Barnacle

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Barnacle
Temporal range: Carboniferous–Recent
Chthamalus stellatus.jpg
Chthamalus stellatus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Thecostraca
Subclass: Cirripedia
Burmeister, 1834
Infraclasses
Synonyms
  • Thyrostraca
  • Cirrhopoda
  • Cirrhipoda
  • Cirrhipedia

Barnacles are arthropods of the subclass Cirripedia in the subphylum Crustacea. They are related to crabs and lobsters, with similar nauplius larvae. Barnacles are exclusively marine invertebrates; many species live in shallow and tidal waters. Some 2,100 species have been described.

Contents

Barnacle adults are sessile; most are suspension feeders with hard calcareous shells, but the Rhizocephala are specialized parasites of other crustaceans, with reduced bodies. Barnacles have existed since at least the mid-Carboniferous, some 325 million years ago.

In folklore, barnacle geese were once held to emerge fully formed from goose barnacles. Both goose barnacles and the Chilean giant barnacle are fished and eaten. Barnacles are economically significant as biofouling on ships, where they cause hydrodynamic drag, reducing efficiency.

Etymology

The word "barnacle" is attested in the early 13th century as Middle English "bernekke" or "bernake", close to Old French "bernaque" and medieval Latin bernacae or berneka, denoting the barnacle goose. [1] [2] Because the full life cycles of both barnacles and geese were unknown at the time, (geese spend their breeding seasons in the Arctic) a folktale emerged that geese hatched from barnacles. It was not applied strictly to the arthropod until the 1580s. The ultimate meaning of the word is unknown. [2] [3]

The name Cirripedia comes from the Latin words cirritus "curly" from cirrus "curl" [4] and pedis from pes "foot". [5] The two words together mean "curly-footed", alluding to the curved legs used in filter-feeding. [6]

Description

Whale barnacles on a humpback whale Anim1032 - Flickr - NOAA Photo Library.jpg
Whale barnacles on a humpback whale

Most barnacles are encrusters, attaching themselves to a hard substrate such as a rock, the shell of a mollusc, or a ship; or to an animal such as a whale (whale barnacles). The most common form, acorn barnacles, are sessile, growing their shells directly onto the substrate, whereas goose barnacles attach themselves by means of a stalk. [7]

Anatomy and physiology

Anatomy of a sessile barnacle Barnacles Sessile anatomy.svg
Anatomy of a sessile barnacle

Barnacles have a carapace made of six hard calcareous plates, with a lid or operculum made of four more plates. Inside the carapace, the animal lies on its stomach, projecting its limbs downwards. Segmentation is usually indistinct; the body is more or less evenly divided between the head and thorax, with little or no abdomen. Adult barnacles have few appendages on their heads, with only a single, vestigial pair of antennae attached to the cement gland. The six pairs of thoracic limbs are called cirri; these are feathery and very long. The cirri extend to filter food, such as plankton, from the water and move it towards the mouth. [8]

Acorn barnacles are attached to the substratum by cement glands that form the base of the first pair of antennae; in effect, the animal is fixed upside down by means of its forehead. In some barnacles, the cement glands are fixed to a long, muscular stalk, but in most they are part of a flat membrane or calcified plate. These glands secrete a type of natural quick cement made of complex protein bonds (polyproteins) and other trace components like calcium. [9] : 2–3  This natural cement can withstand a pulling strength of 5,000 lbf/in2 (30,000 kPa) and a sticking strength of 22–60 lbf/in2 (200–400 kPa). [8]

Barnacles have no true heart, although a sinus close to the esophagus performs a similar function, with blood being pumped through it by a series of muscles. [10] The blood vascular system is minimal. [11] Similarly, they have no gills, absorbing oxygen from the water through the cirri and the surface of the body. [12] The excretory organs of barnacles are maxillary glands. [13]

The main sense of barnacles appears to be touch, with the hairs on the limbs being especially sensitive. The adult has three photoreceptors (ocelli), one median and two lateral. These record the stimulus for the barnacle shadow reflex, where a sudden decrease in light causes cessation of the fishing rhythm and closing of the opercular plates. [14] The photoreceptors are likely only capable of sensing the difference between light and dark. [15] This eye is derived from the primary naupliar eye. [16]

Life cycle

Barnacles pass through two distinct larval stages, the nauplius and the cyprid, before developing into a mature adult.

Nauplius larva

A fertilised egg hatches into a nauplius: a one-eyed larva comprising a head and a telson with three pairs of limbs, lacking a thorax or abdomen. This undergoes six moults, passing through five instars, before transforming into the cyprid stage. Nauplii are typically initially brooded by the parent, and released after the first moult as larvae that swim freely using setae. [17] [18] All but the first instars are filter feeders. [19]

Cypris larva

The cypris larva is the second and final larval stage before adulthood. In Rhizocephala and Thoracica an abdomen is absent in this stage, but the y-cyprids (post-naupliar instar) has three distinct abdominal segments. [21] It is not a feeding stage; its role is to find a suitable place to settle, since the adults are sessile. [17] The cyprid stage lasts from days to weeks. It explores potential surfaces with modified antennules; once it has found a suitable spot, it attaches head-first using its antennules and a secreted glycoproteinous cement. Larvae assess surfaces based upon their surface texture, chemistry, relative wettability, color, and the presence or absence and composition of a surface biofilm; swarming species are more likely to attach near other barnacles. [22] As the larva exhausts its energy reserves, it becomes less selective in the sites it selects. It cements itself permanently to the substrate with another proteinaceous compound, and then undergoes metamorphosis into a juvenile barnacle. [22]

Adult

Typical acorn barnacles develop six hard calcareous plates to surround and protect their bodies. For the rest of their lives, they are cemented to the substrate, using their feathery legs (cirri) to capture plankton. Once metamorphosis is over and they have reached their adult form, barnacles continue to grow by adding new material to their heavily calcified plates. These plates are not moulted; however, like all ecdysozoans, the barnacle moults its cuticle. [23]

Sexual reproduction

Pseudocopulation: the acorn barnacle uses its long penis to reach across to transfer sperm to another individual nearby. Balanus reproduction.svg
Pseudocopulation: the acorn barnacle uses its long penis to reach across to transfer sperm to another individual nearby.

Most barnacles are hermaphroditic, producing both eggs and sperms. A few species have separate sexes, or have both males and hermaphrodites. The ovaries are located in the base or stalk, and may extend into the mantle, while the testes are towards the back of the head, often extending into the thorax. Typically, recently moulted hermaphroditic individuals are receptive as females. Self-fertilization, although theoretically possible, has been experimentally shown to be rare in barnacles. [26] [27]

The sessile lifestyle of acorn barnacles makes sexual reproduction difficult, as they cannot leave their shells to mate. To facilitate genetic transfer between isolated individuals, barnacles have extraordinarily long penises⁠. Barnacles probably have the largest penis-to-body size ratio of the animal kingdom, [26] up to eight times their body length, though on exposed coasts the penis is shorter and thicker. [25] The mating of acorn barnacles is described as pseudocopulation. [24] [28]

The goose barnacle Pollicipes polymerus can alternatively reproduce by spermcasting, in which the male barnacle releases his sperm into the water, to be taken up by females. Isolated individuals always made use of spermcasting and sperm capture, as did a quarter of individuals with a close neighbour. This 2013 discovery overturned the long-held belief that barnacles were limited to pseucocopulation or hermaphroditism. [24]

Rhizocephalan barnacles had been considered hermaphroditic, but their males inject themselves into females' bodies, degrading to little more than sperm-producing cells. [29]

Ecology

Filter feeding

Most barnacles are filter feeders. From within their shell, they repeatedly reach into the water column with their cirri. These feathery appendages beat rhythmically to draw plankton and detritus into the shell for consumption. [7] [30]

Species-specific zones

Although they have been found at water depths to 600 m (2,000 ft), [7] most barnacles inhabit shallow waters, with 75% of species living in water depths less than 100 m (300 ft), [7] and 25% inhabiting the intertidal zone. [7] Within the intertidal zone, different species of barnacles live in very tightly constrained locations, allowing the exact height of an assemblage above or below sea level to be precisely determined. [7]

Since the intertidal zone periodically desiccates, barnacles are well adapted against water loss. Their calcite shells are impermeable, and they can close their apertures with movable plates when not feeding. [31] Their hard shells are assumed by zoologists to have evolved as an anti-predator adaptation. [32]

One group of stalked barnacles has adapted to a rafting lifestyle, drifting around close to the water's surface. They colonize every floating object, such as driftwood, and like some non-stalked barnacles attach themselves to marine animals. The species most specialized for this lifestyle is Dosima fascicularis , which secretes a gas-filled cement that makes it float at the surface. [33]

Parasitism

Sacculina carcini (highlighted) parasitising the crab Liocarcinus holsatus. Sacculina carcini.jpg
Sacculina carcini (highlighted) parasitising the crab Liocarcinus holsatus .

Other members of the class have an entirely different mode of life. Barnacles of the superorder Rhizocephala, including the genus Sacculina , are parasitic castrators of other arthropods, including crabs. The anatomy of these parasitic barnacles is greatly reduced compared to their free-living relatives. They have no carapace or limbs, having only unsegmented sac-like bodies. They feed by extending thread-like rhizomes of living cells into their hosts' bodies from their points of attachment. [34] [15]

Competitors

Barnacles and limpets compete for space in the intertidal zone CornishBarnacles.JPG
Barnacles and limpets compete for space in the intertidal zone

Barnacles are displaced by limpets and mussels, which compete for space. [7] They employ two strategies to overwhelm their competitors: "swamping", and fast growth. In the swamping strategy, vast numbers of barnacles settle in the same place at once, covering a large patch of substrate, allowing at least some to survive in the balance of probabilities. [7] Fast growth allows the suspension feeders to access higher levels of the water column than their competitors, and to be large enough to resist displacement; species employing this response, such as the aptly named Megabalanus , can reach 7 cm (3 in) in length. [7]

Competitors may include other barnacles. Balanoids gained their advantage over the chthalamoids in the Oligocene, when they evolved tubular skeletons, which provide better anchorage to the substrate, and allow them to grow faster, undercutting, crushing, and smothering chthalamoids. [35]

Predators and parasites

Among the most common predators of barnacles are whelks. They are able to grind through the calcareous exoskeleton and eat the animal inside. Barnacle larvae are consumed by filter-feeding benthic predators including the mussel Mytilus edulis and the ascidian Styela gibbsi. [36] Another predator is the starfish species Pisaster ochraceus . [37] [38] A stalked barnacle in the Iblomorpha, Chaetolepas calcitergum, lacks a heavily mineralised shell, but contains a high concentration of toxic bromine; this may serve to deter predators. [39] The turbellarian flatworm Stylochus , a serious predator of oyster spat, has been found in barnacles. [40] Parasites of barnacles include many species of Gregarinasina (alveolate protozoa), a few fungi, a few species of trematodes, and a parasitic castrator isopod, Hemioniscus balani . [40]

History of taxonomy

Balanus improvisus, one of the many barnacle taxa described by Charles Darwin, on a bivalve shell Balanus improvisus on Mya arenaria shell.jpg
Balanus improvisus , one of the many barnacle taxa described by Charles Darwin, on a bivalve shell

Barnacles were classified by Linnaeus and Cuvier as Mollusca, but in 1830 John Vaughan Thompson published observations showing the metamorphosis of the nauplius and cypris larvae into adult barnacles, and noted that these larvae were similar to those of crustaceans. In 1834, Hermann Burmeister reinterpreted these findings, moving barnacles from the Mollusca to Articulata (in modern terms, annelids + arthropods), showing naturalists that detailed study was needed to reevaluate their taxonomy. [41]

Charles Darwin took up this challenge in 1846, and developed his initial interest into a major study published as a series of monographs in 1851 and 1854. [41] He undertook this study at the suggestion of his friend the botanist Joseph Dalton Hooker, namely to thoroughly understand at least one species before making the generalisations needed for his theory of evolution by natural selection. [42] The Royal Society notes that barnacles occupied Darwin, who worked from home, so intensely "that his son assumed all fathers behaved the same way: when visiting a friend he asked, 'Where does your father do his barnacles?'" [43] Upon the conclusion of his research, Darwin declared "I hate a barnacle as no man ever did before." [42] [44]

Evolution

Fossil record

The oldest definitive fossil barnacle is Praelepas from the mid-Carboniferous, around 330-320 million years ago. [45] Older claimed barnacles such as Priscansermarinus from the Middle Cambrian, some 510 to 500 million years ago, [46] do not show clear barnacle morphological traits, though Rhamphoverritor from the Silurian Coalbrookdale Formation of England may represent a stem-group barnacle. Barnacles first radiated and became diverse during the Late Cretaceous. Barnacles underwent a second, much larger radiation beginning during the Neogene and still continuing. [45]

Phylogeny

The following cladogram, not fully resolved, shows the phylogenetic relationships of the Cirripedia within Thecostraca as of 2021. [45]

Thecostraca

Taxonomy

Over 2,100 species of Cirripedia have been described. [45] Some authorities regard the Cirripedia as a full class or subclass. In 2001, Martin and Davis placed Cirripedia as an infraclass of Thecostraca, and divided it into six orders: [47]

In 2021, Chan et al. elevated Cirripedia to a subclass of the Thecostraca, and the superorders Acrothoracica, Rhizocephala, and Thoracica to infraclass. The updated classification with 11 orders has been accepted in the World Register of Marine Species. [45] [48]

Relationship with humans

Biofouling

Barnacles are of economic consequence, as they often attach themselves to man-made structures. Particularly in the case of ships, they are classified as fouling organisms. The number and size of barnacles that cover ships can impair their efficiency by causing hydrodynamic drag. [49]

As food

The flesh of some barnacles is routinely consumed by humans, including Japanese goose barnacles (e.g. Capitulum mitella ), and goose barnacles (e.g. Pollicipes pollicipes ), a delicacy in Spain and Portugal. [50] The Chilean giant barnacle Austromegabalanus psittacus is fished, or overfished, in commercial quantities on the Chilean coast, where it is known as the picoroco. [51]

Technological applications

MIT researchers have developed an adhesive inspired by the protein-based bioglue produced by barnacles to firmly attach to rocks. The adhesive can form a tight seal to halt bleeding within about 15 seconds of application. [52]

The stable isotope signals in the layers of barnacle shells can potentially be used as a forensic tracking method [53] for whales, loggerhead turtles [54] and for marine debris, such as shipwrecks or aircraft wreckage. [55] [56] [57]

In culture

One version of the barnacle goose myth is that the birds emerge fully formed from goose barnacles. [58] [59] The myth, with variants such as that the goose barnacles grow on trees, owes its longstanding popularity to ignorance of bird migration. [60] [61] [62] The myth survived to modern times through bestiaries. [63]

More recently, Barnacle Bill became a "comic folktype" [64] of a seaman, with a drinking song [64] and several films (a 1930 animated short with Betty Boop, [65] a 1935 British drama, [66] a 1941 feature with Wallace Beery, [67] and a 1957 Ealing comedy [68] ) named after him.

The political reformer John W. Gardner likened middle managers who settle into a comfortable position and "have stopped learning or growing" to the barnacle, who "is confronted with an existential decision about where it's going to live. Once it decides... it spends the rest of its life with its head cemented to a rock". [69]

Related Research Articles

<span class="mw-page-title-main">Goose barnacle</span> Type of barnacle

Goose barnacles, also called percebes, turtle-claw barnacles, stalked barnacles, gooseneck barnacles, are filter-feeding crustaceans that live attached to hard surfaces of rocks and flotsam in the ocean intertidal zone. Goose barnacles formerly made up the taxonomic order Pedunculata, but the group has been found to be polyphyletic, with its members scattered across multiple orders of the infraclass Thoracica.

<span class="mw-page-title-main">Thoracica</span> Infraclass of barnacles

Thoracica is an infraclass of crustaceans which contains the most familiar species of barnacles found on rocky coasts, such as Semibalanus balanoides and Chthamalus stellatus. They have six well-developed limbs, and may be either stalked or sessile. The carapace is heavily calcified. The group includes free-living and commensal species.

<span class="mw-page-title-main">Rhizocephala</span> Superorder of barnacles

Rhizocephala are derived barnacles that are parasitic castrators. Their hosts are mostly decapod crustaceans, but include Peracarida, mantis shrimps and thoracican barnacles. Their habitats range from the deep ocean to freshwater. Together with their sister groups Thoracica and Acrothoracica, they make up the subclass Cirripedia. Their body plan is uniquely reduced in an extreme adaptation to their parasitic lifestyle, and makes their relationship to other barnacles unrecognisable in the adult form. The name Rhizocephala derives from the Ancient Greek roots ῥίζα and κεφαλή, describing the adult female, which mostly consists of a network of thread-like extensions penetrating the body of the host.

<span class="mw-page-title-main">Thecostraca</span> Class of crustaceans

Thecostraca is a class of marine invertebrates containing over 2,200 described species. Many species have planktonic larvae which become sessile or parasitic as adults.

<span class="mw-page-title-main">Chthamalidae</span> Family of crustaceans

The Chthamalidae are a family of chthamaloid barnacles, living entirely in intertidal/subtidal habitats, characterized by a primary shell wall of eight, six, or four plates, lacking imbricating plate whorls, and either membraneous or more rarely calcareous basis. They are not found below immediate subtidal habitats, and more likely are found in the highest tier of shallow-water barnacle fauna. They can be found in the most rigorous wave-washed locations, and some species are found in the surf zone above high tide mark, only receiving water from wave action at high tide. The family includes at least 56 recognized species.

<i>Anelasma</i> Species of parasitic barnacles that attack sharks

Anelasma is a monotypic genus of goose barnacles that live as parasites on various shark hosts.

<span class="mw-page-title-main">Acorn barnacle</span> Index of animals with the same common name

Acorn barnacle and acorn shell are vernacular names for certain types of stalkless barnacles, generally excluding stalked or gooseneck barnacles. As adults they are typically cone-shaped, symmetrical, and attached to rocks or other fixed objects in the ocean. Members of the barnacle order Balanomorpha are often called acorn barnacles.

<span class="mw-page-title-main">Whale barnacle</span> Barnacles that attach to whales

Whale barnacles are species of acorn barnacle that belong to the family Coronulidae. They typically attach to baleen whales, and sometimes settle on toothed whales. The whale barnacles diverged from the turtle barnacles about three million years ago.

<span class="mw-page-title-main">Ascothoracida</span> Group of crustaceans

Ascothoracida is a small group of parasitic marine crustaceans, comprising around 100 species and divided into Dendrogastrida and Laurida. They are found throughout the world on cnidarians and echinoderms. Dendrogastrida are parasites on echinoderms, and Laurida are parasites on cnidarians, except from the species Waginella Grygier, which is also a parasite on echinoderms (crinoids). Piercing and sucking mouthparts are used for feeding, and more advanced forms also absorb nutrients through a modified integument of the carapace. More basal forms are ectoparasitic, but most genera are meso- and endoparasitic. The sexes are separate, except from secondary hermaphroditic species of the Petrarcidae. In many species the larger female often have smaller males living inside her mantle cavity.

<i>Pollicipes pollicipes</i> Species of barnacle

Pollicipes pollicipes, known as the goose neck barnacle, goose barnacle or leaf barnacle is a species of goose barnacle, also well known under the taxonomic synonym Pollicipes cornucopia. It is closely related to Pollicipes polymerus, a species with the same common names, but found on the Pacific coast of North America, and to Pollicipes elegans a species from the coast of Chile. It is found on rocky shores in the north-east Atlantic Ocean and is prized as a delicacy, especially in the Iberian Peninsula.

<span class="mw-page-title-main">Acrothoracica</span> Infraclass of barnacles

The Acrothoracica are an infraclass of barnacles.

<i>Dosima fascicularis</i> Genus of barnacles

The buoy barnacle is a species of goose barnacle known for its unique characteristic of hanging downwards from a floating appendage which drifts at the water surface and is carried along by ocean currents. It is regarded as "the most specialised pleustonic goose barnacle" as most other barnacle species are sessile filter feeders that remain fixed onto other firm surfaces.

<span class="mw-page-title-main">Facetotecta</span> Genus of crustaceans

Facetotecta is a poorly known subclass of thecostracan crustaceans. The adult forms have never been recognised, and the group is known only from its larvae, the "y-nauplius" and "y-cyprid" larvae. They are mostly found in the north Atlantic Ocean, neritic waters around Japan, and the Mediterranean Basin, where they also survive in brackish water.

<i>Amphibalanus improvisus</i> Species of barnacle

Amphibalanus improvisus, the bay barnacle, European acorn barnacle, is a species of acorn barnacle in the family Balanidae.

<i>Lepas anserifera</i> Species of barnacle

Lepas anserifera is a species of goose barnacle or stalked barnacle in the family Lepadidae. It lives attached to floating timber, ships' hulls and various sorts of flotsam.

<i>Pollicipes polymerus</i> Species of crustacean

Pollicipes polymerus, commonly known as the gooseneck barnacle or leaf barnacle, is a species of stalked barnacle. It is found, often in great numbers, on rocky shores on the Pacific coasts of North America.

<span class="mw-page-title-main">Multicrustacea</span> Superclass of crustaceans

The clade Multicrustacea constitutes the largest superclass of crustaceans, containing approximately four-fifths of all described non-hexapod crustacean species, including crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, barnacles, copepods, amphipods, mantis shrimp and others. The largest branch of multicrustacea is the class Malacostraca.

Polyascus is a genus of barnacles in infraclass Rhizocephala. It was circumscribed in 2003 by Henrik Glenner, Jørgen Lützen, and Tohru Takahashi. They included three species, all transferred from Sacculina. The generic name polyascus refers to the typical presence of multiple external sac-like female bodies, known as externae. In Polyascus species, these originate from asexual reproduction.

<i>Lepas testudinata</i> Species of barnacle

Lepas testudinata is a species of goose barnacle in the family Lepadidae. First observed in 1834, Lepas testudinata has undergone several reclassifications, and its relationship to other Lepas species is still the subject of ongoing research. L. testudinata is endemic to temperate waters in the China Seas, Australian Sea, and the Indo-West Pacific, and there are two distinct subgroups within the species. This barnacle species exclusively colonizes free-floating debris and tidewrack, and can form colonies of over 1000 members at a time. Due to this colonization habit, L. testudinata plays a role in biofouling and often serves as a foundation species when preyed upon.

<i>Pollicipes elegans</i> Species of barnacles

Pollicipes elegans, the Pacific goose barnacle, is a species of gooseneck barnacle inhabiting the tropical coastline of the eastern Pacific Ocean. Its habitat borders a close relative, Pollicipes polymerus, a gooseneck barnacle covering the coastline of the Pacific Northwest. Other species belonging to the genus Pollicipes are found along the eastern coastlines of the Atlantic Ocean.

References

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