Predators select for defensive adaptations, such as stings, toxins, and camouflage color patterns. Madtoms, Noturus, are diminutive catfishes with dorsal and pectoral stings. Thirteen of the 25 nominal species have serrated spines in the pectoral sting and a contrasting pigment pattern. Behavior of two saddled species, N. miurus and N. hildebrandi, and one uniformly colored species, N. leptacanthus, was investigated to test if the pigment pattern is camouflage. Saddle spacing and crypticity of the saddled species were measured against various substrates and were found to be unevenly spaced, which could be camouflage when viewed against gravel. Given substrate choices, madtoms preferred gravel during daylight conditions. In subsequent experiments, all species were given colored gravel to test color vs. texture-based substrate choice and preferred dark substrates. In the presence of a predator stimulus, madtoms preferred gravel at night and dawn. The pigment pattern likely is camouflage when viewed against gravel substrates.

Velvet ants are a group of parasitic wasps that are well known for a suite of defensive adaptations including bright coloration and a formidable sting. While these adaptations are presumed to function in antipredator defense, observations between potential predators and this group are lacking. We conducted a series of experiments to determine the risk of velvet ants to a host of potential predators including amphibians, reptiles, birds, and small mammals. Velvet ants from across the United States were tested with predator's representative of the velvet ants native range. All interactions between lizards, free-ranging birds, and a mole resulted in the velvet ants survival, and ultimate avoidance by the predator. Two shrews did injure a velvet ant, but this occurred only after multiple failed attacks. The only predator to successfully consume a velvet ant was a single American toad (Anaxyrus americanus). These results indicate that the suite of defenses possessed by velvet ants, including aposematic coloration, stridulations, a chemical alarm signal, a hard exoskeleton, and powerful sting are effective defenses against potential predators. Female velvet ants appear to be nearly impervious to predation by many species whose diet is heavily derived of invertebrate prey.

Many organisms utilize toxic or noxious compounds as a means of deterring predation. Eastern newts (Notopthalmus viridescens), along with other species in the family Salamandridae, possess a potent neurotoxin called tetrodotoxin (TTX). Although TTX can serve as an effective antipredator mechanism in species of newts with high concentrations (e.g., Taricha), eastern newts have relatively low levels of toxicity in comparison to those species, and it may not serve as an effective antipredator mechanism against all threats. In this case, they may benefit rather by utilizing behavioral changes to avoid initial contact with predators. We tested for predator-avoidance behavior in newts by exposing individuals to kairomones from various predators. We recorded activity patterns of newts when they were exposed to cues from potential predators including bullfrogs (Lithobates catesbeiana), water snakes (Nerodia sipedon) and snapping turtles (Chelydra serpentina), as well as a non-predator (bullfrog tadpoles), and a control (deionized water). Newts reduced activity when exposed to snapping turtle stimuli, but did not change activity when exposed to any other chemical cues. We verified that newts interact with this predator by trapping snapping turtles found in ponds from which newts were collected. Finally, we used turtles caught during this sampling to test whether they are an actual predator of newts and whether newts shift microhabitat use when exposed to this predator. In each replicate, turtles consumed newts, and newts spatially avoided the snapping turtle, relative to a control. The results of these experiments indicate newts rely on predator-avoidance behavior to reduce the probability of being consumed by snapping turtles, but do not reduce activity in response to other potential predators that may only consume them rarely.

The principle of competitive exclusion holds that the strongest competitor for a single resource can exclude other species. Yet in many systems, more similar species appear to stably coexist than the small number of limiting resources. Understanding how and when similar species can stably coexist has taken on new urgency in managing biological invasions and their ecological impacts. Recent theoretical advances emphasize the importance of predators in determining coexistence. The effects of predators, however, can be mediated by behavioral changes induced in their prey as well as by their lethality. In this dissertation, I ask how considering multiple trophic levels changes our understanding of how a grass invasion (Pennisetum ciliare) affects species diversity and dynamics in southeastern Arizona. In considering interactions with plant consumers, and with the predators of those consumers, this research reveals more general ecological processes that determine species diversity across biological communities. I first present evidence from a grass removal experiment in the field that shows increased emergence and short-term survival of native perennial plants without grass. This is consistent with Pennisetum ciliare causing the observed concurrent decline in native plant abundance following invasion. I then present results from greenhouse and field studies consistent with that suppression of native plants being driven primarily through resource competition rather than increased rodent granivory. Granivorous rodents do not solely function as consumers, however, because they cache their harvested seeds in shallow scatter-hoards, from which seeds can germinate. Rodents thus act also as seed dispersers in a context-dependent mutualism. The primary granivores in areas invaded by Pennisetum ciliare are pocket mice (genus Chaetodipus), which have a well-studied tendency to concentrate their activity under plant cover to avoid predation by owls. Because the dense canopy of the grass may provide safer refuge, I hypothesized the pocket mice may be directly dispersing native seeds closer to the base of the invasive grass. Such a behavior could increase the competitive effect of the grass on native plant species, further driving the impacts of the invasion. By offering experimental seeds dusted in fluorescent powder and tracking where the seeds were cached, I show that rodents do preferentially cache experimental seeds under the grass. This dispersal interaction may be more general to plant interactions with seed-caching rodents across semi-arid regions that are experiencing plant invasions. Finally, I ask how the predator avoidance behavior exhibited by these rodents affects their ability to coexist with one another. Not only could their diversity affect that of the plant community, but the effects of plant invasions can cascade through other trophic levels. Theoretical understanding of how similar predator avoidance strategy alters coexistence had not yet been developed, however. Instead of a field study, therefore, I modified a general consumer-resource model with three trophic levels to ask whether avoidance behavior by the middle trophic level alters the ability of those species to coexist. I found that more effective avoidance behavior, or greater safety for less cost, increased the importance of resource partitioning in determining overall niche overlap. Lowering niche overlap between two species promotes their coexistence in the sense that their average fitness can be more different and still permit coexistence. These results provide novel understanding of behavioral modifications to population dynamics in multi-trophic coexistence theory applicable to this invasion and more broadly.

Understanding the ways in which animals utilize space and obtain food are central themes in modern ecology. Formulating broad principles and elucidating the factors explaining such patterns are limited, however, by the availability of data from a broad range of species and systems. This problem especially true of snakes, a predator group about which even the most basic natural history data are often entirely lacking, even among abundant, widespread, and ecologically-important species. I studied the natural history and ecosystem role of one such species, the fer-de-lance (Bothrops asper) in lowland rainforest in Costa Rica. B. asper is a large, cryptic pitviper that is highly abundant in many Central American ecosystems and is strongly relevant to human health due to high incidence of snakebite, yet its biology under natural conditions is almost entirely undocumented. I used radiotelemetry to quantify home range, movement patterns, habitat usage, and foraging behavior. B. asper was found to have smaller home ranges and reduced movement patterns than similarly-sized temperate pitvipers, likely due to a greater reliance upon ambush foraging in patches of high prey density. Snakes also demonstrated strong selection for swamp habitat, which may reflect efforts to exploit frogs as a primary food source due to low availability of small mammals at the study site. I subsequently addressed the trophic status of this B. asper population using a supplemental-feeding experiment. In comparison to control snakes, individuals receiving supplemental food had smaller home ranges, shorter and less frequent movements, increased mass acquisition, and shifted to primarily forest rather than swamp habitat. These results support the suggestion that B. asper at the study site are strongly food-limited. Finally, I tested the hypothesis that fer-de-lance mediate local seed-predation rates by influencing habitat usage and foraging behavior of rodents. A series of behavioral experiments conflicted with many existing studies in failing to support this idea, as three rodent species demonstrated little snake avoidance, and none of likely ecological relevance. Collectively, this dissertation represents the first comprehensive field study of Bothrops asper and is among the first for any tropical snake, and suggests several avenues for future research.

The introduction of pen-reared Attwater's prairie-chickens (APC, Tympanuchus cupido attwateri) into the wild to supplement existing populations has met with marginal success. Flight characteristics, predator avoidance behavior, and rearing methods are possible factors contributing to post-release mortality of pen-reared birds. To evaluate flight characteristics and predator avoidance behavior of pen-reared APC's released onto the Attwater Prairie Chicken National Wildlife Refuge, flight characteristics and predator avoidance behavior of pen-reared APC's was compared to wild greater prairie-chickens (GPC, T. c. pinnatus) in Minnesota and Kansas using a radar gun and a trained dog. There was no difference (P = 0.134) in flight speed for pen-reared APC and wild GPC. However, wild GPC had greater (P < 0.001) flight distances than did pen-reared APC. Wild GPC and pen-reared APC that had survived in the wild for at least a year flushed at a greater (P < 0.001) distance from an approaching human than did pen-reared APC that had been released for less than 3 months. A trained dog was able to approach closer (P < 0.001) to APC than GPC before birds flushed, and APC did not fly as far as GPC after being flushed by the dog. Pen-reared APC displayed flight endurance deficiencies and were more approachable by humans and a dog before they flushed when compared to wild GPC, which could explain their increased mortality when released into the wild. To determine if APC chicks could be reared without daily human contact, pelleted food, and water in founts, a greenhouse was used to rear chicks in a semi-natural environment. Planted vegetation and commercial insects provided hiding cover and a food source for the APC chicks. An underground heat source provided chick warmth, and water misters and a sprinkler system simulated dew (a water source for chicks) and rain. The greenhouse provided chicks protection from predators and adverse weather conditions (before they could thermo-regulate) while exposing chicks to natural sunlight, day length, and temperature fluctuations. This technique allowed chicks to be reared in a semi-natural environment which reinforced their natural foraging behavior for food and water, and reinforced their hiding and avoidance behaviors, creating a wilder pen-reared bird.

The introduction of pen-reared Attwater's prairie-chickens (APC, Tympanuchus cupido attwateri) into the wild to supplement existing populations has met with marginal success. Flight characteristics, predator avoidance behavior, and rearing methods are possible factors contributing to post-release mortality of pen-reared birds. To evaluate flight characteristics and predator avoidance behavior of pen-reared APC's released onto the Attwater Prairie Chicken National Wildlife Refuge, flight characteristics and predator avoidance behavior of pen-reared APC's was compared to wild greater prairie-chickens (GPC, T. c. pinnatus) in Minnesota and Kansas using a radar gun and a trained dog. There was no difference (P = 0.134) in flight speed for pen-reared APC and wild GPC. However, wild GPC had greater (P < 0.001) flight distances than did pen-reared APC. Wild GPC and pen-reared APC that had survived in the wild for at least a year flushed at a greater (P < 0.001) distance from an approaching human than did pen-reared APC that had been released for less than 3 months. A trained dog was able to approach closer (P < 0.001) to APC than GPC before birds flushed, and APC did not fly as far as GPC after being flushed by the dog. Pen-reared APC displayed flight endurance deficiencies and were more approachable by humans and a dog before they flushed when compared to wild GPC, which could explain their increased mortality when released into the wild. To determine if APC chicks could be reared without daily human contact, pelleted food, and water in founts, a greenhouse was used to rear chicks in a semi-natural environment. Planted vegetation and commercial insects provided hiding cover and a food source for the APC chicks. An underground heat source provided chick warmth, and water misters and a sprinkler system simulated dew (a water source for chicks) and rain. The greenhouse provided chicks protection from predators and adverse weather conditions (before they could thermo-regulate) while exposing chicks to natural sunlight, day length, and temperature fluctuations. This technique allowed chicks to be reared in a semi-natural environment which reinforced their natural foraging behavior for food and water, and reinforced their hiding and avoidance behaviors, creating a wilder pen-reared bird.

The introduction of nonnative salmonids into Southern Appalachia may pose a threat to resident salamander populations. In recent years, the stocking and encroachment of rainbow trout (Oncorhynchus mykiss) into headwaters where the black-bellied salamander (Desmognathus quadramaculatus) and brook trout (Salvelinus fontinalis) naturally coexist has raised concerns. In aquatic prey, predator-avoidance responses are primarily influenced through the detection of chemical cues released from predators. The objective of this study was to determine how co-occurrence with a predator influences black-bellied salamander predator recognition behavior. To evaluate this, salamander activity metrics (general activity, number of movements, and latency to move) were recorded before and after exposure to either native trout predator cue (brook), introduced trout predator cue (rainbow), or conditioned tap water (control). Larvae were collected from different streams based on their trout predator assemblage with larvae coming from brook, rainbow, rainbow/brook, and no trout stream reaches. Our results show that larvae that co-occur with trout reduced their activity when exposed to brook trout predator cue, but their response to rainbow trout predator cue depended on their previous co-occurrence. Larvae from areas with only brook trout exhibited a weak predator-avoidance when exposed to rainbow trout predator cue. A follow-up test to determine the influence of alarm cue on predator response in these larvae indicated that the alarm cue enhanced the response to the rainbow trout predator.

Intertidal habitats are characterized by dynamic, tidally-driven fluctuations in abiotic and biotic factors. Many of the environmental stressors that vary across the intertidal (e.g., temperature, oxygen, food availability, predation pressure) are strong drivers of metabolic rate in ectotherms. As such, we predicted that there may be pronounced differences in the metabolic and stress physiology of conspecific sessile invertebrates occupying at different relative tidal heights. The common acorn barnacle Balanus glandula represents an ideal model organism in which to investigate the possibility of tidal height-dependent physiological differences, owing to their wide distribution in the intertidal zone and their eurytolerant nature. In the first chapter of my thesis, we investigate the hypothesis that B. glandula anchored in the low intertidal have a greater capacity for anaerobic metabolism than conspecifics in the high intertidal, and that this is due to increased predation pressure during submersion. Further, we explore the temporal and spatial fidelity of certain tidal-height driven trends in lactate dehydrogenase activity previously observed in our lab (i.e., higher LDH activity in low intertidal barnacles; Horn et al., 2021), and attempt to identify environmental variables that drive plasticity in LDH activity. We found that, in general, there were higher densities of B. glandula and gastropod whelk predators in the low intertidal compared to the high intertidal, but follow-up studies in the lab revealed that opercular closure in B. glandula was induced by predator exposure (Acanthinucella spirata) for less than 24h. This time frame for shell closure is unlikely to result in internal hypoxia or enhance capacity for anaerobic metabolism. We were therefore not surprised to find that LDH activity in B. glandula was likewise not affected by predator exposures (48h) carried out in the lab. After failing to find an effect of predators on LDH activity in B. glandula, we attempted to replicate the previous finding that LDH activity was highest in low intertidal populations of B. glandula. We did this at the original location in San Luis Obispo Bay, CA as well as at three novel field sites and across seasons and years. While we did observe variation in LDH activity over time and between sites, we did not consistently observe the same trend in LDH activity whereby low intertidal barnacles had the highest activity. In response to these variable patterns, we attempted to identify what environmental parameters, other than predation, might be responsible for plasticity in LDH activity. Unfortunately, neither temperature nor emersion stress – the two variables we examined – had any significant an effect on LDH activity in B. glandula. These data suggest that there must be multiple, interacting stressors – including tidal position - that influence the anaerobic metabolic capacity of B. glandula. In the second chapter of my thesis, we went on to investigate how the response to thermal stress might differ between populations of B. glandula from different vertical heights in the intertidal zone. To this end, we assessed how aerial temperature stress affected oxygen consumption rates (MO2), superoxide dismutase (SOD) activity, and time to mortality in B. glandula collected from both low and high intertidal positions. We found that barnacles from the low intertidal showed a significant increase in MO2 with higher temperature, while MO2 was unaffected by temperature in B. glandula from the high intertidal. We also observed that SOD activity levels were higher in the high intertidal barnacles compared to the low intertidal barnacles, although neither group was increasing SOD activity under higher temperature. Finally, we observed significantly longer survival times during thermal stress in barnacles from the high intertidal zone (e.g., LT50 = 8.75 h vs 5 h at 33˚C for the high and low barnacles, respectively), although this advantage seemed to be lost with the addition of desiccation stress at these same temperatures. It is evident that life in highest reaches of the intertidal zones is physiologically challenging, and this has resulted in a population of B, glandula barnacles that are less sensitive to and better suited to tolerate temperature extremes than conspecifics in the lowest intertidal regions. Understanding how habitat variation may differentially impact the metabolic and thermal stress physiology of B. glandula is increasingly important as climate change progresses. This is particularly significant considering that organisms in the intertidal already reside within a relatively stressful environment and may be living closer to their thermal tolerance limits than animals from less extreme habitats.

Seed dispersal by caching rodents is a context-dependent mutualism in many systems. Plants benefit when seed remaining in shallow caches germinates before being eaten, often gaining protection from beetles and a favorable microsite in the process. Caching in highly unfavorable microsites, conversely, could undermine the dispersal benefit for the plant. Plant invasions could disrupt dispersal benefits of seed caching by attracting rodents to the protection of a dense invasive canopy which inhibits the establishment of native seedlings beneath it. To determine whether rodents disproportionately cache seed under the dense canopy of an invasive grass in southeastern Arizona, we used nontoxic fluorescent powder and ultraviolet light to locate caches of seed offered to rodents in the field. We fitted a general habitat-use model, which showed that disproportionate use of plant cover by caching rodents (principally Chaetodipus spp.) increased with moonlight. Across all moon phases, when rodents cached under plants, they cached under the invasive grass disproportionately to its relative cover. A greenhouse experiment showed that proximity to the invasive grass reduced the growth and survival of seedlings of a common native tree (Parkinsonia microphylla) whose seeds are dispersed by caching rodents. Biased dispersal of native seed to the base of an invasive grass could magnify the competitive effect of this grass on native plants, further reducing their recruitment and magnifying the effect of the invasion.