Kevin A. Judge

CONDITION DEPENDENCE OF MALE LIFE SPAN AND CALLING EFFORT IN A FIELD CRICKET

Abstract Sexually selected traits are thought to impose survival costs on showy males. Recent empirical work found a negative relationship between male display and survival in a field cricket species (Orthoptera, Gryllidae, Gryllinae) where there is no evidence of a mating bias toward older males. In most species, however, male survival and ornamentation are positively correlated, and older males often have a mating success advantage over younger males. These findings suggest that male quality and survival are positively correlated, but more tests of this hypothesis are needed. We measured the condition dependence of male survival and calling effort in another grylline, Gryllus pennsylvanicus, where older males have previously been shown to have greater mating success. We varied condition by manipulating diet, and measured male life span and calling effort to assess the relative condition dependence of these traits. High- and medium-condition males survived longer than low-condition males, and high-condition males called more than medium- and low-condition males. Differences in calling effort among the condition treatments were not apparent early in life, but emerged as males aged. We discuss possible explanations for the differences between our study and contrasting results such as the previous grylline work.

Male weaponry in a fighting cricket.

Sexually selected male weaponry is widespread in nature. Despite being model systems for the study of male aggression in Western science and for cricket fights in Chinese culture, field crickets (Orthoptera, Gryllidae, Gryllinae) are not known to possess sexually dimorphic weaponry. In a wild population of the fall field cricket, Gryllus pennsylvanicus, we report sexual dimorphism in head size as well as the size of mouthparts, both of which are used when aggressive contests between males escalate to physical combat. Male G. pennsylvanicus have larger heads, maxillae and mandibles than females when controlling for pronotum length. We conducted two experiments to test the hypothesis that relatively larger weaponry conveys an advantage to males in aggressive contests. Pairs of males were selected for differences in head size and consequently were different in the size of maxillae and mandibles. In the first experiment, males were closely matched for body size (pronotum length), and in the second, they were matched for body mass. Males with proportionately larger weaponry won more fights and increasing differences in weaponry size between males increased the fighting success of the male with the larger weaponry. This was particularly true when contests escalated to grappling, the most intense level of aggression. However, neither contest duration nor intensity was related to weaponry size as predicted by models of contest settlement. These results are the first evidence that the size of the head capsule and mouthparts are under positive selection via male-male competition in field crickets, and validate 800-year-old Chinese traditional knowledge.

A lover, not a fighter: mating causes male crickets to lose fights

Both resource-holding potential (RHP) and experience in aggressive contests are known to affect future aggressive behaviour. However, few studies have examined the effects of mating experience on agonistic behaviour, despite the fact that dominant males usually acquire more matings. We investigated the effect of mating experience on male aggressive behaviour including the relationship between RHP and fighting success in the fall field cricket, Gryllus pennsylvanicus. We formed pairs of size- and age-matched males that varied in RHP (relative weapon size) and conducted two experiments. In the first, we varied male mating experience by allowing one male in a pair to either be (a) ‘mated’: court, be mounted and copulate with a virgin female or (b) ‘experienced’: court, be mounted, but prevented from copulating. The second experiment varied postcopulatory experience where the male was allowed (‘contact’) or prevented from (‘no-contact’) continued contact with his recent mate. Following treatment, experimental males engaged in an aggressive contest with the naïve size- and age-matched male. In our first experiment, we found that mated and experienced males were equally likely to escalate contests to combat with a naïve opponent, but mated males were less likely than experienced males to win. There was no effect of mating on the relationship between RHP and fighting success. In our second experiment, we found no effect of maintaining contact with the female on the tendency to escalate or the probability of winning. As in the first experiment, males with relatively larger heads again won more fights and this relationship was unaffected by male experience. These results suggest that mating is itself detrimental to male success in aggressive contests, but that this effect is not sufficient to eliminate the effect of RHP on fighting success.

Do male field crickets, Gryllus pennsylvanicus, signal their age?

Older males are often reported to have higher mating success than younger males. To the extent that male quality and survival are positively correlated, this observation raises the possibility that females use male signals to assess age and thus quality. I tested this hypothesis in the fall field cricket, Gryllus pennsylvanicus, a species in which females are known to prefer older males, and males call to attract females. Tests were both longitudinal (males recorded early and late in life) and cross-sectional (males recorded once, each at different ages). I measured a variety of temporal and spectral calling song parameters and tested the predictions that: 1) calling song changes with age, and 2) variation in calling song correlates with variation in age. I found significant changes with increased age: calls showed decreased pulse period, decreased pulse duration, decreased pulse peak frequency, more pulses per chirp and increased pulse period variability. Although pulse period and pulse duration were negatively correlated with male age in bivariate correlations, canonical correlation failed to detect any significant relationship between male age and any linear combination of song parameters. I also measured a number of male body size traits and found that the majority of information in male song appears to be related to body size. I discuss the results in relation to the auditory sensitivity of G. pennsylvanicus, and suggest a simple mechanism that explains both female preference for older males and female discrimination against heterospecific males.

Condition dependence of female choosiness in a field cricket

Females generally choose mates that produce the loudest, brightest or most elaborate sexual displays, and these costly male displays are predicted to be condition dependent. However, mate choice itself is a costly behaviour also expected to be condition dependent. Male fall field crickets, Gryllus pennsylvanicus, produce a conspicuous long‐distance calling song that attracts females and is condition dependent. In this study, we tested the condition dependence of female preferences (preference function and choosiness) for male calling effort in G. pennsylvanicus. We manipulated female condition by raising crickets from hatching on either a low‐ or high‐quality diet. In a series of two‐speaker phonotaxis trials, both low‐ and high‐condition females preferred playbacks reflecting greater calling effort. However, relative to low‐condition females, high‐condition females took significantly longer to make a choice, were more likely to fail to choose within the time allotted for a phonotaxis trial and significantly increased their latency to choose over the course of multiple trials. We discuss these results with respect to the possibility that female G. pennsylvanicus may be foraging for direct benefits when they choose their mates.

Communicating male size by tremulatory vibration in a Columbian rainforest katydid, Gnathoclita sodalis (Orthoptera, Tettigoniidae)

In the South American rainforest katydid Gnathoclita sodalis (Orthoptera, Tettigonidae), bouts of acoustic (airborne) and vibratory (substrate-borne) signals occur in the context of male agonistic interactions. We characterized the physical form for both sound and substrate signals and evaluated their role in male–male interactions. In a tournament design larger males retained sites against smaller opponents: the probability of winning was predicted by both male size and the incidence of tremulating vibrations. Substrate signaling by longbodied rainforest katydids is a widespread and important modality of communication which embraces both female attraction and male rivalry.

How is a cricket like a rat? Insights from the application of cybernetics to evasive food protective behaviour

Robbing and dodging is a well-documented food protective behaviour in rats. Recently, we demonstrated that a simple cybernetic rule, gaining and maintaining a preferred interanimal distance, can account for much of the variability in dodging by rats. In this paper, the field cricket, Teleogryllus oceanicus, was used to test whether or not the same or similar cybernetic rules are used by animals of different lineages and body plans. Pairs of female crickets were tested in a circular arena with a clear glass surface. A small food pellet was given to one of the crickets and the attempts to rob the food by the other were videotaped from beneath. The results show that, although crickets, unlike rats, use a variety of defensive strategies, all of the cases in which they use evasion to protect a portable food item conform to the same cybernetic rules used by rats.

Polyandry and tibial spur chewing in the Carolina ground cricket (Eunemobius carolinus)

During mating, the female Carolina ground cricket (Eunemobius carolinus (Scudder, 1877)) chews specialized spurs on the male’s hind tibia for access to his hemolymph. One potential benefit to spur ...

Small-male mating advantage in a species of Jerusalem cricket (Orthoptera: Stenopelmatinae: Stenopelmatus)

Abstract Scientific literature often touts the many advantages of large body size, but seldom addresses the value of small body size. Yet selection for large size must be counterbalanced by selection for small size, otherwise, all animals would be large. In this paper, we demonstrate female-biased size dimorphism and a strong copulatory advantage for small males in a Jerusalem cricket (JC) (Stenopelmatus) from central California. We selectively paired male and female JCs of diverse body sizes and recorded their ability to copulate. All copulations were successful for males smaller or equal in size to females. In contrast, when the male was 6.1 mm longer than the female, copulation had only a 50% chance of occurring successfully. In general, as the difference between male and female body length increased (i.e., as males became longer than their mates), the probability of successful copulation decreased. These patterns of mating resulted in net selection for small male size and large female size. We also detected positive linear direct selection on male hind leg length, which may explain why male JCs have longer legs than females. The copulatory disadvantage of large males derives from the odd mating behavior of this group, in which males must contort and precisely align their bodies to couple. We believe that this is the first example of small-male advantage based solely on the physical aspects of copulation. In this species, small, not large, males have a copulatory advantage.

Clinical Evaluation of the BD FACSPresto™ Near-Patient CD4 Counter in Kenya.

Background The BD FACSPresto™ Near-Patient CD4 Counter was developed to expand HIV/AIDS management in resource-limited settings. It measures absolute CD4 counts (AbsCD4), percent CD4 (%CD4), and hemoglobin (Hb) from a single drop of capillary or venous blood in approximately 23 minutes, with throughput of 10 samples per hour. We assessed the performance of the BD FACSPresto system, evaluating accuracy, stability, linearity, precision, and reference intervals using capillary and venous blood at KEMRI/CDC HIV-research laboratory, Kisumu, Kenya, and precision and linearity at BD Biosciences, California, USA. Methods For accuracy, venous samples were tested using the BD FACSCalibur™ instrument with BD Tritest™ CD3/CD4/CD45 reagent, BD Trucount™ tubes, and BD Multiset™ software for AbsCD4 and %CD4, and the Sysmex™ KX-21N for Hb. Stability studies evaluated duration of staining (18–120-minute incubation), and effects of venous blood storage <6–24 hours post-draw. A normal cohort was tested for reference intervals. Precision covered multiple days, operators, and instruments. Linearity required mixing two pools of samples, to obtain evenly spaced concentrations for AbsCD4, total lymphocytes, and Hb. Results AbsCD4 and %CD4 venous/capillary (N = 189/ N = 162) accuracy results gave Deming regression slopes within 0.97–1.03 and R2 ≥0.96. For Hb, Deming regression results were R2 ≥0.94 and slope ≥0.94 for both venous and capillary samples. Stability varied within 10% 2 hours after staining and for venous blood stored less than 24 hours. Reference intervals results showed that gender—but not age—differences were statistically significant (p<0.05). Precision results had <3.5% coefficient of variation for AbsCD4, %CD4, and Hb, except for low AbsCD4 samples (<6.8%). Linearity was 42–4,897 cells/μL for AbsCD4, 182–11,704 cells/μL for total lymphocytes, and 2–24 g/dL for Hb. Conclusions The BD FACSPresto system provides accurate, precise clinical results for capillary or venous blood samples and is suitable for near-patient CD4 testing. Trial Registration ClinicalTrials.gov NCT02396355