The ecology and behaviour of a marine insect, Halobates fijiensis (Hemiptera: Gerridae)

Observations were made in Fiji on the adaptive strategies of a marine gerrid, Halobates fijiensis , a species of the only genus of insects to have successfully colonized the open sea. The immature stages spend most of their time in flotillas on sheltered water close to mangroves, whereas adults usually occur singly and on both sheltered and open water behind the protecting reef systems. Wind has a decisive influence on the distribution of H. fijiensis: in strong onshore winds, all stages, including adults, remain close to the shelter of mangroves, but in calm conditions, older immatures venture considerable distances from the mangroves. Halobates fijiensis feed on floating, dead insects. Adults of the chironomid fly Tanytarsus halophilae , whose larvae live on intertidal sea-grasses, accounted for over 60% of the prey items recorded. Halobates fijiensis feeds at around the time of high water, either as individuals or in small, well-dispersed groups. Eggs are laid on intertidal turtle grass (Syringodium) , green coralline algae (Halimeda) or occasionally coral rubble, at or shortly after the point of low water on days of extreme low spring tides. Halobates fijiensis is preyed upon by the robber-fly Clinopogon scalaris (Diptera: Asilidae). Field observations showed that there is a clear effect of group size on the distance at which an approaching predator model is detected.     

Evolution probing for semiochemicals based on secondary metabolites in the cuticles of three species of Halobates (Heteroptera: Gerridae)

We developed methods for characterizing semiochemicals in the cuticles of the sea-skater Halobates (Gerridae). Such substances may be involved in chemical communication at the sea surface and have been shown to act as sexual pheromones for a related marine insect Trochopus plumbeus (Veliidae). We identified 86 volatile compounds in extracts from three Halobates species ( H. micans Eschscholtz, H. hawaiiensis Usinger and H. sobrinus White) and used them to reconstruct the phylogeny of the clade. Sixty-seven of the compounds were found to be parsimony-informative. Suspected semiochemicals appear to be specific autapomorphies in the phylogram, and our results indicated that chemical profiles may be used to predict the phylogenetic history of Halobates . Moreover, the phylogenetic tree derived from our data is congruent with a previously published one based on morphology and mitochondrial DNA sequence (780 bp in the COI gene). The potential role of each compound to be used for chemical communication was also reviewed. At appropriate concentrations, some of the cuticular compounds, being partly hydrophobic and partly hydrophilic, could disperse on the sea surface and thereby play an energetically efficient role in chemical communication, e.g. mate location. Less dispersible components might act as 'sun-screens' and provide protection from UV-irradiation during daylight hours.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 671–688.     

Genetically separate populations of the ocean‐skater Halobates sericeus (Heteroptera: Gerridae) have been maintained since the late Pleistocene

The oceanic water strider (or ocean‐skater) Halobates sericeus Eschscholtz has a disjunct distribution in the Pacific Ocean, with northern and southern populations widely separated by an equatorial zone. It is sensitive to sea surface conditions and, consequently, its distribution and population structure may provide an insight into environmental changes on the ocean surface on both recent and historical time scales. We assessed the genetic diversity and population structure of H. sericeus in the Pacific Ocean using three gene markers – cytochrome oxidase subunit I (COI), elongation factor 1α and internal transcribed spacer 1 (ITS‐1). These markers indicate that both populations are evolutionarily distinct with limited gene flow, having separated 20 000–50 000 years ago. This suggests that physical conditions and/or biotic interactions on the surface of the Pacific Ocean have provided significant barriers to gene flow since the late Pleistocene or earlier, creating biotic stability over large geographical and temporal scales in spite of a long history of global climate change. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 797–805.     

The sea-skater Halobates (Heteroptera: Gerridae) – probable cause for extinction in the Mediterranean and potential for re-colonisation following climate change

Sea-skaters in the genus Halobates Eschscholtz 1822 include some of the most specialised water striders and are found in tropical and subtropical seas around the world. Even though species of Halobates occur in both the Atlantic Ocean and the Red Sea, no extant sea-skater has been reported from the Mediterranean Sea. A fossil, Halobates ruffoi Andersen et al., 1994, described from Middle–Upper Eocene (45 Ma) Italy indicates that sea skaters were present in this part of the world in the past. Other geological evidence points to dramatic changes in the Mediterranean Sea during the Tertiary and Quaternary that may have led to their later extinction. In this paper we review briefly the distribution, systematics, evolution and ecology of Halobates, and discuss the potential for the Mediterranean to be recolonised following expected environmental changes due to global warming.     

Density estimation for water-striders (Heteroptera: Gerridae)

SUMMARY. Estimates of relative and absolute abundance in field populations of two Gerris species were compared using regression techniques. Separate equations for each gerrid size class allow prediction of population density from timed-catch sampling. Presence or absence of vegetation markedly affected capture rates, but capture rates were not affected by Gerris species or type of emergent vegetation. Availability for capture varied with gerrid leg length; this relationship can be used to estimate availability constants for other water-strider species.     

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.     

Occurrence and density of Halobates micans (Hemiptera: Gerridae) in the eastern South Indian Ocean

Two species of ocean skaters, Halobates germanus and Halobates micans, live in the tropical and subtropical waters of the Indian Ocean. From December 1992 to December 1993, Halobates was intensively sampled in the easternmost region of the South Indian Ocean (13–18.5°S, 114–121E°), from which there have been a small number of records of Halobates. No H. germanus was caught, but a total of 1190 H. micans were collected, with densities estimated at 13 900–28 100 individuals/km². This suggests that H. micans lives in the study area at high densities comparable to those in the Atlantic and the Pacific Oceans. We also discuss the possible effects of ocean currents and winds on the geographic distributions of the two Halobates species in the eastern South Indian Ocean.     

Distribution of the oceanic insects Halobates (Hemiptera: Gerridae) off the south coast of Japan

Specimens of ocean skaters Halobates were collected off the south coast of Japan in the East China Sea in 1995, and from the Kumano‐nada Sea to the East China Sea in 1998 and 1999. Three species were identified: H. micans, H. germanus and H. sericeus. We found two species co‐occurring in comparable densities in different years, a phenomenon not hitherto reported in other regions of the ocean. We discuss distributions of the three Halobates species with special reference to the influence of the Kuroshio Current, temporal variations of sea‐surface temperature, and monsoonal winds.     

Heat coma as an indicator of resistance to environmental stress and its relationship to ocean dynamics in the sea skaters,Halobates (Heteroptera: Gerridae)

Abstract The tolerance to temperature increase was tested for Halobates individuals collected during two cruises in the western tropical Pacific Ocean (MR‐06‐05‐Leg 3, December 21, 2006–January 12, 2007, 0°N‐8°N; KH‐06‐02‐Leg 5, August 18–31, 2006, 12°N–17°N). High temperature coma experiments were conducted on adults and 5th instar larvae. On average, H. sericeus (distributed in the wide latitude zone of 5°N–40°N), H. germanus (distributed in the moderate latitude zone of 0°N–35°N) and H. micans (distributed mainly in the lower latitudes around the equator) were on average paralyzed at 35.6°C (SD: 0.89), 32.9°C (SD: 2.17) and 31.6°C (SD: 2.60), respectively (P= 0.035). According to the current dynamics during the cruise, the colony of H. sericeus at one station (5°N 137°E) may have been transferred from the northern area of 14°N by three currents (North Equatorial Current, Mindanao Current and North Equatorial Counter Current) to the area of 5°N 138°E. Extremely high heat resistance was shown by the adults of H. germanus in the sea area around the equator. Dynamic current and air movements in this area around the equator, that is a “warm seawater pool”, could be hypothesized to be related to the high resistance to heat shown in this study.     

Reproductive behaviour of the ocean skater Halobates robustus (Hemiptera: Gerridae) in the Galapagos Islands

Summary This paper summarizes the recent research on problems of the so-called colonization cycle as one aspect of the migration and flight behaviour of freshwater insects. New results are presented and the applied methods described and discussed. In general the research on this phenomenon shows that the colonization cycle can not only be seen as a type of compensatory flight behaviour, but also as an expression of the winter survival strategy of water insects in high mountainous areas. It opens up the possibility of exploiting ecologically different biotopes such as living in coastal streams and oligohaline brackish water estuaries.     

A fossil Halobates from the Mediterranean and the origin of sea skaters (Hemiptera, Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 38 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, and the phylogeneuc relationships between extant species have recently been analysed (using cladistic methods). In the present paper, we describe the first fossil species of sea skaters, Halobates ruffoi sp. no v. from the Eocene deposit 'Pesciara di Bolca', in the province of Verona, northeastern Italy (geological age about 45 Myr). The significance of this fossil in setting the time scale for the reconstructed phylogeny and anagenesis of adaptive features of sea skaters, and in understanding the evolution and historical zoogeography of these marine insects is discussed.     

Marine insects: genital morphology, phylogeny and evolution of sea skaters, genus Halobates (Hemiptera: Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 36 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, but reliable hypotheses about phylogenetic relationships are required if the biogeography and evolution of sea skaters is to be discussed in a meaningful way. This work presents the results of a study of new characters from the genital segments, especially those of the male phallus and the female gynatrial complex, and a reinterpretation for several other characters. In total 64 characters were scored for 26 species of Halobates , two species ofAsclepios and one species of Metrocoris. With Asclepios and Metrocoris species as outgroups, the character state sets were analysed cladistically using the computer program Hennig86. After critical evaluations of both characters and clades, a phylogeny for Halobates is presented and its taxonomic implications are discussed. A number of monophyletic species groups are delimited. One genus-level synonymy and three species-level synonymies are suggested. The evolution of Halobates is discussed in the light of the reconstructed phylogeny and present knowledge of the ecology and behaviour of sea skaters. A hypothesis of ecological evolution in halobatine water striders is proposed and tested.     

Sources of character conflict in a clade of water striders (Heteroptera: Gerridae)

Incongruence among trees reconstructed with different data may stem from historical (gene tree‐species tree conflict) or process (character change biases) phenomena. Regardless of the source, incongruent data, as determined with “global” measures of homoplasy, have often been excluded from parsimony analysis of the combined data. Recent studies suggest that these homoplasy measures do not predict the contribution of each character to overall tree structure. Branch support measures identify, on a character to node basis, sources of support and conflict resulting from a simultaneous analysis of the data. We implement these branch support measures to identify sources of character conflict in a clade of water striders consisting of Gerris Fabricius, Aquarius Schellenberg, and Limnoporus Stål species. Separate analyses of morphology, mitochondrial cytochrome oxidase I (COI), large mitochondrial ribosomal subunit (16SrRNA), and elongation factor‐1α (EF‐1α) data resulted in cladograms that varied in resolution and topological concordance. Simultaneous analysis of the data resulted in two trees that were unresolved for one node in a strict consensus. The topology agreed with current classification except for the placements of Aquarius chilensis and the Aquarius remigis species group closer to Gerris than to congeneric species. Branch support measures indicated that support derived from each data set varied among nodes, but COI had an overall negative effect on branch support. However, Spearman rank correlation of partitioned branch support values indicated no negative associations of branch support between any data sets and a positive association between EF‐1α and 16SrRNA. Thus incongruence among data sets was not drastic and the gene‐tree versus species tree phenomenon was not implicated. Biases in character change were a more likely reason for incongruence, although saturation curves and incongruence length difference for COI indicated little potential for homoplasy. However, a posteriori inspection of COI nucleotide change with reference to the simultaneous analysis tree revealed AT and codon biases. These biases were not associated with branch support measures. Therefore, it is difficult to predict incongruence or identify its cause. Exclusion of data is ill advised because every character is potentially parsimony informative.     

Dyar's rule and multivariate allometric growth in nine species of waterstriders (Heteroptera: Gerridae)

The constancy of postmoult/premoult ratios of measures of linear size during ontogeny in insect and other arthropods is widely known as Dyar's rule. We tested this rule in nine species of the waterstrider genera Gerris and Aquarius (Heteroptera: Gerridae), using two size variables: head width and a multivariate measure derived from the pattern of multivariate allometry common to the species considered. Allometric patterns were similar in two independent datasets of laboratory-reared and field-caught specimens. Although our data strictly followed Dyar's rule injust a few instances, all growth ratios varied within a limited range only. Growth ratios for head width differed more between moults than those for multivariate size. The relationship between growth ratios for the two size measures conformed to the predictions based on allometry. We discuss hypotheses of the possible adaptive significance of growth ratios, such as their relation to mobility and systematic differences between hemimetabolous and holometabolous insects, and emphasize the importance of allometry. Since Dyar's rule is consistent with available evidence of physiological mechanisms underlying growth and moulting control of insects and crustaceans, it can be used as a general frame of reference to test alternative growth models.     

Correlated traits for migration in the Gerridae (Hemiptera, Heteroptera): a field test

Abstract. 1. The hypothesis that the migratory tendency of macropters is correlated with proportion macropterous in wing-polymorphic insects is tested by comparing the migration of macropters of three species of waterstriders (Heteroptera, Gerridae) under natural conditions.
2. Migration of marked individuals among four permanent and seventeen ephemeral water bodies was recorded between 1 April 1986 and 2 June 1987, within an area of approximately 2 km2.
3. The proportion of macropters migrating was significantly lower for G.buenoi Kirkaldy than for L.dissortis Drake and Harris and G.comatus Drake and Hottes. Similar results were obtained in. comparisons of proportions captured on ephemeral sites.
4. Distances measured in this study appear to be small relative to the migratory capacity of all of the species, and migration distance did not differ significantly among species. All species migrated primarily in early spring, and differences among species were most apparent at this time.
5. The observed differences among species combined with the results of a similar study of macropters from the primarily apterous species, G.remigis Say, support the initial hypothesis. The implications of this for our understanding of the evolution and maintenance of wing polymorphisms are discussed.     

Geographical variation in wing polymorphism of the waterstrider Aquarius najas (Heteroptera,Gerridae)

The waterstrider Aquarius najas is wingless in Northern Europe, while winged individuals occur frequently in Central and Southern Europe. To test if the latitudinal difference is genetically controlled, we collected mature individuals from 10 different populations and raised their offspring in ‘common garden’ laboratory conditions. Half of these populations were from southern and the other half from central Finland. Daylength and temperature do influence wing development among other species of waterstriders, and thus we maintained a similar short daylength and warm conditions for all populations. These conditions should be favourable for wing development in general. Among laboratory-bred individuals several winged individuals appeared, and their proportion varied between populations. The relative frequency of winged individuals was highest in the southern populations. Thus, apart from phenotypic plasticity there seems to be some genetic control over the occurrence of wings, and the latitudinal trend coincides with the direction in natural populations over a larger European scale. Overwinter survival in our laboratory conditions was higher among the wingless individuals. The survival cost may explain why the proportion of winged individuals was lower in the northern populations with more extreme overwintering conditions than in the southern ones.     

Distributional Patterns of Chacoan Water Bugs (Heteroptera: Belostomatidae,Corixidae, Micronectidae and Gerridae)

Distributional patterns of South American species of aquatic and semiaquatic Heteroptera from the Chaco biogeographic province were analyzed. Based on a track analysis of 60 species of Belostomatidae, Corixidae, Micronectidae, and Gerridae, five generalized tracks were found:(1) Bolivia, and northwestern and central Argentina (Belostoma dallasi, Ectemnostega montana, E. quechua, E. stridulata, E. venturii, Sigara tucma, S. yala, Tenagobia pulchra, Eurygerris fuscinervis, and Trepobates taylori); (2) southern Brazil, eastern Bolivia, Paraguay, and northeastern Argentina (Belostoma bosqi, Heterocorixa brasiliensis, Tenagobia selecta tarahui, and T. schadei); (3) southeastern Brazil and northeastern Argentina, determined by Belostoma candidulum, B. testaceopallidum, Heterocorixa nigra, Sigara hungerfordi, Brachymetra furva,Halobatopsis spiniventris, Metrobates plaumanni plaumanni, and M. vigilis; (4) southeastern Brazil, Uruguay, and central western Argentina (Belostoma cummingsi, B. martini, Sigara argentiniensis, Tenagobia fuscata, and T. carapachay); and (5) southern (Trichocorixa milicorum, Sigara santiagiensis, and S. forciceps). Three panbiogeographic nodes have been determined:(1) northeastern Argentina, in the intersection of generalized tracks 2 and 3; (2) central Argentina, in the intersection of generalized tracks 1 and 4; and (3) central Argentina, in the intersection of generalized tracks 4 and 5. In spite of these complex patterns, these results show that the Chaco province appears to be a natural biogeographic area.     

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)