Water Strider Aquarius Remigis Research Articles

Escape jumping by three age-classes of water striders from smooth, wavy and bubbling water surfaces.

Surface roughness is a ubiquitous phenomenon in both oceanic and terrestrial waters. For insects that live at the air-water interface, such as water striders, non-linear and multi-scale perturbations produce dynamic surface deformations which may impair locomotion. We studied escape jumps of adults, juveniles and first-instar larvae of the water strider Aquarius remigis on smooth, wave-dominated and bubble-dominated water surfaces. Effects of substrate on takeoff jumps were substantial, with significant reductions in takeoff angles, peak translational speeds, attained heights and power expenditure on more perturbed water surfaces. Age effects were similarly pronounced, with the first-instar larvae experiencing the greatest degradation in performance; age-by-treatment effects were also significant for many kinematic variables. Although commonplace in nature, perturbed water surfaces thus have significant and age-dependent effects on water strider locomotion, and on behavior more generally of surface-dwelling insects.

An interommatidial exocrine gland with a “nail-headed” structure in the water strider Aquarius remigis (Hemiptera, Gerridae)

The fine structure of the interommatidial exocrine glands, found in the compound eyes of the water strider Aquarius remigis, is described using light, scanning, and transmission electron microscopy. The glandular pores of the glands are specialized into minute “nail-headed” structures (NS), which are described for the first time in arthropod compound eyes. Each NS is composed of two components: a rod-like stalk and a cup-like depression. The TEM study shows that the glands are class 3 epidermal glands as defined by Noirot and Quennedey (1974, 1991). Each gland consists of 3 cells: a gland cell, an intermediary cell, and a duct (canal) cell. The gland cell contains abundant electron-lucent vesicles, while the intermediary cell contains a large number of osmiophilic secretory granules. These two cells might secrete different substances which mix together in the dilated sac-like portion of the conducting canal before final release. The possible functions of the secretions released from these glands are discussed.

Germ-cell hub position in a heteropteran testis correlates with the sequence and location of spermatogenesis and production of elaborate sperm bundles.

In insects, spermatogonial cells undergo several mitotic divisions with incomplete cytokinesis, and then proceed through meiosis and spermatogenesis in synchrony. The cells derived from a single spermatogonial cell are referred to as a cyst. In the water strider Aquarius remigis, spermiogenesis occurs within two bi-lobed testes. In contrast to most insects, in which the germ-cell hub is located apically and sequential stages of spermatogenesis can be seen moving toward the base of the testis, each lobe of the water strider testis contains a single germ-cell hub located medially opposite to the efferent duct of the lobe; the developing cysts are displaced toward the distal ends of the lobe as spermiogenesis proceeds. Water strider sperm have both a long flagellum and an unusually long acrosome. The water strider spermatids elongate most of the flagellum prior to morphogenesis of the acrosome, and exhibit several stages of nuclear remodeling before the final, mature sperm nucleus is formed. The maturing sperm are aligned in register in the cyst, and the flagella fold into a coiled bundle while their acrosomes form a rigid helical process that extends from the cyst toward the efferent duct.

Protease activation and the signal transduction pathway regulating motility in sperm from the water strider Aquarius remigis

Many motile processes are regulated such that movement occurs only upon activation of a signaling cascade. Sperm from a variety of species are initially quiescent and must be activated prior to beating. The signaling events leading to the activation and regulation of sperm motility are not well characterized. Mature seminal vesicle sperm from the water strider Aquarius remigis are immotile in vitro, but vigorous motility is activated by trypsin. Trypsin-activated motility was blocked by pretreatment of the sperm with BAPTA-AM to chelate intracellular Ca(2+) and was partially rescued by subsequent addition of A23187 and Ca(2+). Thapsigargin stimulated motility in the absence of trypsin, suggesting that intracellular Ca(2+) stores are available. In addition, motility could be fully activated by the phosphatase inhibitor calyculin A, suggesting that the immotile state is maintained by an endogenous phosphatase and that kinase activity is required for motility. The MEK1/2 inhibitor U0126 significantly reduced trypsin activated motility, and MPM-2, an antibody which recognizes proline-directed phosphorylation by kinases such as MAPK, recognized components of the water strider sperm flagellum. Antibodies specific for the mouse protease activated receptor PAR2 recognized an antigen on the sperm flagellum. These results suggest that trypsin stimulates a Ca(2+) and MAPK mediated signaling pathway and potentially implicate a PAR2-like protein in regulating motility.

Reproductive allometry does not explain the temperature-size rule in water striders (Aquarius remigis)

Most ectotherms follow the temperature-size rule meaning that individuals growing up under cool conditions are larger as adults than those growing up in warm conditions. This pattern is difficult to explain because growth is usually slower in the cold meaning it takes longer to reach a larger adult size. One potential explanation for this pattern is that the typical increase in fecundity with body size is steeper in cool environments than it is in warm environments. As such, the relative gain in fecundity for being larger in the cold would compensate for the extra time it takes to grow to that size. We present the first empirical test of this model using the water strider Aquarius remigis. Individuals were reared at either 20° or 25°C with subsampling at each instar to estimate growth trajectories. At adult eclosion, half were switched to the alternate temperature and all females isolated, mated, and reproductive output measured for 3 weeks. We found that A. remigis does follow the temperature-size rule but that fecundity was highly plastic with respect to laying temperature such that the slightly greater fecundity of those reared at 20°C was due to their larger size. Overall, those laying at 25°C were more fecund and showed a positive relationship between body size and fecundity. Those laying at 20°C did not show a significant relationship between size and fecundity. As such, the reproductive allometry shows a pattern reverse to what would be needed for a larger size in the cold to be adaptive in this species. Although A. remigis follows the temperature-size rule over this temperature range, this is likely due to a constraint on growth and development rather than being adaptive plasticity.

The effect of ambient humidity and metabolic rate on the gas-exchange pattern of the semi-aquatic insectAquarius remigis

We have examined the effects of temperature on metabolic rate and respiratory pattern in the water strider Aquarius remigis. As temperature was increased from 10 to 30°C, the metabolic rate of the insects increased and the respiratory pattern transitioned from discontinuous, to cyclic, to continuous. The discontinuous gas-exchange cycle (DGC) was observed even in insects standing on water when the respirometry chamber was being perfused with humid (>95% relative humidity) air. Comparisons of insects at 20°C in humid and dry air showed no statistically significant differences in metabolic rate or respiratory pattern (P>0.05). The proportion of time that the spiracles were closed was greater at 10°C than at 20°C (P<0.01), and greater at 20°C than at 30°C (P<0.05). These results are compatible with the hypothesis that the respiratory patterns of insects are determined by the relationship between oxygen supply and oxygen demand. There was no evidence in this insect that humidity had any effect on the respiratory pattern. The results are discussed in the context of the ongoing discussion in the literature of the origin, maintenance and adaptive significance of the DGC in insects.

Divergent juvenile growth and development mediated by food limitation and foraging in the water strider Aquarius remigis (Heteroptera: Gerridae)

AbstractThe factors leading to variation in growth, development and body size are of central importance for the study of life histories. I investigated the effect of food limitation on patterns of growth and development in the water strider Aquarius remigis (Heteroptera: Gerridae). Groups of 10 juveniles were allowed to develop under field conditions at three different food levels. Food limitation prolonged juvenile development, reduced growth rates and led to smaller adult body size and weight, the most common life‐history response in ectotherms; it also increased the variance among individuals in these traits, demonstrating variance accumulation suggesting divergent growth. Evidence that this resulted from competition among individuals for limited food was equivocal. On the one hand, direct observation of foraging nymphs showed that larger instar individuals were more likely to obtain monopolizable prey items than expected by chance, independent of food level. On the other hand, there was only a trend for the variance in moulting dates to increase when striders were raised in groups as opposed to singly at various temperatures in a controlled laboratory experiment. When food limitation prolongs juvenile development, variance accumulation with age can result from food limitation per se, as well as from random or heritable variation in growth trajectories. Foraging competition may amplify this effect, but not necessarily so. Therefore, it must not be assumed that divergent growth is necessarily caused by food competition.

Allometry for Sexual Size Dimorphism: Testing Two Hypotheses for Rensch’s Rule in the Water StriderAquarius remigis

Within any given clade, male size and female size typically covary, but male size often varies more than female size. This generates a pattern of allometry for sexual size dimorphism (SSD) known as Rensch's rule. I use allometry for SSD among populations of the water strider Aquarius remigis (Hemiptera, Gerridae) to test the hypothesis that Rensch's rule evolves in response to sexual selection on male secondary sexual traits and an alternative hypothesis that it is caused by greater phenotypic plasticity of body size in males. Comparisons of three populations reared under two temperature regimes are combined with an analysis of allometry for genital and somatic components of body size among 25 field populations. Contrary to the sexual-selection hypothesis, genital length, the target of sexual selection, shows the lowest allometric slope of all the assayed traits. Instead, the results support a novel interpretation of the differential-plasticity hypothesis: that the traits most closely associated with reproductive fitness (abdomen length in females and genital length in males) are "adaptively canalized." While this hypothesis is unlikely to explain Rensch's rule among species or higher clades, it may explain widespread patterns of intraspecific variation in SSD recently documented for many insect species.

The mix matters: behavioural types and group dynamics in water striders

Summary The effect of an individual’s behavioral type (e.g., its boldness or aggressiveness) on fitness likely depends on the environmental context. In many species, an important component of an individual’s environment is its social environment — the mix of individuals in its social group. Accordingly, much of game theory assumes that the mix of behavioral types (BTs) in a social group influences individual fitness and group dynamics. Few experimental studies, however, have directly investigated how the group’s mix of BTs influences individual and group outcomes. Here we manipulated social group composition in the water strider Aquarius remigis and evaluated the effects of our manipulation on individual behavior and various group outcomes (overall group activity, aggression, mating success). We formed 12 groups that differed substantially in average male BT (activity and aggression level), each with a low variance in BT. That is, one group had only the most active and aggressive males, a second group was made up of the next most active and aggressive males, and on down to a group of all very inactive and unaggressive males. All groups also had females. We found that, on average, groups made up of more active-aggressive males continued to be more active than other groups, but that contrary to predictions, these groups did not tend to enjoy higher mating success. Instead, a major factor affecting group mating activity was the presence of hyper-aggressive males. Hyper-aggressive males drove females out of the group and thereby decreased the group’s overall mating activity. We discuss these findings in terms of their importance to the study of behavioral plasticity in social groups and the potential role of keystone individuals in determining group dynamics.

Functional morphology of sexually selected gentalia in the water strider Aquarius remigis

In the water strider Aquarius remigis (Say), sexual selection favours males with longer genitalia. We used video analysis plus light and scanning electron microscopy to examine the structure, movement, and articulation of the genitalia as a first step in determining the functional basis of this selection. Male A. remigis are characterized by long, robust genital segments; a marked ventral median notch in the posterior margin of the pregenital segment; an enlarged phallus tipped with a uniquely prolonged sclerotized plate; and a large, spine-covered, membranous lobe that inflates within the female's reproductive tract. Detailed examinations of genital interactions prior to and during copulation allow us to deduce the functional significance of these distinguishing traits. We postulate that by increasing the length, mobility, strength, and rigidity of the phallus, the first three traits increase the ability of males to achieve intromission in spite of active female resistance, and to maintain intromission during the prolonged copulations characteristic of this species. Inflation of the large, spinous lobe probably contributes to the latter function, and may also directly affect fertilization success by displacing or damaging sperm from previous males. We discuss these interpretations in the context of current theories of genitalic coevolution and sexual conflict in the Gerridae.

How well do mating frequency and duration predict paternity success in the polygynandrous water strider Aquarius remigis?

The relationship between mating success and paternity success is a key component of sexual selection but has seldom been estimated for species in which both sexes mate with many partners (polygynandry). We used a modification of Parker's sterile male technique to measure this relationship for the water strider Aquarius remigis in 47 laboratory populations simulating natural conditions of polygynandry. We also tested the hypothesis that prolonged copulation, a characteristic of this species, enhances paternity success. Mating behavior and paternity success were assayed for four days while males and females freely interacted. Paternity success was also assayed for an additional 7 days when females were isolated from males. Mating success significantly predicted paternity success and accounted for < or = 36% of the variance. Copulation duration was negatively related to both mating success and paternity success and did not explain any of the residual variance in paternity success. Thus, we found no evidence that prolonged copulation functions as a paternity assurance strategy in this species. Comparisons of sterile and fertile males suggested that paternity success is directly influenced by the quantity of sperm transferred. Our results support previous studies that have used mating success to estimate sexual selection, but also highlight the potential importance of sperm competition and other postinsemination processes.

HOW WELL DO MATING FREQUENCY AND DURATION PREDICT PATERNITY SUCCESS IN THE POLYGYNANDROUS WATER STRIDER AQUARIUS REMIGIS?

The relationship between mating success and paternity success is a key component of sexual selection but has seldom been estimated for species in which both sexes mate with many partners (polygynandry). We used a modification of Parker's sterile male technique to measure this relationship for the water strider Aquarius remigis in 47 laboratory populations simulating natural conditions of polygynandry. We also tested the hypothesis that prolonged copulation, a characteristic of this species, enhances paternity success. Mating behavior and paternity success were assayed for four days while males and females freely interacted. Paternity success was also assayed for an additional 7 days when females were isolated from males. Mating success significantly predicted paternity success and accounted for ≤ 36% of the variance. Copulation duration was negatively related to both mating success and paternity success and did not explain any of the residual variance in paternity success. Thus, we found no evidence that prolonged copulation functions as a paternity assurance strategy in this species. Comparisons of sterile and fertile males suggested that paternity success is directly influenced by the quantity of sperm transferred. Our results support previous studies that have used mating success to estimate sexual selection, but also highlight the potential importance of sperm competition and other postinsemination processes.

LIFETIME SELECTION ON ADULT BODY SIZE AND COMPONENTS OF BODY SIZE IN A WATERSTRIDER: OPPOSING SELECTION AND MAINTENANCE OF SEXUAL SIZE DIMORPHISM

Abstract.— Sexual size dimorphism (SSD), the difference in body size between males and females, is common in almost all taxa of animals and is generally assumed to be adaptive. Although sexual selection and fecundity selection alone have often been invoked to explain the evolution of SSD, more recent views indicate that the sexes must experience different lifetime selection pressures for SSD to evolve and be maintained. We estimated selection acting on male and female adult body size (total length) and components of body size in the waterstrider Aquarius remigis during three phases of life history. Opposing selection pressures for overall body size occurred in separate episodes of fitness for females in both years and for males in one year. Specific components of body size were often the targets of the selection on overall body size. When net adult fitness was estimated by combining each individual's fitnesses from all episodes, we found stabilizing selection in both sexes. In addition, the net optimum overall body size of males was smaller than that of females. However, even when components of body size had experienced opposing selection pressures in individual episodes, no components appeared to be under lifetime stabilizing selection. This is the first evidence that contemporary selection in a natural population acts to maintain female size larger than male size, the most common pattern of SSD in nature.

Lifetime selection on adult body size and components of body size in a waterstrider: opposing selection and maintenance of sexual size dimorphism.

Sexual size dimorphism (SSD), the difference in body size between males and females, is common in almost all taxa of animals and is generally assumed to be adaptive. Although sexual selection and fecundity selection alone have often been invoked to explain the evolution of SSD, more recent views indicate that the sexes must experience different lifetime selection pressures for SSD to evolve and be maintained. We estimated selection acting on male and female adult body size (total length) and components of body size in the waterstrider Aquarius remigis during three phases of life history. Opposing selection pressures for overall body size occurred in separate episodes of fitness for females in both years and for males in one year. Specific components of body size were often the targets of the selection on overall body size. When net adult fitness was estimated by combining each individual's fitnesses from all episodes, we found stabilizing selection in both sexes. In addition, the net optimum overall body size of males was smaller than that of females. However, even when components of body size had experienced opposing selection pressures in individual episodes, no components appeared to be under lifetime stabilizing selection. This is the first evidence that contemporary selection in a natural population acts to maintain female size larger than male size, the most common pattern of SSD in nature.

Evidence of genetic isolation between sexually monomorphic and sexually dimorphic traits in the water strider Aquarius remigis

Estimates of multivariate selection are becoming increasingly common in the literature. Although it is clearly recognized that genetic information is essential to understanding natural selection, few studies include estimates of genetic parameters. Previous studies of the water strider Aquarius remigis have demonstrated selection on body size and components of body size that may favour the evolution and maintenance of sexual size dimorphism. In this paper we present quantitative genetic estimates of heritabilities and genetic correlations for male and female A. remigis. Overall body size (total length) and the majority of components of body size were significantly heritable for both sexes and the shape of the genetic correlation matrix was similar between the sexes. However, patterns of genetic correlations between the sexes and analysis of genetic integration suggest that some traits may be genetically isolated.

Evidence of genetic isolation between sexually monomorphic and sexually dimorphic traits in the water strider Aquarius remigis

Estimates of multivariate selection are becoming increasingly common in the literature. Although it is clearly recognized that genetic information is essential to understanding natural selection, few studies include estimates of genetic parameters. Previous studies of the water strider Aquarius remigis have demonstrated selection on body size and components of body size that may favour the evolution and maintenance of sexual size dimorphism. In this paper we present quantitative genetic estimates of heritabilities and genetic correlations for male and female A. remigis. Overall body size (total length) and the majority of components of body size were significantly heritable for both sexes and the shape of the genetic correlation matrix was similar between the sexes. However, patterns of genetic correlations between the sexes and analysis of genetic integration suggest that some traits may be genetically isolated.

Sexual Size Dimorphism and Selection in the Wild in the Waterstrider Aquarius remigis: Lifetime Fecundity Selection on Female Total Length and Its Components

Sexual Size Dimorphism and Selection in the Wild in the Waterstrider Aquarius remigis: Lifetime Fecundity Selection on Female Total Length and Its Components

SEXUAL SIZE DIMORPHISM AND SELECTION IN THE WILD IN THE WATERSTRIDER AQUARIUS REMIGIS: LIFETIME FECUNDITY SELECTION ON FEMALE TOTAL LENGTH AND ITS COMPONENTS.

Darwin's fecundity advantage model is often cited as the cause of female biased size dimorphism, however, the empirical studies of lifetime selection on male and female body size that would be required to demonstrate this are few. As a component of a study relating sexual size dimorphism to lifetime selection in natural populations of the female size-biased waterstrider Aquarius remigis (Hemiptera: Gerridae), we estimated coefficients for daily fecundity selection, longevity selection, and lifetime fecundity selection acting on female body size and components of body size for two consecutive generations. Daily fecundity was estimated using females confined in field enclosures and reproductive survival was estimated by twice-weekly recaptures. We found that daily fecundity selection favored females with longer total length through direct selection acting on abdomen length. Longevity selection favored females with smaller total length. When daily fecundity and reproductive longevity were combined to estimate lifetime fecundity we found significant balancing selection acting on total length in both years. The relationship between daily fecundity and reproductive longevity also reveals a significant cost of reproduction in one of two years. We relate these selection estimates to previous estimates of sexual selection on male body size and consider the relationship between contemporary selection and sexual size dimorphism.

A New Species of Aquarius from the Southeastern United States, with Electrophoretic Analysis of the Clade Containing Gerris, Limnoporus, and Aquarius (Hemiptera: Gerridae)

The taxonomic status of the common North American stream-dwelling waterstrider Aquarius remigis Say is poorly understood. We describe a new species ( Aquarius remigoides ) based on genetic data and provide 2 characters that allow specific diagnosis. In addition, we furnish a phylogenetic tree based on genetic distance estimates for 11 waterstrider species, and compare these relationships to those of Andersen (1993) based on morphological characters.

Sexual size dimorphism and selection in the wild in the waterstrider Aquarius remigis: Body size, components of body size and male mating success

AbstractSexual size dimorphism is assumed to be adaptive and is expected to evolve in response to a difference in the net selection pressures on the sexes. Although a demonstration of sexual selection is neither necessary nor sufficient to explain the evolution of sexual size dimorphism, sexual selection is generally assumed to be a major evolutionary force. If contemporary sexual selection is important in the evolution and maintenance of sexual size dimorphism then we expect to see concordance between patterns of sexual selection and patterns of sexual dimorphism. We examined sexual selection in the wild, acting on male body size, and components of body size, in the waterstrider Aquarius remigis, as part of a long term study examining net selection pressures on the two sexes in this species. Selection was estimated on both a daily and annual basis. Since our measure of fitness (mating success) was behavioral, we estimated reliabilities to determine if males perform consistently. Reliabilities were measured as ϰ statistics and range from fair to perfect agreement with substantial agreement overall. We found significant univariate sexual selection favoring larger total length in the first year of our study but not in the second. Multivariate analysis of components of body size revealed that sexual selection for larger males was not acting directly on total length but on genital length. Sexual selection for larger male body size was opposed by direct selection favoring smaller midfemoral lengths. While males of this species are smaller than females, they have longer genital segments and wider forefemora. Patterns of contemporary sexual selection and sexual size dimorphism agree only for genital length. For total length, and all other components of body size examined, contemporary sexual selection was either nonsignificant or opposed the pattern of size dimporhism. Thus, while the net pressures of contemporary selection for the species may still act to maintain sexual size dimorphism, sexual selection alone does not.