Conspicuous stripes on prey capture attention and reduce attacks by foraging jumping spiders.

Many animals avoid predation using aposematic displays that pair toxic/dangerous defences with conspicuous achromatic warning patterns, such as high-contrast stripes. To understand how these prey defences work, we need to understand the decision-making of visual predators. Here we gave two species of jumping spiders (Phidippus regius and Habronattus trimaculatus) choice tests using live termites that had their back patterns manipulated using paper capes (solid white, solid black, striped). For P. regius, black and striped termites were quicker to capture attention. Yet despite this increased attention, striped termites were attacked at lower rates than either white or black. This suggests that the termite's contrast with the background elicits attention, but the internal striped body patterning reduces attacks. Results from tests with H. trimaculatus were qualitatively similar but did not meet the threshold for statistical significance. Additional exploratory analyses suggest that attention to and aversion to stripes is at least partially innate and provide further insight into how decision-making played out during trials. Because of their rich diversity (over 6500 species) that includes variation in natural history, toxin susceptibility and degree of colour vision, jumping spiders are well suited to test broad generalizations about how and why aposematic displays work.

Extreme natural size variation in both sexes of a sexually cannibalistic mantidfly.

In sexually cannibalistic animals, the relative sizes of potential mates often predict the outcome of aggressive encounters. Mantidflies are spider egg predators as larvae and generalist predators as adults. Unlike most cannibalistic species, there is considerable individual variation in body size in both sexes. Using preserved collections of Dicromantispa sayi, we focused on three body size metrics that we found to be positively correlated and accurately measured across researchers. We found extreme size variation in both sexes: the largest 10% of females were 1.72× larger than the smallest 10%, and the largest 10% of males were 1.65× larger than the smallest 10%. On average, females were 7.94% larger than males. In exploring possible causes of this variation, we uncovered differences among populations. To explore the effect of spider egg sac size on adult mantidfly size, we reared mantidfly larvae on egg sacs from two jumping spider species with small or large egg sacs. Mantidfly larvae reared on small egg sacs were smaller than those reared on large egg sacs. This study provides the groundwork to design ecologically relevant experiments exploring the causes and consequences of extreme size variation in an understudied system with intriguing natural history.

Lack of neophobic responses to color in a jumping spider that uses color cues when foraging (Habronattus pyrrithrix).

Chemically defended prey often advertise their toxins with bright and conspicuous colors. To understand why such colors are effective at reducing predation, we need to understand the psychology of key predators. In bird predators, there is evidence that individuals avoid novelty-including prey of novel colors (with which they have had no prior experience). Moreover, the effect of novelty is sometimes strongest for colors that are typically associated with aposematic prey (e.g., red, orange, yellow). Given these findings in the bird literature, color neophobia has been argued to be a driving force in the evolution of aposematism. However, no studies have yet asked whether invertebrate predators respond similarly to novel colors. Here, we tested whether naive lab-raised jumping spiders (Habronattus pyrrithrix) exhibit similar patterns of color neophobia to birds. Using color-manipulated living prey, we first color-exposed spiders to prey of two out of three colors (blue, green, or red), with the third color remaining novel. After this color exposure phase, we gave the spiders tests where they could choose between all three colors (two familiar, one novel). We found that H. pyrrithrix attacked novel and familiar-colored prey at equal rates with no evidence that the degree of neophobia varied by color. Moreover, we found no evidence that either prey novelty nor color (nor their interaction) had an effect on how quickly prey was attacked. We discuss these findings in the context of what is known about color neophobia in other animals and how this contributes to our understanding of aposematic signals.

Sexually dimorphic dorsal coloration in a jumping spider: testing a potential case of sex-specific mimicry.

To avoid predation, many animals mimic behaviours and/or coloration of dangerous prey. Here we examine potential sex-specific mimicry in the jumping spider Habronattus pyrrithrix. Previous work proposed that males' conspicuous dorsal coloration paired with characteristic leg-waving (i.e. false antennation) imperfectly mimics hymenopteran insects (e.g. wasps and bees), affording protection to males during mate-searching and courtship. By contrast, less active females are cryptic and display less leg-waving. Here we test the hypothesis that sexually dimorphic dorsal colour patterns in H. pyrrithrix are most effective when paired with sex-specific behaviours. We manipulated spider dorsal coloration with makeup to model the opposite sex and exposed them to a larger salticid predator (Phidippus californicus). We predicted that males painted like females should suffer higher predation rates than sham-control males. Likewise, females painted like males should suffer higher predation rates than sham-control females. Contrary to expectations, spiders with male-like coloration were attacked more than those with female-like coloration, regardless of their actual sex. Moreover, males were more likely to be captured, and were captured sooner, than females (regardless of colour pattern). With these unexpected negative results, we discuss alternative functional hypotheses for H. pyrrithrix colours, as well as the evolution of defensive coloration generally.

Hemipteran defensive odors trigger predictable color biases in jumping spider predators.

Multimodal warning displays often pair one signal modality (odor) with a second modality (color) to avoid predation. Experiments with bird predators suggest these signal components interact synergistically, with aversive odors triggering otherwise hidden aversions to particular prey colors. In a recent study, this phenomenon was found in a jumping spider (Habronattus trimaculatus), with the defensive odor from a coreid bug (Acanthocephala femorata) triggering an aversion to red. Here, we explore how generalizable this phenomenon is by giving H. trimaculatus the choice between red or black prey in the presence or absence of defensive odors secreted from (1) eastern leaf-footed bugs (Leptoglossus phyllopus, Hemiptera), (2) grass stinkbugs (Mormidea pama, Hemiptera), (3) Asian ladybird beetles (Harmonia axyridis, Coleoptera), and (4) eastern lubber grasshoppers (Romalea microptera, Orthoptera). As expected, in the presence of the hemipteran odors, spiders were less likely to attack red prey (compared to no odor). Unexpectedly, the beetle and grasshopper odors did not bias spiders away from red. Our results with the hemipteran odors were unique to red; follow-up experiments indicated that these odors did not affect biases for/against green prey. We discuss our findings in the context of generalized predator foraging behavior and the functions of multimodal warning displays.

Methods for independently manipulating palatability and color in small insect prey.

Understanding how the psychology of predators shapes the defenses of colorful aposematic prey has been a rich area of inquiry, with emphasis on hypothesis-driven experiments that independently manipulate color and palatability in prey to examine predator responses. Most of these studies focus on avian predators, despite calls to consider more taxonomically diverse predators. This taxonomic bias leaves gaps in our knowledge about the generalizability of current theory. Here we have adapted tools that have been successfully used with bird predators and scaled them down and tested them with smaller predators (Habronattus jumping spiders) and small insect prey (termites, milkweed bug nymphs, pinhead crickets, fruit flies). Specifically, we test (1) the application of denatonium benzoate (DB) to the surface of live termites, crickets, and fruit flies, and (2) the effectiveness of manipulating the palatability of milkweed bug nymphs through diet. We also test the effectiveness of combining these palatability manipulations with various color manipulations. Across several experiments, we confirm that our palatability manipulations are not detectable to the spiders before they attack (i.e., they do not produce aversive odors that spiders avoid), and show that unpalatable prey are indeed quickly rejected and spiders do not habituate to the taste with experience. We also investigate limitations of these techniques by assessing possible unintended effects on prey behavior and the risk of contact contamination when using DB-treated prey in experiments. While similar tools have been used to manipulate color and palatability with avian predators and relatively large insect prey, we show how these techniques can be effectively adapted for use with small invertebrate predators and prey.

Influence of seeing a red face during the male-male encounters of mosquito-specialist spiders.

Males of Evarcha culicivora, an East African jumping spider (Salticidae), have bright red faces. Here, we investigated how seeing a red face might influence a male's behaviour during encounters with another male. We applied black eyeliner to conceal the red on a male's face and measured the spectral properties of male faces with and without the eyeliner. Only the faces without eyeliner reflected in the long-wavelength range corresponding to red. In experiments over 2 days, where eyeliner was absent on the first day and present on the second, we compared how two groups of males responded to their mirror images. Face Group: eyeliner concealed their faces. Head Group: eyeliner was applied to the tops of the males' heads instead of on their faces. The males from both groups displayed to their mirror image as if it were a living same-sex conspecific. However, when they could see a red face, males in the face group escalated to higher levels of aggression to their mirror image and initiated displaying from farther away than when the red had been concealed. We also found that the influence of eyeliner was stronger for the face group than for the head group. These findings suggest that, when seeing a red face, E. culicivora males become more confident that the individual in view is another male.

Variation in activity rates may explain sex-specific dorsal color patterns in Habronattus jumping spiders.

In many animals, color pattern and behavior interact to deceive predators. For mimics, such deception can range from precise (near-perfect mimicry) to only subtle resemblance (imperfect mimicry) and such strategies often differ by sex because of differing ecological selection pressures. In this field study, we examine variation in behavior and ecology that may be linked with sex differences in dorsal color pattern in three sympatric species of Habronattus jumping spiders (H. clypeatus, H. hallani, H. pyrrithrix). Males of these species have conspicuous dorsal patterning that is subtly reminiscent of the general color patterns of wasps and bees, while females are cryptic. We show that, compared with females, these conspicuous males exhibited increased leg-waving behavior outside of the context of courtship; such behavior is common in jumping spiders that mimic wasps and bees presumably because a mimic's waving legs resemble antennae. Males of a fourth sympatric species (H. hirsutus) without conspicuous dorsal patterning did not exhibit increased leg-waving. These results are consistent with and offer preliminary support for the idea that male color and behavior may work together to deceive predators. We also examined whether higher movement rates of males (who must wander to find females) and/or different use of the microhabitat by the sexes could explain sexual dichromatism. We found that microhabitat use was similar for males and females, but males of all three conspicuously-colored species spent more time actively moving than females. To our knowledge, this is the first study to speculate that conspicuous male dorsal coloration in Habronattus may have a deceptive function, and to explore why dorsal coloration differs between the sexes.

Odor alters color preference in a foraging jumping spider.

In many prey taxa with aposematic coloration, prey defenses also involve signals in other modalities (odors, sounds, etc.), yet the selective forces that have driven multimodality in warning displays are not well understood. One potential hypothesis that has recently received support in the avian literature (but has yet to be examined in invertebrates) is that different signal components may interact synergistically, such that one component of a signal (odor) may trigger a predator's aversion to another component of a signal (color). Here, we gave jumping spiders (Habronattus trimaculatus) the choice between red or black prey (artificially colored termites) in either the presence or absence of odor from the chemically defended coreid bug (Acanthocephala femorata). When the odor was present, spiders were more likely to avoid the color red compared with when the odor was absent. Interestingly, this pattern only held up when the odor was novel; subsequent exposure to the odor had no effect on color preference. Moreover, this pattern only held for the color red (a color typically used as a warning color and often paired with odor). We replicated this experiment giving spiders the choice between green or black prey, and found that the presence of the odor had no effect on the spiders' responses to the color green. We discuss these findings in the context of predator psychology and the evolution of prey coloration.

Frequent misdirected courtship in a natural community of colorful Habronattus jumping spiders.

Male courtship display is common in many animals; in some cases, males engage in courtship indiscriminately, spending significant time and energy courting heterospecifics with whom they have no chance of mating or producing viable offspring. Due to high costs and few if any benefits, we might expect mechanisms to evolve to reduce such misdirected courtship (or 'reproductive interference'). In Habronattus jumping spiders, males frequently court heterospecifics with whom they do not mate or hybridize; females are larger and are voracious predators, posing a severe risk to males who court indiscriminately. In this study, we examined patterns of misdirected courtship in a natural community of four sympatric Habronattus species (H. clypeatus, H. hallani, H. hirsutus, and H. pyrrithrix). We used direct field observations to weigh support for two hypotheses (differential microhabitat use and species recognition signaling) to explain how these species reduce the costs associated with misdirected courtship. We show that, while the four species of Habronattus do show some differences in microhabitat use, all four species still overlap substantially, and in three of the four species individuals equally encountered heterospecifics and conspecifics. Males courted females at every opportunity, regardless of species, and in some cases, this led to aggression and predation by the female. These results suggest that, while differences in microhabitat use might reduce misdirected courtship to some extent, co-existence of these four species may be possible due to complex communication (i.e. species-specific elements of a male's courtship display). This study is the first to examine misdirected courtship in jumping spiders. Studies of misdirected courtship and its consequences in the field are limited and may broaden our understanding of how biodiversity is maintained within a community.

Specialists and generalists coexist within a population of spider-hunting mud dauber wasps.

Individual foraging specialization describes the phenomenon where conspecifics within a population of generalists exhibit differences in foraging behavior, each specializing on different prey types. Individual specialization is widespread in animals, yet is understudied in invertebrates, despite potential impacts to food web and population dynamics. Sceliphron caementarium (Hymenoptera: Sphecidae) is an excellent system to examine individual specialization. Females of these mud dauber wasps capture and paralyze spiders which they store in mud nests to provision their offspring. Individuals may make hundreds of prey choices in their short lifespan and fully intact prey items can be easily excavated from their mud nests, where each distinct nest cell represents a discrete foraging bout. Using data collected from a single population of S. caementarium (where all individuals had access to the same resources), we found evidence of strong individual specialization; individuals utilized different resources (with respect to prey taxa, prey ecological guild, and prey size) to provision their nests. The extent of individual specialization differed widely within the population with some females displaying extreme specialization (taking only prey from a single species) while others were generalists (taking prey from up to 6 spider families). We also found evidence of temporal consistency in individual specialization over multiple foraging events. We discuss these findings broadly in the context of search images, responses to changing prey availability, and intraspecific competition pressure.

Iridescence: views from many angles.

Iridescent colours have been fascinating to humans throughout history; they are flashy, shimmering, dynamic, and examples surround us, from the commonly seen iridescent sheen of oily street puddles to the exotic, gaudy displays of birds-of-paradise featured in nature documentaries. Iridescent colours and the structures that produce them have unique properties in comparison with other types of colourants found in nature. Scientists from a variety of disciplines study the optics, development, heritability, chemical make-up, origin, evolution, functions and biomimetic technological applications of naturally occurring iridescent colours. For the first time, graduate students at Arizona State University brought together these scientists, along with educators and artists, at 'Iridescence: more than meets the eye', a conference to promote interdisciplinary communication and collaboration in the study of iridescent coloration from all of these perspectives. Here, we summarize the outcomes of this conference, introduce the papers that follow in this special journal issue and briefly review the current status of our understanding of iridescence.

Animal coloration: sexy spider scales.

Many male jumping spiders display vibrant colors that are used in visual communication. A recent microscopic study on a jumping spider from Singapore shows that three-layered 'scale sandwiches' of chitin and air are responsible for producing their brilliant iridescent body coloration.

Blockage of heregulin expression inhibits tumorigenicity and metastasis of breast cancer.

The growth factor heregulin (HRG), expressed in about 30% of breast cancer tumors, activates the erbB-2 receptor via induction of heterodimeric complexes of erbB-2 with erbB-3 or erbB-4. HRG induces tumorigenicity and metastasis of breast cancer cells. Our investigation into whether HRG is a factor likely to promote tumor formation independently of erbB-2 overexpression concludes that blockage of HRG expression suppresses the aggressive phenotype of MDA-MB-231 breast cancer cells by inhibiting cell proliferation, preventing anchorage-independent growth, and suppressing the invasive potential of the cells in vitro. More importantly, we observed a marked reduction in tumor formation, tumor size, and a lack of metastasis in vivo. These studies were achieved by blocking HRG expression in MDA-MB-231 cells using an HRG antisense cDNA. In the search for the mechanism by which blockage of HRG reverts this aggressive phenotype, we discovered that the cells in which HRG is blocked exhibit a marked decrease in erbB activation and a significant reduction in MMP-9 activity, demonstrating a direct causal role in HRG induction of tumorigenicity. Our study is the first report and serves as a proof of the concept that HRG is a key promoter of breast cancer tumorigenicity and metastasis independently of erbB-2 overexpression and should be deemed a potential target in developing therapies for breast cancer.

Substituted 2-(R)-methyl piperazines as muscarinic M(2) selective ligands.

A novel series of 2-(R)-methyl-substituted piperazines (e.g., 2) is described. They are potent M(2) selective ligands that have >100-fold selectivity versus the M(1) receptor. In the rat microdialysis assay, compound 14 showed significantly enchanced levels of acetylcholine after oral administration.