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Gobiidae

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This article is about the fish family Gobiidae. For other fish called goby, seeGoby.

Gobies
Black goby (Gobius niger)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Gobiiformes
Family: Gobiidae
G. Cuvier,1816
Subfamilies

Seetext.

Gobiidaeorgobiesis afamilyofbony fishin theorderGobiiformes,one of the largest fish families comprising more than 2,000speciesin more than 200genera.[1]Most of gobiid fish are relatively small, typically less than 10 cm (3.9 in) in length, and the family includes some of the smallestvertebratesin the world, such asTrimmatom nanusandPandaka pygmaea,Trimmatom nanusare under1 cm (3⁄8in) long when fully grown, thenPandaka pygmaeastandard length are 9 mm (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 arebenthicor bottom-dwellers. Although few are important asfood fishfor humans, they are of great significance aspreyspecies for other commercially important fish such ascod,haddock,sea bassandflatfish.Several gobiids are also of interest asaquarium fish,such as thedartfishof the genusPtereleotris.Phylogeneticrelationships of gobiids have been studied using molecular data.[2][3]

Description

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Racer goby(Babka gymnotrachelus), a member of the formerly valid subfamily Benthophilinae
The monkey goby (Neogobius fluviatilis) and the bighead goby (Ponticola kessleri), members of the formerly valid subfamily Benthophilinae
Clown goby (Microgobius gulosus), a "true goby", formerly a member of the Gobiinae
Common goby(Pomatoschistus microps), a "true goby", formerly a member of the Gobiinae
Blackfin dartfish (Ptereleotris evides), a species from the formerly valid familyPtereleotridae

The most distinctive aspects of gobiidmorphologyare the fusedpelvic finsthat form a disc-shaped sucker. This sucker is functionally analogous to thedorsal finsucker possessed by theremorasor the pelvic fin sucker of thelumpsuckers,but is anatomically distinct; these similarities are the product ofconvergent evolution.The species in this family can often be seen using the sucker to adhere to rocks andcorals,and inaquariumsthey will stick to glass walls of the tank, as well.

Distribution and habitat

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Gobiidae are spread all over the world in tropical and temperate near shore-marine,brackish,and freshwater environments. Their range extends from theOld Worldcoral reefs to the seas of theNew 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 genusGlossogobius) are truecavefish.[5]On coral reefs, species of gobiids constitute 35% of the total number of fishes and 20% of the species diversity.[6]

Subfamilies

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See also:List of Gobiidae genera

The family Gobiidae underwent a major revision in the 5th edition ofFishes of the World.Before the revision the Gobiidae contained six subfamilies:Gobiinae,Benthophilinae,Amblyopinae,Gobionellinae,Oxudercinae,andSicydiinae.The revision retained the first two subfamilies and removed the other four to a separate family, theOxudercidae.In addition, species formerly placed in the familiesKraemeriidae,Microdesmidae,PtereleotridaeandSchindleriidaewere 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 ofFishes of the World:[7]

Benthophilinae

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Main article:Benthophilinae

Members of Benthophilinae are endemic to the Ponto-Caspian region (including theMarmara,Black,Azov,Caspian,andAral 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 aswimbladderin adults and location of the uppermost rays of thepectoral finswithin thefinmembrane.[10]Its members includetadpole gobies,monkey gobies,andbighead gobies.

Gobiinae

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Main article:Gobiinae

Members of the Gobiinae are known astrue 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

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Gobiids are primarily fish of shallow marine habitats, includingtide pools,coral reefs,andseagrass meadows;they are also very numerous inbrackish waterandestuarinehabitats, including the lower reaches of rivers,mangrove swamps,andsalt marshes.A few gobiid species (unknown exactly, but in the low hundreds) are also fully adapted tofreshwaterenvironments. These include theround goby(Neogobius melanostomus), Australian desert goby (Chlamydogobius eremius), and the European freshwater gobyPadogobius bonelli.Most gobies feed on small invertebrates, although some of the larger species eat other fish, and a few eatplanktonicalgae.

Reproduction

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Most species in the Gobiidae attach their eggs to asubstrate,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 fromdetritus.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 manyfreshwatergobiid species are carried downstream to thebrackishwaters, 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 totallifespanof gobiid varies from one to ten years, again with the species in warmer waters generally living longer.[11]

Behavior

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Burrow construction

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Many species in the Gobiidae live in male-female pairs that construct and share burrows, similar to many other fish such asMozambique tilapia.The burrows are used for shelters andspawningplaces. Gobiids use their mouths to dig into the sea bottom, removing dead coral-fragments, rubble, andbenthicalgaein 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 highmoundover 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 thereproductive successis 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

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Kleptogamy refers to a "sneaking behavior" duringreproductionwhere 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 theirspermon the spawning ground as soon as the paired female releases hereggs.[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 givesparental careequally 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.
A 1865 watercolor painting of a Brazilian goby by Jacques Burkhardt

Sex change

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A few species of gobiid, such asblackeye gobyandLythrypnus dalli,can change their sexes. Sex change is possible in these gobies, since theexternal genitaliafor 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 inwrasses,damselfishes,and seabasses.[14]Female-to-male change usually occurs because the resident male of the group is dead. If no male is in the group,reproductionwill 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 manyoffspring.[14][18]

Some gobies have extraordinarily developed sex change ability.Gobiodon histriofrom theGreat Barrier Reefexhibits bidirectional sex changes.G. histriois one of the very few species that can change sex in both ways. When twoG. historiofemales, which used to be males, are on the same coral reef, one of them transforms back into a male goby.[18]

Sex determination

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Sex determination in coral gobyGobiodon erythrospilusdoes not occur until the juveniles meet potential mates.[14]Confronting a potential mate can be difficult forGobiodon erythrospilusjuveniles, 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, andvice versa.This type of sex determination is referred to as socially influenced sex determination.[19]

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Some gobiids rememberlandmarksthat are within short distances, and use them to find their ways. Small frillfin gobies (Bathygobius soporator) live inintertidal zones.They swim through the pools duringhigh tidesand memorize how each pool connects to the others. Then, duringlow 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

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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 betweenforagingand avoidingpredation,made choices that would better their foraging.[22]

Some marine gobies live in symbiosis with shrimp.

Symbiosis

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Species in the Gobiidae sometimes formsymbiotic relationshipswith other species,[23]such as with burrowingshrimps.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 itseggs.Only thealpha maleand 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 ofsymbiosisis demonstrated by the neon gobies (Elacatinusspp.). These gobiids, known as "cleaner gobies", removeparasitesfrom 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 stationswould otherwise treat such small fish as food (for example,groupersandsnapper). 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 ofsymbiosisexists between gobiids and the mushroom coralHeliofungia actiniformis(Fungiidae), in which representatives of the genusEviotaroam among the tentacles possibly hiding from predators.[24][25]

Commercial importance

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Dried gobies for sale on the market inOdesa,Ukraine

Gobiids have commercial importance inRussiaandUkraine.They arefishedin theSea of Azov,northwesternBlack SeaandCaspian Sea.Most important species areround goby,monkey goby,toad goby,andgrass goby.Thegrass gobyis also a commercial fish inItaly.

In the aquarium

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Several species of gobiids are kept in aquaria.[26]Most captive gobies are marine. Perhaps the most popular is the small but colorfulneon 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 includeRandall's shrimp gobyand thewatchman goby.

See also

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  • Sleeper gobiesare a closely related family (Eleotridae) that lack the fused pelvic fin sucker typical of most gobies, but are otherwise very similar in size, shape, and ecology.
  • Blenniesare a group of shallow-water marine fish often confused with gobies.
  • Dragonetsare superficially similar to gobies and sometimes confused with them.
  • Pholidichthys leucotaeniais commonly called the engineer goby or convict goby, but is not a goby.

References

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  1. ^Patzner, R.A.; Van Tassell, J.L.; Kovačić, M.; Kapoor, B.G., eds. (2011).The Biology of Gobies.Enfield, NH: Science Publishers. p. 685.ISBN978-1-57808-436-4.
  2. ^Agorreta, A.; San Mauro, D.; Schliewen, U.; Van Tassell, J.L.; Kovačić, M.; Zardoya, R.; Rüber, L. (2013). "Molecular phylogenetics of Gobioidei and phylogenetic placement of European gobies".Molecular Phylogenetics and Evolution.69(3): 619–633.Bibcode:2013MolPE..69..619A.doi:10.1016/j.ympev.2013.07.017.hdl:10261/123985.PMID23911892.
  3. ^Agorreta, A.; Rüber, L. (2012). "A standardized reanalysis of molecular phylogenetic hypotheses of Gobioidei".Systematics and Biodiversity.10(3): 375–390.Bibcode:2012SyBio..10..375A.doi:10.1080/14772000.2012.699477.
  4. ^Thacker, Christine E.; Dawn M. Roje (2011). "Phylogeny of Gobiidae and identification of gobiid lineages".Systematics and Biodiversity.9(4): 329–347.Bibcode:2011SyBio...9..329T.doi:10.1080/14772000.2011.629011.
  5. ^Romero, A., ed. (2001).The Biology of Hypogean Fishes.Developments in Environmental Biology of Fishes. pp. 35–36.ISBN978-1402000768.
  6. ^Winterbottom, Richard; et al. (2011). "Life span, growth and mortality in the western Pacific goby Trimma benjamini, and comparisons with T. nasa".Environmental Biology of Fishes.91(3): 295–301.Bibcode:2011EnvBF..91..295W.doi:10.1007/s10641-011-9782-6.
  7. ^abJ. S. Nelson; T. C. Grande; M. V. H. Wilson (2016).Fishes of the World(5th ed.). Wiley. p. 752.ISBN978-1-118-34233-6.Archived fromthe originalon 2019-04-08.Retrieved2018-08-11.
  8. ^Simonović, P.D.; Nikolić, V.P.; Skóra, K.E. (1996). "Vertebral number in Ponto-Caspian gobies: phylogenetic relevance".J. Fish Biol.49(5): 1027–1029.Bibcode:1996JFBio..49.1027S.doi:10.1111/j.1095-8649.1996.tb00098.x.
  9. ^Miller P.J. (1986) Gobiidae. In: Whitehead P.J.P., Bauchot M.-L., Hureau J.-C., Nielsen J., Tortonese E. (eds.) Fishes of the North-eastern Atlantic and the Mediterranean, Vol. 3. UNESCO, Paris.
  10. ^Pinchuk, V.I. (1991)."K voprosu o grupirovkakh vidov v predelakh rodaNeogobius(Perciformes) ".Voprosy Ikhtiologii.31:380–393.
  11. ^abHoese, Douglas F. (1998). Paxton, J.R.; Eschmeyer, W.N. (eds.).Encyclopedia of Fishes.San Diego: Academic Press. pp. 218–222.ISBN978-0-12-547665-2.
  12. ^abReebs, Stephan."Can fishes build things?"(PDF).
  13. ^abTakegaki, Takeshi; Akinobu Nakazono (June 1999). "Division of labor in the monogamous goby, Valenciennea longipinnis, in relation to burrowing behavior".Ichthyological Research.46(2): 125–129.Bibcode:1999IchtR..46..125T.doi:10.1007/BF02675430.
  14. ^abcdefghiReebs, Stephan."The sex lives of fishes"(PDF).
  15. ^Svensson, O.; Kvarnemo (2007)."Parasitic spawning in sand gobies: an experimental assessment of nest-opening size, sneaker male cues, paternity, and filial cannibalism".Behavioral Ecology.18(2): 410–419.doi:10.1093/beheco/arl098.
  16. ^Svensson, O; Magnhagen, C.; Forsgren, E.; Kvarnemo, C. (1998). "Parental behaviour in relation to the occurrence of sneaking in the common goby".Animal Behaviour.56(1): 175–179.doi:10.1006/anbe.1998.0769.PMID9710475.S2CID24806138.
  17. ^Lorenzi, V.; Earley, R.L.; Grober, M.S. (2006). "Preventing behavioural interactions with a male facilitates sex change in female bluebanded gobies, Lythrypnus dalli".Behavioral Ecology and Sociobiology.59(6): 715–722.doi:10.1007/s00265-005-0101-0.
  18. ^abMunday, P.L.; Caley, M.J.; Jones, G.P. (1998). "Bi-directional sex change in a coral-dwelling goby".Behavioral Ecology and Sociobiology.43(6): 371–377.doi:10.1007/s002650050504.
  19. ^Hobbs, J.-P. A.; Munday, P.L.; Jones, G.P. (2004)."Social induction of maturation and sex determination in a coral reef fish".Proceedings of the Royal Society of London.B 271 (1553): 2109–2114.doi:10.1098/rspb.2004.2845.PMC1691848.PMID15475329.
  20. ^Reebs, Stephan."How fishes find their ways around"(PDF).
  21. ^Aronson, L.R. (1971). "Further studies on orientation and jumping behaviour in the Gobiid fish, Bathygobius soporator".Annals of the New York Academy of Sciences.188(1): 378–392.Bibcode:1971NYASA.188..378A.doi:10.1111/j.1749-6632.1971.tb13110.x.PMID5288865.
  22. ^Magnhagen, C. (1988)."Changes in foraging as a response to predation risk in two gobiid fish species, Pomatoschistus minutus and Gobius niger".Marine Ecology Progress Series.49:21–26.Bibcode:1988MEPS...49...21M.doi:10.3354/meps049021.
  23. ^G. S. Helfman; B. B. Colette; D. E. Facey (1997). "Chapter 21: Fishes as social animals".The Diversity of Fishes.Blackwell.ISBN978-0-86542-256-8.
  24. ^Bos, Arthur R (2012)."Fishes (Gobiidae and Labridae) associated with the mushroom coralHeliofungia actiniformis(Scleractinia: Fungiidae) in the Philippines ".Coral Reefs.31(1): 133.Bibcode:2012CorRe..31..133B.doi:10.1007/s00338-011-0834-3.
  25. ^Bos AR, Hoeksema BW (2015). "Cryptobenthic fishes and co-inhabiting shrimps associated with the mushroom coral Heliofungia actiniformis (Fungiidae) in the Davao Gulf, Philippines".Environmental Biology of Fishes.98(6): 1479–1489.Bibcode:2015EnvBF..98.1479B.doi:10.1007/s10641-014-0374-0.
  26. ^Schäfer, Frank (2005).Brackish-Water Fishes.Aqualog.ISBN978-3936027822.
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