Thalassoma bifasciatum
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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 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 Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
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 over the reef and are important cleaner fish 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 Mexic ...
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 Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
, small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s (such as
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are ref ...
and krill),
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 wor ...
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 chorda ...
s, and the eggs 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 in the coral reefs they inhabit, they avoid cleaning
piscivores A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evolut ...
such as the
spotted moray The spotted moray (''Gymnothorax moringa'') is a medium to large moray eel. Other common names include conger, spotted eel, red moray, speckled moray, white cong, white jawed moray, white-chinned moray and white-jawed moray eel. Spotted eels h ...
, 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. It ...
, 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 includ ...
. Such species will view them as prey, but will not view gobies, 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, 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 seah ...
. 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 h ...
of ''T. bifasciatum'' is the intracellular
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. ...
'' Kudoa ovivora'', which can be found in the ovaries of females. Infected eggs are
sterile Sterile or sterility may refer to: *Asepsis Asepsis is the state of being free from disease-causing micro-organisms (such as pathogenic bacteria, viruses, pathogenic fungi, and parasites). There are two categories of asepsis: medical and surgi ...
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 nl, In vrijheid verenigd"Unified by freedom" , national_anthem = , common_languages = Dutch English Papiamento , demonym = Netherlands Antillean , capital = Willemstad , year_start = 1954 , year_end = 2010 , date_start = 15 December , ...
. 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. ...
. However, this species shows high
site fidelity Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derives ...
, so coral reef destruction could cause local endangerment. Due to its bright coloration, it is sometimes collected for the aquarium trade, but this is not a threat to the species.


Behavior

Like many other wrasse species, ''Thalassoma bifasciatum'' is a protogynous 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' 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
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sp ...
al transformation with associated decreases in a key steroid hormones ( estradiol 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 Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and va ...
,
dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. Dopamine constitutes about 80% o ...
, 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 released ...
(GnRH)
neurons A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
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 hypothalamus ...
of bluehead wrasses and also with androgen 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 t ...
hormone termed arginine
vasotocin Vasotocin is an oligopeptide homologous to oxytocin and vasopressin found in all non-mammalian vertebrates (including birds, fishes, and amphibians) and possibly in mammals during the fetal stage of development. Arginine vasotocin (AVT), a hormone ...
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 systems 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. ...
in St. Croix, 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 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

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External links

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