Invasive Widow Spiders Perform Differently at Low Temperatures from Conspecifics from the Native Range.

Temperature challenges are one of the leading abiotic causes of success or failure of non-native species in a novel environment, and this is particularly true for low temperatures. Establishing and reproducing in a novel thermal environment can alter survival, behavior, and traits related to fitness. It has been proposed that plasticity or adaptation of thermal tolerance may allow an introduced species to thrive, or that successful invaders may be those with a thermal breadth in their native habitat that encompasses their new environment. Here, we tested these hypotheses using native and invasive populations of Australian redback spiders (Latrodectus hasselti). We measured how exposure to temperatures common to invasive and native range habitats (exposure to 15 and 25°C, respectively) affected behavioral and life-history traits and trade-offs that may underlie fitness in an invasive population detected in 1995 in Japan and a native population from Australia. We found that the critical thermal minimum (CTmin) was higher in the invasive population from Japan than in the native population, but critical thermal maximum (CTmax) did not differ between populations. Compared to the invasive population, eggs from the native population had a longer development time and lower hatching success at 15°C. Both populations performed equally well at 25°C, as measured by egg development time and hatching success. Invasive juveniles tested at 15°C were faster to explore a novel environment and bolder compared to those tested at 25°C. In comparison, the native population showed faster average exploration, with no differences in boldness or exploration at the two development or testing temperatures. Overall, L. hasselti from Japan maintained hatching success and development across a wider temperature range than the native population, indicating greater thermal breadth and higher behavioral plasticity. These results support the importance of plasticity in thermal tolerance and behavior for a successful invasion under novel environmental temperatures.

Does body size predict the buzz-pollination frequencies used by bees?

Body size is an important trait linking pollinators and plants. Morphological matching between pollinators and plants is thought to reinforce pollinator fidelity, as the correct fit ensures that both parties benefit from the interaction. We investigated the influence of body size in a specialized pollination system (buzz-pollination) where bees vibrate flowers to release pollen concealed within poricidal stamens. Specifically, we explored how body size influences the frequency of buzz-pollination vibrations. Body size is expected to affect frequency as a result of the physical constraints it places on the indirect flight muscles that control the production of floral vibrations. Larger insects beat their wings less rapidly than smaller-bodied insects when flying, but whether similar scaling relationships exist with floral vibrations has not been widely explored. This is important because the amount of pollen ejected is determined by the frequency of the vibration and the displacement of a bee's thorax. We conducted a field study in three ecogeographic regions (alpine, desert, grassland) and recorded flight and floral vibrations from freely foraging bees from 27 species across four families. We found that floral vibration frequencies were significantly higher than flight frequencies, but never exceeded 400 Hz. Also, only flight frequencies were negatively correlated with body size. As a bee's size increased, its buzz ratio (floral frequency/flight frequency) increased such that only the largest bees were capable of generating floral vibration frequencies that exceeded double that of their flight vibrations. These results indicate size affects the capacity of bees to raise floral vibration frequencies substantially above flight frequencies. This may put smaller bees at a competitive disadvantage because even at the maximum floral vibration frequency of 400 Hz, their inability to achieve comparable thoracic displacements as larger bees would result in generating vibrations with lower amplitudes, and thus less total pollen ejected for the same foraging effort.

In that vein: inflated wing veins contribute to butterfly hearing.

Insects have evolved a diversity of hearing organs specialized to detect sounds critical for survival. We report on a unique structure on butterfly wings that enhances hearing. The Satyrini are a diverse group of butterflies occurring throughout the world. One of their distinguishing features is a conspicuous swelling of their forewing vein, but the functional significance of this structure is unknown. Here, we show that wing vein inflations function in hearing. Using the common wood nymph, Cercyonis pegala, as a model, we show that (i) these butterflies have ears on their forewings that are most sensitive to low frequency sounds (less than 5 kHz); (ii) inflated wing veins are directly connected to the ears; and (iii) when vein inflations are ablated, sensitivity to low frequency sounds is impaired. We propose that inflated veins contribute to low frequency hearing by impedance matching.

Early erratic flight response of the lucerne moth to the quiet echolocation calls of distant bats.

Nocturnal insects have evolved ultrasound-sensitive hearing in response to predation pressures from echolocating insectivorous bats. Flying tympanate moths take various evasive actions when they detect bat cries, including turning away, performing a steering/zigzagging flight and ceasing flight. In general, infrequent ultrasonic pulses with low sound intensities that are emitted by distant bats evoke slight turns, whereas frequent and loud ultrasonic pulses of nearby bats evoke erratic or rapid unpredictable changes in the flight path of a moth. Flight cessation, which is a freezing response that causes the moth to passively dive (drop) to the ground, is considered the ultimate last-ditch evasive behaviour against approaching bats where there is a high predation threat. Here, we found that the crambid moth Nomophila nearctica never performed passive dives in response to frequent and loud ultrasonic pulses of >60 dB sound pressure level (SPL) that simulated the attacking echolocation call sequence of the predominant sympatric insectivorous bat Eptesicus fuscus, but rather turned away or flew erratically, regardless of the temporal structure of the stimulus. Consequently, N. nearctica is likely to survive predation by bats by taking early evasive action even when it detects the echolocation calls of sympatric bats hunting other insects at a distance. Since aerially hawking bats can track and catch erratically flying moths after targeting their prey, this early escape strategy may be common among night-flying tympanate insects.

How spatial release from masking may fail to function in a highly directional auditory system.

Spatial release from masking (SRM) occurs when spatial separation between a signal and masker decreases masked thresholds. The mechanically-coupled ears of Ormia ochracea are specialized for hyperacute directional hearing, but the possible role of SRM, or whether such specializations exhibit limitations for sound source segregation, is unknown. We recorded phonotaxis to a cricket song masked by band-limited noise. With a masker, response thresholds increased and localization was diverted away from the signal and masker. Increased separation from 6° to 90° did not decrease response thresholds or improve localization accuracy, thus SRM does not operate in this range of spatial separations. Tympanal vibrations and auditory nerve responses reveal that localization errors were consistent with changes in peripheral coding of signal location and flies localized towards the ear with better signal detection. Our results demonstrate that, in a mechanically coupled auditory system, specialization for directional hearing does not contribute to source segregation.

Functional morphology of tegmina-based stridulation in the relict species Cyphoderris monstrosa (Orthoptera: Ensifera: Prophalangopsidae).

Male grigs, bush crickets and crickets produce mating calls by tegminal stridulation: the scraping together of modified forewings functioning as sound generators. Bush crickets (Tettigoniidae) and crickets (Gryllinae) diverged some 240 million years ago, with each lineage developing unique characteristics in wing morphology and the associated mechanics of stridulation. The grigs (Prophalangopsidae), a relict lineage more closely related to bush crickets than to crickets, are believed to retain plesiomorphic features of wing morphology. The wing cells widely involved in sound production, such as the harp and mirror, are comparatively small, poorly delimited and/or partially filled with cross-veins. Such morphology is similarly observed in the earliest stridulating ensiferans, for which stridulatory mechanics remains poorly understood. The grigs, therefore, are of major importance to investigate the early evolutionary stages of tegminal stridulation, a critical innovation in the evolution of the Orthoptera. The aim of this study is to appreciate the degree of specialization on grig forewings, through identification of sound radiating areas and their properties. For well-grounded comparisons, homologies in wing venation (and associated areas) of grigs and bush crickets are re-evaluated. Then, using direct evidence, this study confirms the mirror cell, in association with two other areas (termed 'neck' and 'pre-mirror'), as the acoustic resonator in the grig Cyphoderris monstrosa Despite the use of largely symmetrical resonators, as found in field crickets, analogous features of stridulatory mechanics are observed between C. monstrosa and bush crickets. Both morphology and function in grigs represents transitional stages between unspecialized forewings and derived conditions observed in modern species.

Active auditory mechanics in female black­horned tree crickets (Oecanthus nigricornis).

The acoustic signalling behaviour of many tree cricket species is easily observed and has been well described. Very little is known, however, about the receivers in these communication loops. The exception to this is a single Indian species (Oecanthus henryi) which employs active auditory mechanics to enhance female sensitivity to quiet sounds at male calling frequencies. In most species, male calls have been described, but whether or not sender­receiver matching is present is uncertain. Here we investigate auditory mechanics in females of the North American black-horned tree cricket (Oecanthus nigricornis). The response of the anterior tympanal membrane is nonlinear, exhibiting a lack of tuning at high amplitudes (60 dB and above) but as stimulus amplitude decreases, the membrane becomes tuned to around 4.3 kHz. The tuning of the membrane falls within the frequency range of male calls indicating sender­receiver matching at low amplitudes, which could aid localisation of the highly directional calls of males. The extent of active auditory mechanics in tympanal insects is not yet known, but this paper provides an indication that this may indeed be widespread in at least the Oecanthinae.

Metabolic efficiency in courtship favors males with intermediate mass in the Australian redback spider, Latrodectus hasselti.

Recent studies have suggested that metabolic efficiency may be an important factor in male mating success when females require vigorous and/or prolonged courtship. In capital breeding animals in which a male's resource pool is fixed at adulthood the relationship between energy expenditure and courtship performance may be especially important, as males are expected to utilize their finite resources efficiently when soliciting mates. Males may benefit from being efficient, i.e., achieving a sufficiently high level of courtship signaling at low energetic cost, if it enables them to acquire mates before their limited energy reserves are depleted. We investigated the relationship between metabolic efficiency and courtship vibrational signaling in the Australian redback spider, Latrodectus hasselti, a semelparous capital breeder where males invest heavily in courtship to secure a mating. We assessed metabolic rate in a sample of males and measured two courtship components (duty cycle and amplitude) that reflected the energy content of web-borne vibrations. We then calculated two indices of metabolic efficiency for these courtship properties. There was a quadratic relationship between mass and duty cycle such that the highest duty cycle signals were performed by males having intermediate mass. Furthermore, intermediate-mass males were also the most metabolically efficient. Prolonged courtship is necessary in L. hasselti for successful mating, and the results of this study suggest that intermediate-mass males are superior courters because they utilize their finite resource pool most efficiently to produce high energy vibrational signals.

Lack of correlation between vertical distribution and carrier frequency, and preference for open spaces in arboreal katydids that use extreme ultrasound, in Gorgona, Colombia (Orthoptera: Tettigoniidae) / Falta de correlación entre distribución vertical y frecuencia portadora, y preferencia por espacios abiertos en saltamontes arbóreos que utilizan ultrasonido extremo, en Gorgona, Colombia (Orthoptera: Tettigoniidae)

Male Tettigoniidae emit sound to attract conspecific females. The sound is produced by stridulation. During stridulation the forewings open and close, but it is during the closing stroke that the scraper contacts the file teeth to generate the predominant sound components, which are amplified by adjacent wing cells specialized in sound radiation. The sounds usually exceed the sonic boundary and might occur above 40 kHz, reaching extreme ultrasonic frequencies of 150kHz in some species. Here we test the hypothesis that Tettigoniidae species should prefer microhabitats that favour efficient signal transmission, i.e. that there is a relationship of sound frequency with the vertical distribution of the species (from ground to canopy) at Gorgona National Natural Park, Colombia. We sampled 16 trees and four different altitudinal levels between 1 and 20m above the understory vegetation. We placed collecting blankets separated by vertical distances of 5m, and knocked insects down using the technique known as fogging. We found no correlation between vertical distribution and carrier frequency, but there was a preference for open spaces (below the canopy and above the understory) in species using extreme ultrasound. This is the first quantitative description of the vertical distribution in neotropical species of the family Tettigoniidae and its relationship to the calling song frequency. Rev. Biol. Trop. 62 (Suppl. 1): 289-296. Epub 2014 February 01.

Los machos de Tettigoniidae producen sonido frotando las alas anteriores para atraer a sus hembras coespecíficas con fines reproductivos (mecanismo conocido como estridulación). Un borde esclerotizado (o raspador) en el ala derecha es frotado sobre una vena modificada con dientecillos en el ala izquierda. Durante la estridulación las alas abren y cierran, pero es durante el cierre que el raspador contacta los dientecillos de la lima y produce vibraciones que son amplificadas por celdas adyacentes especializadas para radiar sonido. Sonidos que superen los 20 000Hz, se consideran ultrasónicos, en los Tettigoniidae, la mayoría de los cantos superan el umbral sónico y pueden ocurrir a más de 40 000Hz, con casos extremos de hasta 148 000Hz. La esencia de este trabajo es el estudio de esta gama espectacular de frecuencias de la familia Tettigoniidae y su relación con la distribución vertical de los individuos en las especies que habitan en Gorgona. Se muestrearon 16 árboles a cuatro niveles diferentes de altitud entre 1 y ~20m sobre el sotobosque, instalando sabanas colectoras a intervalos de 4-5m. Los insectos se colectaron mediante el método de nebulización térmica. El análisis indica que no hay correlación entre la distribución vertical de las especies y la frecuencia del canto. Sin embargo se observó una preferencia por espacios abiertos en especies que cantan a frecuencias extremas. Este estudio representa la primera descripción cuantitativa de la distribución vertical de Tettigoniidae Neotropicales y su relación con la frecuencia del canto.

Variability in bumblebee pollination buzzes affects the quantity of pollen released from flowers.

Buzz-pollination is a plant strategy that promotes gamete transfer by requiring a pollinator, typically bees (Hymenoptera: Apoidea), to vibrate a flower's anthers in order to extract pollen. Although buzz-pollination is widespread in angiosperms with over 20,000 species using it, little is known about the functional connection between natural variation in buzzing vibrations and the amount of pollen that can be extracted from anthers. We characterized variability in the vibrations produced by Bombus terrestris bumblebees while collecting pollen from Solanum rostratum (Solanaceae), a buzz-pollinated plant. We found substantial variation in several buzzing properties both within and among workers from a single colony. As expected, some of this variation was predicted by the physical attributes of individual bumblebees: heavier workers produced buzzes of greater amplitude. We then constructed artificial "pollination buzzes" that varied in three parameters (peak frequency, peak amplitude, and duration), and stimulated S. rostratum flowers with these synthetic buzzes to quantify the relationship between buzz properties and pollen removal. We found that greater amplitude and longer duration buzzes ejected substantially more pollen, while frequency had no directional effect and only a weak quadratic effect on the amount of pollen removed. These findings suggest that foraging bumblebees may improve pollen collection by increasing the duration or amplitude of their buzzes. Moreover, given that amplitude is positively correlated with mass, preferential foraging by heavier workers is likely to result in the largest pollen yields per bee, and this could have significant consequences for the success of a colony foraging on buzz-pollinated flowers.

Examination of prior contest experience and the retention of winner and loser effects.

In many animal taxa, prior contest experience affects future performance such that winning increases the chances of winning in the future (winner effect) and losing increases the chances of losing in the future (loser effect). It is, however, not clear whether this pattern typically arises from experience effects on actual or perceived fighting ability (or both). In this study, we looked at winner and loser effects in the jumping spider Phidippus clarus. We assigned winning or losing experience to spiders and tested them against opponents of similar fighting ability in subsequent contests at 1-, 2-, 5-, and 24-h intervals. We examined the strength of winner and loser effects, how long effects persist, as well as how experience affected perceived and actual fighting ability. Our results demonstrate that winner and loser effects are of approximately the same magnitude, although loser effects last longer than winner effects. Our results also demonstrate that previous experience alters actual fighting ability because both the assessment and escalation periods were affected by experience. We suggest that the retention time of experience effects depends on expected encounter rates as well as other behavioral and ecological factors. In systems with short breeding seasons and/or rapidly fluctuating populations, context-dependent retention of experience effects may allow males to track their status relative to the fluctuating fighting ability of local competitors without paying the costs necessary to recall or assess individual competitors.

A precedence effect resolves phantom sound source illusions in the parasitoid fly Ormia ochracea.

Localizing individual sound sources under reverberant environmental conditions can be a challenge when the original source and its acoustic reflections arrive at the ears simultaneously from different paths that convey ambiguous directional information. The acoustic parasitoid fly Ormia ochracea (Diptera: Tachinidae) relies on a pair of ears exquisitely sensitive to sound direction to localize the 5-kHz tone pulsatile calling song of their host crickets. In nature, flies are expected to encounter a complex sound field with multiple sources and their reflections from acoustic clutter potentially masking temporal information relevant to source recognition and localization. In field experiments, O. ochracea were lured onto a test arena and subjected to small random acoustic asymmetries between 2 simultaneous sources. Most flies successfully localize a single source but some localize a 'phantom' source that is a summed effect of both source locations. Such misdirected phonotaxis can be elicited reliably in laboratory experiments that present symmetric acoustic stimulation. By varying onset delay between 2 sources, we test whether hyperacute directional hearing in O. ochracea can function to exploit small time differences to determine source location. Selective localization depends on both the relative timing and location of competing sources. Flies preferred phonotaxis to a forward source. With small onset disparities within a 10-ms temporal window of attention, flies selectively localize the leading source while the lagging source has minimal influence on orientation. These results demonstrate the precedence effect as a mechanism to overcome phantom source illusions that arise from acoustic reflections or competing sources.

Experience affects the outcome of agonistic contests without affecting the selective advantage of size.

In the field, phenotypic determinants of competitive success are not always absolute. For example, contest experience may alter future competitive performance. As future contests are not determined solely on phenotypic attributes, prior experience could also potentially alter phenotype-fitness associations. In this study, we examined the influence of single and multiple experiences on contest outcomes in the jumping spider Phidippus clarus. We also examined whether phenotype-fitness associations altered as individuals gained more experience. Using both size-matched contests and a tournament design, we found that both winning and losing experience affected future contest success; males with prior winning experience were more likely to win subsequent contests. Although experience was a significant determinant of success in future contests, male weight was approximately 1.3 times more important than experience in predicting contest outcomes. Despite the importance of experience in determining contest outcomes, patterns of selection did not change between rounds. Overall, our results show that experience can be an important determinant in contest outcomes, even in short-lived invertebrates, and that experience alone is unlikely to alter phenotype-fitness associations.

Hearing and spatial behavior in Gryllotalpa major Saussure (Orthoptera: Gryllotalpidae).

The prairie mole cricket (Gryllotalpa major Saussure) is a rare orthopteran insect of the tallgrass prairie ecosystem of the south central USA. Populations are known to currently occupy fragmented prairie sites in Oklahoma, Arkansas, Kansas and Missouri, including The Nature Conservancy's Tallgrass Prairie Preserve in north central Oklahoma. Prairie mole cricket populations were surveyed at this site and at another site in Craig County, OK during the spring of 2005 and 2006, using the male cricket's acoustic call to locate advertising aggregations of males. Five males from one large aggregation were removed in a study to describe (1) the hearing thresholds across the call's range of frequencies, (2) the distances over which the higher harmonic components of the male's calls are potentially detectable, (3) the species' sensitivity to ultrasound and (4) the spatio-auditory dynamics of the prairie mole cricket lek. Results indicate that G. major has a bimodal pattern of frequency tuning, with hearing sensitivities greatest at the 2 kHz carrier frequency (41 dB SPL) and declining through the call's frequency range (84 dB at 10 kHz). A second sensitivity peak is evident in the ultrasound range at 25 kHz (62 dB SPL). Spatial analysis of G. major lek sites indicates that approximately 73% of males within the lek are spaced in such a way as to allow acoustic interaction at the species' carrier frequency, while any information in higher harmonic overtones in the call appears to be available only to nearest neighbors.

Hurthle cell neoplasms of the thyroid: sonographic appearance and histologic characteristics.

OBJECTIVE: The purpose of this study was to determine the sonographic features of Hürthle cell neoplasms (HCNs) of the thyroid. METHODS: We retrospectively analyzed the sonographic appearance of 15 histologically proven HCNs in 15 patients aged 16 to 70 years (mean age, 44 years). Sonographic features that were reviewed included the size and echogenicity of the tumors, the presence of cystic areas or calcifications, and detectable blood flow on color Doppler imaging. Correlation of sonographic findings with pathologic results was performed. RESULTS: The tumors ranged from 0.4 to 7 cm in diameter, but most were less than 3 cm in diameter. Four (27%) of the 15 tumors were homogeneously hypoechoic. Two tumors (13%) were predominantly hypoechoic with isoechoic areas to thyroid parenchyma. Two (13%) neoplasms were isoechoic to thyroid parenchyma. Four (27%) tumors were predominantly isoechoic, containing hypoechoic areas, and 3 (20%) tumors were hyperechoic. Three neoplasms contained cystic components. None of the tumors contained calcifications. One tumor was avascular on Doppler examination. One neoplasm showed only peripheral blood flow. Thirteen tumors showed internal vascularity, 7 of them with peripheral blood flow. Twelve HCNs were benign, and 3 were malignant on pathologic examination. CONCLUSIONS: Hürthle cell neoplasms show a spectrum of sonographic appearances from predominantly hypoechoic to hyperechoic lesions and from peripheral blood flow with no internal flow to extensively vascularized lesions. Pathologic criteria differentiating benign and malignant HCNs (absence or presence of a capsular breach, vascular or extrathyroidal tissue invasion, nodal involvement, and distant metastasis) are beyond the resolution of sonography and fine-needle aspiration biopsy and require removal of the entire lesion. This precludes diagnosis and characterization of HCNs by sonography.

Assessment during aggressive contests between male jumping spiders.

Assessment strategies are an important component in game theoretical models of contests. Strategies can be either based on one's own abilities (self assessment) or on the relative abilities of two opponents (mutual assessment). Using statistical methodology that allows discrimination between assessment types, we examined contests in the jumping spider Phiddipus clarus. In this species, aggressive interactions can be divided into 'pre-contact' and 'contact' phases. Pre-contact phases consist of bouts of visual and vibratory signaling. Contact phases follow where males physically contact each other (leg fencing). Both weight and vibratory signaling differences predicted winners with heavier and more actively signaling males winning more contests. Vibratory behaviour predicted pre-contact phase duration, with higher signaling rates and larger differences between contestants leading to longer pre-contact interaction times. Contact phase duration was predicted most strongly by the weight of losing males relative to that of winning males, suggesting that P. clarus males use self-assessment in determining contest duration. While a self-assessment strategy was supported, our data suggest a secondary role for mutual assessment ("partial mutual assessment"). After initial contest bouts, male competitors changed their behaviour. Pre-contact and contact phase durations were reduced while vibratory signaling behaviour in winners was unchanged. In addition, only vibratory signaling differences predicted winners in subsequent bouts suggesting a role of experience in determining contest outcomes. We suggest that the rules and assessment strategies males use can change depending on experience and that assessment strategies are likely a continuum between self- and mutual assessment.

CT findings of normal and inflamed appendix in groin hernia.

Acute appendicitis with the vermiform appendix located in a groin hernia is a rare condition. The preoperative diagnosis is important to decrease morbidity. We describe the computed tomography imaging characteristics of three cases of normal and inflamed appendices in inguinal and femoral hernias. We provide a review of the literature and consider the implications for surgical management.

Generation of extreme ultrasonics in rainforest katydids.

The calling song of an undescribed Meconematinae katydid (Tettigoniidae) from South America consists of trains of short, separated pure-tone sound pulses at 129 kHz (the highest calling note produced by an Arthropod). Paradoxically, these extremely high-frequency sound waves are produced by a low-velocity movement of the stridulatory forewings. Sound production during a wing stroke is pulsed, but the wings do not pause in their closing, requiring that the scraper, in its travel along the file, must do so to create the pulses. We hypothesize that during scraper pauses, the cuticle behind the scraper is bent by the ongoing relative displacement of the wings, storing deformation energy. When the scraper slips free it unbends while being carried along the file and its deformation energy contributes to a more powerful, higher-rate, one-tooth one-wave sound pulse, lasting no more than a few waves at 129 000 Hz. Some other katydid species make pure-tone ultrasonic pulses. Wing velocities and carriers among these pure-tone species fall into two groups: (1) species with ultrasonic carriers below 40 kHz that have higher calling frequencies correlated with higher wing-closing velocities and higher tooth densities: for these katydids the relationship between average tooth strike rate and song frequency approaches 1:1, as in cricket escapement mechanisms; (2) a group of species with ultrasonic carriers above 40 kHz (that includes the Meconematinae): for these katydids closing wing velocities are dramatically lower and they make short trains of pulses, with intervening periods of silence greater than the duration of the pulses they separate. This signal form may be the signature of scraper-stored elastic energy.

Seismic signal production in a wolf spider: parallel versus serial multi-component signals.

Animal signals can consist of multiple parts within or across sensory modalities (multi-component signals or multimodal signals). While recent work has focused on multimodal signals, the production, processing and evolution of multi-component signals has received considerably less attention. Here, using synchronous high-speed video and laser vibrometer recordings followed by experimental manipulations of putative sound-producing structures, we explored the mechanisms of seismic signal production in the courtship display of Schizocosa stridulans Stratton. Two types of seismic courtship signals were observed: 'rev' and 'idle' signals. Revs consist of a high-frequency component produced by flexions of the male pedipalp (stridulation) simultaneous with a low-frequency component produced by movements of the abdomen (tremulation). This multi-component signal is produced by independent structures and represents a parallel multi-component display. By contrast, idle displays consist of a high-intensity component produced by drumming of the forelegs on the substrate (percussion) followed by a high-frequency component produced by flexions of the male pedipalp (stridulation). While the components of the idle display are also produced by independent structures, the leg drumming and palp flexions occur serially and do not overlap in time. We discuss the selective pressures that may drive the evolution of multiple sound-producing structures as well as the selective pressures that drive the evolution of parallel versus serial multi-component signals.

Measuring and quantifying dynamic visual signals in jumping spiders.

Animals emit visual signals that involve simultaneous, sequential movements of appendages that unfold with varying dynamics in time and space. Algorithms have been recently reported (e.g. Peters et al. in Anim Behav 64:131-146, 2002) that enable quantitative characterization of movements as optical flow patterns. For decades, acoustical signals have been rendered by techniques that decompose sound into amplitude, time, and spectral components. Using an optic-flow algorithm we examined visual courtship behaviours of jumping spiders and depict their complex visual signals as "speed waveform", "speed surface", and "speed waterfall" plots analogous to acoustic waveforms, spectrograms, and waterfall plots, respectively. In addition, these "speed profiles" are compatible with analytical techniques developed for auditory analysis. Using examples from the jumping spider Habronattus pugillis we show that we can statistically differentiate displays of different "sky island" populations supporting previous work on diversification. We also examined visual displays from the jumping spider Habronattus dossenus and show that distinct seismic components of vibratory displays are produced concurrently with statistically distinct motion signals. Given that dynamic visual signals are common, from insects to birds to mammals, we propose that optical-flow algorithms and the analyses described here will be useful for many researchers.