Using external morphology as a proxy for stomach size in Hemigrapsus sanguineus.

Stomach morphology can provide insights into an organism's diet. Gut size or length is typically inversely related to diet quality in most taxa, and has been used to assess diet quality in a variety of systems. However, it requires animal sacrifice and time-consuming dissections. Measures of external morphology associated with diet may be a simpler, more cost-effective solution. At the species level, external measures of the progastric region of the carapace in brachyuran crabs can predict stomach size and diet quality, with some suggestion that this approach may also work to examine individual diet preferences and specialization at the individual level; if so, the size of the progastric region could be used to predict trends in diet quality and consumption for individuals, which would streamline diet studies in crabs. Here, we tested whether external progastric region size predicts internal stomach size across latitude and time of year for individuals of the invasive Asian shore crab Hemigrapsus sanguineus. We found that the width of the progastric region increased at a faster rate with body size than stomach width. In addition, the width of the progastric region followed different trends across sites and over time compared to stomach width. Our results therefore suggest that the progastric region may not be used as a proxy for stomach size variation across individuals.

Shift from income breeding to capital breeding with latitude in the invasive Asian shore crab Hemigrapsus sanguineus.

Organisms vary in the timing of energy acquisition and use for reproduction. Thus, breeding strategies exist on a continuum, from capital breeding to income breeding. Capital breeders acquire and store energy for breeding before the start of the reproductive season, while income breeders finance reproduction using energy acquired during the reproductive season. Latitude and its associated environmental drivers are expected to heavily influence breeding strategy, potentially leading to latitudinal variation in breeding strategies within a single species. We examined the breeding strategy of the Asian shore crab Hemigrapsus sanguineus at five sites spanning nearly 10° of latitude across its invaded United States range. We hypothesized that the primary breeding strategy of this species would shift from income breeding to capital breeding as latitude increases. We found that though this species' breeding strategy is dominated by capital breeding throughout much of the range, income breeding increases in importance at lower latitudes. This latitudinal pattern is likely heavily influenced by the duration of the foraging and breeding seasons, which also vary with latitude. We also found that reproductive characteristics at the northern and southern edges of the invaded range were consistent with continued range expansion. We suggest that the reproductive flexibility of the Asian shore crab is a key facilitator of its continued invasion success. Our results highlight the influence of latitude on the breeding strategy of a species and emphasize the need for further research regarding the ecological importance and implications of flexibility in breeding strategies within species.

Speed and degree of functional and compositional recovery varies with latitude and community age.

Rates at which a community recovers after disturbance, or its resilience, can be accelerated by increased net primary productivity and recolonization dynamics such as recruitment. These mechanisms can vary across biogeographic gradients, such as latitude, suggesting that biogeography is likely important to predicting resilience. To test whether community resilience, informed by functional and compositional recovery, hinges on geographic location, we employed a standardized replicated experiment on marine invertebrate communities across four regions from the tropics to the subarctic zone. Communities assembled naturally on standardized substrate while experiencing distinct levels of biomass removal (no removal, low disturbance, and high disturbance), which opened space for new colonizers, thereby providing a pulse of limited resource to these communities. We then quantified functional (space occupancy and biomass) and compositional recovery from these repeated pulse disturbances across two community assembly timescales (early and late at 3 and 12 months, respectively). We documented latitudinal variation in resilience across 47° latitude, where speed of functional recovery was higher toward lower latitudes yet incomplete at late assembly in the tropics and subtropics. The degree of functional recovery did not coincide with compositional recovery, and regional differences in recruitment and growth likely contributed to functional recovery in these communities. While biogeographic variation in community resilience has been predicted, our results are among the first to examine functional and compositional recovery from disturbance in a single large-scale standardized experiment.

Past energy allocation overwhelms current energy stresses in determining energy allocation trade-offs.

Regeneration of lost appendages is a gradual process in many species, spreading energetic costs of regeneration through time. Energy allocated to the regeneration of lost appendages cannot be used for other purposes and, therefore, commonly elicits energetic trade-offs in biological processes. We used limb loss in the Asian shore crab Hemigrapsus sanguineus to compare the strength of energetic trade-offs resulting from historic limb losses that have been partially regenerated versus current injuries that have not yet been repaired. Consistent with previous studies, we show that limb loss and regeneration results in trade-offs that reduce reproduction, energy storage, and growth. As may be expected, we show that trade-offs in these metrics from historic limb losses far outweigh trade-offs from current limb losses, and correlate directly with the degree of historic limb loss that has been regenerated. As regenerating limbs get closer to their normal size, these historical injuries get harder to detect, despite the continued allocation of additional resources to limb development. Our results demonstrate the importance of and a method for identifying historic appendage losses and of quantifying the amount of regeneration that has already occurred, as opposed to assessing only current injury, to accurately assess the strength of energetic trade-offs in animals recovering from nonlethal injury.

Latitudinal and temporal variation in injury and its impacts in the invasive Asian shore crab Hemigrapsus sanguineus.

Nonlethal injury is a pervasive stress on individual animals that can affect large portions of a population at any given time. Yet most studies examine snapshots of injury at a single place and time, making the implicit assumption that the impacts of nonlethal injury are constant. We sampled Asian shore crabs Hemigrapsus sanguineus throughout their invasive North American range and from the spring through fall of 2020. We then documented the prevalence of limb loss over this space and time. We further examined the impacts of limb loss and limb regeneration on food consumption, growth, reproduction, and energy storage. We show that injury differed substantially across sites and was most common towards the southern part of their invaded range on the East Coast of North America. Injury also varied idiosyncratically across sites and through time. It also had strong impacts on individuals via reduced growth and reproduction, despite increased food consumption in injured crabs. Given the high prevalence of nonlethal injury in this species, these negative impacts of injury on individual animals likely scale up to influence population level processes (e.g., population growth), and may be one factor acting against the widespread success of this invader.

Stronger predation intensity and impact on prey communities in the tropics.

The hypothesis that biotic interactions strengthen toward lower latitudes provides a framework for linking community-scale processes with the macroecological scales that define our biosphere. Despite the importance of this hypothesis for understanding community assembly and ecosystem functioning, the extent to which interaction strength varies across latitude and the effects of this variation on natural communities remain unresolved. Predation in particular is central to ecological and evolutionary dynamics across the globe, yet very few studies explore both community-scale causes and outcomes of predation across latitude. Here we expand beyond prior studies to examine two important components of predation strength: intensity of predation (including multiple dimensions of the predator guild) and impact on prey community biomass and structure, providing one of the most comprehensive examinations of predator-prey interactions across latitude. Using standardized experiments, we tested the hypothesis that predation intensity and impact on prey communities were stronger at lower latitudes. We further assessed prey recruitment to evaluate the potential for this process to mediate predation effects. We used sessile marine invertebrate communities and their fish predators in nearshore environments as a model system, with experiments conducted at 12 sites in four regions spanning the tropics to the subarctic. Our results show clear support for an increase in both predation intensity and impact at lower relative to higher latitudes. The predator guild was more diverse at low latitudes, with higher predation rates, longer interaction durations, and larger predator body sizes, suggesting stronger predation intensity in the tropics. Predation also reduced prey biomass and altered prey composition at low latitudes, with no effects at high latitudes. Although recruitment rates were up to three orders of magnitude higher in the tropics than the subarctic, prey replacement through this process was insufficient to dampen completely the strong impacts of predators in the tropics. Our study provides a novel perspective on the biotic interaction hypothesis, suggesting that multiple components of the predator community likely contribute to predation intensity at low latitudes, with important consequences for the structure of prey communities.

Asymmetry of marine invasions across tropical oceans.

Understanding the mechanisms of spatial variation of biological invasions, across local-to-global scales, has been a major challenge. The importance of evolutionary history for invasion dynamics was noted by Darwin, and several studies have since considered how biodiversity of source and recipient regions can influence the probability of invasions. For over a century, the Panama Canal has connected water bodies and biotas with different evolutionary histories, and created a global shipping hot spot, providing unique opportunities to test mechanisms that affect invasion patterns. Here, we test for asymmetry in both the extent of invasions and predation effects, a possible mechanism of biotic resistance, between two tropical oceans at similar latitudes. We estimated nonnative species (NNS) richness for sessile marine invertebrates, using standardized field surveys and literature synthesis, to examine whether invasions are asymmetrical, with more NNS present in the less diverse Pacific compared to the Atlantic. We also experimentally tested whether predation differentially limits the abundance and distribution of these invertebrates between oceans. In standardized surveys, observed total NNS richness was higher in the Pacific (18 NNS, 30% of all Pacific species) than the Atlantic (11 NNS, 13% of all Atlantic species). Similarly, literature-based records also display this asymmetry between coasts. When considering only the reciprocal exchange of NNS between Atlantic and Pacific biotas, NNS exchange from Atlantic to Pacific was eightfold higher than the opposite direction, exceeding the asymmetry predicted by random exchange based simply on differences of overall diversity per region. Predation substantially reduced biomass and changed NNS composition in the Pacific, but no such effects were detected on the Atlantic coast. Specifically, some dominant NNS were particularly susceptible to predation in the Pacific, supporting the hypothesis that predation may reduce the abundance of certain NNS here. These results are consistent with predictions that high diversity in source regions, and species interactions in recipient regions, shape marine invasion patterns. Our comparisons and experiments across two tropical ocean basins, suggest that global invasion dynamics are likely driven by both ecological and evolutionary factors that shape susceptibility to and directionality of invasions across biogeographic scales.