Larger Females Research Articles

Living with males leads to female physical injury in the leaf-footed cactus bug

Males in many species possess sexually selected weapons that they use to fight for mating opportunities. It is well established that male-male competition can lead to physical injuries for males. However, very few studies have looked at the physical consequences for conspecific females. We hypothesized that living with males in a species with male-male competition would result in female injury. Because larger female invertebrates typically have greater reproductive output, they have higher resource value for males and can elicit aggression and fighting. Thus, we further hypothesized that larger females in this context would receive more injuries. For this study, we focused on the leaf-footed cactus bug, Narnia femorata (Hemiptera: Coreidae), a species of insect in which males fight using their spiny and enlarged hindlegs. In just 2 h of observation, we documented males competing with other males in 61% of 103 trials. In 43% of these 63 competitions, females were physically contacted and sometimes attacked with a kick or squeeze. We left insects in social groups for 74 h and found that females living with multiple males had a higher likelihood of obtaining injuries (26.2% of 103 trials) compared to those living only with females (9.7% of 103 trials). In addition, larger females were more likely to be injured compared to smaller females. Our study highlights the harm that females can experience in species with male-male competition.

Ecology and Biogeography of Sexual Size Dimorphism in Squamates

ABSTRACTAimSexual size dimorphism (SSD) is widespread in the animal kingdom. The direction and magnitude of SSD differ considerably across taxa, potentially due to different selective forces acting on female and male sizes. We assembled a comprehensive database of mean body sizes for female and male squamate species. We then tested for associations between the degree and direction of sexual size dimorphism and environmental factors, clutch/litter sizes, reproductive modes, substrate types, and species richness (a common measure of interspecific competition).LocationGlobal.Time PeriodPresent.Major Taxa StudiedSquamata (Reptilia).MethodsWe studied SSD patterns and their correlates for 11792 squamate species. We also tested the effect of the number of putative competitors on SSD within (~9915 km2) grid cells. We applied phylogenetic path analysis and phylogenetic generalised least squares regression (PGLS) at the species level and applied spatial auto‐regressive (SAR) multiple regressions at assemblage levels.ResultsIn general, snake females are larger than males, whereas male lizards are larger, on average, than females. Female squamates in general are larger than males in cold regions, while in warm regions, particularly in deserts, males are usually larger than females. SSD became more female‐biased (i.e., larger females) as clutch size increased, and viviparous taxa had more female‐biased SSD. There was little relationship between the magnitude of SSD and species richness. Sexual size dimorphism did not vary significantly across substrate types.Main ConclusionWe suggest that the mechanisms driving squamate SSD differ between oviparous and viviparous taxa. The more female‐biased SSD in colder regions is likely driven by fecundity selection, while a male bias in warmer regions may be associated with sexual selection. However, we found little evidence to suggest that natural selection for substrates, or resource‐based competition, affects squamate sexual size dimorphism and suspect the underlying hypotheses may be flawed, and/or that species richness is a poor measure of the intensity of interspecific competition.

Interactive effects of body size, food abundance and mating status on reproductive attributes in tortoise beetle

Abstract In numerous studies, it has been seen that the reproductive attributes of an insect may vary according to their body size, food fluctuation in the environment and their mating status. However, studies on the interactive effects of such biotic factors on mating behaviour and reproductive attributes are lacking. Therefore, the present study was designed to investigate the interactive effects of body size variation, different food conditions and the mating status on specific reproductive attributes such as sexual maturity, time to commencement of mating (TCM), latent period (LP), mating duration (MD), fecundity and per cent egg viability of spotted tortoise beetle, Aspidomorpha miliaris (Coleoptera: Chrysomelidae). For this, small‐ and large‐sized females were raised in different feeding conditions (limited and unlimited food) from emergence to sexual maturity and then allowed to mate once, twice, or multiple times with males of intermediate size. Results revealed a significant influence of body size on sexual maturity, as well as mating status on the time to commence mating and showed a significant interactive effect of food conditions and mating status on latent period and mating duration, as well as body size, food conditions, and mating status on fecundity and per cent egg viability of adult females. In the present study, small females attained sexual maturity earlier than large females. Fecundity and per cent egg viability were recorded at maximum in multiply mated large females in the unlimited food treatment. Food conditions substantially influenced fecundity, which was zero in females with limited food conditions. Therefore, this study suggests that multiply mated larger females with unlimited food have better reproductive outputs. It was also concluded that the interactive effect of biotic factors had a significant impact on reproductive outputs together with different sexual behaviours.

Not all is male biased: sexual dimorphism in carapace size and shape of the kelp crab Taliepus dentatus (Decapoda: Majoidea: Epialtdae)

ABSTRACT Different selection pressures acting on males and females generate biases in the cephalothoracic size and shape of many decapod crustaceans. We assessed such patterns in the carapace components (dorsal cephalothorax, ventral cephalothorax, and pleotelson) of the kelp crab Taliepus dentatus, with morphological variables based on Euclidean and Procrustes coordinates of homologous landmarks, which describe variations in size and shape, respectively. The dorsal and ventral cephalothorax exhibited male-biased sexual size dimorphism, differing from the female-biased sexual size dimorphism expressed by the shape and size of the pleotelson. Size allometry was more restricted in the cephalothorax than in the pleotelson. Shape allometry showed greater restriction in larger females, with robust and parallel limits of variation in males. In contrast, the shape of the pleotelson (female-biased) varied within parallel limits in both sexes. The variations in shape and size of both groups of traits are mainly attributable to adaptive factors that favour population fitness in females through natural selection and individual fitness explained through sexual selection in males. However, the variation in the shape of the dorsal and ventral cephalothorax indicates inherent structural complexity, where developmental modules favour the muscular performance of the chelipeds and the remaining pereiopods. Our results suggest that stabilising selection acting differentially on the shape and size of the caparace components directs the expression of this body plan towards optimality, promoting morphological convergences and taxonomic conflicts for this species, which is consistent with highly integrated complex structures.

Reproductive Biology of the Smoothtail Spiny Lobster (Panulirus laevicauda) to Support Fisheries Management in Southeastern Brazil

ABSTRACTThe smoothtail spiny lobster (Panulirus laevicauda) is one of the main target species for fisheries in Brazil. We aimed to estimate reproductive parameters of this species in support of a minimum legal size limit for the region. A sample of 508 individuals ranged 13.75–100 mm in cephalothorax length (CL) from the southern coast of Espírito Santo, Brazil, during 2020–2021. Males were more abundant than females. Females reached morphological maturity before males, at 68.4 mm CL, for both sexes. Reproduction was between November and March, and some large females were capable of spawning more than once during the reproductive period. Because large adult females could spawn more than once per season, we recommend increasing the legal minimum capture size of P. laevicauda to 70 mm CL in Brazil.

Bacillus velezensis as a Novel Species for Mosquito Control: Impacts of Exposure on Key Life History Traits of Culex quinquefasciatus.

Bacillus velezensis is a promising mosquito vector biocontrol agent due to its potent larvicidal properties. This study investigated the lethal and sublethal effects of exposure to crude toxin of B. velezensis strain WHk23 on Culex quinquefasciatus. Toxicity bioassays showed a median lethal concentration of 28.30 µg/mL for third instar larvae. In addition to direct larval mortality, sublethal exposures resulted in concentration-dependent decreases in survival, fecundity, and fertility. Exposure to different concentrations was found to accelerate larval development, resulting in larger females, yet those females exhibited lower lifespan and diminished reproductive success. These results suggest that exposure to B. velezensis not only reduces mosquito populations but also has lasting effects on important life history parameters, ultimately lowering vectorial capacity and disease transmission risk. This study provides valuable insights into the lethal and sublethal consequences of B. velezensis exposure. Given its potent larvicidal properties, B. velezensis represents a promising biocontrol agent for mosquito vectors.

Indirect fitness benefits can reinforce the direct fitness benefits of male mate choice.

Mating preferences frequently evolve because they confer fitness benefits to the chooser. While there is strong evidence for both direct and indirect fitness benefits in the context of female mate choice, the potential for males to receive indirect benefits from mate choice remains understudied. Male mate choice for larger female body size is widespread among ectotherms and is generally attributed to the direct fitness benefits larger females provide due to their higher fecundity. However, when female size is heritable or condition-dependent, males could also receive indirect benefits from mating with larger females in the form of increased offspring fitness. We tested this possibility in Drosophila melanogaster by first confirming the direct fitness benefits associated with large females and then comparing the fitness of offspring produced by large versus small females. Although female body size did not influence offspring juvenile viability, large females produced daughters with higher reproductive success than the daughters of small females. Males only received these indirect benefits through their daughters, however, as reproductive success did not differ significantly between the sons of large and small females. Our findings demonstrate that indirect fitness benefits can serve to reinforce the direct benefits of male mate choice, suggesting that indirect effects may play an underappreciated role in the evolution of male preferences.

A new species of the Phalloceros harpagos species complex (Cyprinodontiformes: Poeciliidae) from the middle and lower Uruguay River floodplains

Phalloceros mimbi sp. nov. is described from specimens collected in the floodplains of the middle and lower Uruguay River in Uruguay (Río Negro and Salto Departments) and Argentina (Entre Ríos Province). This species belongs to the P. harpagos species complex and is diagnosed by the following unique combination of characters: 1) presence of a symmetric hood-like structure located immediately anterior to the urogenital papilla in females, 2) absence of large papillae at the mandibular symphysis of large adult females, 3) possession of well-developed hooks positioned in distal portion of gonopodial appendices of males and4) lateral spot usually present in both sexes. To date, P. mimbi has not been found coexisting in sympatry with other Phalloceros species. The geographically closest species is P. caudimaculatus, who differs from P. mimbi by the absence of hooks in gonopodial appendix in males. Two of the three known populations of P. mimbi are relatively protected from the main threats to the species. However, conservation initiatives should consider its presence and ensure that it receives the necessary attention; considering that its global distribution occupies less than 200.000 km2 and its occurrence area represents less than 10% of the national territory it must be considered a priority for conservation and an Threatened Species for the species list of Sistema Nacional de Áreas Protegidas (SNAP) of Uruguay.

Impact of Male and Female Body Size on Mating Performance and Reproductive Success in Black Soldier Fly, Hermetia illucens L. (Diptera: Stratiomyidae)

Mating and Reproductive success is the most essential phase in establishing efficient maintenance to ensure a sustained supply of Black Soldier Fly/BSF populations. This study aims to ascertain the influence of mating partner body size on successful mating and reproduction in BSF. Adult males and females BSF of the same age (< 24 hours) were paired based on a combination of body size treatments (large, average, small). Semi-outdoor observations were conducted in 30x30x30 cm nylon net cages, with each treatment repeated three times. The observation parameters include mating frequency, oviposition frequency, egg number, egg weight, and egg viability or fertility. The differences in body size of mating partners significantly influence mating and reproductive success in BSF. The combination of a mating pair consisting of a large male and a large female had the highest frequency of mating (35.3 ± 4.64) and oviposition (28 ± 2.04), whereas mating pairs of a small male and a small female had a low mating frequency (13.7 ± 5.18) and lowest oviposition (5.0 ± 0.63). Mating tended to occur more frequently when males were matched with large females than when males were paired with average or small females.

Reproductive biology of the freshwater prawn Macrobrachium rude (Decapoda: Palaemonidae)

Macrobrachium rude is a notable freshwater prawn in Bangladesh owing to its palatability and considerable market value. This study examined the reproductive biology of the species for the first time, encompassing sex ratio, size at sexual maturity, reproductive pattern, egg size, and fecundity. A total of 917 individuals were examined, with males ranging from 4.1 to 7.4 mm carapace length (CL) and females from 4.8 to 14.0 mm CL. The sex ratio exhibited a male bias in the smaller size group and a female bias in the larger size group. The size at which 50% of females reach maturity was estimated to be 8.4 mm CL. Ovigerous females had no mature ovaries right after spawning; however, the percentage of mature ovaries increased as the eggs developed, approaching nearly 100% prior to hatching, suggesting the potential for multiple spawning of females within a reproductive period. Egg size was not correlated with female size but increased with development. The total number of eggs was significantly larger in larger females. The results of this study will help in developing sustainable management and conservation approaches for this significant fishery.

Silver spoons, reproduction, and growth catch-up in eastern Grey kangaroos.

Early growth and environmental conditions can shape life-history trajectories. Long-lived iteroparous species with indeterminate growth face growth-reproduction trade-offs for most of their lives. Poor early conditions can delay primiparity and restrict growth, potentially compensated for by faster growth later in life, at the cost of reduced reproduction. We explored the variation in age at primiparity and early reproduction in eastern Grey kangaroos, based upon 13 yr of monitoring about 100 known-age females. We then examined associations between early reproduction, later reproduction, and lifetime growth. We used a modified von Bertalanffy growth function to model the indeterminate growth of females and to test the effects of early reproduction on lifetime growth. Favorable environmental conditions, large size, and condition as a subadult led to earlier reproduction and higher reproductive success at ages 3 to 5. As females aged, size and condition had diminishing effects on reproductive success. Females with greater early-adulthood reproduction had slightly higher reproduction later in life. We did not detect a growth cost of early reproduction. Large females in good condition favored early reproduction over growth, while those with poor early growth exhibited growth catch-up at the cost of reproduction both early and later in life. As reported for other long-lived iteroparous species with indeterminate growth, female kangaroos prioritize growth over reproduction for much of their lifespan. Eastern Gray kangaroos show heterogeneity in early growth and reproductive strategies. Early primiparity and reproduction are constrained by body condition, size, and environmental conditions when females are pre-reproductive subadults.

Phenotypic effects on male risk-taking behavior in Leptuca beebei from field experiments at Playa El Agallito, Chitre, Panama

Risk-taking behavior in Leptuca beebei was studied through field experiments conducted at Playa El Agallito, Chitré, Panama. This study aimed to determine whether larger males or those with pillar structures take greater risks when courting females and whether female size affects male risk-taking behavior. Females exhibiting mate-searching behavior were tethered 30 cm in front of randomly selected courting males, both with and without pillars. Male behavior was observed using binoculars, recording the maximum distance traveled from their burrow and whether they intercepted the female. Results indicate that male size and pillar presence did not significantly affect the distance traveled or the likelihood of intercepting the tethered female. Males traveled an average of 18.8 cm from their burrows, but this distance was not influenced by female size. Instead, males' risk-taking behavior may have been influenced by factors such as predation risk and potential burrow loss, rather than female size. The findings suggest that the dangers of leaving their burrows outweigh the benefits of courting distant or larger females. Thus, their behavior is influenced more by ecological risks and reproductive costs than by the characteristics of potential mates.

Solar radiation alters heat balance and thermoregulation in a flying desert bee.

Solar radiation is an important environmental variable for terrestrial animals, but its impact on the heat balance of large flying insects has been poorly studied. Desert bees are critical to ecosystem function through their pollination services, and are exposed to high radiant loads. We assessed the role of solar radiation in the heat balance of flying desert Centris pallida bees by calculating heat budgets for individuals in a respirometer in shaded versus sunny conditions from 16 to 37°C air temperatures, comparing the large and small male morphs and females. Solar radiation was responsible for 43 to 54% of mean total heat gain. Bees flying in the sun had thorax temperatures 1.7°C warmer than bees flying in the shade, storing a very small fraction of incident radiation in body tissues. In most cases, flight metabolic rate was not suppressed for bees flying in the sun, but evaporative water loss rates more than doubled. The most dramatic response to solar radiation was an increase in convection, mediated by a more than doubling of convective conductance, allowing thermoregulation while conserving body water. In large morph males and females, the increased convective conductance in the sun was mediated by increased heat transfer from the thorax to abdomen. Because convection is limited as body temperatures approach air temperatures, solar radiation combined with warming air temperatures may cause endothermic flying bees to reach a tipping point at which increases in non-sustainable evaporation are necessary for survival.

Genetic Variation in Male Mate Choice for Large Females in Drosophila melanogaster.

Males in many species show courtship and mating preferences for certain females over others when given the choice. One of the most common targets of male mate choice in insects is female body size, with males preferring to court and mate with larger, higher-fecundity females and investing more resources in matings with those females. Although this preference is well-documented at the species level, less is known about how this preference varies within species and whether there is standing genetic variation for male mate choice within populations. We used hemiclonal analysis in the fruit fly, Drosophila melanogaster, to test for heritable genetic variation in pre- and postcopulatory components of male mate choice for large females. We found additive genetic variation for both forms of male choice: Males from different hemiclone lines varied in the strength of their courtship preferences for large females and the degree to which they extended matings with large females. Although males from hemiclone lines with stronger courtship preferences for large females were more likely to mate with those females, there was no genetic correlation between pre- and postcopulatory components of male mate choice, suggesting that they are under independent genetic control. Genetic variation in male mate choice may be widespread, potentially impacting the fitness of both sexes and the adaptive evolution of populations.

Assessing spawning behavior at the northern latitudinal extreme of Pacific halibut

Many species have reduced reproductive potential at the poleward extreme of their range, where they exhibit unique spawning dynamics. However, recent poleward range expansions have resulted in many species being unstudied in these regions, such as the Pacific halibut ( Hippoglossus stenolepis) in the northern Bering Sea (NBS). To characterize Pacific halibut spawning dynamics at the northern extreme of their range, we attached pop-up satellite telemetry tags to large females in the NBS, with time series data and tag reporting locations being used to infer spawning behavior and to identify occupied spawning habitat conditions, location, and timing. Pacific halibut in the NBS occupied spawning habitat later and farther north than previously described, where spawning habitat was occupied from January to May and reached as far north as the Russian continental shelf edge. Additionally, 42% of mature individuals never occupied presumed spawning habitat, suggesting the presence of skip spawning behavior. These findings suggest that Pacific halibut exhibit unique spawning dynamics in the NBS, which may result in a reduced reproductive potential within this northern population component.

A vain attempt: why a change in the reproductive strategy of a rodent population failed to restore ecological balance

Based on long-term observations of a bank vole (Myodes glareolus Schreber, 1780) population, the preconditions, mechanisms, and consequences of changes in reproductive strategy to restore ecological balance are analysed. The studies were conducted in the Kaniv Nature Reserve during different periods of the ecosystem’s existence affected by anthropogenic impact. The characteristics of two cycles of population density dynamics are compared. The first cycle covers the destabilisation stage of the dynamics, which precedes changes in reproductive strategy, whereas the second cycle demonstrates changes in breeding parameters. It was found that in order to compensate the high levels of mortality, reproductive processes intensify. This is achieved by the simultaneous launch of various mechanisms, from increasing individual fecundity to expanding the proportion of individuals involved in reproduction. The first scenario turned out to be unsuccessful and, without adequate ecological support in the form of sufficient feeding resources, caused even higher mortality rates. The size and number of litters dropped to minimum values, and reproduction parameters indicated only a slight success in realising the breeding potential. The most vulnerable was the group of large females, whose fecundity and, consequently, contribution to population growth are the highest. It was found that such processes changed not only the size and weight composition of the population (the proportion of large individuals decreased), but also the sex and age structure (the number of adult females decreased). The decline in the number of large breeding females only exacerbated the misbalance. Population collapse was avoided by expanding the number of females involved in reproduction (extensive reproduction). It is shown that restoration of ecological balance did not follow the change in reproductive strategy. This is supported by the following facts: preservation of signs of a destabilised population dynamics and absence of the previous interdependencies between demographic indices and parameters of feeding resources. As a result, after about a decade of intensive but ineffective reproduction, the population was ‘forced’ to switch to another strategy to restore ecological balance.

QUANTITATIVE VARIABLES OF ARTEMIA IN SALT LAKE TUZKOL (BALKHASH-ALAKOL BASIN)

Studies of salt waterbodies are topical for understanding their role in preserving and producing cysts of the valuable bioresource Artemia. A comprehensive analysis of the salt lake Tuzkol carried out in May and August 2022. Standard methods performed sampling and processing of samples of Artemia. Artemia population represented by a bisexual race, but abundance of males is insignificant (0.02% of total population). The total abundance of artemia reached 34.0 thousand individuals/m3 in May and 13.0 thousand individuals/m3 in August. Nauplii and individuals at the juvenile stage formed the basis of quantitative variables of artemia. Artemia biomass reached 31.0 mg/m3 in May and 13.0 mg/m3 in August. Large egg-carrying females created the population biomass. One-way analysis of variance (ANOVA) did not reveal significant differences between the average values ​​of the abundance and biomass of Artemia in different years. During research period, a slight increase in the biomass of Artemia noted, which might be due to a twofold increase in the level of TDS in lake in 2022 compared to the previous year of study. The productivity (by cysts) of Artemia varied from 10.3 to 190.8 kg/ha. The main portion of Artemia cysts concentrated in female ovisacs and the water column in May, by August the cysts prevailed (98.1%) in bottom sediment. In both research periods, cysts were not found in the coastal zone where their catching carried out. Based on benthic cysts, Tuzkol Lake classified as a highly productive waterbody in summer 2022. Bottom cysts, being at the dormant stage, provide the abundance of the first generation of Artemia and are a reserve for preserving the population. Based on this, Lake Tuzkol recommended to be used as a habitat for preserving the unique gene pool of Artemia and for studying the adaptation of halobionts to unstable conditions of the aquatic environment.

Selection for increased sexual bimaturism in the common housefly, Musca domestica

Abstract The growing demand for insect mass production requires methodologies that allow for easy separation of males and females as the two sexes can have very different economic values. The common housefly, Musca domestica (L.) (Diptera: Muscidae), is one of the candidate species for industrial production. In this species, males typically develop faster and are smaller than females. Exploiting housefly sexual bimaturism (sexual difference in developmental time) could allow the maintenance of colonies with a higher proportion of females, which is favourable in terms of egg yield. Here, we report an artificial selection experiment of breeding early‐emerging males with late‐emerging females for six generations, with the aim to increase the time interval between emergence of males and females. The level of sexual bimaturism was not increased after six generations of selection. The life‐history traits fecundity, egg‐to‐adult survival and hatchability were not affected by the selection procedure. Wing size measurement revealed that early‐ and late‐emerging males did not differ in size, suggesting that developmental time and body size are decoupled in this sex. Still, large females consistently emerged later than small females, suggesting an intricate sex‐specific genetic architecture for size and developmental time in the housefly. Our results are relevant in the context of mass production, providing new insight into the manipulation and maintenance of female‐biased sex ratios in production colonies of the common housefly.

Sexual size dimorphism affecting mate choice and reproduction in the pine sawyer beetle, Monochamus alternatus.

Monochamus alternatus is a notorious pest of the pine forests across East Asian countries. Both large and small individuals of either sex coexist in the wild. However, whether and how body size affects sexual selection outcomes and longevity remains unknown. In this study, we allowed a male or a female of small or large size to choose between two mates of varying sizes, which were from a wild-caught population. Our results revealed that mating latency and frequency of mating attempts varied across different social environments. Both large males and large females preferred larger mates, whereas this preference was not observed for the small males and small females. Male-male competition had no significant effect on ejaculation duration and frequency or sperm number in ejaculates. When a single male had an opportunity to choose between two females of different sizes, the reproductive investment of small males towards the small females was almost doubled compared to that of the large males, i.e., longer ejaculation duration, higher ejaculation frequency, and greater numbers of sperm transferred. However, the greater mating efforts of small males did not translate to greater fecundity/fertility of small females. We also showed that regardless of their body size, females always had greater fertility when they mated with large males but evidence for a trade-off between fecundity and longevity was not found. Alternatively, female fecundity was positively associated with longevity, and fertility was negatively associated with longevity and fecundity. We discussed these results placing it in a broader context of mate choice evolutionary dynamics.

Phenology and social status of the endemic Sable Island Sweat Bee, Lasioglossum sablense

Lasioglossum (Dialictus) sablense Gibbs, 2010 (Hymenoptera: Halictidae) is endemic to Sable Island, an isolated sandbar located about 160 km east of Nova Scotia, Canada. Lasioglossum sablense is currently listed as Threatened due to its restricted geographic distribution, and promoting its conservation requires detailed information about nesting phenology, behaviour, and other life history details. We combined measurements of body size, wear, and ovarian status from adult females collected from flowers in 2016 and 2017 with measurements and behavioural observations of foragers collected at the Sable Island Station in 2019 and 2022, to compile a preliminary description of flight phenology and social behaviour. Like many members of its genus, L. sablense exhibited a diphasic life history. Phase 1 began when large, fertilized adult females emerged from hibernation, began nest and brood cell construction, and foraged to provision Brood 1, which comprised both daughters and sons. Most nests initiated during Phase 1 reactivated during Phase 2, with adult Brood 1 daughters foraging to provision Brood 2. Observations of bees at nest entrances demonstrated that some reactivated nests contained multiple adult females. Phase 1 foragers were larger, less worn, and had marginally higher levels of ovarian development than Phase 2 foragers. Five large females collected during Phase 2 had high levels of wear, suggesting they were long-lived Phase 1 foundresses that resumed foraging during Phase 2. Taken together, these observations suggest a mix of univoltine (single generation) and bivoltine (two generation) life histories, as well as a mix of solitary and social behaviour in L. sablense.