Water Strider Aquarius Paludum Research Articles

Morph-specific life-history correlations in a wing-dimorphic water strider.

Life-history theory predicts a negative correlation between reproduction and survival because individuals differ in their investment in early reproduction at the expense of survival. However, life-history trade-offs can be masked when individual differences in resource allocation are smaller than those in resource acquisition. In polymorphic species, as distinct morphs exhibit differences in intrinsic physiology, the relative effects of resource acquisition and allocation on life-history traits will differ between morphs, contributing to morph-specific life-history correlations. Here, in the wing-dimorphic water strider Aquarius paludum, we found that wing morphs differed in within-morph individual-level life-history correlations. Longer-lived flight-capable long-winged females produced fewer eggs per day and matured later, whereas life-history trade-offs were not observed in short-winged flightless females. The survival-reproduction trade-off observed in long-winged females may be a result of individual differences in the timing of wing muscle histolysis. Individuals that underwent wing histolysis early would have increased reproduction at the expense of a shorter life, whereas individuals with late wing histolysis would have reduced reproduction but a longer life. Short-winged females, who never develop wings, effectively have more resources to allocate to both survival and reproduction, masking any life-history trade-offs. Thus, we suggest that morph-specific effects of resource allocation trade-offs can shape the morph-specific extent of individual variation in life-history strategies, which may contribute to the evolution and maintenance of within-species polymorphism.

Does Population Density Determine the Mating Strategy in Males of the Water Strider Aquarius paludum?

Mating behavior of a large Aquarius paludum paludum (Fabricius 1794) (Hemiptera: Gerridae: Gerrini) population was investigated on several field surveys during the mating season. Both types of mating behavior known for semi-aquatic true bugs were observed in the studied population: aggressive Type I mating for non-territorial males and unaggressive Type II mating for the territorial males. Some males establish individual territories at the lake shores that they defend for around one hour and switch to aggressive mating if they fail to attract a female to mate, while most males remain non-territorial. Radial 1–1.5 meter territories were aggressively defended from other males, conflicts being always won by the male already occupying the territory. The presence of Type II males is conditioned by the absence of sufficient space for all males to establish territories. Contact guarding was much longer than documented before for A. paludum paludum, lasting for at least 12–13 hours in some cases. Based on the observations and on the existing information on the group, a more comprehensive classification of mating behavior of true bugs is proposed: Type I - non-territorial, unsignaled grasping, with contact guarding; Type II - territorial, signaled mounting with noncontact guarding; Type III - non-territorial, signaled mounting with contact guarding.

Parasitoid avoidance behavior is not triggered by airborne cues in a semi-aquatic bug

Until recently, studies on host-parasitoid interactions have rarely reported the detection and resistance of parasitoids by hosts prior to experiencing a direct attack. Several recent studies have indicated that some hosts have an ability to detect parasitoids without being attacked directly. However, the particular cues used for parasitoid detection have received little attention. We investigate the use of airborne cues (such as volatile chemicals or sound) by the water strider Aquarius paludum insularis (Heteroptera: Gerridae) for detecting its egg parasitoid wasp. Since A. paludum females with previous exposure to the parasitoids oviposit at deeper positions relative to unexposed females, we were able to infer whether A. paludum had detected the parasitoid from oviposition depth. We allowed A. paludum to oviposit after one of the three treatments: exposure to the parasitoid directly, airborne cues (the parasitoid was enclosed in the tube of which ends were sealed by mesh), or wasp absent. The oviposition depth was shallower in A. paludum with exposure to airborne cues than in A. paludum with exposure to the parasitoids directly. The depth did not differ between airborne cues and wasp absent treatments. These results indicate that A. paludum were not able to detect the parasitoid from airborne cues alone.

Potential Costs of Selecting Good Sites For Offspring: Increased Risk of Drowning And Negative Effects on Egg Production

AbstractMany studies have shown the benefits of selecting suitable sites for offspring survival and growth. However, costs of selecting suitable sites have been little covered. If the female's costs associated with selecting suitable sites exceed the benefit from improvement of offspring performance, selecting such sites can have a negative effect on the ongoing reproductive success for ovipositing females. We investigate the potential costs of selecting suitable sites in the water strider Aquarius paludum insularis. Where there exists a risk of egg parasitism, the female A. paludum will submerge and select a deep site, more suitable for offspring survival, for oviposition. By forcing A. paludum underwater once a day for 10 d, we investigated the potential costs associated with oviposition at deep sites, firstly relating to the performance of submergence (latency to asphyxiation or proportion of buoyancy loss) and, secondly, in the number of eggs laid. Buoyancy became weaker and the latency to asphyxiation became shorter with the number of submergences. The number of eggs laid in the period of forced submergence was smaller than both before and after the period. Selecting deep sites can therefore increase the mortality risk of ovipositing females and also decrease the number of eggs laid in their lives. Selecting suitable sites in all the oviposition bouts can decrease ovipositing females' reproductive success. Costs, not only benefits, should be taken into consideration for understanding oviposition site selection during the lifespan of an ovipositing female.

Effect of adult females' predation risk on oviposition site selection in a water strider

AbstractIn many egg‐laying species, females avoid ovipositing at sites where the predation risk is high. Previous studies have mainly focused on the risk for offspring. The effect of predation risk for the females has been considered in some taxa in which parents spend much time at an oviposition site for parental care or mating (e.g., birds, amphibians). In species in which females do not perform activities other than oviposition at sites, the effect of predation risk for females on oviposition site selection has been rarely investigated. We examined whether the predation risk for ovipositing females affects the decision on oviposition in the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae). Adult A. paludum are preyed upon by a backswimmer, Notonecta triguttata (Motschulsky) (Heteroptera: Notonectidae), but N. triguttata does not prey upon eggs or early instars of A. paludum. We allowed female A. paludum to oviposit under one of three conditions: in presence of N. triguttata, in presence of its chemicals, and in absence of the predator or its cues (control). Female A. paludum less frequently oviposited in presence of N. triguttata than in its absence. Oviposition frequency did not differ between females in presence of chemicals of N. triguttata vs. those in the control. Female A. paludum recognized the predation risk upon themselves from the presence of N. triguttata and avoided ovipositing. This study is the first to directly show that the predation risk upon ovipositing females changes oviposition site selection in species in which the time spent at an oviposition site is short.

Comparing salinity tolerance in embryonic and larval development of two species of water strider, Aquarius paludum and Gerris latiabdominis (Hemiptera: Gerridae)

Water strider Aquarius paludum (Fabricius) is a cosmopolitan species colonizes mainly freshwater but occasionally brackish habitats throughout the Palearctic and Oriental regions. Water strider Gerris latiabdominis (Miyamoto) is a common species in Japan lives in temporary habitats as freshwater paddy fields. These two species often occur syntopically. We investigated differences in the developmental response to brackish water during embryonic and larval stages between the two species. Eggs were exposed to 0-1.8% NaCl solutions within 24 h of oviposition. Larvae of G. latiabdominis were exposed to salinities of 0, 0.5%, and 0.9% from the first instar until adult emergence. Limits of NaCl concentration for hatching were 1.3% and 1.0% for A. paludum and G. latiabdominis, respectively. The hatching rate of G. latiabdominis was lower than that of A. paludum at salinities ≥ 0.9%. The period of embryonic development of G. latiabdominis was more prolonged than that of A. paludum at a given salinity. Although the salinity tolerance of G. latiabdominis was lower than that of A. paludum, our results suggest G. latiabdominis has the physiological capacity to expand into brackish waters. High and low salinity tolerances of A. paludum and G. latiabdominis, respectively, reflect the relatively wide range of habitat salinities utilized by A. paludum and the relatively restricted habitats preferred by G. latiabdominis. The high salinity tolerance of A. paludum could be an important factor contributing to their cosmopolitan distribution because high tolerance to salinity means the possibility of them to be dispersed via ocean or sea to other continents and islands.

Changes in life‐history traits of the water strider Aquarius paludum in accordance with global warming

Seasonal records of nymphs of the water strider Aquarius paludum Fabricius (Heteroptera: Gerridae) at a waterway in Kochi prefecture, Japan (33°00′N to 33°30′N), in 2007, show that populations of new generation adults occur four or more times during the year, in mid and late May; late July to early August; September; and October to November. In laboratory experiments, more than 50% of adults are reproductive, irrespective of photoperiod, and this ratio increases to 80% even under a short‐day LD 11.5 : 13.5 h photocycle, which corresponds to the winter season at Kochi (33°N). There is a diminishing photoperiodic response in relation to the induction of diapause compared with measurements made in previous years (1995–1997), suggesting that reproduction may be possible over a much longer period in the near future (and even in winter) if local climate warming is sustained. A high proportion of males sampled in 2008 have small testes and a high proportion of females do not have mature oöcytes even in summer. This contrasts with earlier data showing that the testis volume index for the same period is greater in 1995–1997, and that 75% of females have mature oöcytes. These changes suggest that the Kochi‐Nankoku population of A. paludum is beginning to show partial aestivation. Overwintering adults of both sexes are found to have no mature flight muscles in October/November 2008. This is in marked contrast with earlier data from 1995 and 2004. Adults that are reared in the laboratory also show a lower flight propensity in response to shorter day lengths. The absence of mature flight muscles in the autumn and lower flight propensity under shorter days may indicate a cessation of dispersal between the summer habitats and overwintering sites on land far from the waterways. Taken together, these data suggest that populations of A. paludum in the Kochi‐Nankoku region are continuing to show adaptive changes, apparently in relation to global warming.

Cost of oviposition site selection in a water strider Aquarius paludum insularis: Egg mortality increases with oviposition depth

Females generally avoid selecting sites for oviposition which have a high predation risk to increase offspring survival. Previous studies have focused on costs to ovipositing females. However, although offspring may also incur costs by being oviposited at low predation risk sites, no studies have focused on costs to offspring. Such costs to offspring were examined by using Aquarius paludum insularis, females of which avoid eggs parasitism by ovipositing at deep sites. Deep sites are safe from egg parasitism but may be unsuitable for hatching due to environmental factors. We examined the costs to offspring at deep sites by comparing the hatching rate, the duration to hatching and the proportion of drowned larvae between eggs that were set at three levels of water depth (0 cm, 25 cm and 50 cm depth). While the hatching rate at 50 cm was lower than that at 0 cm, the rate at 25 cm did not differ from that at 0 cm. Duration to hatching and the proportion of drowned larvae did not differ between the three depths. It is suggested that the declining survival rate of A. paludum eggs was due to increased water pressure at greater depth. Such a cost may exist in other species and such an observation may aid in understanding oviposition site selection.

Role of body size in dominance interactions between male water striders, Aquarius paludum

Water striders are a model system for the study of sexual size dimorphism, but the effect of body size on the dominance relationship between individuals has not been experimentally tested. In 34 staged contests between males of the water strider Aquarius paludum, we determined the effect of body size difference between contestants on the outcome of the aggressive interactions. In contests between a large and a small male, the larger individuals won the interactions significantly more often than expected by chance. This is the first experimental evidence for the importance of body size in pair-wise contests among water striders.

Polyandry, multiple mating, and female fitness in a water strider Aquarius paludum

Contrary to Bateman’s principle, polyandry appears to be a common female mating strategy. Several hypotheses have been proposed to explain the evolution of polyandry. It is assumed that females gain either material or genetic benefits from multiple matings, or that they are coerced into mating by males. In water striders, mating is generally assumed to be costly to females, and they are thought to mate for reasons of convenience, adjusting their resistance to mating according to male harassment. Here, we tested the effect of number of matings (with the same male) and number of partners on female fitness in a water strider Aquarius paludum. In the first experiment, we regulated the time females spent with a male and found that females’ egg production increased with multiple matings up to a point. The result supports the existence of an optimal female mating frequency. In the second experiment, we tested how polyandry affects the number of eggs laid and egg hatching success. We conducted three different trials: females mated four times with either a single male, two different males, or with four different males. Females that mated with four different males laid the lowest number of eggs and had the lowest egg hatching success, suggesting that polyandry reduces females’ egg production and egg hatching success in A. paludum. We conclude that A. paludum females probably gain material benefits from mating but no genetic benefits were found in this study.

Oviposition depth in response to egg parasitism in the water strider: high-risk experience promotes deeper oviposition

In many species, predation risk at oviposition sites causes females to avoid high-risk sites. Previous studies, however, have focused on assessment of current risk at the site during oviposition. Whether memory of previous risk plays a role has not been considered. We examined whether memorized risk affects oviposition site selection in the water strider Aquarius paludum insularis. Aquarius paludum eggs are laid on vegetation on or in the water and are frequently parasitized by a parasitoid wasp. Eggs at deeper positions are less frequently parasitized than those near the surface. After one of three treatments (no, low or high wasp density), A. paludum females were allowed to oviposit in parasitoid-free aquaria. The depth of oviposition of females that had previously experienced high wasp density was greater than that of those that had not experienced wasps. However, the difference in oviposition depth between low-density and no-wasp treatments was slight although significant. This indicates that A. paludum makes the decision on oviposition depth in response to the egg parasitism risk experienced before oviposition. The effect of the memorized risk was largest just after the treatment and became weaker over time. A measurable effect persisted for 28 days.

Morphology and neurophysiology of tarsal vibration receptors in the water strider Aquarius paludum (Heteroptera: Gerridae)

Substrate vibratory information receptors are extensively studied in insects and spiders, however for water surface dwelling species little data is available. We studied the vibration receptive organs in tarsi of the water strider Aquarius paludum, using light, transmission and scanning electron microscopes, and recorded the neural activity of the organs in response to vibrational stimuli, which were afterwards analysed with a custom made spike sorting program. We found that the tarsal chordotonal organ has one set of three scoloparia: one in the tarsomere I and two in the tarsomere II, all of which consisted of a few scolopidia. The chordotonal organ clearly responded to vibratory stimulation. Furthermore, we found that a pair of large subapical emergent dorsal setae, which had been deemed mechanosensory by previous authors, are not so. In turn, four ventral subapical trichobothria that are in direct contact with the water surface during locomotion, proved to be mechanosensory. The anatomical and ultrastructural observations support these electro-physiological results.

Responses of life-history traits of brackish- and freshwater populations of the water strider to NaCl Aquarius paludum (Hemiptera: Gerridae)

The water strider Aquarius paludum inhabits the surfaces of a wide variety of freshwater habitats both temporary and permanent. It can also live on the surface brackish-water around river mouths. Exposure to a brackish rather than freshwater environment may affect a range of adult and offspring life-history traits. In a two-stage experiment A. paludum offspring from fresh- (F) and brackish-water (B) populations were obtained and their offspring reared in either a fresh- or brackish-water (0.45% NaCl) environments. The four offspring treatment groups (F-F, F-B, B-F, B-B) varied in terms of the parental and offspring rearing environments. The effect of parental and offspring rearing environment on longevity, fecundity and flight was assessed. Offspring reared in a brackish environment throughout their larval and adult stages had a reduced longevity and egg production. The flight activity of the offspring originated from the brackish-water population was maintained when they were reared in a brackish environment, but inhibited when they were exposed to freshwater. Our results suggest that the life-history strategies depend critically on the degree of salinity in both the current environment and that of their parents.

Avoidance of egg parasitism through submerged oviposition by tandem pairs in the water strider, Aquarius paludum insularis (Heteroptera: Gerridae)

Abstract 1. Females of the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae) carry males on their backs and oviposit under water after copulation. This study focuses on the benefit A. paludum insularis receives by ovipositing in tandem.2. Males guarded females in tandem through to the end of oviposition in 85% of copulations. Females in tandem dived deeper than single females, and the density of A. paludum insularis eggs increased with water depth. The proportion of eggs parasitized by a scelionid wasp, Tiphodytes gerriphagus Marchal (Hymenoptera: Scelionidae) decreased with increasing water depth.3. These results suggest that during oviposition guarding by males is beneficial for females, because it enables pairs to dive and lay eggs deeper and in oviposition sites where the risk of egg parasitism is lower.