ABSTRACT
We used computer simulations of growth, mating and death of cephalopods and fishes to explore the effect of different life-history strategies on the relative prevalence of alternative male mating strategies. Specifically, we investigated the consequences of single or multiple matings per lifetime, mating strategy switching, cannibalism, resource stochasticity, and altruism towards relatives. We found that a combination of single (semelparous) matings, cannibalism and an absence of mating strategy changes in one lifetime led to a more strictly partitioned parameter space, with a reduced region where the two mating strategies co-exist in similar numbers. Explicitly including Hamilton's rule in simulations of the social system of a Cichlid led to an increase of dominant males, at the expense of both sneakers and dwarf males ("super-sneakers"). Our predictions provide general bounds on the viable ratios of alternative male mating strategies with different life-histories, and under possibly rapidly changing ecological situations.
Subject(s)
Cephalopoda , Computer Simulation , Fishes , Models, Biological , Sexual Behavior, Animal , Animals , Male , Sexual Behavior, Animal/physiology , Cephalopoda/physiology , Fishes/physiology , Female , Reproduction/physiology , Cannibalism , Mathematical Concepts , Cichlids/physiologyABSTRACT
BACKGROUND: Male courtship investment may evolve in response to the male's expectation of future mating opportunities or the degree of female control during mating interactions. We used a comparative approach to test this hypotheses by assessing the courtship and mating behaviors of five widow spider species (genus Latrodectus) under common laboratory conditions. We predicted male investment in courtship would be higher in species where males mate only once because of high cannibalism rates (monogyny, L. geometricus, L. hasselti, L. mirabilis), compared to species with rare cannibalism (L. mactans, L. hesperus) in which males should reserve energy for future mating opportunities. Increased male investment, measured as courtship duration, might also evolve with increased female control over mating outcomes if females prefer longer courtships. We tested this by assessing the frequency of copulations, timing of sexual cannibalism, and the degree of female-biased size dimorphism, which is expected to be negatively correlated with the energetic cost of rebuffing male mating attempts. RESULTS: Copulation frequency was consistently lower in species with extreme female-skewed size dimorphism, and where sexual cannibalism was more prevalent, suggesting the importance of female control for mating outcomes. We confirmed significant interspecific variation in average courtship duration, but contrary to predictions, it was not predicted by male mating system, and there was no consistent link between courtship duration and sexual size dimorphism. CONCLUSION: We show that the degree of sexual dimorphism is not only correlated with sexual cannibalism, but also with mating success since restriction of male copulation frequency by female Latrodectus affects paternity. However, predictions about male mating system or female control affecting courtship duration were not supported. We propose that the form of female control over mating and cannibalism, and male responses, might be more informative for understanding the evolution of courtship duration. For example, male tactics to avoid female aggression may drive lower courtship duration in species like L. mirabilis. Nonetheless, our results differ from inferences based on published studies of each species in isolation, illuminating the need for standardized data collection for behavioural comparative studies.
Subject(s)
Cannibalism , Sexual Behavior, Animal , Spiders , Animals , Male , Female , Spiders/physiology , Sexual Behavior, Animal/physiology , Courtship , Sex CharacteristicsABSTRACT
Ecological interactions among hosts are critical to consider when predicting disease dynamics. Most theory predicts that intraguild predation (IGP) and cannibalism negatively impact parasite populations, but this is based primarily on assumptions of simple or single-host life cycles. Here we investigate the effects of cannibalism in a size-structured host population on two digenean trematodes that have complex, multihost life cycles. A high incidence of cannibalism among paratenic hosts produced higher parasite infection loads and abundance, whereas cannibalism among obligate hosts reduced parasite abundances. We attributed this difference to trophic transmission aggregating parasites in larger, potentially fitter hosts and also to transmission among paratenic hosts via cannibalism. Moreover, we found evidence of indirect competitive interactions between parasites that can also increase infections at small scales. Our results show there are multiple mechanisms through which high cannibalism environments can benefit parasites that use paratenic hosts and trophic transfer to complete their life cycles.
Subject(s)
Cannibalism , Host-Parasite Interactions , Animals , Trematoda/physiology , Food Chain , Fish Diseases/parasitologyABSTRACT
Parasites are known to have direct effects on host dispersal ability and motivation. Yet, parasites have a variety of impacts on host populations, including shaping predation and cannibalism rates, and therefore may also have indirect effects on host dispersal; these indirect pathways have not been studied. We tested the hypothesis that parasites influence host dispersal through effects on cannibalism using backswimmers (Notonecta undulata) and Hydrachnidia freshwater mites. Mite parasitism impedes swimming in backswimmers, which we found increased their vulnerability to cannibalism. We imposed a manipulation that varied cannibalism rates across experimental populations consisting of a mix of backswimmers with and without simulated parasites. Using simulated parasites allowed us to examine the effects of cannibalism without introducing infection risk. We found that the odds of dispersal for infected backswimmers increased by 2.25× with every 10% increase in the risk of being cannibalized, and the odds of dispersal for healthy backswimmers increased by 2.34× for every additional infected backswimmer they consumed. Our results suggest that cannibalism was used as an energy source for dispersal for healthy individuals, while the risk of being eaten motivated dispersal in infected individuals. These results elucidate the complex ways that parasites impact host populations and strengthen our understanding of host-parasite interactions, including host and parasite population stability and spread. This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.
Subject(s)
Cannibalism , Host-Parasite Interactions , Mites , Animals , Mites/physiology , Animal Distribution , Heteroptera/physiology , Heteroptera/parasitologyABSTRACT
Factors that increase reproductive variance among individuals act to reduce effective population size (Ne), which accelerates the loss of genetic diversity and decreases the efficacy of purifying selection. These factors include sexual cannibalism, offspring investment and mating system. Pre-copulatory sexual cannibalism, where the female consumes the male prior to mating, exacerbates this effect. We performed comparative transcriptomics in two spider species, the cannibalistic Trechaleoides biocellata and the non-cannibalistic T. keyserlingi, to generate genomic evidence to support these predictions. First, we estimated heterozygosity and found that genetic diversity is relatively lower in the cannibalistic species. Second, we calculated dN/dS ratios as a measure of purifying selection; a higher dN/dS ratio indicated relaxed purifying selection in the cannibalistic species. These results are consistent with the hypothesis that sexual cannibalism impacts operational sex ratio and demographic processes, which interact with evolutionary forces to shape the genetic structure of populations. However, other factors such as the mating system and life-history traits contribute to shaping Ne. Comparative analyses across multiple contrasting species pairs would be required to disentangle these effects. Our study highlights that extreme behaviours such as pre-copulatory cannibalism may have profound eco-evolutionary effects.
Subject(s)
Cannibalism , Genetic Variation , Selection, Genetic , Sexual Behavior, Animal , Spiders , Animals , Spiders/genetics , Spiders/physiology , Male , Female , Biological EvolutionABSTRACT
Using a recursion model with real parameters of Nabis pseudoferus, we show that its filial cannibalism is an optimal foraging strategy for life reproductive success, but it is not an evolutionarily optimal foraging strategy, since it cannot maximize the descendant's number at the end of the reproductive season. Cannibalism is evolutionarily rational, when the number of newborn offspring produced from the cannibalized offspring can compensate the following two effects: (a) The cannibalistic lineage wastes time, since the individuals hatched from eggs produced by cannibalism start to reproduce later. (b) Cannibalism eliminates not only one offspring, but also all potential descendants from the cannibalized offspring during the rest of reproductive season. In our laboratory trials, from conspecific prey Nabis pseudoferus did not produce newborn nymphs enough to compensate the above two effects.
Subject(s)
Cannibalism , Reproduction , Humans , Infant, NewbornABSTRACT
AbstractThe adaptive value of routinely laying more eggs than can be successfully fledged has intrigued evolutionary biologists for decades. Extra eggs could, for instance, be adaptive as insurance against hatching failures. Moreover, because recent literature demonstrates that sibling cannibalism is frequent in the Eurasian hoopoe (Upupa epops), producing extra offspring that may be cannibalized by older siblings might also be adaptive in birds. Here, directed to explore this possibility in hoopoes, we performed a food supplementation experiment during the laying period and a clutch size manipulation during the hatching stage. We found that females with the food supplement laid on average one more egg than control females and that the addition of a close-to-hatch egg at the end of the hatching period increased the intensity of sibling cannibalism and enhanced fledging success in hoopoe nests. Because none of the extra nestlings from the experimental extra eggs survived until fledging, these results strongly suggest that hoopoes obtain fitness advantages by using temporarily abundant resources to produce additional nestlings that will be cannibalized. These results therefore suppose the first experimental demonstration of the nutritive adaptive function of laying extra eggs in vertebrates with parental care.
Subject(s)
Birds , Reproduction , Animals , Female , Humans , Clutch Size , Cannibalism , SiblingsABSTRACT
Animal welfare has evolved during the past decades to improve not only the quality of life of laboratory rodents but also the quality and reproducibility of scientific investigations. Bibliometric analysis has become an important tool to complete the current knowledge with academic databases. Our objective was to investigate whether scientific research on cannibalism/infanticide is connected with maternal aggression towards the offspring in laboratory rodents. To carry out our research, we performed a specific search for published articles on each concept. Results were analyzed in the open-source environment RStudio with the package Bibliometrix. We obtained 253 and 134 articles for the first search (cannibalism/infanticide) and the second search (maternal aggression towards the pups) respectively. We observed that the interest in infanticide/cannibalism started in the 1950s, while researchers started showing interest in maternal aggression towards the pups 30 years later. Our analyses indicated that maternal aggression had better citations in scientific literature. In addition, although our results showed some common features (e.g. oxytocin or medial preoptic area in the brain), we observed a gap between cannibalism/infanticide and maternal aggression towards the pups with only 14 published articles in common for both the searches. Therefore, we recommend researchers to combine both concepts in further investigations in the context of cannibalism for better dissemination and higher impact in laboratory rodents' welfare research.
Subject(s)
Aggression , Bibliometrics , Cannibalism , Animals , Female , Maternal Behavior , Rats/physiology , Animals, Laboratory/physiology , Rodentia/physiology , Animal Welfare , Mice/physiology , Behavior, AnimalABSTRACT
Female spiders and praying mantises are renowned for their cannibalism of male partners before, during, or after mating. While several hypotheses have been proposed to explain species-specific examples of sexual cannibalism, much variation remains unexplained, including why the timing of cannibalism varies across taxa. Here, I outline how sexually cannibalistic behavior could evolve via sexually antagonistic selection as a type of behavioral resistance to male-imposed mating costs, and how such a generalizable interpretation provides a framework for understanding the evolution of both sexual cannibalism in females and anti-cannibalistic traits in males. I discuss how differences between mating systems that physiologically constrain males to mate only once (monogyny) or twice (bigyny) and systems where the sexes can potentially mate multiply (polygyny and polyandry) are likely to influence how sexual conflict shapes cannibalistic behavior. I review key examples from the literature that suggest how sexually cannibalistic behavior might function as a female resistance trait and provide comprehensive predictions for testing this hypothesis empirically.
Subject(s)
Sexual Behavior, Animal , Spiders , Animals , Female , Male , Sexual Behavior, Animal/physiology , Cannibalism , Reproduction , Sex , Spiders/physiologyABSTRACT
El dragón rojo (1981) de Thomas Harris es la carta de presentación del perverso psiquiatra Hannibal Lecter, un psicópata asesino que brilla singularmente en el firmamento de los homicidas más abyectos de la historia del cine. Esta colección se completó con la publicación de El silencio de los corderos (1988), Hannibal (1999) y Hannibal: El origen del mal (2006), hasta ahora su última secuela. Fuente de inspiración cinematográfica, la saga se inició con la galardonada El silencio de los corderos (1991) de Jonathan Demme, continuada por Hannibal (2001) de Ridley Scott, El dragón rojo (2002) de Brett Ratner y Hannibal: El origen del mal (2007) de Peter Webber, si bien además existe una primera adaptación de la novela El dragón rojo titulada Hunter/Cazador de hombres (1986) de Michael Mann, donde los agentes del FBI se enfrentan con el temible asesino en serie Dr. Hannibal Lecktor (Brian Cox). Asimismo, el escritor y productor estadounidense Bryan Fuller creó la serie televisiva Hannibal (2013-2015), 39 episodios en los que el actor danés Mads Mikkelsen encarna al perverso antropófago. Independiente de sus connotaciones antropológicas, morales y culturales, la mayor parte de las conductas de canibalismo están relacionadas con trastornos psicóticos, como por ejemplo la esquizofrenia paranoica, cuando estos enfermos asesinan y se comen parte de los cuerpos de sus víctimas dentro de sus cuadros delirantes patológicos.(AU)
The Red Dragon (1981) by Thomas Harris is the introduction of the perverse psychiatrist Hannibal Lecter, a murdeous psychopath who shines singularly in the firmament of the most heinous murderers in the history of cinema. This collection was completed with the publication of The Silence of the Lambs (1988), Hannibal (1999) and Hannibal: The Origin of Evil (2006), until now its last sequel. A source of cinematographic inspirations, the saga began with the Oscar-winning The Silence of the Lambs (1991) by Jonathan Demme, followed by Hannibal (2001) by Ridley Scott, The Red Dragon (2002) by Brett Rainer, and Hannibal: The Origin of Evil (2007) by Peter Webber, although there is also a first adaptation of the novel The Red Dragon titled Manhunter (1986) by Michael Mann, where FBI agents confront the fearsome serial killer Dr. Hannibal Lecktor (Bryan Cox). Likewise, the American writer and producer Bryan Fuller created the television series Hannibal (2013-2015), 39 episodes in which the danish actor Mads Mikkelsen embodies the perverse cannibal. Regardless of its anthropological, moral and cultural connotations, most cannibalism behaviors are related to psychotic disorders, such as paranoid schyzophrenia, when this patients kill an d eat part of the bodies of their victims in their delusional pictures.(AU)
Subject(s)
Humans , Male , Female , Motion Pictures , Medicine , Mental Disorders , Mental Health , Cannibalism , AnthropologyABSTRACT
The 21kD GTPase Rac is an evolutionarily ancient regulator of cell shape and behavior. Rac2 is predominantly expressed in hematopoietic cells where it is essential for survival and motility. The hyperactivating mutation Rac2E62K also causes human immunodeficiency, although the mechanism remains unexplained. Here, we report that in Drosophila, hyperactivating Rac stimulates ovarian cells to cannibalize neighboring cells, destroying the tissue. We then show that hyperactive Rac2E62K stimulates human HL60-derived macrophage-like cells to engulf and kill living T cell leukemia cells. Primary mouse Rac2+/E62K bone-marrow-derived macrophages also cannibalize primary Rac2+/E62K T cells due to a combination of macrophage hyperactivity and T cell hypersensitivity to engulfment. Additionally, Rac2+/E62K macrophages non-autonomously stimulate wild-type macrophages to engulf T cells. Rac2E62K also enhances engulfment of target cancer cells by chimeric antigen receptor-expressing macrophages (CAR-M) in a CAR-dependent manner. We propose that Rac-mediated cell cannibalism may contribute to Rac2+/E62K human immunodeficiency and enhance CAR-M cancer immunotherapy.
Subject(s)
Immunologic Deficiency Syndromes , Neoplasms , Receptors, Chimeric Antigen , Animals , Mice , Humans , rac GTP-Binding Proteins/genetics , rac GTP-Binding Proteins/metabolism , rac1 GTP-Binding Protein/metabolism , Cannibalism , Macrophages/metabolism , Immunologic Deficiency Syndromes/genetics , Cell DeathABSTRACT
As an abundant element in the Earth's crust, sodium plays an unusual role in food webs. Its availability in terrestrial environments is highly variable, but it is nonessential for most plants, yet essential for animals and most decomposers. Accordingly, sodium requirements are important drivers of various animal behavioural patterns and performance levels. To specifically test whether sodium limitation increases cannibalism in a gregarious lepidopteran herbivore, we hydroponically manipulated Helianthus annuus host plants' tissue-sodium concentrations. Gregarious larvae of the bordered patch butterfly, Chlosyne lacinia, cannibalized siblings when plant-tissue sodium concentrations were low in two separate experiments. Although cannibalism was almost non-existent when sodium concentrations were high, individual mortality rates were also high. Sodium concentration in host plants can have pronounced effects on herbivore behaviour, individual-level performance, and population demographics, all of which are important for understanding the ecology and evolution of plant-animal interactions across a heterogeneous phytochemical landscape.
Subject(s)
Butterflies , Cannibalism , Animals , Herbivory , Larva , Sodium , PlantsABSTRACT
The care-kill response determines whether a sick individual will be treated or eliminated from an insect society, but little is known about the physiological underpinnings of this process. We exploited the stepwise infection dynamics of an entomopathogenic fungus in a termite to explore how care-kill transitions occur, and identify the chemical cues behind these shifts. We found collective responses towards pathogen-injected individuals to vary according to severity and timing of pathogen challenge, with elimination, via cannibalism, occurring sooner in response to a severe active infection. However, injection with inactivated fungal blastospores also resulted in increased albeit delayed cannibalism, even though it did not universally cause host death. This indicates that the decision to eliminate an individual is triggered before pathogen viability or terminal disease status has been established. We then compared the surface chemistry of differently challenged individuals, finding increased amounts of long-chained methyl-branched alkanes with similar branching patterns in individuals injected with both dead and viable fungal blastospores, with the latter showing the largest increase. This coincided with the highest amounts of observed cannibalism as well as signs of severe moribundity. Our study provides new mechanistic insight into the emergent collective behaviors involved in the disease defense of a termite society.
Subject(s)
Isoptera , Humans , Animals , Isoptera/physiology , CannibalismSubject(s)
Animals, Wild , Cannibalism , Animals , Dogs , Animals, Wild/genetics , DNA/genetics , Forensic MedicineABSTRACT
A 6-years and 6-months-old, neutered male mongrel dog had a skin mass between the left eye and nose. Fine needle aspiration biopsy of the mass revealed giant epithelial cells containing many neutrophils in the cytoplasm. The mass was excised and histopathologically diagnosed as an infundibular cyst with giant epithelial-like cells containing neutrophils in part of the lumen. The giant cells were immunopositive for cytokeratin and immunonegative for vimentin. Cell cannibalism is a cell engulfing phenomenon in which a cell incorporates either the same or a different type of cell and the incorporated cells are degenerate or necrotic. In this case, epithelial cells in the lumen of the cyst incorporated neutrophils, which corresponds to neutrophil cannibalism by squamous epithelial cells.
Subject(s)
Cysts , Dog Diseases , Male , Dogs , Animals , Neutrophils , Cannibalism , Epithelial Cells/pathology , Cysts/veterinary , Dog Diseases/pathologyABSTRACT
Feed quality influences insect cannibalistic behavior and gut microbial communities. In the present study, Spodoptera exigua larvae were fed six different artificial diets, and one of these diets (Diet 3) delayed larval cannibalistic behavior and reduced the cannibalism ratio after ingestion. Diet 3-fed larvae had the highest gut bacterial load (1.396 ± 0.556 × 1014 bacteria/mg gut), whereas Diet 2-fed larvae had the lowest gut bacterial load (3.076 ± 1.368 × 1012 bacteria/mg gut). The gut bacterial composition and diversity of different diet-fed S. exigua larvae varied according to the 16S rRNA gene sequence analysis. Enterobacteriaceae was specific to the Diet 3-fed larval gut. Fifteen culturable bacterial isolates were obtained from the midgut of Diet 3-fed larvae. Of these, ten belonged to Escherichia sp. After administration with Diet 1- or 2-fed S. exigua larvae, two bacterial isolates (SePC-12 and -37) delayed cannibalistic behavior in both tested larval groups. Diet 2-fed larvae had the lowest Juvenile hormone (JH) concentration and were more aggressive against intraspecific predation. However, SePC-12 loading increased the JH hormone levels in Diet 2-fed larvae and inhibited their cannibalism. Bacteria in the larval midgut are involved in the stabilization of JH levels, thereby regulating host larval cannibalistic behavior.
Subject(s)
Cannibalism , Escherichia , Animals , Spodoptera/genetics , Larva/physiology , RNA, Ribosomal, 16S/genetics , BacteriaABSTRACT
Biological control using natural enemies with additional food resources is one of the most adopted and ecofriendly pest control techniques. Moreover, additional food is also provided to natural enemies to divert them from cannibalism. In the present work, using the theory of dynamical system, we discuss the dynamics of a cannibalistic predator prey model in the presence of different harvesting schemes in prey (pest) population and provision of additional food to predators (natural enemies). A detailed mathematical analysis and numerical evaluations have been presented to discuss the pest free state, coexistence of species, stability, occurrence of different bifurcations (saddle-node, transcritical, Hopf, Bogdanov-Takens) and the impact of additional food and harvesting schemes on the dynamics of the system. It has been obtained that the multiple coexisting equilibria and their stability depend on the additional food (quality and quantity) and harvesting rates. Interestingly, we also observe that the pest population density decreases immediately even when small amount of harvesting is implemented. Also the eradication of pest population (stable pest free state) could be achieved via variation in the additional food and implemented harvesting schemes. The individual effects of harvesting parameters on the pest density suggest that the linear harvesting scheme is more effective to control the pest population rather than constant and nonlinear harvesting schemes. In the context of biological control programs, the present theoretical work suggests different threshold values of implemented harvesting and appropriate choices of additional food to be supplied for pest eradication.
Subject(s)
Cannibalism , Population DensityABSTRACT
Under high-density culture, cannibalism occurs frequently during the molting of the Chinese mitten crabs Eriocheir sinensis, resulting in a large reduction in production. We found that the leakage of molting fluid from sexually immature crabs informs conspecifics that they are in a molting process. This hypothesis was verified through metabolomics analyses combined with behavioral experiments. The GlcNAc-6-P was identified as a molting biomarker from the differential metabolites by non-targeted metabolomics. In addition, we found that the concentration of GlcNAc-6-P in the molting fluid was significantly higher than other molting metabolites at different molting stages, reaching 5.84 µmol L-1, indicating that the molting fluid was the source of GlcNAc-6-P. Moreover, the behavioral experiments showed that crabs were actively approached to high concentrations of GlcNAc-6-P (1 µmol L-1), but had no obvious choice tendency at different concentrations of UTP, 20-HE and low concentrations of GlcNAc-6-P (0.1 µmol L-1, 0.01 µmol L-1) compared with the control groups. In conclusion, that E. sinensis by sensing the concentration change of GlcNAc-6-P can locate the source of GlcNAc-6-P release and actively approach the high concentration GlcNAc-6-P area and attack the molting crab, causing cannibalism. Blocking the reception pathway of molting chemical cues in E. sinensis, thereby preventing the perception of signals originating from conspecifics' molting in the vicinity, could lead to a reduction in cannibalistic behavior and an increase in overall production. Additionally, this method presents a prospective solution for addressing cannibalism in other crustacean species where such behavior is prevalent.
Subject(s)
Brachyura , Water Pollutants, Chemical , Animals , Molting , Cues , Acetylglucosamine , Cannibalism , Prospective Studies , Water Pollutants, Chemical/toxicity , PhosphatesABSTRACT
Adopting cannibalism substantially affects individual fitness, and recognizing the presence of other cannibals provides additional benefits such as the opportunity to prepare for hunting or defense. This recognition can be facilitated by perceiving conspecific chemical cues. Their role in cannibalistic interactions is less studied than in interspecific predation and it is unclear whether these cues inform individuals of danger or of food availability. Interpretation of these cues is crucial to balance the costs and benefits of anti-predator and feeding strategies, which can directly influence individual fitness. In this study we aimed to test whether damselfly larvae shift towards bolder and more exploratory (cannibalistic) behavior, or become more careful to avoid potential cannibals (as prey) in response to such cues. We conducted behavioral and respiratory experiments with Ischnura elegans larvae to investigate their response to chemical cues from older and larger conspecific larvae. We found that I. elegans larvae decrease their activity and shift their respiratory-related behavior, indicating activation of anti-predator defense mechanisms in response to conspecific chemical cues. Our findings indicate that individuals exposed to conspecific chemical cues balance catching prey with staying safe.
Subject(s)
Cannibalism , Cues , Humans , Animals , Exercise , Food , LarvaABSTRACT
AbstractIn cannibalistic species, selection to avoid conspecifics may stem from the need to avoid being eaten or to avoid competition. Individuals may thus use conspecific cues to modulate their behavior to such threats. Yet the nature of variation for such cues remains elusive. Here, we use a half-sib/full-sib design to evaluate the contribution of (indirect) genetic or environmental effects to the behavioral response of the cannibalistic wolf spider Lycosa fasciiventris (Dufour, 1835) toward conspecific cues. Spiders showed variation in relative occupancy time, activity, and velocity on patches with or without conspecific cues, but direct genetic variance was found only for occupancy time. These three traits were correlated and could be lumped in a principal component: spiders spending more time in patches with conspecific cues moved less and more slowly in those areas. Genetic and/or environmental components of carapace width and weight loss in the social partner, which may reflect the quality and/or quantity of cues produced, were significantly correlated with this principal component, with larger partners causing focal individuals to move more slowly. Therefore, environmental and genetic trait variation in social partners may maintain trait diversity in focal individuals, even in the absence of direct genetic variation.