Behavior and Ethology Commons^™

A Dual Function Of White Coloration In A Nocturnal Spider Dolomedes Raptor (Araneae: Pisauridae), Tai-Shen Lin, Shichang Zhang, Chen-Pan Liao, Eileen A. Hebets, I-Min Tso

Eileen Hebets Publications

Nocturnal animals frequently possess seemingly conspicuous color patterns that can function in a variety of ways (e.g. prey attraction, camouflage, predator avoidance, etc.). The use of color patterns in intraspecific signaling, especially reproductive activities, in nocturnal animals has received relatively little attention. This study tested for a dual function of color in the nocturnal fishing spider, Dolomedes raptor (Araneae: Pisauridae), whose males develop dimorphic white stripes at sexual maturation. We tested for a role in foraging as well as mate assessment. First, quantifications of the natural variation of male stripes indicated a correlation between stripe area and male body size …

Embracing Multiple Definitions Of Learning, Andrew B. Barron, Eileen A. Hebets, Thomas A. Cleland, Courtney L. Fitzpatrick, Mark E. Hauber, Jeffrey R. Stevens

Eileen Hebets Publications

Definitions of learning vary widely across disciplines, driven largely by different approaches used to assess its occurrence. These definitions can be better reconciled with each other if each is recognized as coherent with a common conceptualization of learning, while appreciating the practical utility of different learning definitions in different contexts.

Learning is a major focus of research in psychology, neuro- science, behavioral ecology, evolutionary theory, and computer science, as well as in many other disciplines. Despite its conceptual prevalence, definitions of learning differ enormously both within and between these disciplines, and new definitions continue to be proposed [1]. Ongoing disputes …

Temporal Patterns Of Nutrition Dependence In Secondary Sexual Traits And Their Varying Impacts On Male Mating Success, Malcolm F. Rosenthal, Eileen A. Hebets

Eileen Hebets Publications

Variation in the quantity of nutrients ingested over an individual’s lifetime is likely to differentially affect distinct male secondary sexual traits and courtship signals, potentially providing females with information about a male’s past and present foraging history. We hypothesize that female choice is thus influenced by a male’s lifetime foraging history. To test this, we manipulated the quantity of nutrients (i.e. prey items) available to male wolf spiders, Schizocosa stridulans, using a fully crossed 2 × 2 design with low versus high prey quantity across juvenile and adult life stages, and assessed the impact of these diet treatments on male …

The Complexities Of Female Mate Choice And Male Polymorphisms: Elucidating The Role Of Genetics, Age, And Mate-Choice Copying, Kasey D. Fowler-Finn, Laura Sullivan-Beckers, Amy M. Runck, Eileen A. Hebets

Eileen Hebets Publications

Genetic, life history, and environmental factors dictate patterns of variation in sexual traits within and across populations, and thus the action and outcome of sexual selection. This study explores patterns of inheritance, diet, age, and mate-choice copying on the expression of male sexual signals and associated female mate choice in a phenotypically diverse group of Schizocosa wolf spiders. Focal spiders exhibit one of two male phenotypes: ‘ornamented’ males possess large black brushes on their forelegs, and ‘non-ornamented’ males possess no brushes. Using a quantitative genetics breeding design in a mixed population of ornamented/non-ornamented males, we found a strong genetic basis …

Octopamine Levels Relate To Male Mating Tactic Expression In The Wolf Spider Rabidosa Punctulata, Eileen A. Hebets, Matthew Hansen, Thomas C. Jones, Dustin J. Wilgers

Eileen Hebets Publications

In the wolf spider Rabidosa punctulata, upon encountering a female, males use one of two distinct strategies: (1) they court the female in an attempt to elicit a mating, or (2) they engage in a direct-mount tactic that involves extensive grappling with the female until a mating is achieved. The latter tactic appears more sexually aggressive, and both tactics come with the risk of being cannibalized. We explored the physiological mechanisms underlying this behavioral variation by assessing the relationship between circulating levels of the biogenic amine octopamine (OA), a neuromodulator suggested to play a role in “fight or flight” responses …

Functional Approach To Condition, Dustin J. Wilgers, Eileen A. Hebets

Eileen Hebets Publications

Animal signaling is commonly thought to be costly. Signaling costs can arise via a variety of avenues, including energy expenditure, predator attraction, and so on (reviews in Zuk and Kolluru, 1998; Kotiaho, 2001), and are predicted to increase with signal expression (e.g., size, amplitude, and intensity; Johnstone, 1997). Due to these costs, signaler condition, which is hypothesized to be a reflection of a signaler’s genetic quality, is expected to influence the level of signal expression one can afford (Zahavi, 1975), resulting in a positive correlation between signaler condition and signal expression – that is, condition-dependent signaling (Zahavi, 1977; West-Eberhard, 1979; …

Eight Legged Encounters, Eileen A. Hebets

Eileen Hebets Publications

This program was funded in part by a National Science Foundation grant (DRL–1241482 to EAH). Material was developed in collaboration with Marie- Claire Chelini, Jessie Rose Storz, Cody Storz, and Malcolm Rosenthal. Steven Schwartz, Jason Stafstrom, Kathy French, Priscilla Grew, and Judy Diamond were all extremely helpful in grant writing, facilitating the first live event, and/or discussions. Pawl Tisdale (artist) was phenomenal to work with on all aspects of the project!

TABLE OF CONTENTS

CLASSIFICATION & TAXONOMY STATIONS
I. WHAT IS AN ARTHROPOD? page 4 a. The goal of this station is to introduce the audience to some basic information …

Dangerous Mating Systems: Signal Complexity, Signal Content And Neural Capacity In Spiders, Marie E. Herberstein, Anne E. Wignall, Eileen Hebets, Jutta M. Schneider

Eileen Hebets Publications

Spiders are highly efficient predators in possession of exquisite sensory capacities for ambushing prey, combined with machinery for launching rapid and determined attacks. As a consequence, any sexually motivated approach carries a risk of ending up as prey rather than as a mate. Sexual selection has shaped courtship to effectively communicate the presence, identity, motivation and/or quality of potential mates, which help ameliorate these risks. Spiders communicate this information via several sensory channels, including mechanical (e.g. vibrational), visual and/or chemical, with examples of multimodal signaling beginning to emerge in the literature. The diverse environments that spiders inhabit have further shaped …

Multimodal Sensory Reliance In The Nocturnal Homing Of The Amblypygid Phrynus Pseudoparvulus (Class Arachnida, Order Amblypygi)?, Eileen A. Hebets, Alfonso Aceves-Aparicio, Samuel Aguilar-Argüello, Verner P. Bingman, Ignacio Escalante, Eben J. Gering, David R. Nelson, Jennifer Rivera, José Ángel Sánchez-Ruiz, Laura Segura-Hernández, Virginia Settepani, Daniel D. Wiegmann, Jay A. Stafstrom

Eileen Hebets Publications

Like many other nocturnal arthropods, the amblypygid Phrynus pseudoparvulus is capable of homing. The environment through which these predators navigate is a dense and heterogeneous tropical forest understory and the mechanism(s) underlying their putatively complex navigational abilities are presently unknown. This study explores the sensory inputs that might facilitate nocturnal navigation in the amblypygid P. pseudoparvulus. Specifically, we use sensory system manipulations in conjunction with field displacements to examine the potential involvement of multimodal—olfactory and visual—stimuli in P. pseudoparvulus’ homing behavior. In a first experiment, we deprived individuals of their olfactory capacity and displaced them to the opposite side of …

Obligate Male Death And Sexual Cannibalism In Dark Fishing Spiders, Steven Schwartz, William E. Wagner, Eileen A. Hebets

Eileen Hebets Publications

Male dark fishing spiders (Dolomedes tenebrosus Araneae, Pisauridae) always die during their first and only copulation, making all males monogynous. Such obligate male death can be adaptive if it facilitates sexual cannibalism, and if sexual cannibalism results in male reproductive benefits, such as an advantage in sperm competition through reduced female remating. We first conducted an experiment to determine the extent to which D. tenebrosus (1) males are cannibalized by females and (2) females engage in remating, both of which are prerequisites for several adaptive hypotheses of male self-sacrifice. We then conducted an experiment to test the hypothesis that …

A Male Spider’S Ornamentation Polymorphism Maintained By Opposing Selection With Two Niches, Bo Deng, Alex Estes, Brett Grieb, Douglas Richard, Brittney Hinds, Eileen Hebets

Eileen Hebets Publications

The Levene mechanism to maintain genotypic polymorphism by opposing selection on genotypes in multiple niches was proposed 60 years ago, and yet no systems were found to satisfy the mechanisms rather restrictive conditions. Reported here is such an example that a wolf spider population lives in a habitat of mixed rocks and leafy litter for which the females are phenotypically indistinguishable and the males have two distinct phenotypes subject to opposing selection with respect to the substrates. Census data is best-fitted to a population genetics model of the Levene type. A majority of the best fit support polymorphism, with many …

Tactical Adjustment Of Signaling Leads To Increased Mating Success And Survival, Laura Sullivan-Beckers, Eileen A. Hebets

Eileen Hebets Publications

Most sexually reproducing animals overcome the challenge of searching for and attracting mates by utilizing signals that are broadcast through a spatially and temporally varying environment. A diverse suite of behavioral solutions exist for overcoming such environmental variability, including the adjustment of signaling behavior based upon receiver feedback. Few studies have directly examined the relationship between such tactical signaling adjustments and proxies of male fitness; the few that have, failed to find a relationship. Using the wolf spider, Schizocosa rovneri, we set out to first quantify among-male variation in the form and degree of responsiveness to female feedback. Following …

Nocturnal Homing In The Tropical Amblypygid Phrynus Pseudoparvulus (Class Arachnida, Order Amblypygi), Eileen Hebets, E.J. Gering, Verner P. Bingman, Daniel D. Wiegmann

Eileen Hebets Publications

Arthropods are renowned for their navigational capabilities, with numerous examples known from insects and crustaceans. Early studies of amblypygids (Class Arachnida, Order Amblypygi) also suggest complex nocturnal navigation, despite their apparent lack of visual adaptations to the low-light conditions of a tropical understory. In a series of two studies, we use the tropical amblypygid, Phrynus pseudoparvulus, to assess their nocturnal homing ability. Our first experiment displaced and tracked resident and nonresident individuals. Resident individuals, displaced up to 4.5 m from their home refuges and released onto their home tree, were more likely to return to their previously occupied refuge than …

Detection Of Predator Cues Alters Mating Tactics In Male Wolf Spiders, Dustin J. Wilgers, Daniel Wickwire, Eileen Hebets

Eileen Hebets Publications

Males of the wolf spider, Rabidosa punctulata, exhibit condition-dependent alternative mating tactics, whereby small, poor condition males engage in multimodal courtship while large, good condition males adopt a direct mount tactic that forgoes courtship. This study explores the possibility that tactic-specific costs can help explain this unintuitive pattern of mating tactic expression. Specifically, we hypothesize that courtship signaling is costly with respect to eavesdropping by predators and that males can alter their tactic expression based upon the perceived environmental predation risk. We test this by first examining the risk of predation associated with different mating tactics. We use a …

Female Mate Choice For Multimodal Courtship And The Importance Of The Signaling Background For Selection On Male Ornamentation, Jay A. Stafstrom, Eileen A. Hebets

Eileen Hebets Publications

Conspicuous visual ornaments are frequently incorporated into complex courtship displays that integrate signal components from multiple sensory modalities. Mature male Schizocosa crassipes (Walckenaer, 1837) wolf spiders wave, arch, and tap their ornamented forelegs in a visual courtship display that simultaneously incorporates seismic components. To determine the importance of modality-specific signal components in female mate choice, we used a signal ablation design and compared the mating frequency of female-male pairs across signaling environments with manipulated modality-specific transmission properties. We found that the successful transmission of isolated visual or seismic signaling was sufficient for mating success; neither signaling modality was necessary. Additionally, …

Diversification Under Sexual Selection: The Relative Roles Of Mate Preference Strength And The Degree Of Divergence In Mate Preferences, Rafael L. Rodríguez, Janette W. Boughman, David A. Gray, Eileen A. Hebets, Gerlinde Höbel, Laurel B. Symes

Eileen Hebets Publications

The contribution of sexual selection to diversification remains poorly understood after decades of research. This may be in part because studies have focused predominantly on the strength of sexual selection, which offers an incomplete view of selection regimes. By contrast, students of natural selection focus on environmental differences that help compare selection regimes across populations. To ask how this disparity in focus may affect the conclusions of evolutionary research, we relate the amount of diversification in mating displays to quantitative descriptions of the strength and the amount of divergence in mate preferences across a diverse set of case studies of …

No Evidence For A Relationship Between Hemolymph Ecdysteroid Levels And Female Reproductive Behavior In Schizocosa Wolf Spiders, Reed M. Stubbendieck, Anthony J. Zera, Eileen A. Hebets

Eileen Hebets Publications

This study used radioimmunoassay (RIA) to explore the relationship between levels of hemolymph ecdysteroids and female reproductive behavior in Schizocosa wolf spiders. Specifically, we investigated the relationship between circulating ecdysteroid concentrations in females and 1) likelihood to copulate, or female receptivity [Experiment I— Schizocosa avida (Walckenaer 1837)], 2) time post copulation (Experiment 2— Schizocosa rovneri Uetz & Dondale 1979) and 3) exposure to conspecific male courtship (Experiment 3— Schizocosa uetzi Stratton 1997). In Experiment 1, we expected higher levels of circulating ecdysteroids in receptive versus unreceptive females, based upon prior research demonstrating an increase in receptivity following injections of 20-hydroexyedysone …

Enigmatic Ornamentation Eases Male Reliance On Courtship Performance For Mating Success, Eileen Hebets, Jay A. Stafstrom, Rafael L. Rodriguez, Dustin J. Wilgers

Eileen Hebets Publications

Female preferences are frequently invoked to explain the widespread occurrence of elaborate male ornaments, yet empirical data demonstrating such preferences are sometimes equivocal or even contradictory. In the wolf spider Schizocosa stridulans, despite evidence of strong female choice, prior research has been unable to link the conspicuous sexually dimorphic foreleg ornamentation of males to their mating success. We conducted three experiments aimed at determining the function of this previously enigmatic ornamentation. Our first two experiments used males with phenotypically modified foreleg phenotypes in simple and complex mating environments in order to examine the relationship between the presence/absence of ornamentation …

More Ornamented Males Exhibit Increased Predation Risk And Antipredatory Escapes, But Not Greater Mortality, Kasey D. Fowler-Finn, Eileen Hebets

Eileen Hebets Publications

Secondary sexual traits not only confer benefits to their bearer through increased mate acquisition, but may also have inherent costs, including the attraction of predators. Here, we examined the relationship between conspicuous secondary sexual traits and predation costs using two male morphs of Schizocosa wolf spiders: brushlegged and non-ornamented. In the field, we ran two predation experiments using artificial enclosures to directly test mortality costs of predation on the two male morphs. Using a natural predator, a larger wolf spider in the genus Hogna, we found no difference in predation on brush-legged vs. non-ornamented males. However, predation was depends …

Current Status And Future Directions Of Research In Complex Signaling, Eileen Hebets

Eileen Hebets Publications

The term ‘complex signaling’ reflects a recent scientific focus on the multiple elements frequently incorporated into animal signals (reviewed in Candolin, 2003; Hebets and Papaj, 2005; Partan and Marler, 2005; Bro-Jorgensen, 2010). It embodies a new appreciation that signals used in communication are regularly composed of numerous components and that each component may individually vary in a number of ways. For example, signal components may vary in their time or mechanism of production, in the efficacy of their transmission, and/or in their mechanism of reception, among others. Employing the term ‘complex signaling’ reminds us of the need to maintain a …

Chemical Communication In A Multimodal Context, Eileen A. Hebets, Aaron S. Rundus

Eileen Hebets Publications

All animals are equipped with multiple sensory systems (e.g., visual, chemical, acoustic, tactile, electrical, thermal), and signals perceived via these sensory systems facilitate communication. Such communication often involves displays that incorporate more than one signal from more than one sensory modality, resulting in multimodal signaling. The number of empirical and theoretical studies addressing issues of multimodal signaling is ever-increasing and this chapter highlights why crustaceans, as a taxonomic group, are ideal for advancing such studies. Early classifications of multimodal signaling sought to categorize signal components as either redundant or nonredundant, while more recent classifications lay out specific hypotheses relating to …

Females Are Choosier In The Dark: Environment-Dependent Reliance On Courtship Components And Its Impact On Fitness, Aaron S. Rundus, Laura Sullivan-Beckers, Dustin J. Wilgers, Eileen Hebets

Eileen Hebets Publications

A broad understanding of multimodal courtship function necessitates knowledge of the potential information content of signal components, the efficacy of signal components in eliciting the appropriate receiver response, and the fitness consequences of mating decisions based upon various signal components. We present data addressing each of these requirements for the multimodal-signaling wolf spider, Schizocosa floridana Bryant. Using diet manipulations, we first demonstrate that both visual and seismic courtship signals are condition-dependent. Next, using high- and low-quantity diet individuals in mate choice trials across manipulated signaling environments, we demonstrate that the seismic signal is crucial for mating success and further show …

A Sticky Situation: Solifugids (Arachnida, Solifugae) Use Adhesive Organs On Their Pedipalps For Prey Capture, Rodrigo H. Willemart, Roger D. Santer, Andrew J. Spence, Eileen Hebets

Eileen Hebets Publications

Solifugids (Arachnida, Solifugae) have unique evertable adhesive organs on the tips of their pedipalps, named ‘suctorial’ or ‘palpal’ organs. Previous studies have shown that these organs enable solifugids to climb smooth glass-like surfaces and have hypothesized that these structures facilitate prey capture. Here, we use high-speed videography to demonstrate that the suctorial organs of Eremochelis bilobatus are its primary means of capturing insect prey. We also present calculations of the adhesive pressure exerted by these suctorial organs during real prey capture events.

Complex Courtship Displays Facilitate Male Reproductive Success And Plasticity In Signaling Across Variable Environments, Dustin Wilgers, Eileen Hebets

Eileen Hebets Publications

Effective signal transmission is essential for communication. In environments where signal transmission is highly variable, signalers may utilize complex signals, which incorporate multiple components and modalities, to maintain effective communication. Male Rabidosa rabida wolf spiders produce complex courtship signals, consisting of both visual and seismic components. We test the hypothesis that the complex signaling of R. rabida contributes to male reproductive success in variable signaling environments. We first examine the condition-dependence of foreleg ornamentation (a presumed visual signal) and seismic signal components and find that both may provide potentially redundant information on foraging history. Next, we assessed reproductive success across …

The Degree Of Response To Increased Predation Risk Corresponds To Male Secondary Sexual Traits, Kasey Fowler-Finn, Eileen Hebets

Eileen Hebets Publications

Secondary sexual traits are expected to reflect a balance between sexual selection and natural selection. We test the hypothesis that plasticity in sexual advertisement behaviors can influence this trade-off, allowing showier traits than expected for a given level of predation risk. Specifically, we tested whether the degree of behavioral plasticity exhibited in response to chemical cues of a co-occurring predatory wolf spider corresponds to courtship rate and the degree of ornamentation in male wolf spiders. Both ornamented (brush-legged) males and non-ornamented males decreased locomotion, decreased their likelihood to court, and increased their time to initiate courtship in response to predator …

Female Mate Choice Based Upon Male Motor Performance, John Byers, Eileen Hebets, Jeffrey Podos

Eileen Hebets Publications

Our goal in this essay is to review the hypothesis that females choose mates by the evaluation of male motor performance. We define motor performance as vigor, the ability to perform energetically expensive acts repeatedly, or as skill, the ability to perform difficult motor tasks well. Motor performance reflects most aspects of whole-organism performance that relate to survival, and thus should indicate, more reliably than ornaments do, individual male genetic quality and/or developmental history. Male sexual displays in many animal taxa contain elements of vigor and/or skill, and accumulating evidence suggests that females choose mates in nature based upon their …

A Signal-Substrate Match In The Substrate-Borne Component Of A Multimodal Courtship Display, Damian O. Elias, Andrew C. Mason, Eileen Hebets

Eileen Hebets Publications

The environment can impose strong limitations on the efficacy of signal transmission. In particular, for vibratory communication, the signaling environment is often extremely heterogeneous at small scales. Nevertheless, natural selection is expected to select for signals well-suited for effective transmission. Here, we test for substrate-dependent signal efficacy in the wolf spider Schizocosa stridulans Stratton 1991. We first explore the transmission characteristics of this important signaling modality by playing recorded substrate-borne signals through three different substrates (leaf litter, pine litter, and red clay) and measuring the propagated signal. We found that the substrate-borne signal of S. stridulans attenuates the least on …

Multimodal Courtship Efficacy Of Schizocosa Retrorsa Wolf Spiders: Implications Of An Additional Signal Modality, Aaron S. Rundus, Roger D. Santer, Eileen Hebets

Eileen Hebets Publications

Here, we simultaneously examine both content and efficacy-based sources of selection on the visual and seismic multimodal courtship display of the wolf spider Schizocosa retrorsa. Immature field-collected S. retrorsa were reared in the laboratory on either high-quantity diet (HD) or low-quantity diet (LD) treatments. On maturation, females of each diet treatment were run in simultaneous mate choice trials with both a HD and an LD male (content-based selection). Simultaneous mate choice trials were conducted across different signaling environments (efficacy-based selection) in a fully crossed 2 × 2 design with visual treatments of light/dark (visual signal present/absent) and seismic treatments …

Mate Choice And Learning, Eileen Hebets, Laura Sullivan-Beckers

Eileen Hebets Publications

While an individual’s genetic framework is a major contributor in determining its eventual mate choice, the role of the environment in further influencing mating decisions has long been recognized. Animals gather information from the environment throughout life, and in some cases, may apply this information to increase their odds of obtaining a high-quality mate. In short, these individuals learn. Moreover, such learning can have a social component. “Social learning” is a general term that describes any learning based on observing, interacting with, and/or imitating others in a social context. Social learning can transmit information vertically, generation to generation (e.g., parent …

Prey Capture By The Whip Spider Phrynus Marginemaculatus C.L. Koch, Roger D. Santer, Eileen Hebets

Eileen Hebets Publications

Whip spiders (Arachnida, Amblypygi) are little-studied arachnids with enlarged spiny pedipalps and elongated ‘‘antenniform’’ forelegs. These antenniform legs contain at least seven giant sensory neurons with no known behavioral function. Here we use high-speed cinematography to describe the prey capture behavior of the whip spider Phrynus marginemaculatus C.L. Koch 1840, in order to examine how these giant neurons might be involved. When presented with a prey item (a cricket), a whip spider first accurately aims one of its antenniform legs in the prey’s direction. Next, the whip spider orients its body to the prey item and approaches, placing one antenniform …