Thalassoma bifasciatum (Bluehead Wrasse) juvenile yellow stage over Bispira brunnea (Social Fea
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''Thalassoma bifasciatum'', the bluehead, bluehead wrasse or blue-headed wrasse, is a species of marine
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
, a
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 81 genera, which are divided into 9 subgroups or tribes. They are typically small, most of them le ...
from the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Labridae. It is native to the
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups. C ...
s of the tropical waters of the western
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
. Individuals are small (less than 110 mm standard length) and rarely live longer than two years. They form large
schools A school is an educational institution designed to provide learning spaces and learning environments for the teaching of students under the direction of teachers. Most countries have systems of formal education, which is sometimes compulsor ...
over the reef and are important
cleaner fish Cleaner fish are fish that show a specialist feeding strategy by providing a service to other species, referred to as clients, by removing dead skin, ectoparasites, and infected tissue from the surface or gill chambers. This example of cleaning ...
in the reefs they inhabit.


Distribution and habitat

''Thalassoma bifasciatum'' is found in coral reefs of the
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
. Its main range includes the
Caribbean Sea The Caribbean Sea ( es, Mar Caribe; french: Mer des Caraïbes; ht, Lanmè Karayib; jam, Kiaribiyan Sii; nl, Caraïbische Zee; pap, Laman Karibe) is a sea of the Atlantic Ocean in the tropics of the Western Hemisphere. It is bounded by Mexico ...
and the southeast area of the
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an ocean basin and a marginal sea of the Atlantic Ocean, largely surrounded by the North American continent. It is bounded on the northeast, north and northwest by the Gulf Coast of the United ...
.


Description

Young/small females and males have yellow upper bodies and white lower bodies, often with green or black lateral stripes and occasionally dark vertical bars. This coloration is known as the initial phase. They can rapidly alter the presence or intensity of their yellow color, stripes, and bars, and these color changes appear to correspond to behavioral changes. Large females and some males can permanently change coloration and/or sex and enter the terminal phase coloration, which has a blue head, black and white bars behind the head, and a green body. This color phase gives the species its name. Terminal phase males are larger (70 to 80 mm) than the initial phase males (60 mm).


Ecology


Diet

''Thalassoma bifasciatum'' forages for
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, mollusks, small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s (such as shrimp and
krill Krill are small crustaceans of the order Euphausiacea, and are found in all the world's oceans. The name "krill" comes from the Norwegian word ', meaning "small fry of fish", which is also often attributed to species of fish. Krill are consi ...
),
worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always). Worms vary in size from microscopic to over in length for marine polychaete wo ...
s, other motile
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s, and the
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s of smaller fish,https://sta.uwi.edu/fst/lifesciences/sites/default/files/lifesciences/documents/ogatt/Thalassoma_bifasciatum%20-%20Bluehead%20Wrasse.pdf as well as ectoparasites on other fish. Initial phase males eat primarily zooplankton from currents, and females and initial phase males have certain hunting times during the day.


Predators and parasites

Though ''Thalassoma bifasciatum'' is a common
cleaner fish Cleaner fish are fish that show a specialist feeding strategy by providing a service to other species, referred to as clients, by removing dead skin, ectoparasites, and infected tissue from the surface or gill chambers. This example of cleaning ...
in the coral reefs they inhabit, they avoid cleaning piscivores such as the spotted moray, the
graysby The graysby (''Cephalopholis cruentata'') is a species of marine ray-finned fish, a grouper from the subfamily Epinephelinae which is in the family Serranidae which also includes the anthias and sea basses. It is found in the western Atlantic ...
, and the
red hind The red hind (''Epinephelus guttatus''), also known as the koon or lucky grouper in Caribbean vernacular, is a species of marine ray-finned fish, a grouper from the subfamily Epinephelinae which is part of the family Serranidae, which also in ...
. Such species will view them as prey, but will not view
gobies Gobiidae or gobies 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. Most of gobiid fish are relatively small, typically less than in length, and the ...
, another kind of cleaner fish, as prey. Other predators include the
greater soapfish Greater may refer to: *Greatness, the state of being great *Greater than, in inequality * ''Greater'' (film), a 2016 American film *Greater (flamingo), the oldest flamingo on record * "Greater" (song), by MercyMe, 2014 *Greater Bank, an Australian ...
,
roughtail stingray The roughtail stingray (''Bathytoshia centroura'') is a species of stingray in the family Dasyatidae, with separate populations in coastal waters of the northwestern, eastern, and southwestern Atlantic Ocean. This bottom-dwelling species typica ...
, and the
trumpetfish The trumpetfishes are three species of highly specialized, tubularly-elongated marine fishes in the genus ''Aulostomus'', of the monogeneric family Aulostomidae. The trumpetfishes are members of the order Syngnathiformes, together with the sea ...
. A significant
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
of ''T. bifasciatum'' is the
intracellular This glossary of biology terms is a list of definitions of fundamental terms and concepts used in biology, the study of life and of living organisms. It is intended as introductory material for novices; for more specific and technical definitions ...
myxozoan Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. O ...
''
Kudoa ovivora ''Kudoa'' is a genus of Myxozoa and the only genus recognized within the monotypic family Kudoidae. There are approximately 100 species of ''Kudoa'' all of which parasitize on marine and estuarine fish. ''Kudoa'' are most commonly known and studi ...
'', which can be found in the ovaries of females. Infected eggs are sterile and are also larger than uninfected eggs, and contain more organic and inorganic material. This implies the parasite causes a shift in resources from the mother to the eggs and decreases the fitness of the mother. Infected bluehead wrasses have been found to change sex earlier than uninfected females, possibly in response to the parasite.


Conservation status

''Thalassoma bifasciatum'' is widespread in the northwestern Atlantic region and is one of the most abundant species in coral reefs near
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and unincorporated ...
, the
Virgin Islands The Virgin Islands ( es, Islas Vírgenes) are an archipelago in the Caribbean Sea. They are geologically and biogeographically the easternmost part of the Greater Antilles, the northern islands belonging to the Puerto Rico Trench and St. Cro ...
, and the Netherlands Antilles. It is listed under
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
. However, this species shows high site fidelity, so coral reef destruction could cause local endangerment. Due to its bright coloration, it is sometimes collected for the
aquarium trade Fishkeeping is a popular hobby, practiced by aquarists, concerned with keeping fish in a home aquarium or garden pond. There is also a piscicultural fishkeeping industry, serving as a branch of agriculture. Origins of fishkeeping Fish ha ...
, but this is not a threat to the species.


Behavior

Like many other
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 81 genera, which are divided into 9 subgroups or tribes. They are typically small, most of them le ...
species, ''Thalassoma bifasciatum'' is a
protogynous Sequential hermaphroditism (called dichogamy in botany) is a type of hermaphroditism that occurs in many fish, gastropods, and plants. Sequential hermaphroditism occurs when the individual changes its sex at some point in its life. In particular, ...
sequential hermaphrodite; individuals may begin life either as males or females, but females can change sex later in life and become males.


Social system

A specific social system exists within the males - terminal phase males (which are the most aggressive and have the "highest" ranking among the males) and initial phase males (which mate when they can get a chance in a larger group). Color change of the ''T. bifasciatum'' indicates their motive or role. When terminal phase males chase initial phase males, their color changes to a metallic green, whereas when they are courting a female, they become pink/grey and form black circles on their fins.


Sex change

Initial phase females and initial phase males both can change into terminal phase males. This change can be relative quick, taking around 8 days. However, this change in sex is permanent: once an initial phase female or male changes into a terminal phase male, it cannot change back. An experiment removing terminal phase males from a population showed that more females changed into terminal phase males, and even aggression levels increased and coloration changes occurred to signify the female's change to terminal phase male. The initial phase males have comparatively larger
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoste ...
than the larger, terminal phase males. This enables the initial phase males to produce more sperm for the snatched opportunities they must take when trying to fertilize the eggs of the females in the guarded harem. Initial phase (IP) males also achieve fertilizations through participating in group spawns. These groups consist of 20–50 or more IP males that congregate at specific sites during the daily spawning period on medium and larger sized reefs. Females visit these groups to spawn and release eggs in a 'spawning rush'. IP males attempt to position themselves next to a female when she releases her eggs, as this maximizes their probability of fertilizing these eggs. Releasing large numbers of sperm also increases this probability and this is thought to also help explain the large testis size observed in IP males. This type of mating competition is referred to as '
sperm competition Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and ...
' and is seen in many species.


Biological research on sex change

''Thalassoma bifasciatum'' and its congener, the saddle wrasse (''T. duperrey'') have become important models for understanding the physiological and neurobiological bases of sex change. Sex change can be induced socially in both species by making large females the largest members of social groups. Sex change in experimental pens by saddle wrasses involves complete gonadal transformation with associated decreases in a key steroid hormones (
estradiol Estradiol (E2), also spelled oestradiol, is an estrogen steroid hormone and the major female sex hormone. It is involved in the regulation of the estrous and menstrual female reproductive cycles. Estradiol is responsible for the development o ...
and 11-ketotestosterone) and steroid hormone synthesizing enzymes in the gonads. Sex changing saddle wrasses also show substantial changes in brain levels of the monoamine neurotransmitters serotonin, dopamine, and
norepinephrine Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad' ...
. Initial phase males, terminal phase males, and females have the capability of reproducing. This is due to the protogyny, or the female's ability to become a male. The density of the type of male depends on the size of the reef. There are more terminal phase males than initial phase males on smaller reefs, on which they guard a small number of females. However, on larger reefs, there are equal proportions of initial phase and terminal phase males. This increases the chances of initial phase males to mate because they are less aggressive compared to terminal phase males. Sex change has been studied in bluehead wrasses primarily using field manipulations, where it can be induced in large females by removing dominant terminal phase males from small reefs.
Gonadotropin-releasing hormone Gonadotropin-releasing hormone (GnRH) is a releasing hormone responsible for the release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary. GnRH is a tropic peptide hormone synthesized and release ...
(GnRH) neurons differ across sexual phenotypes in the
hypothalamus The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamu ...
of bluehead wrasses and also with
androgen An androgen (from Greek ''andr-'', the stem of the word meaning "man") is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This in ...
implants that induce sex change. Behavioral sex change is very rapid in ''T. bifasciatum'' under field conditions, with male-typical behaviors being observed within minutes to hours after dominant terminal phase males are removed. Behavioral sex change occurs even in females whose gonads (ovaries) have been surgically removed prior to becoming socially dominant. Behavioral sex change is associated with increases in expression of a
neuropeptide Neuropeptides are chemical messengers made up of small chains of amino acids that are synthesized and released by neurons. Neuropeptides typically bind to G protein-coupled receptors (GPCRs) to modulate neural activity and other tissues like the ...
hormone termed arginine vasotocin or AVT and these increases occur regardless of whether sex changing females have gonads or not. Injections of AVT can induce sexual and aggressive behaviors in terminal phase male bluehead wrasses while injections of fluoxetine (tradename: Prozac) can reduce aggressive behaviors by terminal phase males.


Mating systems

Large terminal phase males will defend breeding sites to which females migrate on a daily basis. A study was done to estimate the relative roles of each sex in choosing the location of such sites. All terminal phase males or all females were replaced in local isolated populations, and the resulting site use was monitored. After males were replaced, the mating system was not affected. On the other hand, after the females were replaced, half of the old sites were lost and the same number of new sites came into use, and continued to be occupied for over a year after these manipulations. This occurred although large males originally continued to defend and display at the original sites. Therefore, this shows the importance of female choice in the feeding system of the blue-headed wrasse, and that males will respond to the females' site preferences. ''T. bifasciatum'' do not have distinct territories and their populations roam freely. Also, the females usually do not leave their original spawning spots. The males are known be taken away or leave. A plausible reason for why the females stay could be that they are most accustomed to those areas, and also because the predator threat is constant. Most of the literature on
mating system A mating system is a way in which a group is structured in relation to sexual behaviour. The precise meaning depends upon the context. With respect to animals, the term describes which males and females mate under which circumstances. Recognised ...
s of the blue-headed wrasse was described in small patches of concentrated reef habitats. In a large, linear
barrier reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups. Co ...
in
St. Croix Saint Croix; nl, Sint-Kruis; french: link=no, Sainte-Croix; Danish and no, Sankt Croix, Taino: ''Ay Ay'' ( ) is an island in the Caribbean Sea, and a county and constituent district of the United States Virgin Islands (USVI), an unincor ...
, U.S. Virgin Islands, very large aggregations of group-matings form daily in a single area near the foreside of the reef. Tagging studies have shown that fish are generally faithful to particular feeding schools that are assorted throughout the forereef, and that they tend to migrate to spawning grounds over 1.5 kilometers away. There is no mating that appears to happen in other upcurrent areas of the forereef. Despite large differences in the times that are spent on the migration, there are no significant differences in the
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
or frequency of spawning among females that live at different distances from the mating aggregation. The higher growth rate corresponded to a higher general feeding rate in the location, suggesting that food intake may outweigh the costs of the long migration.


References


Further reading

* * * * * * * * * * * * * * * * *


External links

* {{Taxonbar, from=Q1971551 bifasciatum Fish of the Caribbean Fish of the Western Atlantic bluehead wrasse