Cheilostomatida, also called Cheilostomata, is an order of

Bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...

in the class

Gymnolaemata Gymnolaemata are a class of Bryozoans. Gymnolaemata are sessile, mostly marine organisms and grow on the surfaces of rocks, kelp, and in some cases on animals, like fish. Zooids are cylindrical or flattened. The lophophore is protruded by ac ...

. They are exclusively marine, colonial invertebrate animals. Cheilostome colonies are composed of

calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...

and grow on a variety of surfaces, including

rock Rock most often refers to: * Rock (geology), a naturally occurring solid aggregate of minerals or mineraloids * Rock music, a genre of popular music Rock or Rocks may also refer to: Places United Kingdom * Rock, Caerphilly, a location in Wale ...

s, shells,

seagrass Seagrasses are the only flowering plants which grow in marine (ocean), marine environments. There are about 60 species of fully marine seagrasses which belong to four Family (biology), families (Posidoniaceae, Zosteraceae, Hydrocharitaceae and ...

and

kelp Kelps are large brown algae or seaweeds that make up the order (biology), order Laminariales. There are about 30 different genus, genera. Despite its appearance and use of photosynthesis in chloroplasts, kelp is technically not a plant but a str ...

s. The colony shapes range from simple encrusting sheets to erect branching and even unattached forms. As in other

bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...

n groups, each colony is composed of a few to thousands of individual

polypide The polypide in bryozoans encompasses most of the organs and tissues of each individual zooid. This includes the tentacles, tentacle sheath, U-shaped digestive tract, musculature and nerve cells. It is housed in the zooidal exoskeleton, which in ...

s. Each individual has a U-shaped gut, and no

respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gr ...

circulatory In vertebrates, the circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the body. It includes the cardiovascular system, or vascular system, that consists of the heart an ...

system. Unique among bryozoans, cheilostome polypides are housed in a box-shaped

zooids A zooid or zoöid is an animal that is part of a colonial animal. This lifestyle has been adopted by animals from separate unrelated taxa. Zooids are multicellular; their structure is similar to that of other solitary animals. The zooids can ...

, which do not grow larger once the zooid is mature. The opening through which the polypide protrudes is protected by a

calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime (mineral), lime or being chalky. The term is used in a wide variety of Science, scientific disciplines. In zoology ''Calcare ...

chitin Chitin (carbon, C8hydrogen, H13oxygen, O5nitrogen, N)n ( ) is a long-chain polymer of N-Acetylglucosamine, ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is the second most abundant polysaccharide in nature (behind only cell ...

ous lidlike structure, an operculum. Cheilostomes possess avicularia, which have modified the operculum into a range of mandibles (possibly for defense) or hair-like setae (possibly for cleaning). The cheilostomes are the most abundant and varied of modern bryozoans. The classification in suborders is based upon frontal calcification and the mechanism of lophophore protrusion.

Evolution

Cheilostomes first appeared in the Late

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...

('' Pyriporopsis'') but diversified very slowly during the Early

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

, with only 1 family known up to the

Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...

. During the Late Cretaceous, cheilostomes diversified rapidly to reach a level of more than 20 families in the

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

, replacing

cyclostomes Cyclostome is a biological term (from the Greek for "round mouth") used in a few different senses: * for the taxon Cyclostomi, which comprises the extant jawless fishes: the hagfish (Myxini) and the lampreys (Petromyzontidae). This was thought for ...

as the dominant group of bryozoans. At the same time new forms evolved which partly or fully used

aragonite Aragonite is a carbonate mineral and one of the three most common naturally occurring crystal forms of calcium carbonate (), the others being calcite and vaterite. It is formed by biological and physical processes, including precipitation fr ...

instead of

calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...

in their exoskeleton. This diversification is thought to be a consequence of the evolution of a new larval type. Though the

Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...

had some impact on genetic diversity, the rapid diversification continued into the Eocene, then apparently reaching a plateau of about 50 families up to the

Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...

. Most species incubate their offspring in brood chambers which has evolved independently at least 10 times in the order.

References

* Hayward, P.J. (2001). Bryozoa, in: Costello, M.J. et al. (Ed.) (2001). European register of marine species: a check-list of the marine species in Europe and a bibliography of guides to their identification. Collection Patrimoines Naturels, 50: pp. 325–333 * Clarke, A.; Johnston, N.M. (2003). Antarctic marine benthic diversity. Oceanography and Marine Biology: an Annual Review. 41: 47-114. {{Taxonbar, from1=Q56315980, from2=Q2309550 Bryozoan orders Extant Late Jurassic first appearances