Amazons Are Back: Absence of Males in a Praying Mantis from Uruguayan Savannas.

With a broad distribution throughout South America, Brunneria subaptera (Saussure) (Coptopterygidae) is the most abundant praying mantis species in Uruguay, mostly associated with grassland vegetation. Their body coloration can vary between green or brown sepia. The recent catalog of Uruguayan mantids showed that all specimens of B. subaptera from the collections in Uruguay were females, suggesting an absence of males in natural populations. The aim of this study was to determine the occurrence of female-only populations of B. subaptera in Uruguay. Moreover, we analyzed the genetic variation and estimated the time of the most recent common ancestor of the species. We performed bimonthly samplings in two localities in Uruguay for 1 year and we sexed the individuals. We extracted DNA from the abdominal tissue of adult females and sequenced a fragment of the mitochondrial cox1 gene to assess the genetic diversity and to estimate diversification times. We did not find any males in either of the studied localities. The age estimates revealed a recent origin of the species (2.33 Ma), and there was no genetic differentiation between the individuals from Uruguayan localities. This absence of males of B. subaptera documented in the sampled locations suggests that the species reproduces by thelytokous parthenogenesis. This study opens promising venues for future research into reproductive strategies and polymorphism in South-American praying mantises that inhabit one of the largest areas of grasslands in the world, currently under dramatic deterioration and reduction.

Reproductive diapause influences spermatogenesis and testes' size in the diplochronous wolf spider Allocosa senex (Lycosidae, Araneae) - A case study using a non-experimental approach.

Seasonality considerably impacts on the life of organisms and leads to numerous evolutionary adaptations. Some species face seasonal changes by entering a diapause during different life stages. During adulthood, a diapause in the non-reproductive period can affect male gametogenesis as, for example, it occurs in insects. Spiders are distributed worldwide and show a variety of life cycles. However, data on spiders' life cycles and seasonal adaptations are limited. Here, we explored the effect of reproductive diapause in a seasonal spider for the first time. We used the South American sand-dwelling spider Allocosa senex as a model as this species is diplochronous, meaning that individuals live two reproductive seasons, with juveniles and adults overwintering in burrows. It has been observed that individuals of this species reduce their metabolism during the non-reproductive season, diminishing prey consumption and locomotion to a minimum. This species is also well-known for exhibiting wandering and courting females and sedentary males. We analyzed spermatogenesis throughout the male's life cycle and described the male's reproductive system and spermiogenesis using light and transmission electron microscopy. We found that spermatogenesis in A. senex is asynchronous and continuous. However, when males face the non-reproductive season, the late spermatogenic stages and spermatozoa decline, causing an interruption but not a total arrest of this process. This seasonality is also reflected in smaller testes' size in males from the non-reproductive season than in other periods. The mechanisms and constraints are unknown, but they could be related to the metabolic depression during this life cycle period. Since sex-role reversal apparently sets a low-intensity sperm competition scenario compared with other wolf spiders, surviving two reproductive seasons may balance mating opportunities by distributing them between both periods. Thus, the partial interruption of spermatogenesis during diapause could allow new mating encounters during the second reproductive season.

Breaking the cliché: sex reversal in size dimorphism and mobility in South American Allocosinae (Lycosidae) spiders

Typically, females and males are expected to have characteristic sexual strategies and patterns of size dimorphism, but these generalizations are subject to exceptions. The occurrence of atypical cases has been related to species or populations from environments under strong physical, ecological and/or social constraints. Allocosa marindia and Allocosa senex are two coastal spiders (Lycosidae: Allocosinae) with reversal in sex roles and sexual size dimorphism. Males are larger than females, and females are the mobile sex that initiates courtship. It is unclear whether the occurrence of non-typical sexual traits in Allocosinae spiders is correlated with coastal habitats. Our aim was to study sexual size dimorphism and surface mobility in Allocosinae spiders from different habitats throughout South America. We revised specimens from scientific collections and performed 3-day samplings to collect individuals and determine nocturnal surface mobility. We analysed a total of 1071 Allocosinae adult individuals from 18 species and/or morphotypes. Our results revealed new species inhabiting coastal habitats with reversal in sexual size dimorphism and higher nocturnal surface activity in females; however, not all coastal species shared those characteristics. Future studies will focus on studying other ecological, physiological and/or phylogenetic factors that could be shaping the origin and maintenance of sex role reversal in Allocosinae.

Challenging monogamy in a spider with nontraditional sexual behavior.

Each species and sex can develop different reproductive strategies to optimize their fitness while assigning reproductive effort. Allocosa senex is a sex-role reversed spider whose males construct long burrows in the sand. They wait for wandering females to approach, assess their sexual partners and donate their constructions to females after copulation. Females stay in the burrow and lay their egg-sac. When offspring are ready for dispersion, females leave the burrow and gain access to new mating opportunities. Males are choosy during mate courtship, preferring to mate with virgin females over copulated ones, which can even be cannibalized if males reject them. This situation turns new mating opportunities dangerous for copulated females. We wondered whether a copulated female inside the previous mate's burrow responds to courtship from a new male and if this new male can copulate, avoiding burrow construction costs. We also explored whether courtship and copulation behaviors during the first sexual encounter affected the probability of occurrence of a second copulation. For that purposes we exposed copulated females inside male burrows to new males (non-donor males). Males could locate and court females inside the previous male's burrow, and females accepted a second copulation. Hence, A. senex females are not monogamous as was expected but increase their reproductive success by copulating with non-donor males. Also, males can develop opportunistic tactics, suggesting a more dynamic mating system for this sex-role reversed spider than assumed.

Size matters: Antagonistic effects of body size on courtship and digging in a wolf spider with non-traditional sex roles.

Body size, nuptial gift characteristics and courtship behaviour, among other traits, can reflect the quality of a potential mate and, thus, might be under sexual selection. To maximize their mating success, individuals can show behavioural plasticity in sexual context. Allocosa senex is a burrow-digging wolf spider that exhibits reversal in courtship roles and in sexual size-dimorphism expected for spiders. Males construct the mating refuge and females prefer males that build longer burrows, which are considered as nuptial gifts because they are delivered to them after mating. This study aims to determine whether male body size and female reproductive status influence burrow dimensions, courtship displays, female preferences and cannibalism rate in A. senex. For that purpose, we allowed males to construct burrows and performed sexual trials under laboratory conditions. Larger males were more courted by females, and in turn, they expressed more vibratory behaviours during courtship. However, and contrary to our expectations, smaller males constructed longer burrows. We suggest that males of A. senex exhibit size-dependent behavioural plasticity, and when they are smaller, they invest more in burrow construction to compensate their lower opportunities of courting intensively as larger males. In addition, females courted differentially according to their reproductive status, overriding male preferences for virgin females. This study opens several doors to future research regarding mutual choice in A. senex and the traits assessed by males and females during courtship, as well as about the influence of intrinsic and extrinsic factors shaping reproductive decision-making in this and other wandering spider species.

Sex and burrowing behavior and their implications with lytic activity in the sand-dwelling spider Allocosa senex.

The immune response can be costly. Studies in several arthropod species have indicated a trade-off between immunity and other life-history traits, including reproduction. In sexually dimorphic species in which females and males largely differ in their life history strategies and related energetic demands, we can expect to find sex differences in immune functions. Sex differences in immunity are well documented in vertebrates; however, we largely lack data from invertebrate systems. Lytic activity, the immune system's ability to lysate bacteria and viruses, has been widely used as a proxy for the strength of the immune response in several invertebrates. With this in mind, we used the burrowing wolf spider Allocosa senex to test differences in lytic activity between females and males. We also studied whether digging behavior affects the immune responses in this species. While females of A. senex construct simple refuges where they stay during the day, males construct deep burrows, which they donate to females after copulation. In accordance with our hypothesis, females showed higher lytic activity compared with males, and those males who dug showed higher levels of lytic activity than those that did not dig. Furthermore, male body condition and lytic activity did not correlate with burrow length, a trait under female choice in this species. Our results show sexual dimorphism in lytic activity responses, which are likely related to differences in life-history strategies and energetic requirements of each sex in A. senex spiders.

No pain, no gain: Male plasticity in burrow digging according to female rejection in a sand-dwelling wolf spider.

Behavioral plasticity allows individuals to reversibly respond to short-term variations in their ecological and social environment in order to maximize their fitness. Allocosa senex is a burrow-digging spider that inhabits the sandy coasts of South America. This species shows a reversal in typical sex roles expected in spiders: females are wanderers that visit males at their burrows and initiate courtship. They prefer males with long burrows for mating, and males prefer virgin over mated females. We tested whether female sexual rejection induced males to enlarge their burrows and if female reproductive status affected males' responses. We exposed males who had constructed burrows to: a) virgin females or b) mated females, (n=16 for each category). If female rejection occurred, we repeated the trial 48h later with the same female. As control, we maintained a group of males without female exposure (unexposed group, n=32). Rejected males enlarged their burrows more frequently and burrows were longer compared to unexposed males. However, frequency and length of enlargement did not differ according to female reproductive status. Males of A. senex showed plasticity in digging behavior in response to the availability of females, as a way to maximize the possibilities of future mating.

Geographical variation in sexual behavior and body traits in a sex role reversed wolf spider.

Mating partners need to recognize, assess each other, and exchange information through behavioral events that occur before, during, and after mating. Sexual signals, as well as life history traits, are influenced by selective pressures and environmental factors that can vary across distant geographical areas. Allocosa senex is a sand-dwelling wolf spider which constructs burrows along the sandy coasts of Argentina, Brazil, and Uruguay. Females are the mobile sex that searches for males and initiates courtship. They prefer males which construct longer burrows, and males prefer virgin females in good body condition. The objective of this study was to compare sexual behavior patterns, as well as body characteristics and burrow dimensions, between two geographically distant locations of A. senex, one in Uruguay (Uruguayan location) and the other from central Argentina (Argentinean location). We found differences in the number of male abdominal vibrations, male and female touches during mating, and number of erections of male leg spines, which all were higher in matings of Argentinean pairs. On the other hand, male body mass and female body condition were higher in Uruguayan individuals. The wide distribution of A. senex could be determining variations in the biotic and abiotic features that affect the species, generating differences in the strength of selective forces acting on individuals from the two studied locations.

Communication during copulation in the sex-role reversed wolf spider Allocosa brasiliensis: Female shakes for soliciting new ejaculations?

Traditional studies on sexual communication have focused on the exchange of signals during courtship. However, communication between the sexes can also occur during or after copulation. Allocosa brasiliensis is a wolf spider that shows a reversal in typical sex roles and of the usual sexual size dimorphism expected for spiders. Females are smaller than males and they are the roving sex that initiates courtship. Occasional previous observations suggested that females performed body shaking behaviors during copulation. Our objective was to analyze if female body shaking is associated with male copulatory behavior in A. brasiliensis, and determine if this female behavior has a communicatory function in this species. For that purpose, we performed fine-scaled analysis of fifteen copulations under laboratory conditions. We video-recorded all the trials and looked for associations between female and male copulatory behaviors. The significant difference between the time before and after female shaking, in favor of the subsequent ejaculation is analyzed. We discuss if shaking could be acting as a signal to accelerate and motivate palpal insertion and ejaculation, and/or inhibiting male cannibalistic tendencies in this species.

Sexual dimorphism in immune response, fat reserves and muscle mass in a sex role reversed spider.

In sex role reversed species, females and males adopt behaviors that are not the traditional ones for that animal group. Furthermore, this reversal can translate into physiological differences between the sexes in characteristics such as energetic demands or immune response. Allocosa brasiliensis shows a reversal in the sex roles and sexual size dimorphism that would be expected for spiders. Males are larger than females and are sedentary, while females are the mobile sex that looks for males and initiates courtship. Our objective was to explore the occurrence of sexual dimorphism in immune response, fat content and muscular mass in A. brasiliensis, and relate the results to the reproductive strategies of the species. An encapsulation response was used as an estimate of the immune response. Abdominal fat content and leg muscular mass were quantified and the results were compared between females (N = 19) and males (N = 21). Males showed higher values of the three characteristics as compared to females. Life history divergences between the sexes regarding size, mobility and foraging opportunities could be factors driving these differences in immune response and energy requirements.

Sexual behavior, cannibalism, and mating plugs as sticky traps in the orb weaver spider Leucauge argyra (Tetragnathidae).

Unpublished field observations in Leucauge argyra, a tropical orb weaver spider, suggest the occurrence of conspicuous mating plugs that could reduce or prevent remating attempts. Otherwise, the sexual behavior of this species remains unknown. The aims of this study were to describe the courtship behavior and copulation in L. argyra and investigate mating plug formation in this species. Fourteen virgin females and 12 plugged females were exposed to up to three males and checked for mating plug formation. Of the 12 virgins that copulated, nine produced plugs (five immediately after copulation), and the five plugged females that copulated produced another mating plug immediately after copulation. We did not detect the transfer of any male substance during copulation but observed a whitish liquid emerging from female genital ducts. Plug formation was positively associated with male twanging during courtship. One virgin and four plugged females cannibalized males. In seven trials with virgins and in three trials with plugged females, the male's palp adhered to a substance that emerged from female genital ducts and spread on her genital plate. The male had to struggle energetically to free his glued palp; two of these males were cannibalized while trying to release their palps. Females seem to determine copulation duration by altering the timing of mating plug formation and through sexual cannibalism. This is the first case reported of a mating plug as a sticky trap for males.

Behavioural evidence of male volatile pheromones in the sex-role reversed wolf spiders Allocosa brasiliensis and Allocosa alticeps.

The use of chemical signals in a sexual context is widespread in the animal kingdom. Most studies in spiders report the use of female pheromones that attract potential sexual partners. Allocosa brasiliensis and Allocosa alticeps are two burrowing wolf spiders that show sex-role reversal. Females locate male burrows and initiate courtship before males perform any detectable visual or vibratory signal. So, females of these species would be detecting chemical or mechanical cues left by males. Our objective was to explore the potential for male pheromones to play a role in mate detection in A. brasiliensis and A. alticeps. We designed two experiments. In Experiment 1, we tested the occurrence of male contact pheromones by evaluating female courtship when exposed to empty burrows constructed by males or females (control). In Experiment 2, we tested the existence of male volatile pheromones by evaluating female behaviour when exposed to artificial burrows connected to tubes containing males, females or empty tubes (control). Our results suggest the occurrence of male volatile pheromones that trigger female courtship in both Allocosa species. The sex-role reversal postulated for these wolf spiders could be driving the consequent reversal in typical pheromone-emitter and detector roles expected for spiders.