Gobiidae

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Gobies
Gobius niger (Stefano Guerrieri) 1.jpg
Black goby ( Gobius niger )
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Gobiiformes
Family: Gobiidae
G. Cuvier, 1816
Subfamilies

See text.

Gobiidae is a family of bony fish in the order Gobiiformes, one of the largest fish families comprising more than 2,000 species in more than 200 genera, sometimes referred to as the "true gobies". [1] Most of them are relatively small, typically less than 10 cm (3.9 in) in length. The Gobiidae includes some of the smallest vertebrates in the world, such as Trimmatom nanus and Pandaka pygmaea , Trimmatom nanus are under 1 cm (38 in) long when fully grown, then Pandaka pygmaea standard length are 9mm (0.35 in),maximum known standard length are 11 mm (0.43 in). Some large gobies can reach over 30 cm (0.98 ft) in length, but that is exceptional. Generally, they are benthic, or bottom-dwellers. Although few are important as food for humans, they are of great significance as prey species for commercially important fish such as cod, haddock, sea bass, and flatfish. Several gobiids are also of interest as aquarium fish, such as the dartfish of the genus Ptereleotris . Phylogenetic relationships of gobiids have been studied using molecular data. [2] [3]

Contents

Description

Racer goby from the Bug River 4.jpg
Racer goby Babka gymnotrachelus, a member of the formerly valid subfamily Benthophilinae.
Neogobiusfluviatilisandkessleri.JPG
the monkey goby ( Neogobius fluviatilis ) and the bighead goby ( Ponticola kessleri ), members of the formerly valid subfamily Benthophilinae.
Microgobius gulosus2.jpg
clown goby ( Microgobius gulosus ), a "true goby", formerly a member of the Gobiinae
046-011 Pomatoschistus microps (cropped).JPG
Common goby (Pomatoschistus microps), a "true goby", formerly a member of the Gobiinae
Ptereleotris evides 2557.jpg
Blackfin dartfish ( Ptereleotris evides ) a species from the formerly valid family Ptereleotridae

The most distinctive aspects of gobiid morphology are the fused pelvic fins that form a disc-shaped sucker. This sucker is functionally analogous to the dorsal fin sucker possessed by the remoras or the pelvic fin sucker of the lumpsuckers, but is anatomically distinct; these similarities are the product of convergent evolution. The species in this family can often be seen using the sucker to adhere to rocks and corals, and in aquariums they will stick to glass walls of the tank, as well.

Distribution and habitat

Gobiidae are spread all over the world in tropical and temperate near shore-marine, brackish, and freshwater environments. Their range extends from the Old World coral reefs to the seas of the New World, and includes the rivers and near-shore habitats of Europe and Asia. [4] Gobies are generally bottom-dwellers. Although many live in burrows, a few species (e.g. in the genus Glossogobius ) are true cavefish. [5] On coral reefs, species of gobiids constitute 35% of the total number of fishes and 20% of the species diversity. [6]

Subfamilies

The family Gobiidae underwent a major revision in the 5th edition of Fishes of the World . Before the revision the Gobiidae contained six subfamilies: Gobiinae, Benthophilinae, Amblyopinae, Gobionellinae, Oxudercinae, and Sicydiinae. The revision retained the first two subfamilies and removed the other four to a separate family, the Oxudercidae. In addition, species formerly placed in the families Kraemeriidae, Microdesmidae, Ptereleotridae and Schindleriidae were added to the revised Gobiidae, although no subfamilies were described. [7]

The two formerly recognised subfamilies where the species have been retained in Gobiidae in the 5th Edition of Fishes of the World: [7]

Benthophilinae

Members of Benthophilinae are endemic to the Ponto-Caspian region (including the Marmara, Black, Azov, Caspian, and Aral Seas). [8] The representatives of the subfamily have fused pelvic fins and elongated dorsal and anal fins. [9] They are distinguished from the closely related subfamily Gobiinae by the absence of a swimbladder in adults and location of the uppermost rays of the pectoral fins within the fin membrane. [10] Its members include tadpole gobies, monkey gobies, and bighead gobies.

Gobiinae

Members of the Gobiinae are known as true gobies. It is the most widespread and most diverse of the subfamilies formerly recognised under the Gobiidae, containing around 2000 species and 150 genera.

Ecology and biology

Gobiids are primarily fish of shallow marine habitats, including tide pools, coral reefs, and seagrass meadows; they are also very numerous in brackish water and estuarine habitats, including the lower reaches of rivers, mangrove swamps, and salt marshes. A few gobiid species (unknown exactly, but in the low hundreds) are also fully adapted to freshwater environments. These include the round goby (Neogobius melanostomus), Australian desert goby ( Chlamydogobius eremius ), and the European freshwater goby Padogobius bonelli . Most gobies feed on small invertebrates, although some of the larger species eat other fish, and a few eat planktonic algae.

Reproduction

Most species in the Gobiidae attach their eggs to a substrate, such as vegetation, coral, or a rock surface. They lay from five to a few thousand eggs, depending on the species. After fertilizing the eggs, the male guards the eggs from predators and keep them free from detritus. The male fans the eggs, thereby providing them with oxygen. The female maintains the burrow. The eggs hatch after a few days. The larvae are born transparent, and they develop coloration after spreading to find a suitable habitat. The larvae of many freshwater gobiid species are carried downstream to the brackish waters, or even to the sea. They return to fresh water weeks or months later. [11]

Gobiids in warmer waters reach adulthood in a few months, while gobies in cooler environments reach adulthood in two years. The total lifespan of gobiid varies from one to ten years, again with the species in warmer waters generally living longer. [11]

Behavior

Burrow construction

Many species in the Gobiidae live in male-female pairs that construct and share burrows, similar to many other fish such as Mozambique tilapia. The burrows are used for shelters and spawning places. Gobiids use their mouths to dig into the sea bottom, removing dead coral-fragments, rubble, and benthic algae in order to build their burrows. [12] Gobiids maintain their burrows by fanning away sand inside the burrows. Furthermore, gobies use coral rubble to block burrow entrance. A single goby carry as many as nine pieces of coral rubble per minute. Gobiids also build a 6–13 cm high mound over the entrance of their spawning burrow. [12] The mound lets the water flow fast over the mound. The water flow created by the mound helps to provide oxygen to the eggs. While burrow building is a cooperative behavior done by both sexes, males usually put more effort in burrow maintenance than females. Females feed more instead, because the reproductive success is optimal when females put more energy in preparing for the reproduction. [13] After spawning eggs, the roles of male and female changes. Females primarily maintain the burrow, and males mainly care for the eggs by fanning them, thereby providing oxygen. When females leave the burrow, however, the mounds lose their heights. The males then give up on the eggs and eat them, preparing for future mating opportunities. Gobiid burrows vary in size depending on the size of the species. [13]

Kleptogamy

Kleptogamy refers to a "sneaking behavior" during reproduction where an unpaired male fertilises the eggs of a paired female and the paired male cares for the eggs. Females prefer male gobies with large bodies. Since not all males have large bodies, the smaller ones may cheat instead of expending energy to find mates. [14] The sneakers wait near the spawning ground of paired fish. The sneakers then release their sperm on the spawning ground as soon as the paired female releases her eggs. [15] Though sneakers’ sperm fertilizes some eggs, the paired male cannot distinguish the eggs fertilized by the sneakers from those fertilized by his own sperm. Therefore, the paired male gives parental care equally to all the eggs. [16]

Kleptogamy is a good strategy in many ways. First, the sneakers do not need their own territories, indicating that they do not need to spend energy in protecting territories, as most other males do. Most male gobies need their own territories, since females do not choose to mate with a male that does not own his own territory. [14] Secondly, the sneakers do not provide parental care to their eggs. The paired males provide parental care instead of the sneakers. Therefore, the sneakers can save energy, and they can put more effort into finding new targets for cheating. [14]

The cost of kleptogamy is that the sneakers can receive aggressive attacks from the paired males that are usually much larger and stronger than the sneakers. For small sneakers, the attacks by the paired males can be detrimental and often lead to death. [14]

The sneakers are also referred to as pseudo-females, since they are small and hardly distinguishable from females. This small body size makes cheating easier. Most of the time the paired males mistake the sneakers for females and thus do not chase the sneakers away. The paired males are called “bourgeois” males, because they are larger, stronger, and most importantly, paired. [14]

An 1865 watercolor painting of a Brazilian goby by Jacques Burkhardt. Gobius (Rio de Janeiro, Brazil, 15 July 1865).jpg
An 1865 watercolor painting of a Brazilian goby by Jacques Burkhardt.

Sex change

A few species of gobiid, such as blackeye goby and Lythrypnus dalli , can change their sexes. Sex change is possible in these gobies, since the external genitalia for males and females do not differ much. [14] Sex changes can take from days to weeks. Most sex changes in gobies are from female to male (protogyny) rather than male to female (protandry). Female-to-male changes are observed not only in gobiids but also in wrasses, damselfishes, and sea basses. [14] Female-to-male change usually occurs because the resident male of the group is dead. If no male is in the group, reproduction will be impossible. Therefore, the dominant female turns into male, allowing mating to happen. [17] Male-to-female change occurs when the females have preference for specific features in males. For example, females prefer large males, and a few large males mate with multiple females, whereas small males lose their chance to mate. Small males either choose to become sneakers (kleptogamy) or choose to transform into females because all females technically have high mating opportunities. By turning into females, males can ensure that they produce many offspring. [14] [18]

Some gobies have extraordinarily developed sex change ability. Gobiodon histrio from the Great Barrier Reef exhibits bidirectional sex changes. G. histrio is one of the very few species that can change sex in both ways. When two G. historio females, which used to be males, are on the same coral reef, one of them transforms back into a male goby. [18]

Sex determination

Sex determination in coral goby Gobiodon erythrospilus does not occur until the juveniles meet potential mates. [14] Confronting a potential mate can be difficult for Gobiodon erythrospilus juveniles, since most coral resources, crucial for attracting mates, are occupied by pre-existing paired gobies. Juveniles can only meet potential mates when one member of the pre-existing pairs dies. Juveniles’ sexes are determined according to the sexes of their potential mates. When a juvenile meets a female, it becomes a male, and vice versa. This type of sex determination is referred to as socially influenced sex determination. [19]

Some gobiids remember landmarks that are within short distances, and use them to find their ways. Small frillfin gobies ( Bathygobius soporator ) live in intertidal zones. They swim through the pools during high tides and memorize how each pool connects to the others. Then, during low tides, they can exhibit accurate jumping behaviors, as they have memorized the paths. [20] In a new environment, these fish do not show jumping behaviors or jump into wrong pools. Nevertheless, after one night, they show the same accurate jumping behaviors. [21]

Habitat choice

A study was done to understand how gobiids react to changing habitat. The fish were given two choices: a safe habitat with less food and a dangerous habitat with more food. Results from both the full and hungry fish revealed that gobiids, when confronted with the trade-off between foraging and avoiding predation, made choices that would better their foraging. [22]

Some marine gobies live in symbiosis with shrimp. Goby fish with shrimp.jpg
Some marine gobies live in symbiosis with shrimp.

Symbiosis

Species in the Gobiidae sometimes form symbiotic relationships with other species, [23] such as with burrowing shrimps. The shrimp maintains a burrow in the sand in which both the shrimp and the fish live. The shrimp has poor eyesight compared to the gobiid, but if it sees or feels the fish suddenly swim into the burrow, it will follow. The fish and shrimp keep in contact with each other, the shrimp using its antennae, and the fish flicking the shrimp with its tail when alarmed. These gobiids are thus sometimes known as "watchmen gobies" or "prawn gobies". Each party gains from this relationship: the shrimp gets a warning of approaching danger, and the fish gets a safe home and a place to lay its eggs. Only the alpha male and female reproduce, other fish in the colony eat sparingly to resist being eaten by the alpha male or female. This way, only the largest and fittest are able to reproduce.[ citation needed ]

Another example of symbiosis is demonstrated by the neon gobies ( Elacatinus spp.). These gobiids, known as "cleaner gobies", remove parasites from the skin, fins, mouth, and gills of a wide variety of large fish. The most remarkable aspect of this symbiosis is many of the fish that visit the cleaner gobies' cleaning stations would otherwise treat such small fish as food (for example, groupers and snapper). Again, this is a relationship where both parties gain: the gobies get a continual supply of food as bigger fish visit their cleaning stations, and the bigger fish leave the cleaning stations healthier than they were when they arrived.

Another form of symbiosis exists between gobiids and the mushroom coral Heliofungia actiniformis (Fungiidae), in which representatives of the genus Eviota roam among the tentacles possibly hiding from predators. [24] [25]

Commercial importance

Dried gobies for sale on the market in Odessa, Ukraine Gobies for sale.jpg
Dried gobies for sale on the market in Odessa, Ukraine

Gobiids have commercial importance in Russia and Ukraine. They are fished in the Sea of Azov, northwestern Black Sea and Caspian Sea. Most important species are round goby, monkey goby, toad goby, and grass goby. The grass goby is also a commercial fish in Italy.

In the aquarium

Several species of gobiids are kept in aquaria. [26] Most captive gobies are marine. Perhaps the most popular is the small but colorful neon goby. Most gobies stay toward the lower portion of the aquarium, hiding in the rockwork, but some species (most notably the shrimp gobies) prefer to dig themselves little burrows. Aquarists typically provide them with a fine-grained substrate to prevent damage to their delicate undersides. Commonly kept saltwater species include Randall's shrimp goby and the watchman goby.

See also

Related Research Articles

Pomacentridae Family of fishes

Pomacentridae is a family of ray-finned fish, comprising the damselfishes and clownfishes. This family were formerly placed in the order Perciformes but are now regarded as being incertae sedis in the subseries Ovalentaria in the clade Percomorpha. They are primarily marine, while a few species inhabit freshwater and brackish environments. They are noted for their hardy constitutions and territoriality. Many are brightly colored, so they are popular in aquaria.

Alpheidae Family of crustacean

Alpheidae is a family of caridean snapping shrimp, characterized by having asymmetrical claws, the larger of which is typically capable of producing a loud snapping sound. Other common names for animals in the group are pistol shrimp or alpheid shrimp.

Eleotridae Family of fishes

Eleotridae is a family of fish commonly known as sleeper gobies, with about 34 genera and 180 species. Most species are found in the tropical Indo-Pacific region, but there are also species in subtropical and temperate regions, warmer parts of the Americas and near the Atlantic coast in Africa. While many eleotrids pass through a planktonic stage in the sea and some spend their entire lives in the sea; as adults, the majority live in freshwater streams and brackish water. One of its genera, Caecieleotris, is troglobitic. They are especially important as predators in the freshwater stream ecosystems on oceanic islands such as New Zealand and Hawaii that otherwise lack the predatory fish families typical of nearby continents, such as catfish. Anatomically, they are similar to the gobies (Gobiidae), though unlike the majority of gobies, they do not have a pelvic sucker.

<i>Thalassoma bifasciatum</i> Species of fish

Thalassoma bifasciatum, the bluehead, bluehead wrasse or blue-headed wrasse, is a species of marine ray-finned fish, a wrasse from the family Labridae. It is native to the coral reefs of the tropical waters of the western Atlantic Ocean. Individuals are small and rarely live longer than two years. They form large schools over the reef and are important cleaner fish in the reefs they inhabit.

<span class="mw-page-title-main">Mudskipper</span> Subfamily of fishes

Mudskippers are amphibious fish. They are of the family Oxudercidae and the subfamily Oxudercinae. There are 23 living species of mudskipper. They are known for their unusual appearance and their ability to survive both in and out of water. They can grow up to 30 centimetres (12 in) long and most are a brownish green colour that range anywhere from dark to light. During mating season the males will also develop brightly coloured spots in order to attract females. The spots can be red, green or blue. A mudskipper's eyes protrude from the tops of its flat head. Their most noticeable feature however is their side pectoral fins that are located more forward and under their elongated body. These fins function similarly to legs in that they allow the mudskipper to move from place to place. Although having the typical appearance of any other fish, these forward fins allow the mudskipper to “skip” across muddy surfaces and even give them the ability to climb trees and low branches. Because of these fins, mudskippers have also been found to be able to leap distances of up to two feet.

<span class="mw-page-title-main">Black-ray goby</span> Species of fish

Stonogobiops nematodes, the Filament-finned prawn-goby, the Antenna goby, the high-fin goby, the red-banded goby, the high-fin red-banded goby, the striped goby, the barber-pole goby, or the black-ray Goby, is a species of marine goby native to the Indian Ocean and western Pacific Ocean from the Seychelles to the Philippines and Bali.

<span class="mw-page-title-main">Yellow clown goby</span> Species of fish

The yellow clown goby, Gobiodon okinawae, also known as the Okinawa goby or yellow coral goby, is a member of the goby family native to the western Pacific from southern Japan to the southern reaches of the Great Barrier Reef. As the name implies, they are bright yellow in color, save for a whitish patch on each cheek.

<span class="mw-page-title-main">Steinitz' prawn goby</span> Species of fish

Amblyeleotris steinitzi, Steinitz' prawn goby or simply Steinitz' goby, is a species of small fish in the family Gobiidae. It lives in association with an alpheid shrimp and is found from the Red Sea through the Indian Ocean to the western Pacific Ocean.

<i>Elacatinus</i> Genus of fishes

Elacatinus is a genus of small marine gobies, often known collectively as the neon gobies. Although only one species, E. oceanops, is technically the "neon goby," because of their similar appearance, other members of the genus are generally labeled neon gobies, as well. Except for a single East Pacific species, all reside in warmer parts of the West Atlantic, including the Caribbean and Gulf of Mexico. They are known for engaging in symbiosis with other marine creatures by providing them cleaning service that consists of getting rid of ectoparasites on their bodies. In return, Elacatinus species obtain their primary source of food, ectoparasites.

Gobiiformes Order of fishes

The Gobiiformes are an order of fish that includes the gobies and their relatives. The order, which was previously considered a suborder of Perciformes, is made up of about 2,211 species that are divided between seven families. Phylogenetic relationships of the Gobiiformes have been elucidated using molecular data. Gobiiforms are primarily small species that live in marine water, but roughly 10% of these species inhabit fresh water. This order is composed chiefly of benthic or burrowing species; like many other benthic fishes, most gobiiforms do not have a gas bladder or any other means of controlling their buoyancy in water, so they must spend most of their time on or near the bottom. Gobiiformes means "Goby-like".

<i>Elacatinus puncticulatus</i> Species of fish

Elacatinus puncticulatus is a species of goby from the eastern central Pacific Ocean, where it is found on reefs from the Gulf of California to Ecuador. This species occurs at depths ranging from 1 to 21m, and usually in association with the sea urchin Eucidaris thouarsii. The size of the goby varies depending on sex, with females being typically smaller than males, and their geographical location as well as their role as a cleaner goby also has impacts on their morphology. Due to their bright coloration and lack of aggression, the species is commonly found in the aquarium trade.

<i>Rhinogobiops</i> Genus of fishes

Rhinogobiops nicholsii, the blackeye goby, is a species of true goby in the family Gobiidae. It is the sole species classified under the genus Rhinogobiops. They are common inhabitants of coral reefs and rocky habitats along the Eastern Pacific Ocean coasts of Mexico, the United States, and Canada, although they are hardly noticed, as they often rest motionless near their shelters. Other common names for the species include bluespot goby and crested goby.

Trypauchen vagina, commonly known as the burrowing goby, is a species of eel goby found in the Indo-Pacific region. It has an elongated body about 20 to 22 cm in length. It is reddish-pink in color and possesses distinctive pouches in the upper edges of its gill covers. It lives in burrows in the silty and muddy bottoms of its marine and brackish habitats. It has reduced eyes that are entirely covered with skin and the anterior portion of its head is protected by thick flesh. Both adaptations aid it in digging its burrows.

<i>Elacatinus evelynae</i> Species of fish

Elacatinus evelynae, the sharknose goby, Caribbean cleaner goby, Caribbean cleaning goby, is a species of goby native to the Western Atlantic Ocean from the Bahamas and the Lesser Antilles to the northern coast of South America, as well as the Antilles and western Caribbean.

<i>Bryaninops yongei</i> Species of fish

Bryaninops yongei, the wire-coral goby or whip coral goby, is a benthic species of goby widely distributed from the tropical and subtropical waters of the Indian Ocean to the islands in the center of the Pacific Ocean.

Quietula is a genus of fish in the goby subfamily, Gobionellinae. There are two species, both native to the Gulf of California in Mexico. One is endemic to the Gulf, and the other also occurs along the western coast of Baja California and the coast of California. The fish were first described from Guaymas in Sonora, Mexico. The genus name Quietula is from the Latin quietus, meaning "quiet".

<i>Trimma nasa</i> Species of fish

Trimma nasa, commonly called the nasal dwarfgoby or nasal pygmy goby, is a species of goby from the Western Pacific. They are small fish, averaging at around 2 cm (0.79 in) in length. They are bright orange and transparent yellow in life, with a white stripe running down from between the eyes to the upper lip and a dark brown spot at the base of the tail fin. They are usually found in large schools in the sloping or vertical drop-offs at coral reef edges.

Trimma tevegae, commonly known as the bluestripe pygmygoby or blue-striped cave goby among other names, is a species of goby from the western Pacific. They are small fish, averaging at 2 cm (0.79 in), orange-brown with white undersides in life, with characteristic iridescent blue or lavender stripes on the sides and on top of the body. They are usually found in large schools in the sloping or vertical drop-offs at coral reef edges. They are sometimes caught for the aquarium trade, and are also known by hobbyists under the name blue line flagtail goby. The species is named in honor of the schooner Te Vega.

<i>Trimma caudomaculatum</i> Species of fish

Trimma caudomaculatum, the blotch-tailed pygmygoby , is a species of goby from the Western Pacific. Like other members of the genus, they are usually found in large schools in the sloping or vertical drop-offs at coral reef edges. Similar to other species of Trimma this species consists of multiple cases of bidirectional sex change, meaning that if a group is lacking in a specific sex a partial amount of the group can change their undeveloped gonad structure of the opposite sex in order to accommodate. This sex change is made possible due to the females having a developed set of ovaries with female hormones that are developed, and a set of testis and male hormones that are underdeveloped; The males follow a similar set up in vice versa, so their testis and male hormones are developed, while the ovaries and female hormones are underdeveloped.

<i>Vanderhorstia mertensi</i> Species of fish

Vanderhorstia mertensi, Mertens' shrimp goby or the slender shrimp goby, is a ray-finned fish species native to the Red Sea, Japan, Papua-New Guinea and the Great Barrier Reef. Male individuals can reach a length of 11 cm in total. In 2008 a specimen was collected in the Mediterranean Sea, in Gulf of Fethiye on the coast of southern Turkey, where it was found on sandy bottoms in the vicinity of beds of sea grass such as Zostera spp., Posidonia oceanica and Cymodocea nodose in the burrows of the alpheid shrimps Alpheus glaber and Alpheus rapacida. It is thought that the gobies most likely entered the Mediterranean through the Suez Canal by Lessepsian migration from the Red Sea but the possibility of transportation in ships' ballast waters cannot be excluded. It was recorded off Israel in 2013, confirming that its origin was as a Lessepsian migrant. The specific name honours the German herpetologist Robert Mertens (1894-1975), the former director of the Naturmuseum Senckenberg in Frankfurt, from whom the author, Klausewitz, learnt about the biological and ecological view of modern systematics and taxonomy.

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