Multiple maternal risk-management adaptations in the loggerhead sea turtle (Caretta caretta) mitigate clutch failure caused by catastrophic storms and predators.

Maternal risk-management, an extension of r/K selection, is an indispensable tool for understanding the natural selection pressures that shape the evolution of reproduction. Central to the construct of maternal risk-management is its definition of reproductive success as replacement fitness (w = 2), the survival of one breeding daughter to replace the female and one outbreeding son to replace her mate. Here, I apply maternal risk-management as a theoretical framework to explain multiple reproductive adaptations by loggerhead sea turtles nesting on a barrier island off the southern coast of Florida, US, from 1988 to 2004. Extrapolated over a 30-year reproductive span, nesting females averaged 4000-4500 eggs. I show that, rather than "putting all their eggs in one basket," females divided eggs into 40 clutches of variable size (50-165 eggs). To deposit clutches, females migrated to the barrier island 10-12 times at unpredictable intervals of 2-8 years. Each nesting season, females deposited 1-7 clutches over diversified time intervals at diversified locations on the beach. Despite devastating clutch losses caused by ten catastrophic hurricanes, hundreds of erratic thunderstorms and dozens of predation events during this study, 72% of clutches produced by nesting females on this barrier island were undisturbed-median hatching success for these clutches was an astonishing 92%. I conclude that diversified maternal investments over time and space by nesting females are reproductive adaptations that have successfully offset clutch losses, thus enabling populations of loggerhead females to meet or exceed their reproductive goal of replacement fitness.

Extending r/K selection with a maternal risk-management model that classifies animal species into divergent natural selection categories.

Reproduction is a defining process of biological systems. Every generation, across all species, breeding females repopulate ecosystems with offspring. r/K selection was the first theory to classify animal species by linking the rates with which breeding females repopulated ecosystems, to the stability of ecosystems. Here, I introduce a species classification scheme that extends the reach of r-K selection and CSR selection by linking breeder investments in offspring quantity, quality, and diversity to specific natural selection pressures. The species classification scheme is predicated on the assumption that high rates of predation favor breeders that invest more in offspring quantity than quality; and that spatiotemporal scarcity favors breeders that investment more in offspring quality than quantity. I present equations that convert the species classification scheme into a maternal risk-management model. Thereafter, using the equations, I classify eighty-seven animal species into the model's natural selection categories. Species of reptiles, fish, and marine invertebrates clustered in the predation selection category. Species of birds and mammals clustered in the scarcity selection category. Several species of apex predators clustered in the weak selection category. Several species of social insects and social mammals clustered in the convergent selection category. In summary, by acknowledging breeding females as the individuals upon which natural selection acts to repopulate ecosystems with offspring, the proposed maternal risk-management model offers a testable, theoretical framework for the field of ecology.

Virus discovery in all three major lineages of terrestrial arthropods highlights the diversity of single-stranded DNA viruses associated with invertebrates.

Viruses encoding a replication-associated protein (Rep) within a covalently closed, single-stranded (ss)DNA genome are among the smallest viruses known to infect eukaryotic organisms, including economically valuable agricultural crops and livestock. Although circular Rep-encoding ssDNA (CRESS DNA) viruses are a widespread group for which our knowledge is rapidly expanding, biased sampling toward vertebrates and land plants has limited our understanding of their diversity and evolution. Here, we screened terrestrial arthropods for CRESS DNA viruses and report the identification of 44 viral genomes and replicons associated with specimens representing all three major terrestrial arthropod lineages, namely Euchelicerata (spiders), Hexapoda (insects), and Myriapoda (millipedes). We identified virus genomes belonging to three established CRESS DNA viral families (Circoviridae, Genomoviridae, and Smacoviridae); however, over half of the arthropod-associated viral genomes are only distantly related to currently classified CRESS DNA viral sequences. Although members of viral and satellite families known to infect plants (Geminiviridae, Nanoviridae, Alphasatellitidae) were not identified in this study, these plant-infecting CRESS DNA viruses and replicons are transmitted by hemipterans. Therefore, members from six out of the seven established CRESS DNA viral families circulate among arthropods. Furthermore, a phylogenetic analysis of Reps, including endogenous viral sequences, reported to date from a wide array of organisms revealed that most of the known CRESS DNA viral diversity circulates among invertebrates. Our results highlight the vast and unexplored diversity of CRESS DNA viruses among invertebrates and parallel findings from RNA viral discovery efforts in undersampled taxa.

Modeling temperature-mediated fluctuation in colony size in the fire ant, Solenopsis invicta.

In the social insects, colony size is central to the survival of the queen. Two endogenous factors, worker longevity and queen's daily egg production, are known to determine maximum colony size. A third endogenous factor, duration of worker development from egg to adult, regulates the rate of colony growth. In this paper, we report findings from a simulation quantifying the effects of temperature on colony size in the fire ant, Solenopsis invicta. The monthly average temperature over a six year period for the panhandle of north Florida was interpolated to determine the effects of daily temperature on a queen's egg production, worker developmental time and worker longevity. Additional daily temperatures were simulated: 7°C higher and 7°C lower than daily temperatures for north Florida. As expected, colony size was the largest when annual temperatures were the highest across seasons, ranging from 57,000 to 187,000. Colony size at intermediate daily temperatures ranged from 14,000 to 103,000; small colonies recovered rapidly as temperatures warmed. Colony size at lower daily temperatures ranged from 14,000 to 21,000. Extended worker longevity at lower temperatures compensated for low egg production and longer developmental time. And vice versa, the queen's high rate of egg production and the shorter developmental time compensated for shorter worker longevity at high temperatures. Because the fire ant nest consists of a heat-collecting dome in which to incubate brood during cold weather, and deep chambers in which to cool workers during hot weather, colony size is likely to be higher and more stable than our simulation showed. The extended longevity of workers and queens at low temperatures, and perhaps their ability to hibernate below the permafrost, might explain the ability of ants to colonize habitats worldwide.

Young fire ant workers feign death and survive aggressive neighbors.

Feigning death is a method of self-defense employed among a wide range of prey species when threatened by predator species. This paper reports on death-feigning behavior by the fire ant, Solenopsis invicta, during intraspecific aggression among neighboring fire ant workers. Days-old workers responded to aggression by death feigning, weeks-old workers responded by fleeing and months-old workers responded by fighting back. By feigning death, days-old workers were four times more likely to survive aggression than older workers. From a proximate perspective, retaliation by young workers against aggressive older workers is certain to fail. With their relatively soft exoskeleton, young workers would be prone to injury and death and unable to execute an effective attack of biting or stinging older workers with harder exoskeletons. From an ultimate perspective, death feigning allows young workers to survive and contribute to brood care and colony growth, both of which are essential to queen survival and fitness.