Seahorse

Seahorses Fossil range: 23–0 Ma PreЄ Є O S D C P T J K Pg N Lower Miocene to Present
Hippocampus sp.
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Syngnathiformes
Family:	Syngnathidae
Subfamily:	Hippocampinae
Genus:	Hippocampus Rafinesque, 1810[1]
Species
See text for species.

Seahorses compose the fish genus Hippocampus within the family Syngnathidae, in order Syngnathiformes. Syngnathidae also includes the pipefishes. "Hippocampus" comes from the Ancient Greek hippos meaning “horse” and kampos meaning “sea monster”.^[2].

There are about 40 species of seahorse are mainly found in shallow tropical and temperate waters throughout the world. They prefer to live in sheltered areas such as seagrass beds, coral reefs, or mangroves. Colonies have been found in European waters such as the Thames Estuary.^[3] From North America down to South America there are approximately four species, ranging from the very small (dwarf seahorses are only about 2.5 centimeters (1 in) to much larger specimens off the Pacific Coast of Central America (the foot-long H. ingens). H. erectus are larger seahorses that range from Nova Scotia to around Uruguay. Three species live in the Mediterranean Sea: H. hippocampus (long snout), H. brevirostris (short snout) and H. fuscus (immigrated from the Red Sea). These fish form territories, with males staying in about 1 square metre (11 sq ft) of their habitat while females range about one hundred times that area. They bob around in sea grass meadows, mangrove stands, and coral reefs where they adopt murky brown and gray patterns to camouflage themselves among the sea grass. During social moments or in unusual surroundings, seahorses turn bright colors.

Description

Spiny seahorse Hippocampus hystrix from East Timor holding on to soft coral with its prehensile tail

Seahorses are named for their equine profile. Although they are bony fish, they do not have scales, but rather a thin skin stretched over a series of bony plates arranged in rings throughout their body. Each species has a distinct number of rings. Seahorses swim upright, another characteristic that is not shared by their close pipefish relatives, which swim horizontally. Seahorses have a coronet on their head, which is distinct to each individual, much like a human fingerprint. They swim very poorly by using a dorsal fin, which they rapidly flutter and pectoral fins, located behind their eyes, which they use to steer. Seahorses have no caudal fin. Since they are poor swimmers, they are most likely to be found resting, with their prehensile tails wound around a stationary object. They have long snouts, which they use to suck up food, and eyes that can move independently of each other, much like a chameleon. Seahorses eat small shrimp, tiny fish and plankton.

Evolution and fossil record

Anatomical evidence, supported by molecular, physical, and genetic evidence, demonstrates that seahorses are highly modified pipefish. The fossil record of seahorses, however, is very sparse. The best known and best studied fossils are specimens of H. guttulatus (though literature more commonly refers to them under the synonym of H. ramulosus), from the Marecchia River Formation of Rimini Province, Italy, dating back to the Lower Pliocene, about 3 million years ago. The earliest known seahorse fossils are of two pipefish-like species, H. sarmaticus and H. slovenicus from the coprolitic horizon of Tunjice Hills, a middle Miocene lagerstätte in Slovenia dating back about 13 million years.^[4] Molecular dating finds that pipefish and seahorses separated during the Late Oligocene. This has led to speculation that seahorses evolved in response to large areas of shallow-water, newly created as the result of tectonic events. The shallow water allowed the expansion of seagrass habitats that selected for the camouflage offered by the seahorses’ upright posture.^[5] These tectonic changes occurred in the Western Pacific Ocean suggesting an origin there with molecular data suggesting two later and separate invasions of the Atlantic Ocean.^[6]

Reproduction

Pregnant seahorse, New York Aquarium

Seahorses are often thought of as being monogamous, though recent research shows this may not be true.^[7] The male seahorse is equipped with a brood pouch on the ventral, or front-facing, side. When mating, the female seahorse deposits up to 1,000 eggs in the male's pouch,^[8] which the male then internally fertilizes. The male carries the eggs for about two months until they emerge,^[8] expelling fully-developed, miniature seahorses in the water. The father continues to protect the young until they are able to live on their own, however they have been known to eat a few offspring while at it.^[8]

Courtship

Before breeding, pairs court for several days, even while others try to interfere. Scientists believe the courtship behavior synchronizes the animals' movements so that the male can receive the eggs when the female is ready to deposit them. During this time they may change color, swim side by side holding tails or grip the same strand of sea grass with their tails and wheel around in unison in what is known as a “pre-dawn dance". They eventually engage in a “true courtship dance" lasting about 8 hours, during which the male pumps water through the egg pouch on his trunk which expands and open to display its emptiness. When the female’s eggs reach maturity, she and her mate let go of any anchors and snout-to-snout, drift upward out of the seagrass, often spiraling as they rise. The female inserts her ovipositor into the male’s brood pouch and deposits dozens to thousands of eggs. As the female releases her eggs, her body slims while his swells. Both animals then sink back into the seagrass and she swims away.

Gestation

West Australian seahorse, H. subelongatus

The male fertilizes the eggs, which embed in the pouch wall and become enveloped by tissue.^[9] The male supplies the eggs with prolactin, the same hormone responsible for milk production in pregnant mammals. The pouch provides oxygen as well as a controlled environment incubator. The eggs then hatch in the pouch where the salinity of the water is regulated; this prepares the newborns for life in the sea.^[10][11] Throughout gestation, which in most species requires two to four weeks, his mate visits him daily for “morning greetings”. They interact for about 6 minutes, reminiscent of courtship. The female then swims away until the next morning, and the male returns to vacuuming up food through his snout.^[10]

Research published in 2007 indicates the male releases sperm into the surrounding sea water during fertilization, and not directly into the pouch as previously thought.^[12]

Birth

White's seahorse, H. whitei

The number of young released by the male seahorse averages 100-200 for most species, but may be as low as 5 for the smaller species, or as high as 1,500. When the fry are ready to be born, the male expels them with muscular contractions. He typically gives birth at night and is ready for the next batch of eggs by morning when his mate returns. Like almost all other fish species, seahorses do not nurture their young after birth. Infants are susceptible to predators or ocean currents which wash them away from feeding grounds or into temperatures too extreme for their delicate bodies. Fewer than .5% of infants survive to adulthood, explaining why litters are so large. These survival rates are actually fairly high compared to other fish, because of their protected gestation, making the process worth the great cost to the father. The eggs of most other fish are abandoned immediately after fertilization.^[11]

Questions surrounding reproductive roles

Reproduction is energetically costly to the male. This brings into question why the sexual role reversal even takes place. In an environment where one partner incurs more energy costs than the other, Bateman's principle suggests that the lesser contributor takes the role of the aggressor. Male seahorses are more aggressive and sometimes “fight” for female attention. According to Amanda Vincent of Project Seahorse, only males tail-wrestle and snap their heads at each other. This discovery prompted further study of energy costs. To estimate the female’s direct contribution, researcher Heather D. Masonjones, associate professor of biology at the University of Tampa, chemically analyzed the energy stored in each egg. To measure the burden on the male, Masonjones measured its oxygen consumption. By the end of incubation, the male consumed almost 33% more oxygen than before mating. The study concluded that the female's energy expenditure while generating eggs is twice that of males during incubation^[10] confirming the standard hypothesis.

Why the male seahorse (and other members of Syngnathidae) carries the offspring through gestation is unknown, though some researchers believe it allows for shorter birthing intervals, in turn resulting in more offspring. Given an unlimited number of ready and willing partners, males have the potential to produce 17 percent more offspring than females in a breeding season. Also, females have “time-outs” from the reproductive cycle that are 1.2 times longer than those of males. This seems to be based on mate choice, rather than physiology. When the female’s eggs are ready, she must lay them in a few hours or eject them into the water column. Making eggs is a huge cost to her physically, since they amount to about a third of her body weight. To protect against losing a clutch, the female demands a long courtship. The daily greetings help to cement the bond between the pair.

Monogamy

A study conducted by Amanda Vincent of Project Seahorse demonstrates the importance of the daily greetings ritual in establishing seahorses' monogamous relationships. Vincent kept a female seahorse in a tank with two males. After the female filled one male’s pouch with eggs Vincent removed him and she was left with the unimpregnated male. During the weeks of her mate’s pregnancy, the female and her tankmate greeted each other daily, clinging to the same bit of grass and changing color, but did not display signs of courtship. After the original mate gave birth he was returned to the tank. Both males competed for her attention, but in six of six tests the female presented the next clutch of eggs to the other male.^[10]

Although monogamy within species is not common, it does appear to exist for some. In this case, the mate guarding hypothesis may be an explanation. This hypothesis states that “males remain with a single female because of ecological factors that make male parental care and protection of offspring especially advantageous.”^[13] Because the rates of survival for newborn seahorses are so low, incubation is essential. Though not proven, males could have taken on this role because of the lengthy period the females require to produce their eggs. If males incubate while females prepare the next clutch (amounting to 1/3 of body weight), they can reduce the interval between clutches.

Feeding habits

Seahorses feed on small crustaceans floating in the water or crawling on the bottom. With excellent camouflage and a lot of patience, seahorses ambush prey that float within striking range. Mysid shrimp and other small crustaceans are favorites, but some seahorses have been observed eating other kinds of invertebrates and even larval fish.

In aquaria

Seahorses (Hippocampus erectus) at the New England Aquarium, USA.

While many aquarium hobbyists keep seahorses as pets, seahorses collected from the wild tend to fare poorly in home aquaria. Many eat only live foods such as brine shrimp and are prone to stress, which damages their immune systems and makes them susceptible to disease.

In recent years, however, captive breeding has become more popular. Such seahorses survive better in captivity, and are less likely to carry diseases. They eat frozen mysidacea (crustaceans) that are readily available from aquarium stores,^[14] and do not experience the stress of moving out of the wild. Although captive-bred seahorses are more expensive, they take no toll on wild populations.

Seahorses should be kept in an aquarium to themselves, or with compatible tank-mates. Seahorses are slow feeders, and fast, aggressive feeders will leave them without food.^[14]

Seahorses can co-exist with many species of shrimp and other bottom-feeding creatures. Gobies also make good tank-mates. Avoid eels, tangs, triggerfish, squid, octopus, and sea anemones.^[15]

Animals sold as "freshwater seahorses" are usually the closely related pipefish, of which a few species live in the lower reaches of rivers. The supposed true "freshwater seahorse" called H. aimei was not a real species, but a name sometimes used for Barbour's and Hedgehog seahorses. The latter is a species that can be found in brackish waters, but not actually a freshwater fish.^[16]

Use in Chinese medicine

Medicinal seahorse.

Seahorse populations are thought to have been endangered in recent years by overfishing and habitat destruction. The seahorse is used in traditional Chinese herbology, and as many as 20 million seahorses may be caught each year and sold for this purpose.^[17] Medicinal seahorses are not readily bred in captivity as they are susceptible to disease and have somewhat different energetics from aquarium seahorses. Seahorses are also used as medicines by the Indonesians, the Central Filipinos, and many other ethnic groups around the world.

Import and export of seahorses has been controlled under CITES since May 15, 2004. However, Indonesia, Japan, Norway, and South Korea have chosen to opt out of the trade rules set by CITES.

The problem may be exacerbated by the growth of pills and capsules as the preferred method of ingesting medication as they are cheaper and more available than traditional, individually tailored prescriptions of raw medicinals but the contents are harder to track. Seahorses once had to be of a certain size and quality before they were accepted by TCM practitioners and consumers. But declining availability of the preferred large, pale and smooth seahorses has been offset by the shift towards prepackaged medicines, which make it possible for TCM merchants to sell previously unused juvenile, spiny and dark-coloured animals. Today almost a third of the seahorses sold in China are prepackaged. This adds to the pressure on the species.^[18]

Taxonomy

Common seahorse, Hippocampus kuda

• Genus Hippocampus
  • Big-belly seahorse, H. abdominalis Lesson, 1827 (New Zealand and south and east Australia)
  • Winged seahorse, H. alatus Kuiter, 2001
  • West African seahorse, H. algiricus Kaup, 1856
  • Narrow-bellied seahorse, H. angustus Günther, 1870
  • Barbour's seahorse, H. barbouri Jordan & Richardson, 1908
  • Pygmy seahorse, H. bargibanti Whitley, 1970 West Pacific area (Indonesia, Philippines, Papua New Guinea, Solomon Islands, etc)
  • False-eyed seahorse, H. biocellatus Kuiter, 2001
  • Réunion seahorse, H. borboniensis Duméril, 1870
  • Short-head seahorse or knobby seahorse, H. breviceps Peters, 1869 (south and east Australia)
  • Giraffe seahorse, H. camelopardalis Bianconi, 1854
  • Knysna seahorse, H. capensis Boulenger, 1900
  • H. colemani Kuiter, 2003
  • Tiger tail seahorse, H. comes Cantor, 1850
  • Crowned seahorse, H. coronatus Temminck & Schlegel, 1850
  • Denise's pygmy seahorse, H. denise Lourie & Randall, 2003
  • Lined seahorse, H. erectus Perry, 1810 (east coast of the Americas, between Nova Scotia and Uruguay)
  • Fisher's seahorse, H. fisheri Jordan & Evermann, 1903
  • Sea pony, H. fuscus Rüppell, 1838 (Indian Ocean)
  • Big-head seahorse, H. grandiceps Kuiter, 2001
  • Long-snouted seahorse, H. guttulatus Cuvier, 1829
  • Eastern spiny seahorse, H. hendriki Kuiter, 2001
  • Short-snouted seahorse, H. hippocampus (Linnaeus, 1758) (Mediterranean Sea and Atlantic Ocean)
  • Thorny seahorse, H. histrix Kaup, 1856 (Indian Ocean, Persian Gulf, Red Sea, and the Far East)
  • Pacific seahorse, H. ingens Girard, 1858 (Pacific coast of North, Central and South America)
  • Jayakar's seahorse, H. jayakari Boulenger, 1900
  • Collared seahorse, H. jugumus Kuiter, 2001
  • Great seahorse, H. kelloggi Jordan & Snyder, 1901
  • Common seahorse, H. kuda Bleeker, 1852
  • Lichtenstein's seahorse, H. lichtensteinii Kaup, 1856
  • Bullneck seahorse, H. minotaur Gomon, 1997
  • Japanese seahorse, H. mohnikei Bleeker, 1854
  • Monte Bello seahorse, H. montebelloensis Kuiter, 2001
  • Northern spiny seahorse, H. multispinus Kuiter, 2001
  • H. pontohi Lourie and Kuiter, 2008
  • High-crown seahorse, H. procerus Kuiter, 2001
  • Queensland seahorse, H. queenslandicus Horne, 2001
  • Longsnout seahorse, H. reidi Ginsburg, 1933 (Caribbean coral reefs)
  • Satomi's pygmy seahorse, H. satomiae Lourie and Kuiter, 2008
  • Half-spined seahorse, H. semispinosus Kuiter, 2001
  • H. severnsi Lourie and Kuiter, 2008
  • Shiho's seahorse, H. sindonis Jordan & Snyder, 1901
  • Hedgehog seahorse, H. spinosissimus Weber, 1913
  • West Australian seahorse, H. subelongatus Castelnau, 1873
  • Longnose seahorse, H. trimaculatus Leach, 1814
  • White's seahorse, H. whitei Bleeker, 1855 (east Australia)
  • Zebra seahorse, H. zebra Whitley, 1964
  • Dwarf seahorse, H. zosterae Jordan & Gilbert, 1882 (Gulf of Mexico and the Caribbean)

Pygmy seahorses

Hippocampus satomiae attached to a coral

Pygmy Seahorses are less than 15 millimeters (0.6 in) tall and 17 millimeters (0.7 in) wide members of the genus. Previously the term was applied exclusively to the species H. bargibanti but since 1997, discoveries have made this term obsolete. The species H. minotaur, H. denise, H. colemani, H. pontohi, H. severnsi and H. satomiae have been described. Other species that are believed to be unclassified have also been reported in books, dive magazines and on the Internet. They can be distinguished from other species of seahorse by their 12 trunk rings, low number of tail rings (26–29), the location in which young are brooded in the trunk region of males and their extremely small size.^[19] Molecular analysis (of ribosomal RNA) of 32 Hippocampus species found that H. bargibanti belongs in a separate clade from other members of the genus and therefore that the species diverged from the other species in the "ancient" past.^[20]

Most pygmy seahorses are well camouflaged and live in close association with other organisms including colonial hydrozoans (Lytocarpus and Antennellopsis), coralline algae (Halimeda) sea fans (Muricella, Annella, Acanthogorgia). This combined with their small size accounts for why most species have only been noticed in recent years.^[19]

Cultural references

'Fucus like seahorse' from Lydekker's The Royal Natural History

A sculpture of a heraldic seahorse that adorned an 18th or 19th century French naval vessel

• In heraldry, a seahorse is depicted as a creature with the foreparts of a horse and the hindparts of a fish. See, for example, the right supporter of the Isle of Wight Arms, the supporters on either side of the crest of the city of Newcastle upon Tyne, or the coincidental arms of the University of Newcastle, Australia.

References

Further reading

• Amanda C.J. Vincent and Laila M. Sadler. "Faithful pair bonds in wild seahorse, Hippocampus whitei." Animal Behaviour 50 (1995): 1557-1569.
• Amanda C.J. Vincent. "A role for daily greetings in maintaining seahorse pair bonds." Animal Behaviour 49 (1995): 258-260.
• Amanda C.J. Vincent. "A seahorse father makes a good mother." Natural History, 12 (1990): 34-43.
• Amanda C.J. Vincent and Rosie Woodroffe. "Mothers little helpers: patterns of male care in mammals." Trends in Ecology and Evolution, 9 (1994): 294-297.
• John Sparks: Battle of the Sexes: The Natural History of Sex. London: BBC Books, 1999
• Sara A. Lourie, Amanda C.J. Vincent and Heather J. Hall: Seahorses: An Identification Guide to the World's Species and their Conversation. London: Project Seahorse, 1999
• Teske P. R., Hamilton H., Matthee C. A. & Barker N. P.: Signatures of seaway closures and founder dispersal in the phylogeny of a circumglobally distributed seahorse lineage. BMC Evolutionary Biology 2007, 7:138. BiomedCentral.com

External links

• Vimeo.com, Video of various seahorse species at the Monterey Bay Aquarium
• ARKive—images and movies of the pygmy seahorse (Hippocampus bargibanti)
• FusedJaw.com, Information about seahorses
• ProjectSeahorse.org, Research and Conservation Group
• Seahorse.org, Aquarium Hobby
• DiveGallery.com, Seahorse Photos
• RobertoSozzani.it, Hippocampus denise Pygmy seahorse photo galleries
• RobertoSozzani.it, Pontohi Pygmy Seahorse
• RobertoSozzani.it, Walea Pygmy Seahorse
• Arnes.si, Fossil Seahorses from Tunjice hills (Slovenia)
• QataVisitor.com n Sea horses in Qatar