest success such as male–male rivalry. There were site-specific morphological differences in hatchlings, including overall body size, tail, inter-limb, thigh, and hindlimb lengths; however, there were no sex-specific differences indicating the body size differences present in the adult form occur during ontogeny.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 699–709.     

Saline stress tolerance partly matches with habitat preference in ground‐living wolf spiders

Salinity interacts with many physiological functions and therefore probably influences the distribution of terrestrial fauna in tidal flooded salt marshes. The present study tests the hypothesis that the physiological tolerance of stenotopic wolf spiders for saline conditions at least partially determines their occurrence throughout salt‐marsh and nonsaline habitats. The duration of survival of three stenotopic wolf spider species (Araneae: Lycosidae) with different habitat preferences is compared in a controlled laboratory experiment. The forest‐dwelling Pardosa saltans, the salt‐marsh resident Pardosa purbeckensis and its sister species the inland‐living Pardosa agrestis are exposed to experimental conditions with different levels of salinity. Individuals (45 males and 20–45 females per treatment) are placed in individual air‐tight boxes filled with water‐saturated sand. Three levels of salinity are tested: nonsaline (0‰), medium saline (33–35‰) and highly saline (66–70‰). Contents of carbon, hydrogen and nitrogen and the molar ration carbon/nitrogen remain constant over time and do not differ among salinity treatments, indicating that starvation effects on survival, if any, are similar for all treatments. Conversely, body water significantly decreases over time and differs among salinity treatments, in accordance with patterns of survival. Conforming to their habitat preference, the survival of P. saltans and P. agrestis decreases quickly under highly saline conditions. Pardosa purbeckensis, however, has a high survival under both saline and nonsaline conditions. The duration of survival of females is significantly lower than that of males of P. saltans and P. purbeckensis. Durations of survival of ground‐living wolf spiders exposed to salinity partly match their habitat distribution but do not explain the restriction of salt‐marsh species to saline habitats.     

Sex‐Specific Response of Pardosa milvina (Araneae: Lycosidae) to Experience with a Chemotactile Predation Cue

Predators frequently leave behind chemical information (i.e., semiochemicals such as pheromones or kairomones) that can be detected by their prey and used to avoid areas where predators are likely present. Prey that have interacted indirectly with predators via chemical information thus may gain insight into their risk of being consumed that naïve individuals lack. Pardosa milvina (Araneae: Lycosidae) is a chemosensitive wolf spider that shows adaptive responses to chemotactile cues deposited by the larger wolf spider Tigrosa helluo. We raised offspring from P. milvina to examine the effect of experience with a predation cue on activity, foraging, and antipredator behavior. Spiders differed in activity and foraging behavior across ontogeny and between sexes, but there was no effect of experience with a predation cue. However, a sex‐specific effect of experience was found in antipredator behavior. Male spiders, but not females, used experience with a predator cue to increase their survival in the presence of a live predator. Specifically, naïve males were attacked sooner than experienced males, indicating that prior exposure to predator cues can modify Pardosa antipredator behavior. Intersexual differences in how spiders respond to experience with a predation cue likely reflect the risk of predation faced by males and females in nature.     

Divergent Sex-Specific Plasticity in Long-Lived Vertebrates with Contrasting Sexual Dimorphism

Sex-specific plasticity can profoundly affect sexual size dimorphism (SSD), but its influence in female-larger-SSD vertebrates remains obscure. Theory predicts that sex-specific plasticity may drive SSD evolution if the larger sex benefits from optimal-growth conditions when available (condition-dependent hypothesis), or if attaining a suboptimal size is penalized by selection (adaptive canalization hypothesis). Sex-specific plasticity enhances the size of the larger sex in male-larger-SSD turtles but whether the same occurs in female-larger species is unknown. Sexual shape dimorphism (SShD) is also widespread in nature but is understudied, and whether SShD derives from sex-specific responses to identical selective pressures or from sex-specific selection remains unclear. Here we tested whether sex-specific growth plasticity underlies the development of sexual size and shape dimorphism in the female-larger-SSD turtle, Podocnemis expansa. Individuals hatched from several incubation temperatures and were raised under common-garden conditions with varying temperature and resources. Body size and shape were plastic and sexually dimorphic, but plasticity did not differ between the sexes, opposite to the male-larger turtle Chelydra serpentina. Maternal effects (egg size) were significant on size and shape, suggesting that females increase their fitness by allocating greater energy to enhance offspring growth. Results ruled out the sex-specific plasticity hypotheses in P. expansa, indicating that SSD and SShD do not derive form differential responses to identical drivers but from sex-specific selective pressures. Our results indicate that differential plasticity does not favor males inherently, nor the larger sex, as would be expected if it was a pervasive driver of macroevolutionary patterns of sexual dimorphism across turtle lineages.     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

Interacting Effects of Leg Autotomy and Exposure to Predator Cues on Survival in A Wolf Spider (<Emphasis Type="Italic">Pardosa valens</Emphasis>)

Chemical cues from predators (kairomones) are used by many aquatic and terrestrial animals when deciding on behavioral responses to predation threats. These responses may also be affected by the animal’s physiological state (e.g., nutrition level, parasitism, or prior injury), which could alter normal responses to kairomones. In this study, we examined effects of leg autotomy (the voluntary sacrifice of a leg) on subsequent responses to predator chemical cues in females of the riparian-zone wolf spider Pardosa valens. In a fully-crossed design, spiders with all legs intact or with one leg removed were exposed to one of two cue treatments for 90 min: a control (no predator cue) or one with chemical cues (silk and excreta) from a larger sympatric wolf spider, Rabidosa santrita. We then introduced an R. santrita into each container, and recorded subsequent survival of P. valens. Survivorship was significantly higher for individuals previously exposed to predator cues than for those in the control group; however, autotomy had no effect on survivorship, which was similar for both intact and autotomized spiders in both the predator-cue and control treatments. In addition, although P. valens were more likely to be found off the substrate than on it when the predator was added in each of the four treatment pairings, this initial position did not influence survivorship. These results therefore indicate that P. valens can behaviorally respond to predator kairomones in ways that reduce their risk of predation, but that this response is unaffected by the prior sacrifice of a leg.     

Two ecologically-divergent generalist predators have different responses to landscape fragmentation

We conducted three replicated field experiments to test the population response of two ecologically-divergent wolf spider species ( Hogna helluo and Pardosa milvina ) to three correlates of landscape fragmentation: area reduction, spatial subdivision, and increased edge to core ratio. We selected these two species because they differ in vagility and habitat selectivity. Hogna helluo is relatively large, averse to disturbed substrata, and has poor colonization abilities. Conversely, Pardosa is small, vagile, and will use barren, disturbed areas. In a test for the effect of area reduction on populations of the two wolf spiders, we destroyed 0%, 20% or 80% of randomly selected habitat islands in replicated experimental landscapes. We found that population densities of Hogna declined significantly, even at the lowest level of area reduction (20%), and that there was an increase in numbers of Pardosa . In a test for the response to an increase in landscape subdivision, we created four levels of habitat fragmentation in replicate plots. We found a significant decline in Hogna populations with increasing fragmentation. Pardosa populations did not respond to the fragmentation. In the third experiment we kept landscape area and subdivision constant, but manipulated the edge-to-core ratio. We found that populations of Hogna declined sharply with increasing edge, and that populations of Pardosa did not respond. These two syntopic wolf spiders have distinctly different responses to landscape fragmentation.     

Influence of landscape structure and farming practice on body condition and fecundity of wolf spiders

Structurally complex landscapes and organic management have been shown to augment natural pest enemy populations on arable land. Here, body condition and reproductive capacity of wolf spiders were studied to see if these traits can explain the larger populations in these environments. Females of Pardosa spiders (Araneae: Lycosidae) were collected at 7 organically and 7 conventionally managed fields situated in landscapes with different proportions of perennial crops, annual crops, forest, and different numbers and sizes of fields. Body condition (relative female weight) and fecundity (number of offspring and relative egg sac weight) were measured for each captured spider. In contrast to the hypothesis, Pardosa females caught in fields situated in landscapes dominated by large fields of annual crops had superior body condition. Farming practice had no effect on either body condition or fecundity measures. It is suggested that increased spider body condition in homogeneous landscapes may be due to less competition for available resources, although temporal variation should be included before drawing final conclusions on spiders’ body condition in the agricultural landscape.     

Hypoxic coma as a strategy to survive inundation in a salt-marsh inhabiting spider

Spiders constitute a major arthropod group in regularly inundated habitats. Some species survive a flooding period under water. We compared survival during both submersion and a recovery period after submersion, in three stenotopic lycosids: two salt-marsh species Arctosa fulvolineata and Pardosa purbeckensis, and a forest spider Pardosa lugubris. Both activity and survival rates were determined under controlled laboratory conditions by individually surveying 120 females kept submerged in sea water. We found significant differences between the three species, with the two salt-marsh spiders exhibiting higher survival abilities. To our knowledge, this study reports for the first time the existence of a hypoxic coma caused by submersion, which is most pronounced in A. fulvolineata, the salt-marsh spider known to overcome tidal inundation under water. Its ability to fall into that coma can therefore be considered a physiological adaptation to its regularly inundated habitat.     

Morphology and function of the spinning apparatus of the wolf spider Pardosa amentata (Cl.) (Araneae,Lycosidae)

The histology of the four types of silk gland, occurring in the wolf spider Pardosa amentata are given. The changes in epithelium height, lumen breadth and size of the granules were studied in relation to silk production in the sub-adult and adult stage of the life-cycle.