Overview

Comprehensive Description

Diversity

Wrasses (the family Labridae), are the most abundant and conspicuous fishes on tropical reefs around the world. Wrasses also comprise an important element of the coldwater fish population on temperate reefs. They are second largest family of marine fishes and the third largest family in the Perciformes order, containing approximately 60 genera and roughly 500 species. Wrasses appear in a diverse range of colors, shapes, and sizes, often varying considerably within individual species (see Physical Description). This morphological diversity is matched by the wide variety of prey consumed. Wrasses fill the roles of piscivores, zooplanktivores, molluscivores, herbivores, planktivores, polychaete predators, decapod crab predators, and coral predators, as well as many others (see Food Habits). Many wrasses are organized into harem-based social systems and hermaphroditism is common (see Reproduction: Mating Systems). Finally, as suggested by their diverse food habits, wrasses fill many important ecological roles on reefs of tropical and temperate regions around the world.

  • Choat, H., D. Bellwood. 1998. Wrasses & Parrotfishes. Pp. 209-213 in W Eschmeyer, J Paxton, eds. Encyclopedia of fishes – second edition. San Diego, CA: Academic Press.
  • Wainwright, P., D. Bellwood. 2002. Ecomorphology of Feeding in Coral Reef Fishes. Pp. 33-55 in P Sale, ed. Coral Reef Fishes: Dynamics and Diversity in a Complex Ecosystem. San Diego, CA: Academic Press.
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Distribution

Geographic Range

Wrasses occupy all tropical seas and penetrate considerable distances into temperate waters, reaching as far north as Norway. Many temperate species in the genera Oxyjulius, Tautoga, Tautogolabrus, Semicossyphus, and Labrus can be found in both the Atlantic and Pacific Oceans. Wrasses are most highly concentrated off the coasts of Australia where about 165 species and 42 genera are represented.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native ); oceanic islands (Native ); indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native ); mediterranean sea (Native )

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Physical Description

Morphology

Physical Description

Most wrasses are quite small, usually below 20 cm. The smallest species, Minilabrus striatus of the Red Sea, reaches a maximum length of only 4.5 cm. The genera Pseudocheilinus and Doratonotus contain several other dwarf wrasses. One species, Conniella apterygia, is so small that it lacks even pelvic fins and a supporting skeleton. The largest wrasse, Cheilinus undulatus, can reach a length of about 2.3 m and weighs more than 150 kg. Wrasses are most easily identified by their pointed snouts and prominent canine teeth in the front of the jaws, which often project forward. Wrasses characteristically have a protractile mouth,   cycloid scales , and a single continuous dorsal fin lacking an obvious notch between the soft and spiny portions. The lateral line may be continuous or interrupted. (Click here to see a   fish diagram).

Wrasses display myriad colors and shapes. Razorfishes are elongate and laterally compressed, while members of Cheilinus, Choerodon, and many of Bodianus are large and stocky. However, most are elongate and tapered at both ends, often referred to as “cigar-shaped.” Cigar-shaped fishes are found in the genera Thalassoma, Halichoeres, and Labroides. Often, there is considerable diversity of colors and shapes within individual species. As in parrotfishes, some wrasses progress through “phases” (see Reproduction: Mating Systems), and each phase corresponds with a change in morphology (shape and color). Dominant males (and sometimes females) are the most distinctly colored, with complex patterns of red, yellow, green, blue and black. Subordinate males and females are smaller than dominant individuals and are often drab-colored with cryptic patterns. Juveniles range in coloration from bright yellow and orange to drab gray and brown, and some have camouflaging patterns. (See Reproduction: Mating Systems for details). Some wrasses exhibit   sexual dimorphism.

Other Physical Features: ectothermic ; bilateral symmetry ; polymorphic

Sexual Dimorphism: male larger; sexes colored or patterned differently; male more colorful; sexes shaped differently

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Ecology

Habitat

Habitat

Wrasses can be found in a wide variety of habitats, such as tidal pools, grass beds, rocky or coral reefs, or open sand bottoms. Many wrasses prefer specific environments. Doratonotus, for example, prefer turtle grass beds, Hemipteronotus, mixed turtle grass and sandy patch areas, and hogfishes, weed-covered rocky flats. Plankton feeders, such as Clepticus, often concentrate in large schools at reef fronts, reef gaps, or other areas where plankton is concentrated in the water column. However, some species, such as the slippery dick, can be found in a broad range of habitats.

Habitat Regions: temperate ; tropical ; saltwater or marine

Aquatic Biomes: benthic ; reef ; coastal ; brackish water

Other Habitat Features: estuarine ; intertidal or littoral

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Trophic Strategy

Food Habits

Many wrasses are specialized and voracious feeders, as reflected by the highly variable skull and body shape, modified   pharyngeal jaw, and prominent canines. The type of nourishment ranges widely: fish, ectoparasites, mollusks, polychaete worms, decapod crabs, corals, coral mucous, amphipods, various echinoderms, plankton, and several types of vegetation. Many small wrasses follow larger fishes and exploit any benthic (reef bottom) disturbances that help to reveal the well-camouflaged invertebrates. A considerable number are plankton feeders, forming schools in reef gaps, reef fronts or other areas with current. The food habits of cleaner wrasses are probably most well known. Cleaner wrasses remove mucous, parasites and scales from the bodies of larger fishes. Cleaning is not limited to the Labroides genus however; young bluehead and young Spanish hogfish in the Bahamas have also been observed cleaning larger fishes. Finally, some piscivorous (fish-eating) wrasses mimic harmless fishes (Randall and Kuiter, 1989 in Nelson, 1994).

Primary Diet: carnivore (Piscivore , Eats eggs, Eats body fluids, Eats non-insect arthropods, Molluscivore ); herbivore ; omnivore

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Associations

Ecosystem Roles

The ecological role of cleaner wrasses of the Indo-Pacific region provides a good example of the complexity of seemingly mutualistic relationships between fishes. Typically, cleaner fishes are elaborately colored and perform displays over a patch of reef while larger fish approach and assume a relaxed posture. Cleaner fishes are commonly thought to benefit the host by removing dead or damaged tissue and ectoparasites. Accordingly, investigators reported higher recovery rates for wounded fish in the presence of cleaners. However, in experiments where all cleaners were removed from an environment there was no incidence of fishes leaving the area or becoming particularly unhealthy. Further, when levels of parasitic infections are high the host benefits from cleaning but when infection levels are low, which they usually are, some cleaners feed on healthy tissue, such as scales, pieces of fin, mucous, or in some cases the eggs of other reef fishes. Despite these parasitic qualities of the relationship, fishes being cleaned have a positive response to the tactile stimulation from cleaners, suggesting that some cleaners are mildly beneficial while others have taken advantage of the cleaning arrangement.

The relationship between wrasse species and their invertebrate prey is a spectacular example of coevolution. As invertebrates have developed anti-predator adaptations, such as spines, toxins, heavy armor, and adherence to the substrate, wrasses have evolved simultaneously. Some physical changes include the development of strong, hard beaks and a second set of strong teeth in the throat (  pharyngeal jaw), making it possible to crush hard-shelled invertebrates. A conspicuous behavioral adaptation is “following behavior.” As larger fish disturb the substrate, some wrasses follow close behind to capture exposed invertebrates. Other small wrasses have become adept at combing the reef for invertebrates too small for most fishes to prey upon. Finally, some wrasses use their snouts to flip rocks and pieces of coral to expose hidden invertebrates.

Ecosystem Impact: parasite

Species Used as Host:

  • some fishes

Mutualist Species:

  • many fish species

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Predation

Many juvenile wrasses are cryptically colored to avoid predation while others find protection among the tentacles of sea anemones. Nearly all adult wrasses bury themselves in sand at night to avoid predators. A few species seek out reef crevices and produce a foul-smelling   mucous bag to deter predators while sleeping. Razorfishes (Hemipteronotus, Xyrichtys) also use the sand for protection during the day by diving into the bottom. Razorfishes are apparently quite agile in this environment, sometimes resurfacing several meters from the point of entry.

Known Predators:

Anti-predator Adaptations: mimic; cryptic

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Known predators

Labridae is prey of:
Actinopterygii

This list may not be complete but is based on published studies.
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Known prey organisms

Labridae preys on:
non-insect arthropods
Amphiprion percula

This list may not be complete but is based on published studies.
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Life History and Behavior

Behavior

Communication and Perception

Most wrasses rely on vision to find their prey. Visual recognition may also be important for terminal phase (TP) males to identify harem members. Although TP males are susceptible to streaking attempts by initial phase (IP) males (see Reproduction: Mating Systems), no IP males have been found in harem-forming species. This suggests that IP males are unable to mimic IP females, despite very similar morphology.

Communication Channels: visual

Other Communication Modes: mimicry

Perception Channels: visual

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Life Cycle

Development

Wrasses occupy a wide range of water temperatures and incubation time is directly affected by water temperature. In laboratory experiments incubation took approximately 24 hours at 27˚C. The planktonic stage is estimated to be around one month, although very little is known about this stage. The age or size at which individuals reach sexual maturity depends on the maximum size of the species.

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Life Expectancy

Lifespan/Longevity

No information was found concerning the lifespan of wrasses but, in general, reef species live between three and five years.

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Reproduction

Reproduction

Like parrotfishes, many wrasses utilize some of the most complex and unusual reproduction systems known to fishes. Males can be either primary (born male), or secondary (females that have undergone sex change). In some species there are no secondary males while in others all individuals are born female (monandric) and change sex when necessary. In the most complex systems, species are diandric – both primary and secondary males exist in the population. In these species, individuals proceed through three distinct phases, marked by color differences. In fact, the color differences are so pronounced that for over 200 years researchers regarded some phases as distinct species. Sexually immature juveniles represent the first phase. The second, known as the initial, phase (IP) can include sexually mature males or females, which are impossible to tell apart without internal examination or observation during spawning. IP males and females may group spawn in some species. The terminal phase (TP) includes only mature males, which display brilliant colors. TP males usually dominate reproductive activity through a harem-based social system. The death of a TP male serves as a social cue for an IP female to change sex and behavior. The morphology of IP males may also change in response to the death of a TP male. In some cases, IP males attempt to fertilize IP females by following a TP male and IP female pair during spawning. In this behavior, called “streaking,” IP males follow the pairs at peak spawning and release a large cloud of gametes in an attempt to overwhelm fertilization by the TP male. This is thought to increase the fecundity (ability to produce offspring) of IP males. IP males are well equipped to perform streaking as they have larger gonads and so are able to produce more gametes, while TP males have smaller testes and rely on aggression to deter other males. The larger volume of milt (gametes) produced by IP males is related to group spawning events with IP females, in which competition for fertilization is intense and more milt is needed.

Some specific examples of wrasse mating systems demonstrate the complexity and variation of the phase system described above. For instance, the cleaner wrasse, which is monandric (all individuals are born female), forms harems that are held together by male aggression towards subordinate females. With the death of the dominant male, subordinate females jockey for position and the newly dominant female adopts aggressive male behavior within a few hours. Each individual moves a step up in the dominance hierarchy and the last position is filled by a juvenile. If the newly dominant female is able to withstand attempts by neighboring males to take over the vacant harem, she will become a fully functional male within a two to four days. Some other harem-forming species are Cirrhilabrus temminckii, Cirrhilabrus jordani, Labroides bicolor, Hemipteronotus splendens, Pseudocheilinus hexataenia and Macropharyngodon moyeri. The Caribbean species Halichoeres garnoti is also monandric, but individuals do not exhibit territoriality or conspicuous dominance relationships, nor do they use aggressive actions to maintain sexual state. Instead, size or some size-related factor determines which individual will fill the male role. In Halichoeres garnoti males are larger than females and both sexes behave similarly. While these examples focus on the mating extremes of wrasses, most species fall between the systems of the cleaner wrasse and Halichoeres garnoti in terms of the influence of social control on sex reversal. Other hermaphroditic but non-harem-forming species include Halichoeres bivittatus and Halichoeres poeyi, Halichoeres maculipinna and possibly Thalassoma lunare. Finally, some species, such as Oxyjulis californica and Crenilabrus melops, do not follow the phase system at all as they are not hermaphroditic, and there are probably more non-hermaphroditic species yet to be found.

Mating System: polygynous ; polygynandrous (promiscuous)

In tropical wrasses spawning occurs year-round but some temperate species seem to restrict spawning to warmer parts of the year. Spawning typically occurs along the outer edge of patch reefs or along the outer edges of more extensive reef complexes. The correlation between spawning and lunar periodicity (the lunar cycle) is sketchy in some species and non-existent in most that have been investigated. Spawning in several species corresponds with outgoing tides, however, many species spawn at a particular time in the day, regardless of tidal patterns. This variation may be due to local conditions. For instance, in areas where tidal forces are weak, factors like time of day or light intensity may have more influence. However, evidence from different species on the same reef suggests that temporal (measured time) differences in spawning evolved to decrease the probability of hybridization with other species.

Wrasses may spawn in groups or pairs depending on the species or phase of individuals. Typically, group or aggregate spawning occurs between initial phase (IP) individuals, which are diandric (containing male and female IP individuals). However, in some species, such as Thalassoma cupido, Thalassoma lucasanum, and Halichoeres bivitattus, terminal phase (TP) males have been observed participating in group spawning. The size of the spawning groups ranges from a dozen to several hundred individuals. Males outnumber females, sometimes by as much as ten to one. Paired spawning is found in many, if not all, tropical wrasses and involves a TP male and IP female. In rare cases, IP individuals also spawn in pairs. Most species defend small territories only during spawning. Currently Anampses cuvieri is the only known species of tropical wrasse to produce demersal eggs (eggs laid on the bottom as opposed to being released in the water column). Demersal spawning of Anampses cuvieri was only observed in captivity and still needs to be confirmed, but work on other species of this genus seems to support this observation.

Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sequential hermaphrodite (Protogynous ); sexual ; fertilization (External ); oviparous

Some temperate wrasse species, such as the ballan wrasse and Anampses cuvieri, are demersal nest builders. The nests are usually made out of plant material and the male guards the eggs after they are deposited.

Parental Investment: male parental care

  • Choat, H., D. Bellwood. 1998. Wrasses & Parrotfishes. Pp. 209-213 in W Eschmeyer, J Paxton, eds. Encyclopedia of fishes – second edition. San Diego, CA: Academic Press.
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Molecular Biology and Genetics

Barcode

Locations of barcode samples

Collection Sites: world map showing specimen collection locations for Labridae
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Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
                                                             
Specimen Records:2,698
Specimens with Sequences:2,696
Specimens with Barcodes:2,380
Public Records:535
Species:327
Species With Barcodes:317
  
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Barcode data

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Conservation

Conservation Status

Conservation Status

Four labrid species are listed as vulnerable: Cheilinus undulates, Lachnolaimus maximus, Thalassoma ascensionis, and Xyrichtys virens.

  • The World Conservation Union, 2002. "IUCN 2002" (On-line). 2002 IUCN Red List of Threatened Species. Accessed August 22, 2003 at http://www.iucnredlist.org/.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

No specific information was found concerning any negative impacts to humans.

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Economic Importance for Humans: Positive

Wrasses from the Coris genera are popular aquarium fishes and two species from the Atlantic coast of North America, the cunner and the tautog, are valued as commercial and sport fish. Some other medium to large wrasses are popular food fishes as well.

Positive Impacts: pet trade ; food ; research and education

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Wikipedia

Wrasse

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 nine subgroups or tribes.[1] They are typically small fish, most of them less than 20 centimetres (7.9 in) long, although the largest, the Humphead wrasse, can measure up to 2.5 metres (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.[2]

The word "wrasse" comes via Cornish from the Welsh word gwrach meaning an old woman or hag.[3]

Contents

Distribution

Wrasses inhabit 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

Wrasses have protractile mouths, usually with separate jaw teeth that jut outwards.[4] 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.)[5] The dorsal fin has 8–21 spines and 6–21 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 or IP individuals) but the largest adults become territory-holding (Terminal Phase or TP) males.[4]

The wrasses have become a primary study species in fish-feeding biomechanics due to their jaw structure. 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 4 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.[4]

Reproductive behavior

Most labroids are protogynous hermaphrodites within a haremic mating system.[6] [7] Hermaphroditism allows for complex mating systems. Labroids exhibit three different mating systems: polygynous, lek-like, and promiscuous mating systems.[8] Group spawning and pair spawning occur within mating systems. The type of spawning that occurs depends on male body size.[7] Labroids typically exhibit broadcast spawning, releasing high amounts of planktonic eggs, which are broadcast by tidal currents; adult labroids have no interaction with offspring.[9] Wrasse of a particular subgroup of the Labridae family Labrini do not exhibit broadcast spawning.

Broodcare behavior of Labrine tribe

The subgroup Labrini arose from a basal split within family Labridae during the Eocene period.[1] Subgroup Labrini is composed of eight genera, wherein 15 out of 23 species exhibit broodcare behavior.[9] 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.[9] In species that express this behavior, eggs cannot survive without parental care.[10] Species of Symphodus, Centrolabrus, and Labrus genera exhibit broodcare behavior.

Cleaner wrasse

Photo of two small wrasses cleaning large wrasse's gills
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" fishes 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.[11]

Other wrasse species,[which?] rather than inhabiting fixed locations, make "house calls" — that is, their "clientele" are too territorial or shy to go to a cleaning station.[citation needed]

Significance to humans

Humans eat wrasse in many places. In the western Atlantic, the most common food species is the tautog.[5][dubious ]

Wrasse are common in both public and home aquaria. Some species are small enough to be considered reef safe.

Wrasse may be employed as cleaner fish to combat sea-lice infestation in salmon farms.[12]

Gallery

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
Ctenolabrus
Cymolutes
Decodon

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

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

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

Timeline

References

  1. ^ a b 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. 
  2. ^ Choat, J.H. & Bellwood, D.R. (1998). Paxton, J.R. & Eschmeyer, W.N.. ed. Encyclopedia of Fishes. San Diego: Academic Press. pp. 211. ISBN 0-12-547665-5. 
  3. ^ [1]
  4. ^ a b c Wainwright et al. (2005). "Many-to-One Mapping of Form to Function: A General Principle in Organismal Design?". Integrative & comparative biology 45: 256–262. 
  5. ^ a b  Chisholm, Hugh, ed. (1911). "wrasse". Encyclopædia Britannica (11th ed.). Cambridge University Press. 
  6. ^ 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. 
  7. ^ a b Kazancioglu, E.; S.H. Alonzo. "A comparative analysis of sex change in Labridae supports the size advantage hypothesis". Evolution 64: 2254–2264. 
  8. ^ 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. 
  9. ^ a b c Hanel, R.; M. W. Westneat and C. Sturmbauer. "Phylogenetic relationships, evolution of broodcare behavior, and geographic speciation in the Wrasse tribe Labrini.". Journal of Molecular Evolution 55: 776–789. 
  10. ^ Taborsky, M.; B. Hudde and P. Wirtz. "Reproductive-behavior and ecology of Symphodus (Crenilabrus) ocellatus, a European wrasse with 4 types of male-behavior.". Behaviour 102: 82–118. 
  11. ^ Trivers, R. L. 1971
  12. ^ "Sea Lice". Scottish Salmon Producers' Organisation. http://www.scottishsalmon.co.uk/science/sea_lice%284%29.aspx. Retrieved 8 June 2011. 
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