Olfactory self-recognition in two species of snake.

Mark tests, in which an animal uses a mirror to locate and examine an otherwise unnoticeable mark on its own body, are commonly used to assess self-recognition, which may have implications for self-awareness. Recently, several olfactory-reliant species have appeared to pass odour-based versions of the mark test, though it has never been attempted in reptiles. We conducted an odour-based mark test on two species of snakes, Eastern gartersnakes and ball pythons, with widely divergent ecologies (i.e. terrestrial foragers that communally brumate versus semi-arboreal ambush predators that do not). We find that gartersnakes, but not ball pythons, pass the test, and a range of control tests suggest this is based on self-recognition. Gartersnakes are more social than ball pythons, supporting recent suggestions that social species are more likely to self-recognize. These results open the door to examination of the ecology of self-recognition, and suggest that this ability may evolve in response to species-specific ecological challenges, some of which may align with complexity of social structures.

Bolder together: conformity drives behavioral plasticity in eastern gartersnakes.

Personality traits drive individual differences in behaviour that are consistent across time and context. Personality limits behavioural plasticity, which could lead to maladaptive choices if animals cannot adapt their behavior to changing conditions. Here, we assessed consistency and flexibility in one personality trait, boldness, across non-social and social contexts in eastern gartersnakes (Thamnophis sirtalis sirtalis). Snakes explored a novel open arena either alone or in a pair. Pairs were assigned based on the data from the solo trials, such that each snake was paired once with a bolder and once with a less bold partner. We predicted that snakes would conform when in a social context, displaying plasticity in their personality, and causing boldness scores to converge. We found that snakes were consistent within contexts (solo or paired), but changed their behavior across contexts (from solo to paired). Plasticity in boldness resulted from an interaction between conformity and repeatable individual differences in plasticity. In line with some data on other species, snakes conformed more when they were the less bold partner. Personality reflects a consistent bias in decision-making, but our results highlight that the cognitive processes that drive the expression of personality traits in behavior are flexible and sensitive to social context. We show that both consistency and plasticity combine to shape snake social behavior in ways that are responsive to competition. This pattern of behavior may be particularly beneficial for species in which group-living is seasonal.

Social networks reveal sex- and age-patterned social structure in Butler's gartersnakes (Thamnophis butleri).

Sex- and age-based social structures have been well documented in animals with visible aggregations. However, very little is known about the social structures of snakes. This is most likely because snakes are often considered non-social animals and are particularly difficult to observe in the wild. Here, we show that wild Butler's Gartersnakes have an age and sex assorted social structure similar to more commonly studied social animals. To demonstrate this, we use data from a 12-year capture-mark-recapture study to identify social interactions using social network analyses. We find that the social structures of Butler's Gartersnakes comprise sex- and age-assorted intra-species communities with older females often central and age segregation partially due to patterns of study site use. In addition, we find that females tended to increase in sociability as they aged while the opposite occurred in males. We also present evidence that social interaction may provide fitness benefits, where snakes that were part of a social network were more likely to have improved body condition. We demonstrate that conventional capture data can reveal valuable information on social structures in cryptic species. This is particularly valuable as research has consistently demonstrated that understanding social structure is important for conservation efforts. Additionally, research on the social patterns of animals without obvious social groups provides valuable insight into the evolution of group living.

Socially-mediated activation in the snake social-decision-making network.

Brain areas important for social perception, social reward, and social behavior - collectively referred to as the social-decision-making network (SDN) - appear to be highly conserved across taxa. These brain areas facilitate a variety of social behaviors such as conspecific approach/avoidance, aggression, mating, parental care, and recognition. Although the SDN has been investigated across taxa, little is known about its functioning in reptiles. Research on the snake SDN may provide important new insights, as snakes have a keen social perceptual system and express a relatively reduced repertoire of social behaviors. Here, we present the results of an experiment in which ball pythons (Python regius) interacted with a same-sex conspecific for one hour and neural activation was investigated through Fos immunoreactivity. Compared to controls, snakes that interacted socially had higher Fos counts in brain areas implicated in social behavior across taxa, such as the medial amygdala, preoptic area, nucleus accumbens, and basolateral amygdala. Additionally, we found differential Fos immunoreactivity in the ventral amygdala, which facilitates communication between social brain areas. In many of these areas, Fos counts differed by sex, which may be due to increased competition between males. Fos counts did not differ in early sensory (i.e., vomeronasal) processing structures. As ball python social systems lack parental care, cooperation, or long-term group living, these results provide valuable insight into the basal functions of the vertebrate social decision-making network.

An egg-sabotaging mechanism drives non-Mendelian transmission in mice.

Selfish genetic elements drive in meiosis to distort their transmission ratio and increase their representation in gametes, violating Mendel's law of segregation. The two established paradigms for meiotic drive, gamete killing and biased segregation, are fundamentally different. In gamete killing, typically observed with male meiosis, selfish elements sabotage gametes that do not contain them. By contrast, killing is predetermined in female meiosis, and selfish elements bias their segregation to the single surviving gamete (i.e., the egg in animal meiosis). Here, we show that a selfish element on mouse chromosome 2, Responder to drive 2 (R2d2), drives using a hybrid mechanism in female meiosis, incorporating elements of both killing and biased segregation. We propose that if R2d2 is destined for the polar body, it manipulates segregation to sabotage the egg by causing aneuploidy, which is subsequently lethal in the embryo, ensuring that surviving progeny preferentially contain R2d2. In heterozygous females, R2d2 orients randomly on the metaphase spindle but lags during anaphase and preferentially remains in the egg, regardless of its initial orientation. Thus, the egg genotype is either euploid with R2d2 or aneuploid with both homologs of chromosome 2, with only the former generating viable embryos. Consistent with this model, R2d2 heterozygous females produce eggs with increased aneuploidy for chromosome 2, increased embryonic lethality, and increased transmission of R2d2. In contrast to typical gamete killing of sisters produced as daughter cells in a single meiosis, R2d2 prevents production of any viable gametes from meiotic divisions in which it should have been excluded from the egg.

An egg sabotaging mechanism drives non-Mendelian transmission in mice.

During meiosis, homologous chromosomes segregate so that alleles are transmitted equally to haploid gametes, following Mendel's Law of Segregation. However, some selfish genetic elements drive in meiosis to distort the transmission ratio and increase their representation in gametes. The established paradigms for drive are fundamentally different for female vs male meiosis. In male meiosis, selfish elements typically kill gametes that do not contain them. In female meiosis, killing is predetermined, and selfish elements bias their segregation to the single surviving gamete (i.e., the egg in animal meiosis). Here we show that a selfish element on mouse chromosome 2, R2d2, drives using a hybrid mechanism in female meiosis, incorporating elements of both male and female drivers. If R2d2 is destined for the polar body, it manipulates segregation to sabotage the egg by causing aneuploidy that is subsequently lethal in the embryo, so that surviving progeny preferentially contain R2d2. In heterozygous females, R2d2 orients randomly on the metaphase spindle but lags during anaphase and preferentially remains in the egg, regardless of its initial orientation. Thus, the egg genotype is either euploid with R2d2 or aneuploid with both homologs of chromosome 2, with only the former generating viable embryos. Consistent with this model, R2d2 heterozygous females produce eggs with increased aneuploidy for chromosome 2, increased embryonic lethality, and increased transmission of R2d2. In contrast to a male meiotic driver, which kills its sister gametes produced as daughter cells in the same meiosis, R2d2 eliminates "cousins" produced from meioses in which it should have been excluded from the egg.

Western Hognose Snakes (Heterodon nasicus) Prefer Environmental Enrichment.

The environmental enrichment needs of snakes are often disregarded. Using preference testing, we aimed to shed light on the enrichment preferences of a popular pet species, the western hognose snake (Heterodon nasicus). Snakes' enclosures were divided into enriched and standard sides. The enriched half had substrate for burrowing, interactive stimuli, and a large water dish. The standard half had paper towel substrate and a small water dish. Each side also contained a single shelter. We provided belly heat to create a thermal gradient on one side of the cage. Snakes were observed for 6 days, four times daily. We predicted a preference for enriched conditions and, as snakes are ectothermic, a preference for the warmer side. Snakes were additionally given an exploration assay, to explore whether differences in preference for environmental enrichment interact with boldness levels. We found that hognose snakes preferred enrichment, and the strength of this preference increased over time. Preference for enrichment was stronger when the enriched side was cooler. This may be due to the burrowing tendencies of these snakes. We found no relationship between preference and boldness. These findings emphasise the importance of preference testing in establishing research-informed enrichment opportunities for reptiles.