Wrasse

Wrasses
Moon wrasse, Thalassoma lunare, a typical wrasse
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Perciformes
Suborder:	Labroidei
Family:	Labridae G. Cuvier, 1816
Genera
See text.

The wrasses are a family, Labridae, of marine fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 82 genera, which are divided into 9 subgroups or tribes.^[1][2][3] They are typically small fish, most of them less than 20 cm (7.9 in) long, although the largest, the humphead wrasse, can measure up to 2.5 m (8.2 ft). They are efficient carnivores, feeding on a wide range of small invertebrates. Many smaller wrasses follow the feeding trails of larger fish, picking up invertebrates disturbed by their passing.^[4] Juveniles of some representatives of the genera Bodianus, Epibulus,Cirrhilabrus, Oxycheilinus, and Paracheilinus hide among the tentacles of the free-living mushroom coral Heliofungia actiniformis.^[5][6]

The word "wrasse" comes from the Cornish word wragh, a lenited form of gwragh, meaning an old woman or hag, via Cornish dialect wrath. It is related to the Welsh gwrach and Breton gwrac'h.^[7]

Distribution

Wrasses inhabit the tropical and subtropical waters of the Atlantic, Indian, and Pacific Oceans; usually in shallow-water habitats such as coral reefs and rocky shores where they live close to the substrate.

Anatomy

Lips of Labrus festivus

Wrasses have protractile mouths, usually with separate jaw teeth that jut outwards.^[8] Many species can be readily recognized by their thick lips, the inside of which is sometimes curiously folded, a peculiarity which gave rise the German name of "lip-fishes" (Lippfische.)^[9] and the Dutch name of "lipvissen". The dorsal fin has eight to 21 spines and six to 21 soft rays, usually running most of the length of the back. Wrasse are sexually dimorphic. Many species are capable of changing sex. Juveniles are a mix of males and females (known as initial phase individuals), but the largest adults become territory-holding (terminal phase) males.^[8]

The wrasses have become a primary study species in fish-feeding biomechanics due to their jaw structures. The nasal and mandibular bones are connected at their posterior ends to the rigid neurocranium, and the superior and inferior articulations of the maxilla are joined to the anterior tips of these two bones, respectively, creating a loop of four rigid bones connected by moving joints. This "four-bar linkage" has the property of allowing numerous arrangements to achieve a given mechanical result (fast jaw protrusion or a forceful bite), thus decoupling morphology from function. The actual morphology of wrasses reflects this, with many lineages displaying different jaw morphology that results in the same functional output in a similar or identical ecological niche.^[8]

Reproductive behavior

Most labroids are protogynous hermaphrodites within a haremic mating system.^{[10] [11]} A good example of this reproductive behavior is seen in the California sheephead. Hermaphroditism allows for complex mating systems. Labroids exhibit three different mating systems: polygynous, lek-like, and promiscuous mating systems.^[12] Group spawning and pair spawning occur within mating systems. The type of spawning that occurs depends on male body size.^[11] Labroids typically exhibit broadcast spawning, releasing high numbers of planktonic eggs, which are broadcast by tidal currents; adult labroids have no interaction with offspring.^[13] Wrasse of a particular subgroup of the Labridae family Labrini do not exhibit broadcast spawning.

Sex change in wrasse is generally female-to-male, but experimental conditions have allowed for male-to-female sex change. Placing two male Labroides dimidiatus wrasses in the same tank will result in the smaller of the two becoming female again.^[14] Additionally, while the individual to change sex is generally the largest female,^[15] evidence also exists of the largest female instead "choosing" to remain female in situations in which she can maximize her evolutionary fitness by refraining from changing sex.^[16]

Broodcare behavior of the labrine tribe

The subgroup Labrini arose from a basal split within family Labridae during the Eocene period.^[3] Subgroup Labrini is composed of eight genera, wherein 15 of 23 species exhibit broodcare behavior.^[13] Broodcare behavior ranges from simple to complex parental care of spawn; males build algae nests or crude cavities, ventilate eggs, and defend nests against conspecific males and predators.^[13] In species that express this behavior, eggs cannot survive without parental care.^[17] Species of Symphodus, Centrolabrus, and Labrus genera exhibit broodcare behavior.

Cleaner wrasse

Cleaner wrasses working on dragon wrasse Novaculichthys taeniourus, Labroides sp. on a reef in Hawaii

Cleaner wrasse are the best-known of the cleaner fish. They live in a cleaning symbiosis with larger, often predatory fish, grooming them and benefiting by feeding on what they remove. "Client" fish congregate at wrasse "cleaning stations" and wait for the cleaner fish to remove gnathiid parasites, even swimming into their open mouths and gill cavities.

Cleaner wrasses are best known for feeding on dead tissue and scales and ectoparasites, although they are also known to 'cheat', consuming healthy tissue and mucus, which is energetically costly for the client fish to produce. The bluestreak cleaner wrasse, Labroides dimidiatus, is one of the most common cleaners found on tropical reefs. Few cleaner wrasses have been observed being eaten by predators, possibly because parasite removal is more important for predator survival than the short-term gain of eating the cleaner.^[18]

Significance to humans

In the western Atlantic, in the past the most common food species for humans has been the tautog.^[9] Wrasses are common in both public and home aquaria. Some species are small enough to be considered reef safe. They may be employed as cleaner fish to combat sea-lice infestation in salmon farms.^[19] Commercial fish farming of cleaner wrasse for sea-lice pest control in commercial salmon farming has developed in Scotland as lice busters, with apparent commercial benefit and viability.

Parasites

As all fish, labrids are the hosts of a number of parasites. A list of 338 parasite taxa from 127 labrid fish species was provided by Muñoz & Diaz in 2015.^[20] An example is the nematode Huffmanela ossicola.

Gallery

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  Giant Napoleon Wrasse in Apo Reef, Philippines

• 

  Bird wrasse, Gomphosus varius, Kona (Hawaii)

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  A yellowtail coris wrasse, Coris gaimard, is being cleaned by Labroides phthirophagus in Hawaii.

• 

  Bluehead wrasse, Belize Barrier Reef

• 

  Clown wrasse, Coris aygula, Red Sea

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  Yellowtail wrasse, Coris gaimard, Hawaii

• 

  Pearl wrasse, Anampses cuvieri, Hawaii

• 

  Humphead wrasse, Cheilinus undulatus, Melbourne Aquarium

• 

  Bluestreak wrasse, Labroides dimidiatus

• 

  Gomphosus varius

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  Gomphosus caeruleus swimming with a yellow goatfish

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  Six-line wrasse (Pseudocheilinus hexataenia)

Subgroups and tribes

Hypsigenyines Labrines Cheilines Scarines Pseudocheilines Novaculines Pseudolabrines Labrichthyines Julidines

Genera

Acantholabrus Achoerodus Ammolabrus Anampses Anchichoerops Austrolabrus Bodianus Centrolabrus Cheilinus Cheilio Choerodon Cirrhilabrus Clepticus Conniella Coris Ctenolabrus Cymolutes Decodon

Diproctacanthus Doratonotus Dotalabrus Epibulus Eupetrichthys Frontilabrus Gomphosus Halichoeres Hemigymnus Hologymnosus Iniistius Labrichthys Labroides Labropsis Labrus Lachnolaimus Lappanella Larabicus

Leptojulis Macropharyngodon Malapterus Minilabrus Notolabrus Novaculichthys Novaculoides Novaculops Ophthalmolepis Oxycheilinus Oxyjulis Paracheilinus Parajulis Pictilabrus Polylepion Pseudocheilinops Pseudocheilinus Pseudocoris

Pseudodax Pseudojuloides Pseudolabrus Pteragogus Sagittalarva Semicossyphus Stethojulis Suezichthys Symphodus Tautoga Tautogolabrus Terelabrus Thalassoma Wetmorella Xenojulis Xiphocheilus Xyrichtys

Timeline

References

1. Parenti, Paolo; John E. Randall (15 April 2011). "Checklist of the species of the families Labridae and Scaridae: an update". Smithiana Bulletin 13: 29–44.
2. Parenti, Paolo; John E. Randall (June 2000). "An annotated checklist of the species of the labroid fish families Labridae and Scaridae". Ichthyological Bulletin 68: 1–97. ISSN 0073-4381.
3. Cowman, P.F.; D.R. Bellwood and L. van Herwerden. "Dating the evolutionary origins of wrasse lineages (Labridae) and the rise of trophic novelty on coral reefs". Molecular Phylogenetics and Evolution 52: 621–631. doi:10.1016/j.ympev.2009.05.015. PMID 19464378. Cite uses deprecated parameter |coauthors= (help)
4. Choat, J.H. & Bellwood, D.R. (1998). Paxton, J.R. & Eschmeyer, W.N., ed. Encyclopedia of Fishes. San Diego: Academic Press. p. 211. ISBN 0-12-547665-5.
5. Bos, Arthur R (2012). "Fishes (Gobiidae and Labridae) associated with the mushroom coral Heliofungia actiniformis (Scleractinia: Fungiidae) in the Philippines". Coral Reefs 31: 133. doi:10.1007/s00338-011-0834-3.
6. 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. doi:10.1007/s10641-014-0374-0.
7. "Wrasse | Define Wrasse at Dictionary.com". Dictionary.reference.com. Retrieved 2012-06-28.
8. Wainwright; et al. (2005). "Many-to-One Mapping of Form to Function: A General Principle in Organismal Design?". Integrative & comparative biology 45: 256–262. doi:10.1093/icb/45.2.256.
9.  Chisholm, Hugh, ed. (1911). "wrasse". Encyclopædia Britannica (11th ed.). Cambridge University Press.
10. Robertson, D.R.; R.R. Warner. "Sexual patterns in the labroid fishes of the Western Caribbean II: the parrotfishes (Scaridae)". Smithsonian Contributions to Zoology 255: 1–26. doi:10.5479/si.00810282.255.
11. Kazancioglu, E.; S.H. Alonzo. "A comparative analysis of sex change in Labridae supports the size advantage hypothesis". Evolution 64: 2254–2264. doi:10.1111/j.1558-5646.2010.01016.x. PMID 20394662.
12. Colin, P.L.; L. J. Bell. "Aspects of the spawning of labrid and scarid fishes (Pisces, Labroidei) at Enewetak Atoll, Marshall Islands with notes on other families (corrected reprint.)". Environmental Biology of Fishes 33: 330–345. doi:10.1007/BF00005881.
13. Hanel, R.; M. W. Westneat; C. Sturmbauer (December 2002). "Phylogenetic relationships, evolution of broodcare behavior, and geographic speciation in the Wrasse tribe Labrini.". Journal of Molecular Evolution 55 (6): 776–789. doi:10.1007/s00239-002-2373-6. PMID 12486536.
14. Kuwamura, T.; N. Tanaka; Y. Nakashima; K. Karino; Y Sakai (2002). "Reversed sex-change in the protogynous reef fish Labroides dimidiatus.". Ethology 108: 443–450. doi:10.1046/j.1439-0310.2002.00791.x.
15. Munday, P. L.; C. A. Ryen; M. I. McCormick; S. P. W. Walker (2009). "Growth acceleration, behaviour and otolith check marks associated with sex change in the wrasse Halichoeres miniatus.". Coral Reefs 28: 623–634. doi:10.1007/s00338-009-0499-3.
16. Munoz, R. C.; R. R. Warner (2003). "A new version of the size-advantage hypothesis for sex change: incorporating sperm competition and size-fecundity skew.". American Naturalist 161: 749–761. doi:10.1086/374345.
17. Taborsky, M.; B. Hudde; P. Wirtz. "Reproductive-behavior and ecology of Symphodus (Crenilabrus) ocellatus, a European wrasse with 4 types of male-behavior.". Behaviour 102: 82–118. doi:10.1163/156853986x00063.
18. Trivers, R. L. 1971
19. "Sea Lice". Scottish Salmon Producers' Organisation. Retrieved 8 June 2011.
20. Muñoz G., Diaz P.E. 2015: Checklist of parasites of labrid fishes (Pisces: Labridae). Viña del Mar, Chile. PDF.

• Sepkoski, Jack (2002). "A compendium of fossil marine animal genera". Bulletins of American Paleontology 364: 560. Retrieved 2011-05-18.

External links

• FishBase info for Labridae
• How Fish Hire a Cleaning Service
• Male and Female Images or Rock Wrasse Fish
• Smith, J.L.B. 1957. List of the fishes of the Family Labridae in the Western Indian Ocean. Ichthyological Bulletin; No. 7. Department of Ichthyology, Rhodes University, Grahamstown, South Africa.