Mate disruption in the water strider Aquaris remigis Say, 1832 (Hemiptera: Gerridae)

This report describes the behaviour of a previously uncharacterised mating tactic amongst male water striders (Aquarius remigis Say, 1832), involving the physical break up of existing mating pairs. Using data from four separate laboratory observations on the water strider mating system, we show that this behaviour represents roughly 12.6% of all mating attempts by males. Furthermore, we demonstrate that males are successful in breaking up the existing mating pair in 15.6% of the cases, resulting in secure matings with the disrupted female 6.2% of time. We suspect that mate disruption may serve as an alternative means for acquiring mates by males with low mating success using conventional behaviours. Further research should be performed to determine the prevalence of this behaviour in various natural populations and the specific contexts in which this behaviour occurs.     

Genital morphology and taxonomy of the water strider Aquarius remigis (Say) (Insecta, Hemiptera-Heteroptera: Gerridae)

The shape of the dorsal vesical plate was used to determine the extent of morphological differentiation among populations of the water strider Aquarius remigis (Say) (Hemiptera-Heteroptera: Gerridae), and between species assigned to the A. remigis species group. Populations were sampled throughout North America and Mesoamerica, and included paratypes of A. remigoides Gallant & Fairbairn, and material collected from the type localities of A. amplus (Drake & Harris) and A. nyctalis (Drake & Hottes). If taxonomy is to be inferred from the shape of the dorsal vesical plate, as suggested by previous authors, several taxonomic issues need to be reconsidered. First, there was one species distributed widely across North America and deep into Mexico that is very likely to represent A. remigis , meaning that populations from the western USA previously assigned to ' A. nyctalis ' should be synonymized with this species. Second, A. remigoides as currently defined also includes some populations of A. remigis , and is probably much less widely distributed in the south-eastern USA than previously thought. A hybridization zone in Pennsylvania between A. remigis and A. remigoides , as inferred from measurements of body size and allozyme electrophoresis, is not congruent with data from the male genitalia, which indicated a transition zone between the two forms in Virginia much further to the south. Finally, A. amplus was widespread in Mexico, reaching into Guatemala to the south and into Arizona to the north.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 381–398.     

A test of the hypothesis of compensatory upstream dispersal using a stream-dwelling waterstrider,Gerris remigis Say

Summary This paper reports the results of 2 experiments designed to determine if adult Gerris remigis (Hemiptera, heteroptera), a stream-dwelling waterstrider, tend to disperse preferentially upstream, as predicted by the colonization cycle hypothesis summarized by Muller (1982). Markrecapture observations and experimental removals were used to assess the distance and direction of movement of adult G. remigis along a small mountain stream, over a full year. These experiments indicated that adult G. remigis show a significant upstream bias in movement distance, but not in numbers of animals moving. This upstream bias is characteristic of pre-reproductive, sexually immature adults of both sexes, and occurs primarily in association with movements to and from diapause sites. Although the existence of a significant upstream bias in movement distances tends to support the colonization cycle hypothesis, the data from the removal experiments clearly show that upstream dispersal is not sufficient to compensate entirely for downstream drift.     

The origins of allometry: size and shape polymorphism in the common waterstrider, Gerris remigis Say (Heteroptera, Gerridae)

Changes in size, whether ontogenetic or phylogenetic, tend to be associated with changes in shape. This allometry can arise through two different evolutionary mechanisms: (1) selection acting primarily on overall size may be associated with changes in shape because of physiological and mechanical constraints or differential responses of different body components; or (2) selection acting primarily on shape (on the size of specific body components) may be associated with changes in overall size because of genetic correlations, and thus correlated responses, of other body components. To assess the relative importance of these two mechanisms, shape polymorphism is examined along two axes of size dimorphism (sex and wing morphology) in the common waterstrider, Gerris remigis Say. Eight measurements were made of body and appendage components of 234 adults, from three independent populations. Univariate and multivariate analyses reveal that both sexes and wing morphs differ significantly in size and shape. Shape differentiation along the two axes of size dimorphism is found to be dissimilar, partially independent of size, and strongly correlated with the ecological specialization of the various morphs. These observations suggest that selection is acting directly on shape, and thus that allometry in this species primarily reflects shape-mediated changes in size (mechanism 2), rather than size-mediated changes in shape. The role of developmental processes in facilitating this shape differentiation is discussed.     

Assembly of the fluorescent acrosomal matrix and its fate in fertilization in the water strider,Aquarius remigis

Animal sperm show remarkable diversity in both morphology and molecular composition. Here we provide the first report of intense intrinsic fluorescence in an animal sperm. The sperm from a semi‐aquatic insect, the water strider, Aquarius remigis, contains an intrinsically fluorescent molecule with properties consistent with those of flavin adenine dinucleotide (FAD), which appears first in the acrosomal vesicle of round spermatids and persists in the acrosome throughout spermiogenesis. Fluorescence recovery after photobleaching reveals that the fluorescent molecule exhibits unrestricted mobility in the acrosomal vesicle of round spermatids but is completely immobile in the acrosome of mature sperm. Fluorescence polarization microscopy shows a net alignment of the fluorescent molecules in the acrosome of the mature sperm but not in the acrosomal vesicle of round spermatids. These results suggest that acrosomal molecules are rearranged in the elongating acrosome and FAD is incorporated into the acrosomal matrix during its formation. Further, we followed the fate of the acrosomal matrix in fertilization utilizing the intrinsic fluorescence. The fluorescent acrosomal matrix was observed inside the fertilized egg and remained structurally intact even after gastrulation started. This observation suggests that FAD is not released from the acrosomal matrix during the fertilization process or early development and supports an idea that FAD is involved in the formation of the acrosomal matrix. The intrinsic fluorescence of the A. remigis acrosome will be a useful marker for following spermatogenesis and fertilization. J. Cell. Physiol. 226: 999–1006, 2011. © 2010 Wiley‐Liss, Inc.     

Presence of the water lily,Trapa natans L., increases fitness of the water strider,Gerris nepalensis

Abstract Exposure of the waterstrider, Gerris nepalensis, to leaves of the water lily, Trapa natans, during either the larval or adult stage increases the proportion of reproductive females (60.0%) and increases the number of eggs laid by G. nepalensis (25.1 ± 8.1) compared with leaves of another floating plant, Hydrocharis dubia (20.2%, 6.7 ± 17.8), and mimic leaves made of polystyrene (24.2%, 20.0 ± 16.9). The larval period at 25 °C is significantly shorter when larvae are reared together with water lily leaves than when reared with mimic leaves made of thin styrene. A significantly higher percentage (76.4 ± 39.9) of eggs laid by females that are reared with lily leaves in larval and adult stages develop successfully to the first instar compared with those reared with ‘mimic‐leaves’ (% hatched‐out successfully: 53.9 ± 39.3). The effect of T. natans on G. nepalensis demonstrated in the present study is to increase the number of G. nepalensis in the habitat and likely increase foraging pressure on the lily leaf beetle, Galerucella nipponensis. Possible mechanisms of this relationship between T. natans and G. nepalensis are discussed.     

Fitness consequences of foraging success in water striders (Gerris remigis; Heteroptera: Gerridae)

Foraging theory's central premise assumes that behavioral actionsthat affect the acquisition of food also ultimately affect fitness.However, very few investigations in behavioral ecology actuallydemonstrate ultimate fitness consequences of any particularbehavior. Many studies focus on short-term estimates of fitnessin animals with considerable life spans. I investigated whetherdifferences in foraging performance, as related to patch choiceand behavioral dominance, show consistent patterns over extendedperiods of an animal's life such that they translate into differencesin reproductive success. In one experiment, I focused on theability of single female water strider (Gerris remigis) to discriminatebetween foraging patches of differing prey abundances in laboratorystreams simulating natural conditions. In another experiment,I focused on competitive interactions within groups of threefemale water striders foraging in laboratory streams where resourceavailability varied with position in a patch. The first experimentshowed that, overall, the individuals detected and preferredto forage in the richer patch, which directly increased theirforaging success and their lifetime fecundity. However, therewas marked variability among individuals in their ability torespond to the differences in food availability, ranging fromno tracking to fast tracking when the patch qualities were switched.In the second experiment, both the foraging position of individualsin the stream and their dominance rank were reasonably consistentover their entire reproductive life. Foraging position and dominancerank were significant predictors of lifetime fecundity, theformer being the better predictor due to an imperfect correlationbetween the two variables. In both experiments, the increasein fecundity was achieved by higher oviposition rates ratherthan by extending the oviposition period or by inducing firstreproduction at an earlier age. [Behav Ecol 1991; 2: 46-55)     

Predation of frog eggs by the water strider Gerris latiabdominis Miyamoto (Hemiptera: Gerridae)

Water striders (Hemiptera: Gerridae) are predators found on the water surface that prey mainly on arthropods. The feeding on other organisms (dead vertebrates, fishes and tadpoles) is a rare event. To our knowledge, predation of frog eggs by water striders has not yet been reported. We observed that adult water striders, Gerris latiabdominis Miyamoto, 1958 (Hemiptera: Gerridae), preyed on the eggs of three frog species, Pelophylax nigromaculata Hallowell, 1861 (Anura: Ranidae), Rana japonica Boulenger, 1879 (Anura: Ranidae), and Rhacophorus schlegelii Günther, 1858 (Anura: Rhacophoridae). We found predation by the water striders affects the survival rate of frog eggs floating on the water surface. We suggest that this hunting event would occur in water bodies in which water striders and frogs coexist, especially the region where their phenology overlaps.     

Mate guarding and sperm displacement in the water strider Gerris lateralis Schumm. (Heteroptera: Gerridae)

SUMMARY. 1. Field and laboratory observations on the mating behaviour of Gerris lateralis Schumm. allowed three distinct phases to be distinguished: (i) a precopulatory phase (1.5min, SD=1.3), (ii) copulation (16.2min, SD=12.9), and (iii) a postcopulatory phase. The duration of the postcopulatory phase, during which the male rode passively on the back of the female without genital contact, varied considerably (11 min to > 48h). Females appeared reluctant in all matings, and matings were forced by males.
2. Laboratory experiments showed that the females were able to store sperm for more than 30 days without decrease in fertilization rate. In double mating experiments, where partially sterilized males were used, it was demonstrated that sperm displacement was extensive. The last male to mate fertilized approximately 80% of the eggs.
3. It is concluded that the postcopulatory behaviour is beneficial to males in terms of paternity assurance, and it is interpreted as a mate guarding behaviour.     

Conflict between sexes in the water strider, Gerris lacustris: a test of two hypotheses for male guarding behavior

We studied the effect of operational sex ratio on female reluctanceand male persistence to mate as well as on the length of copulationand postcopulatory guarding in Gerris lacustris by adding fivesurplus males or females to the basin with a pair in tandem.In the control treatment, a pair alone was tested. Accordingto the copulatory guarding hypothesis (CGH), males should prolongmating and guard females in the presence of surplus males. Accordingto the convenience polyandry hypothesis (CPH), females shouldshow lower levels of resistance to prolonged mating in the presenceof surplus males because the mating male protects the femaleagainst harassment from other males. As expected on the basisof both the CGH and CPH, mating (copulation + guarding) averagedlonger in the male-biased treatment. The behavior of males andfemales during mating suggested that both hypotheses hold true:females showed less resistance to prolonged mating (as predictedfrom CPH), and male behavior suggested stronger efforts to stayon the female when surplus males were present (as predictedfrom CGH). Comparisons of the treatment with surplus femaleswith the results from the mating pair without surplus individualssuggested that the capabilities of water striders in tandemto assess the sex of nearby nonmating striders are limited.     

Experiments on the effects of food and density on voltinism in a stream-dwelling water strider (Aquarius remigis)

1. The stream water strider Aquarius remigis shows a latitudinal pattern of variation in voltinism. In general, populations with shorter growing seasons (e.g. in eastern Canada) tend to be univoltine (animals that reach adulthood in the summer overwinter before reproducing in the following spring), whereas populations with somewhat longer growing seasons (e.g. in the north-eastern United States) tend to be bivoltine. 2. This pattern was broken at our study site in the south-eastern United States (Kentucky) where A. remigis had a long growing season, but was almost always univoltine. In summer 1993, however, adult A. remigis in central Kentucky displayed a bivoltine reproductive cycle; i.e. individuals in some pools began breeding shortly after maturing to the adult stage. 3. A field survey documented a negative relationship between local water strider density and reproductive activity in prediapause adults. A laboratory experiment manipulating food availability and density, revealed that animals held at low density with high food levels displayed greater mating activity and egg production than did their counterparts at higher density or lower food levels. 4. A laboratory experiment also showed that high water strider density resulted in a greater frequency of very short pair durations (< 10 min). 5. Although the observed effects of density and food availability on mating activity of prediapause adults seem intuitively reasonable, they differ from the patterns observed in overwintered adults. The difference in reproduction patterns might be due to differences in selective pressures on prediapause vs. post-diapause adults.     

Influence of gut parasites on growth performance in the water strider Gerris buenoi (Hemiptera: Gerridae)

Water striders harbor a diverse assemblage of symbionts in their digestive tract We used a field experiment, in which water striders were reared in enclosures in their natural pond habitat, to assess the effects of gut symbionts on growth Trypanosomatid flagellates had significant adverse effects on both development time and adult size, and therefore are clearly parasitic Yet because of their low prevalences (2% or less), trypanosomatids cannot be a major factor in the dynamics of our study population Gregannes occurred in 36% of the water striders, often in high numbers, and filling the entire midgut of some bugs Nevertheless, infected and uninfected gerrids did not differ in their growth, and gregarine loads were uncorrelated with development time and adult size attained We also did not find effects of gregarines in a second experiment with different rearing conditions, including a treatment with food stress We used a quantitative genetic approach to test if resistance against gregarines has a heritable component There was no evidence for any genetic variation, suggesting that variability in gregarine loads is the result of environmental heterogeneity Comparison with published reports from water striders shows that there is a great variability in the diversity and prevalence of symbionts among different populations and species, of gerrids     

Dispersion and spatial orientation of Gerris remigis in response to water current: a comparison of pre- and post-diapause adults

ABSTRACT. The movements and orientation of adult Gerris remigis Say (Hemiptera; Gerridae) were examined in an artificial stream to determine the mechanisms by which these stream-dwelling waterstriders compensate for and/or avoid downstream drift, and to compare the movement tendency of pre- and post-diapause adults. G. remigis respond to current by turning to face directly into the current (rotational orientation), intermittently rowing against the current (translational orientation), and moving onto land to avoid the current. These rotational and translational responses enable G. remigis to maintain their positions in currents =S10 cm s^-1. However, avoidance increases with current speed, and few animals are found where currents exceed 10 cm s^-1. These observations correlate well with the observed distribution of adult G. remigis in a natural stream.
Comparisons between life stages reveal that post-diapause adults move more, and are more susceptible to drift than pre-diapause adults. Post-diapause adults also avoid even slight currents (<5 cm s^_1) by seeking land, while pre-diapause adults do not show a significant preference for land until currents exceed 10 cm s^-1. These data suggest that previously reported differences in the movement tendency of pre- and post-diapause adults in the field may reflect physiologically-based behavioural differences between these two life stages rather than merely direct responses to environmental cues.     

Allometry for sexual size dimorphism: testing two hypotheses for Rensch's rule in the water strider Aquarius 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.     

Life history adaptation along a latitudinal cline in the water strider Aquarius remigis (Heteroptera: Gerridae)

We addressed the general hypothesis that life history differences among eastern populations of the North American water strider, Aquarius remigis (Heteroptera: Gerridae), along a north-south gradient are manifestations of genetic differentiation due to natural selection. We raised offspring of two field-caught populations from each of three latitudes in a common laboratory environment at 20° C and two photoperiods. Nearly all Quebec (PQ) individuals (ca. 46° N) entered diapause to reproduce the following spring (univoltine life cycle), while intermediate proportions of New York (NY; ca. 43° N) and New Jersey (NJ; ca. 41° N) individuals reproduced directly, producing a second generation (bivoltine life cycle). PQ females were smaller, developed faster, and laid smaller eggs than NY and NJ individuals; NY and NJ populations differed little in these variables. NY females had longer life spans than either PQ or NJ females, but lower oviposition rates. Total reproductive output did not differ across latitudes. Photoperiod affected body length, development time, and reproductive pathway, resulting in a latitude by environmental interaction. PQ individuals reproduced directly under 15L : 9D (summer) conditions only, while the NY and NJ populations exhibited more direct reproduction under 13L : 11D (spring or fall) conditions. Some life history characters of the NY and NJ populations displayed the higher variability indicative of phenological transition zones. These results indicate local adaptation of populations to long-term climatic patterns. Water striders appear to adapt to longer seasons by extending development, growing larger, and breeding directly. Larger body size and extended or rapid development are associated with bivoltinism and increase in egg size, but not necessarily with higher fecundity or oviposition rate. The phenological transition zone appears to be unrelated to a transition zone a little further south established by allozyme data and morphology, as all populations studied here could be electrophoretically identified as northern "type".