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".     

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.     

Fitness consequences of alternative life histories in water striders,Aquarius remigis (Heteroptera: Gerridae)

Using field and laboratory observations and experiments over 3 years, I investigated whether reproductive trade-offs shape individual life histories in two natural populations of the water strider, Aquarius remigis, in which univoltine and bivoltine life cycles coexist. Both later eclosion dates and food shortages, even after adult eclosion, induced diapause in females, thus deferring reproduction to the following spring. Adult body size was positively affected by food availability during juvenile development. Higher food levels also increased the reproductive output of females, but not their longevity or oviposition period. When compared to spring breeders (univoltine life cycle), direct (summer) breeders (bivoltine life cycle) experienced reduced lifetime egg numbers and longevity, as well as reduced survivorship of their second-summer-generation offspring; these reproductive costs offset, at least in part, the advantage in non-decreasing populations of having two generations per year. Fecundity was correlated with body size, and among summer-generation females direct breeders were larger than non-breeders. The time remaining before the onset of winter and/or the time since adult eclosion augmented cumulative energy uptake, and consequently the lipid reserves and winter survival probability of non-breeding (diapausing) summer adults approaching hibernation. Overwintered spring reproductives died at faster rates than non-reproductive summer individuals despite greater food availability in spring, indicating a mortality cost of reproduction. Body length correlated with absolute and not with proportional lipid content but showed no consistent relationship with survivorship in the field. These results are in agreement with current theory on the evolution of insect voltinism patterns, and further indicate high degrees of life history flexibility (phenotypic plasticity) in the study populations in response to variable environmental factors (notably photoperiod and food availability). This may be related to their location in a geographic transition zone from uni- to bivoltine life cycles.     

The histology and histochemistry of oogenesis in the water strider, Gerris remigis Say

In each ovariole of Gerris remigis, nurse cells arise by mitotic divisions at the anterior end of the germarium. These cells enlarge as they move posteriorly. This size increase is possibly caused by fusion of cells, but probably by endopolyploidy as well. The nurse cells then establish connections with a central trophic core, which receives the products of subsequent nurse cell degradation. Two possible pathways of nuclear degradation are suggested: one involves the condensation of chromatin within the nucleus; the other, the release of DNA as fine granules into the cytoplasm. Cytoplasmic areas containing such DNA are also rich in proteinaceous granules, but have a meager content of RNA. The remainder of the cytoplasm of the mature nurse cells contains a high concentration of RNA, as do the nucleoli. Posteriorly the trophic core connects via nutritive cords with each developing oocyte in the prefollicular region and in the anterior vitellarium. RNA is apparently contributed to the ooplasm via the trophic stream. Patches of cytoplasmic DNA are present in the young oocytes; the origin and fate of this DNA is uncertain. During early oocyte maturation chromosomal stainability decreases, and the nucleolus enlarges. In previtellogenic stages, numerous proteinaceous bodies appear in association with the nucleolus-chromosome complex. These bodies, like the nucleolus, have only a low RNA content. They may pass to the cytoplasm, but cannot be traced with certainty. During the latter part of this period a complex population of small proteinaceous and lipid preyolk bodies accumulates peripherally in the oocyte. Definitive protein and lipid yolk are probably derived by the enlargement and inward migration of these bodies. The oocytes are each surrounded by a layer of follicle cells proliferated in the prefollicular region. These become binucleate and enlarge as the enclosed oocytes grow and elongate. RNA also increases in the nucleoli and cytoplasm of the follicle cells as they move posteriorly in the vitellarium. There is no evidence of transfer of nucleic acids or protein from the follicle cells to the oocyte. The nurse cells are therefore implicated as the major source of nucleic acids for the maturing oocyte.     

Costs of loading associated with mate-carrying in the waterstrider, Aquarius remigis

When one sex carries the other during some phase of courtshipor mating, the associated loading may entail a significant costto the carrier. This paper presents a series of laboratory experimentsdesigned to identify the costs of mate-carrying in Aquariusremigis. Female A. remigis mate repeatedly and carry each matefor several hours. Dead males and lead weights were used tosimulate normal mating and loading associated with mate-carrying,respectively. Females carrying weights equivalent to the weightof an average male showed no detectable reductions in survival,lipid reserves, or foraging success, and maintained themselveson the water surface for more than 10 days without access toresting sites. Weights equivalent to two males were supportedfor 6.1 ± 4.9 days. Thus, female A. remigis appear tobe very well adapted to carrying their mates and are unlikelyto be near their load limits when carrying a single mate. However,females carrying males or equivalent weights suffered a significantreduction in maximum mobility (stride length and speed), andan increased risk of predation by frogs (Rana clamitans). Femalescarrying weights were more susceptible to predation than unburdenedfemales but were less susceptible than females carrying males,suggesting that loading contributes significantly to, but doesnot fully explain, the increased predation risk. This risk probablyresults from both reduced mobility due to loading and greatervisibility (size). Possible influences of the costs of loadingon mating behavior and sexual size dimorphism are discussed.     

The sexual arms race and phenotypic correlates of mating success in the waterstrider,Aquarius remigis (Hemiptera: Gerridae)

Intersexual conflicts over mating decisions may have an important impact on the evolution of mating behaviours and strategies and may develop into an arms race between the sexes. In waterstrider species, where intersexual conflict is known to occur, the evolution of male traits that allow them to overcome female reluctance to mate is expected. Reproductively active waterstriders,Aquarius remigis, were videotaped in the laboratory to examine the influence of total body length and front femoral width of males, male: female body size ratio, and female reproductive condition (number of mature eggs) on three variables associated with mating success: duration of the premating struggle, duration of mating, and number of successful matings. None of these behaviors was significantly correlated with the size ratio of the mating pair. However, total body length was negatively correlated with premating struggle duration, male front femoral width was positively correlated with number of successful matings, and number of mature eggs in females was positively correlated with duration of both the premating struggle and the mating itself. The relative influence of male sexual armaments and female choice on the outcome of mating interactions is discussed.     

Body size and fecundity in the waterstrider Aquarius remigis: a test of Darwin's fecundity advantage hypothesis

The general female bias in body size of animals is usually attributed to fecundity selection. While many studies have demonstrated a positive relationship between body size and fecundity, the most common interpretation of fecundity selection is that larger females have larger abdomens and can hold more eggs, yet the relationship between abdomen size and fecundity has rarely been examined. For the waterstrider, Aquarius remigis, we find a significant relationship between body size and fecundity and demonstrate that the target of fecundity selection is abdomen size. Thus, larger females have higher fecundities because they have larger abdomens and not because of their total size per se. The rate at which fecundity increases with increasing abdomen size exceeds that which would be expected due to a simple volume constraint and suggests that other factors, such as increased ability to obtain resources, may contribute to the increase in fecundity with body size. Selection intensities estimated from our data indicate that fecundity selection could be a significant selective force on both total and abdomen lengths. Previous studies have found that abdomen size increased faster than body size and thus, larger females had relatively larger abdomens. The relationship of abdomen length and thorax length in A. remigis is hypoallometric and indicates that larger females have relatively smaller abdomens. We hypothesize that this may reflect conservation of abdomen size in females developing under poor conditions. Finally, while egg size is not directly related to body size, we find a trade-off between egg size and number when female abdomen length is held constant, suggesting that selection on egg size may influence abdomen length only indirectly through its effects on fecundity.     

Glinal variation in morphology among eastern populations of the waterstrider, Aquarius remigis Say (Hemiptera: Gerridae)

We examined morphological variation among populations of the stream-dwelling waterstrider, Aquarius remigis. A previous analysis of allozyme variation along a transect from southern Quebec to southern North Carolina revealed two distinct 'types' of A. remigis, north and south of a transition zone in southern New York and northern Pennsylvania. To assess the concordance of morphological and allozyme differentiation, adults were sampled from 15 populations along the original transect, as well as 11 additional populations in New York, Connecticut and New Jersey. Eight measurements of leg and body size were taken of preserved individuals. Univariate and multivariate analyses (multiple regression, PCA, discriminant analysis) reveal significant differentiation between northern and southern populations with a transition zone between latitudes 39^oand 42^onorth. Strong clines within each region and a clear transition between regions support previous suggestions that there are two distinct 'types' of this species in eastern North America. The transition zone, in morphology coincides with that revealed by the allozyme variation, but extends further south. The covariance of allozyme frequency and morphometric traits suggests that differentiation has occurred through vicariance and secondary contact, and that in this system, cladistic and adaptive differentiation are coupled.     

Vibratory signals enhance mate-guarding in a water strider (Hemiptera: Gerridae)

Mating males of the water strider Gerris remigisproduce vibratory signals when-single males grasp mating pairs. When played through live females with dead males on their backs, these signals repelled mating attempts by single males. A previous study showed that male mate-guarding enhances female foraging effectiveness in this species. Thus male mate-guarding signals also enhance female foraging effectiveness.     

Direct and indirect effects of multiple enemies on water strider mating dynamics

Many different measures of range size are used for both empirical and conservation purposes. The possible consequences of the particular methods used in determining observed patterns of results are seldom considered. Using species of butterflies and freshwater molluses in Britain, we investigate the relationship between the range sizes measured by nine different methods and the sets of rare species they distinguish. A comparison of range sizes measured at different scales. Britain and Europe, is also made for the butterflies. We find that for many studies involving range size the various measures of range size are interchangeable. With respect to the identification of rare species the results are not as clear.     

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.     

Phylogeny of the water strider genus Rheumatobates (Heteroptera: Gerridae)

The genus Rheumatobates comprises thirty‐seven species and subspecies of New World water striders belonging to subfamily Rhagadotarsinae. Among species, males vary dramatically in the degree and nature of modifications of the antennae, three pairs of legs and abdominal and genital segments. Characters describing this modification have traditionally been used to differentiate and group species. The general assumption has been that modified species belong to one group and unmodified species to another. These two ‘species groups’ are subdivided into ‘subgroups’, but little effort has been made to resolve relationships among them. We conduct the first numerical cladistic analysis of Rheumatobates using a data set comprised of 102 characters, primarily describing modification of male external morphology. To address concerns about the inclusion of characters to be optimized on the phylogeny, characters describing modification of antennae and hind legs were included and then excluded in separate analyses. A preferred phylogeny was chosen from the four equally parsimonious cladograms found after successive reweighting of characters. There was good resolution at all levels of the phylogeny. Most of the major clades and terminal relationships were moderately to strongly supported, whereas the basal relationships were less well supported. The general assumption that unmodified and modified species form two monophyletic groups was not supported. However, traditionally recognized ‘subgroups’ within the modified species group were largely upheld. The analysis also suggested several major clades and relationships among these clades that were not previously recognized. The exclusion of characters describing modification of antennae and hind legs did not change the resolved major clades of the reconstructed phylogeny.     

Fishing spiders, green sunfish, and a stream-dwelling water strider: male-female conflict and prey responses to single versus multiple predator environments

Many studies have experimentally addressed the effects of a particular predator species on prey behavior. In nature, however, prey frequently face multiple species of predators that often vary in their predatory mode and in their level of predation risk. Relatively few studies have considered prey responses under these complex conditions. In Kentucky, the stream-dwelling water strider (Aquariusremigis) coexists with many potentially dangerous predators, two of which are the green sunfish (Lepomiscyanellus) and the fishing spider (Dolomedesvittatus). Green sunfish occupy stream pools and attack water striders from below. In contrast, fishing spiders hunt along stream shorelines where they perch on overhanging vegetation or rocks and attack water striders near shore. We compared how A. remigis individuals respond to these two very different predators in pools with one or both predators. The presence of sunfish in pools had strong effects on male water strider behavior, including increased use of three types of refuge from sunfish (riffles, climbing out of the water, sitting on the water but at the edges of pools), decreased activity and a decreased number of aggressive males on the water. Spiders also influenced water strider behavior; male water striders avoided spiders by shifting away from the edges of pools. Comparisons of the effects of the two predator species showed that in general, antipredator responses by male water striders were stronger in pools with fish alone than in those with spiders alone. In the presence of both predators, male water strider behavior (microhabitat use and activity) was generally similar to behavior in the presence of fish alone. In contrast, female water striders showed no significant response to the presence of sunfish, and little response to the presence of spiders. This lack of response could be because females spent much of their time in refuges even in the absence of predators (apparently hiding from harassment by males). Both spiders and fish caused decreases in water strider mating activity. The presence of fish reduced both the number of matings per pool (mating frequency), and mean mating durations. Spiders induced a decrease in mean mating duration, but not in mating frequency. The largest reductions in mating activity occurred in pools with both predators present. Pools with either spiders or fish alone suffered 15–20% water strider mortality during our experiment (versus no mortality in predator-free pools). Extant theory suggests that when prey face conflicting microhabitat responses to two predators (as in this study), the predators should have facilitative effects on predation rates (i.e., prey that avoid one predator are often killed by the other and vice versa). Mortality rates in pools with both predators present, however, were not significantly different from that predicted by a null model of multiple predator effects. The lack of predator facilitation can be explained by the compensatory reductions in water strider activity and mating activity in the presence of both predators. Received: 26 August 1996 / Accepted: 12 June 1998     

The realized food niches of three species of stream-dwelling triclads (Turbellaria)

SUMMARY. 1. The diet of Phagocata vitta, Crenobia alpina and Polycelis felina in a Welsh stream was examined, using a serologicat technique, at monthly intervals over 18 months.
2. For each triclad species, dietary composition changed little with time or increasing body size, and overall feeding intensity was greatest in summer and least in winter.
3. Alt species fed most heavily on oligochaetes, chironomids and Gammarus. Phagocata vitta fed lightly on Plecoptera, Trichoptera and Ephemeroptera whilst the other two triclad species fed substantially on the first two of these prey categories.
4. Food niche breadth indices indicated a wide diet for all species with some specialization in some months. Electivity indices showed positive electivity for oligochaetes and chironomids by all species and for Plecoptera by C. alpina and P. felina .
5. Diets overlapped and none had a food refuge: therefore the potential for interspecific competition for food exists.     

Antagonistic pre- and post-copulatory sexual selection on male body size in a water strider (Gerris lacustris)

A crucial question in sexual selection theory is whether post-copulatory sexual selection reinforces or counteracts conventional pre-copulatory sexual selection. Male body size is one of the traits most generally favoured by pre-copulatory sexual selection; and recent studies of sperm competition often suggest that large male size is also favoured by post-copulatory sexual selection. In contrast to this general pattern, this study shows that pre- and post-copulatory sexual selection act antagonistically on male body size in Gerris lacustris. One large and one small male were kept together with two females in this experiment. Large males had a significant mating advantage, but small males copulated longer and gained higher fertilization success from each mating. Large and small males, however, gained similar reproductive success, and there was no overall correlation between mating success and reproductive success. These results suggest that estimates of male fitness based solely on mating success should be viewed with caution, because of potentially counteracting post-copulatory selection.     

Signalling Asymmetry in the Communication of the Water Strider Aquarius remigis in the Context of Dominance and Spacing in the Non-mating Season

The role of high-frequency ripple signals (HF signals) made by males of the water strider Aquarius remigis was studied in the contexts of competition for food and general spacing behaviour during the non-mating season. HF signals were played back through the ripple-producing legs of males during dyadic agonisdc encounters, using a signal-driven wire coil to oscillate a magnet glued to a leg. These signals were in addition to normal signals. The additional signals significantly increased the number of retreats by non-magneted males, showing that the signals increased the dominance of a non-territorial male. Hence, our results increase the number of contexts in which HF signals of A. remigis function. Males, but not females, avoided a site occupied by a magneted dead male through which HF signals were played. Thus, the communication system used by A. remigis males during competition for food seemed to be ignored by females, suggesting sex-specific signals even in a non-mating context. Evolutionary models of signalling often assume that contestants have evolved the same repertoire of signals in order to resolve conflicts peacefully. This water strider system thus poses an interesting challenge for future theoretical and empirical research on communication asymmetry.     

Comparative effects of food and water deprivation on movement patterns in the pigeon (Columba Livia)

Groups of naive pigeons were either food or water deprived and exposed to response-independent fixed time schedules of reinforcement. Reinforcers were always appropriate to the deprivation state. After 15 Phase One sessions under the original deprivation state, an additional 15 Phase Two sessions were run under the alternate deprivation regimen. The Phase One results indicated that the frequency of movement over time, as measured by floorboard panels, had a characteristic distribution depending on deprivation state. After state-switch in Phase Two, both groups exhibited changes in frequency distribution over time, but the new distributions were different from those produced in subjects without any deprivation history. There appeared to be sustained effects on movement patterns due to prior exposure to alternate deprivation and periodic feeding regiments, results which confirm earlier experimental work. The findings are discussed in light of research on the effect of prior response-independent schedule training, the matching law, and the effects of contextual conditioning.     

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.