Gerridae


The Gerridae are a family of insects in the order Hemiptera, commonly known as water striders, water skeeters, water scooters, water bugs, pond skaters, water skippers, water gliders, water skimmers or puddle flies. They are true bugs of the suborder Heteroptera and have mouthparts evolved for piercing and sucking. A distinguishing feature is the ability to move on top of the water's surface, making them pleuston animals. They can be found on most ponds, rivers or lakes, and over 1,700 species of gerrids have been described, 10% of them being marine.
While 90% of gerrids are freshwater bugs, the oceanic Halobates makes the family quite exceptional among insects. The genus Halobates was first heavily studied between 1822 and 1883 when Francis Buchanan White collected several different species during the Challenger expedition. Around this time, Eschscholtz discovered three species of the Gerridae, bringing attention to the species, though little of their biology was known. Since then, the Gerridae have been continuously studied due to their ability to walk on water and unique social characteristics.

Description

The family Gerridae is physically characterized by having hydrofuge hairpiles, retractable preapical claws, and elongated legs and body.
Hydrofuge hairpiles are small, hydrophobic microhairs. These are tiny hairs with more than one thousand microhairs per mm. The entire body is covered by these hairpiles, providing the water strider resistance to splashes or drops of water. These hairs repel the water, preventing drops from weighing down the body.

Size

They are generally small, long-legged insects and the body length of most species is between. A few are between. Among widespread genera, the North Hemisphere Aquarius includes the largest species, generally exceeding, at least among females, and the largest species averaging about. Females are typically larger than males of their own species, but this appears to be reversed in the largest species, the relatively poorly known Gigantometra gigas of streams in northern Vietnam and adjacent southern China. It typically reaches a body length of about in wingless males and in winged females. In this species each middle and hind leg can surpass.

Antennae

Water striders have two antennae with four segments on each. Antennal segments are numbered from closest to the head to farthest. The antennae have short, stiff bristles in segment III. Relative lengths of the antennae segments can help identify unique species within the family Gerridae, but in general, segment I is longer and stockier than the remaining three. The four segments combined are usually no longer than the length of the water strider head.

Thorax

The thorax of water striders is generally long, narrow, and small in size. It generally ranges from 1.6 mm to 3.6 mm long across the species, with some bodies more cylindrical or rounder than others. The pronotum, or outer layer of the thorax, of the water strider can be either shiny or dull depending on the species, and covered with microhairs to help repel water. The abdomen of a water strider can have several segments and contains both the metasternum and omphalium.

Appendages

Gerridae have front, middle, and back legs. The front legs are shortest and have preapical claws adapted to puncture prey. Preapical claws are claws that are not at the end of the leg, but rather halfway through, like mantises. For most species of Gerridae the middle legs are longest, an adaption for propulsion through the water, spreading weight over a large surface area, as well as steering the bug across the surface of the water. The front legs are attached just posterior to the eyes, while the middle legs are attached closer to the back legs which attach midthorax but extend beyond the terminal end of the body.

Wings

Some water striders have wings present on the dorsal side of their thorax, while other species of Gerridae do not, particularly Halobates. Water striders experience wing length polymorphism that has affected their flight ability and evolved in a phylogenetic manner where populations are either long-winged, wing-dimorphic, or short-winged. Wing dimorphism consists of summer gerrid populations evolving different length wings than winter populations within the same species. Habitats with rougher waters are likely to hold gerrids with shorter wings, while habitats with calm waters are likely to hold long-winged gerrids. This is due to potential for damage of the wings and ability for dispersal.

Evolution

Cretogerris, from the Cretaceous Charentese amber of France, was initially suggested as a gerrid. However, it was later interpreted as an indeterminate member of Gerroidea. The Gerridae are morphologically similar to the unrelated Chresmoda, an enigmatic genus of insect known from the Late Jurassic to the Mid Cretaceous with a presumably similar lifestyle.
Molecular analysis suggest an origin of the family Gerridae about 128 Million years ago in the Cretaceous, splitting from the sister group Veliidae, with whom they share a single origin of rowing as a locomotive mechanism. According on the transcriptome-based phylogeny, Gerridae is a monophyletic group.

Wing polymorphism

Wing polymorphism has independently evolved multiple times in Gerridae, as well as complete wing loss, something that has been important for the evolution of the variety in species we see today, and dispersal of Gerridae. The existence of wing polymorphism in a given species can be explained as a particular case oogenesis-flight syndrome. Following this rationale, which is commonly applied in insects, developing short wings provides the individual with the capacity to dedicate the energy stores that would usually be used for wing and wing muscle development to increasing egg production and reproducing early, ultimately enhancing the individual's fitness. The ability for one brood to have young with wings and the next not allows water striders to adapt to changing environments. Long, medium, short, and nonexistent wing forms are all necessary depending on the environment and season. Long wings allow for flight to a neighboring water body when one gets too crowded, but they can get wet and weigh a water strider down. Short wings may allow for short travel, but limit how far a gerrid can disperse. Nonexistent wings prevent a gerrid from being weighed down, but prevent dispersal.
Wing polymorphism is common in the Gerridae despite most univoltine populations being completely apterous or macropterous. Apterous populations of gerrids would be restricted to stable aquatic habitats that experience little change in environment, while macropterous populations can inhabit more changing, variable water supplies. Stable waters are usually large lakes and rivers, while unstable waters are generally small and seasonal. Gerrids produce winged forms for dispersal purposes and macropterous individuals are maintained due to their ability to survive in changing conditions. Wings are necessary if the body of water is likely to dry since the gerrid must fly to a new source of water. However, wingless forms are favored due to competition for ovarian development and wings and reproductive success is the main goal due to the selfish gene theory. Overwintering gerrids usually are macropterous, or with wings, so they can fly back to their aquatic habitat after winter. An environmental switch mechanism controls seasonal dimorphism observed in bivoltine species, or species having two broods per year. This switch mechanism is what helps determine whether or not a brood with wings will evolve. Temperature also plays an important role in photoperiodic switch. Temperatures signify the seasons and thus when wings are needed since they hibernate during winter. Ultimately, these switching mechanisms alter genetic alleles for wing characteristics, helping to maintain biological dispersal.

Ability to move on water's surface

Water striders are able to move on top of water due to a combination of several factors. Water striders use the high surface tension of water and long, hydrophobic legs to help them stay above water.
Gerridae species use this surface tension to their advantage through their highly adapted legs and distributed weight.
The legs of a water strider are long and slender, allowing the weight of the water strider body to be distributed over a large surface area. The legs are strong, but have flexibility that allows the water striders to keep their weight evenly distributed and flow with the water movement. Hydrofuge hairs line the body surface of the water strider. There are several thousand hairs per square millimeter, providing the water strider with a hydrofuge body that prevents wetting from waves, rain, or spray, which could inhibit their ability to keep their entire body above the water surface if the water stuck and weighed down the body. This position of keeping the majority of the body above the water surface, called epipleustonic, is a defining characteristic of water striders. If the body of the water strider were to accidentally become submerged, for instance by a large wave, the tiny hairs would trap air. Tiny air bubbles throughout the body act as buoyancy to bring the water strider to the surface again, while also providing air bubbles to breathe from underwater. Despite their success in overcoming submergence in water, however, water striders are not as competent in oil, and experimental oil spills have suggested that oil spilled in freshwater systems can drive water strider immobility and death.
The tiny hairs on the legs provide both a hydrophobic surface as well as a larger surface area to spread their weight over the water. The middle legs used for rowing have particularly well developed fringe hairs on the tibia and tarsus to help increase movement through the ability to thrust. The hind pair of legs are used for steering. When the rowing stroke begins, the middle tarsi of gerrids are quickly pressed down and backwards to create a circular surface wave in which the crest can be used to propel a forward thrust. The semicircular wave created is essential to the ability of the water strider to move rapidly since it acts as a counteracting force to push against. As a result, water striders often move at 1 meter per second or faster.