Reproductive interference via interspecific pairing in an amphipod species complex

Many species experience reproductive interference defined as interspecific mating interactions that reduce the fitness of individuals in at least one of the species. We examined reproductive interference among three cryptic amphipod species that co-occur in the vegetated zone of lakes. Because these animals form precopulatory pairs and males commonly have indiscriminant mating behavior, we predicted that (1) there would be interspecific pairing, (2) interspecific pairing would be more common when conspecifics were rare, and (3) interspecific pairing would be more common in species combinations where males are larger than females. Using color phenotypes to distinguish species, a novel discovery reported here, we conducted experiments to evaluate patterns of interspecific pairing. In a no-choice experiment (i.e., amphipods had access to mates of only one species), we observed a low rate (5 %) of interspecific pairing and 80 % of these cases involved males that were larger than females. In a second experiment where individuals had access to conspecific mates at varying relative abundance, only 0.04 % of the pairs observed were between heterospecifics, suggesting that the interspecific pairing observed in the no-choice experiment was an artifact of not having access to conspecific mates. Our results suggest that at least one sex has sufficient species-recognition abilities to circumvent the formation of interspecific pairs; therefore, reproductive interference may be minimal in these species.     

An unforeseen chain of events: lethal effects of pesticides on frogs at sublethal concentrations

The field of toxicology has traditionally assessed the risk of contaminants by using laboratory experiments and a range of pesticide concentrations that are held constant for short periods of time (1-4 days). From these experiments, one can estimate the concentration that causes no effect on survival. However, organisms in nature frequently experience multiple, applications of pesticides over time rather than a single constant concentration. In addition, organisms are embedded in ecological communities that can propagate indirect effects through a food web. Using outdoor mesocosms, we examined how low concentrations (10-250 microg/L) of a globally common insecticide (malathion) applied at various amounts, times, and frequencies affected aquatic communities containing zooplankton, phytoplankton, periphyton, and larval amphibians (reared at two densities) for 79 days. All application regimes caused a decline in zooplankton, which initiated a trophic cascade in which there was a bloom in phytoplankton and, in several treatments, a subsequent decline in the competing periphyton. The reduced periphyton had little effect on wood frogs (Rana sylvatica), which have a short time to metamorphosis. However, leopard frogs (Rana pipiens) have a longer time to metamorphosis, and they experienced large reductions in growth and development, which led to subsequent mortality as the environment dried. Hence, malathion (which rapidly breaks down) did not directly kill amphibians, but initiated a trophic cascade that indirectly resulted in substantial amphibian mortality. Importantly, repeated applications of the lowest concentration (a "press treatment" consisting of seven weekly applications of 10 microg/L) caused larger impacts on many of the response variables than single "pulse" applications that were 25 times as great in concentration. These results are not only important because malathion is the most commonly applied insecticide and is found in wetlands, but also because the mechanism underlying the trophic cascade is common to a wide range of insecticides, offering the possibility of general predictions for the way in which many insecticides impact aquatic communities and the populations of larval amphibians.     

"Cryptic" direct benefits of mate choice: choosy females experience reduced predation risk while in precopula

Despite the central role that female mate choice plays in the production of biological diversity, controversy remains concerning its evolution and maintenance. This is particularly true in systems where females are choosy but do not receive obvious direct benefits such as nuptial gifts that increase a female's survival and fecundity. In the absence of such direct benefits, indirect benefits (i.e., the production of superior offspring) are often invoked to explain the evolution of mate choice. However, females may receive less obvious, or "cryptic," direct benefits, particularly in species with prolonged pre-mating interactions (e.g., precopulatory mate guarding). We assessed the “cryptic” direct benefits of female choice for large male size in two species of freshwater amphipods that do not receive obvious direct benefits. Females paired with large males experienced decreased predation from fish. However, we found that the size of a female’s mate did not affect her predation risk against predatory dragonflies or the harassment she received by single males while paired. Our results demonstrate that even when females receive no traditional direct benefits, female choice for large male size can still provide important direct benefits. Such “cryptic” direct benefits may be common, especially in species with prolonged mating interactions, and are likely important for fully understanding the evolution of mate choice.     

Development and Validation of an Aquatic Fine Sediment Biotic Index

The Fine Sediment Biotic Index (FSBI) is a regional, stressor-specific biomonitoring index to assess fine sediment (<2 mm) impacts on macroinvertebrate communities in northwestern US streams. We examined previously collected data of benthic macroinvertebrate assemblages and substrate particle sizes for 1,139 streams spanning 16 western US Level III Ecoregions to determine macroinvertebrate sensitivity (mostly at species level) to fine sediment. We developed FSBI for four ecoregion groupings that include nine of the ecoregions. The grouping were: the Coast (Coast Range ecoregion) (136 streams), Northern Mountains (Cascades, N. Rockies, ID Batholith ecoregions) (428 streams), Rockies (Middle Rockies, Southern Rockies ecoregions) (199 streams), and Basin and Plains (Columbia Plateau, Snake River Basin, Northern Basin and Range ecoregions) (262 streams). We excluded rare taxa and taxa identified at coarse taxonomic levels, including Chironomidae. This reduced the 685 taxa from all data sets to 206. Of these 93 exhibited some sensitivity to fine sediment which we classified into four categories: extremely, very, moderately, and slightly sensitive; containing 11, 22, 30, and 30 taxa, respectively. Categories were weighted and a FSBI score calculated by summing the sensitive taxa found in a stream. There were no orders or families that were solely sensitive or resistant to fine sediment. Although, among the three orders commonly regarded as indicators of high water quality, the Plecoptera (5), Trichoptera (3), and Ephemeroptera (2) contained all but one of the species or species groups classified as extremely sensitive. Index validation with an independent data set of 255 streams found FSBI scores to accurately predict both high and low levels of measured fine sediment.     

Differences in sensitivity to the fungal pathogen Batrachochytrium dendrobatidis among amphibian populations

Contributing to the worldwide biodiversity crisis are emerging infectious diseases, which can lead to extirpations and extinctions of hosts. For example, the infectious fungal pathogen Batrachochytrium dendrobatidis (Bd) is associated with worldwide amphibian population declines and extinctions. Sensitivity to Bd varies with species, season, and life stage. However, there is little information on whether sensitivity to Bd differs among populations, which is essential for understanding Bd‐infection dynamics and for formulating conservation strategies. We experimentally investigated intraspecific differences in host sensitivity to Bd across 10 populations of wood frogs (Lithobates sylvaticus) raised from eggs to metamorphosis. We exposed the post‐metamorphic wood frogs to Bd and monitored survival for 30 days under controlled laboratory conditions. Populations differed in overall survival and mortality rate. Infection load also differed among populations but was not correlated with population differences in risk of mortality. Such population‐level variation in sensitivity to Bd may result in reservoir populations that may be a source for the transmission of Bd to other sensitive populations or species. Alternatively, remnant populations that are less sensitive to Bd could serve as sources for recolonization after epidemic events.     

New effects of Roundup on amphibians: predators reduce herbicide mortality; herbicides induce antipredator morphology

The use of pesticides is important for growing crops and protecting human health by reducing the prevalence of targeted pest species. However, less attention is given to the potential unintended effects on nontarget species, including taxonomic groups that are of current conservation concern. One issue raised in recent years is the potential for pesticides to become more lethal in the presence of predatory cues, a phenomenon observed thus far only in the laboratory. A second issue is whether pesticides can induce unintended trait changes in nontarget species, particularly trait changes that might mimic adaptive responses to natural environmental stressors. Using outdoor mesocosms, I created simple wetland communities containing leaf litter, algae, zooplankton, and three species of tadpoles (wood frogs [Rana sylvatica or Lithobates sylvaticus], leopard frogs [R. pipiens or L. pipiens], and American toads [Bufo americanus or Anaxyrus americanus]). I exposed the communities to a factorial combination of environmentally relevant herbicide concentrations (0, 1, 2, or 3 mg acid equivalents [a.e.]/L of Roundup Original MAX) crossed with three predator-cue treatments (no predators, adult newts [Notophthalmus viridescens], or larval dragonflies [Anax junius]). Without predator cues, mortality rates from Roundup were consistent with past studies. Combined with cues from the most risky predator (i.e., dragonflies), Roundup became less lethal (in direct contrast to past laboratory studies). This reduction in mortality was likely caused by the herbicide stratifying in the water column and predator cues scaring the tadpoles down to the benthos where herbicide concentrations were lower. Even more striking was the discovery that Roundup induced morphological changes in the tadpoles. In wood frog and leopard frog tadpoles, Roundup induced relatively deeper tails in the same direction and of the same magnitude as the adaptive changes induced by dragonfly cues. To my knowledge, this is the first study to show that a pesticide can induce morphological changes in a vertebrate. Moreover, the data suggest that the herbicide might be activating the tadpoles' developmental pathways used for antipredator responses. Collectively, these discoveries suggest that the world's most widely applied herbicide may have much further-reaching effects on nontarget species than previous considered.     

How flexible is phenotypic plasticity? Developmental windows for trait induction and reversal

Inducible defenses allow prey to modulate their phenotypic responses to the level of predation risk in the environment and reduce the cost of constitutive defenses. Inherent in this statement is that prey must alter their phenotypes during development in order to form these defenses. This has lead many ecologists and evolutionary biologists to call for studies that examine developmental plasticity to provide insights into the importance of development in controlling the trajectories of trait formation, the integration of phenotypes over ontogeny, and the establishment of developmental windows for trait formation and reversal. By moving away from studies that focus on a single point in development, we can obtain a more complete understanding of the phenotypic decisions and limitations of prey. We exposed freshwater snails (Helisoma trivolvis) to environments in which predatory water bugs (Belostoma flumineum) were always absent, always present, or added and removed at different points in development. We discovered that snails formed morphological defenses against water bugs. Importantly, after the initial induction of defenses, snails showed similar developmental trajectories as snails reared without predators. Further, the snails possessed wide developmental windows for inducible defenses that extended past sexual maturity. However, being induced later in development appeared to have an associated cost (i.e., decreased shell thickness) that was not found when water bugs were always present. This epiphenotype (i.e., new shell formation as an extension of the current shell) suggests that resource limitation plays an important role in responses to temporal variation in predation risk and may have critical ecological costs that limit the benefits of the inducible defense. Lastly, the ability of snails to completely reverse their defenses was limited to early in ontogeny due to the constraints associated with modular growth of shell material. In sum, we demonstrate that taking a developmental perspective is extremely valuable for understanding the ecology of inducible defenses.