Chaetogaster Diaphanus
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''Chaetogaster'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
belonging to the segmented worms (
Annelida The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to variou ...
). It is classified in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Naididae The Naididae (including the former family Tubificidae) are a family (biology), family of clitellate oligochaete worms like the sludge worm, ''Tubifex tubifex''. They are key components of the benthic communities of many freshwater and ocean, mar ...
(subfamily Naidinae) in the order
Oligochaeta Oligochaeta () is a subclass of soft-bodied animals in the phylum Annelida, which is made up of many types of aquatic and terrestrial worms, including all of the various earthworms. Specifically, oligochaetes comprise the terrestrial megadril ...
. These are ca. 2–25 mm long transparent
worm Worms are many different distantly related bilateria, bilateral animals that typically have a long cylindrical tube-like body, no limb (anatomy), limbs, and usually no eyes. Worms vary in size from microscopic to over in length for marine ...
s that are very common in fresh water and often form chains of individuals through asexual multiplication.


Morphology

Externally, the animals correspond to the general building plan of the
Naididae The Naididae (including the former family Tubificidae) are a family (biology), family of clitellate oligochaete worms like the sludge worm, ''Tubifex tubifex''. They are key components of the benthic communities of many freshwater and ocean, mar ...
, however, in distinction from the other
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
, they bear
setae In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae ...
(bristles) only on their ventral side, which is what gives them their name. The size of the worms, and of their setae too, is very variable even within single species.


Distribution and feeding

These worms occur mostly in stagnant or flowing
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...
, but the largest species, C''haetogaster diaphanus'', also lives in
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuary ...
. All species seem to be widely distributed within Europe and in addition, many species are found on multiple continents. The animals are usually very active but cannot swim purposefully. They therefore live on solid substrates like plants, stones and in the case of one
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
(''Chaetogaster limnaei limnaei''), on snails or (occasionally) mussels. They feed by sucking in small organisms of adequate size. The composition of the food items differs between species, as the size of the sucked in food components depends on each species' body size. While the biggest species ''Ch. diaphanus'' consumes organisms in the size class of about 0.3–3 mm, which mostly contains zooplankton (
rotifer The rotifers (, from Latin 'wheel' and 'bearing'), sometimes called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic Coelom#Pseudocoelomates, pseudocoelomate animals. They were first describ ...
s,
water flea The Diplostraca or Cladocera, commonly known as water fleas, is a superorder of small, mostly freshwater crustaceans, most of which feed on microscopic chunks of organic matter, though some forms are predatory. Over 1000 species have been recog ...
s etc.), ''Ch. limnaei'' e.g. selects organisms in the order of 0.03-0.3 mm, which are mainly
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
(
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s and
green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
) or very small
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
. In principle, all species are therefore omnivores, the bigger species taking in mainly animal food and the smaller ones rather plant particles. The unselective nature of their feeding is clearly shown by the fact that sand grains and other indigestible particles (e.g.
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
pollen) are always found in their stomachs. These particles are also in the indicated size range. Physiologically, the ''Chaetogaster'' species have already long been known as
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s with acid stomach content, just like
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s.


The special case of ''Chaetogaster limnaei''

The species ''Ch. limnaei'' represents a peculiar case. It occurs in two subspecies that are clearly distinct in their way of life. One subspecies lives parasitic inside
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s (''Ch. l. vaughini'') and only attacks snails above a certain minimum size. It apparently feeds mainly on the host's kidney cells. The other subspecies (''Ch. l. limnaei'') attaches itself externally to the body of snails or to the inside of their shell (''
Lymnaea ''Lymnaea'' is a genus of small to large-sized air-breathing freshwater snails, aquatic pulmonate gastropod mollusks in the subfamily Lymnaeinae of the family Lymnaeidae, the pond snails.Bouchet, P.; Rosenberg, G. (2013). Lymnaea Lamarck, 1799. ...
'', ''
Physa ''Physa'' is a genus of small, left-handed or sinistral, air-breathing freshwater snails, aquatic pulmonate gastropod molluscs in the subfamily Physinae of the family Physidae.MolluscaBase eds. (2021). MolluscaBase. Physa Draparnaud, 1801. Acces ...
'', ''
Ancylus ''Ancylus'' is a genus of very small, freshwater, air-breathing limpets. They are aquatic pulmonate gastropod mollusks in the tribe Ancylini within the family Planorbidae, the ram's horn snails and their allies.Bouchet, P.; Rosenberg, G. (20 ...
'', '' Australorbis'' and many other genera) and can freely move (so-called
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit f ...
ectosymbiosis Ectosymbiosis is a form of symbiotic behavior in which an organism lives on the body surface of another organism (the host), including internal surfaces such as the lining of the digestive tube and the ducts of glands. The ectosymbiotic species, ...
). It consumes various tiny organisms that it manages to suck in from its vantage point on its host. Apart from, e.g., rotifers and algae,
cercariae A cercaria (plural cercariae) is a larval form of the trematode class of parasites. It develops within the germinal cells of the sporocyst or redia. A cercaria has a tapering head with large penetration glands. It may or may not have a long swim ...
and
miracidia The miracidium is the second stage in the life cycle of trematodes. When trematode eggs are laid and come into contact with fresh water, they hatch and release miracidium. In this phase, miracidia are ciliated and free-swimming. This stage is comp ...
(two types of
trematode Trematoda is a Class (biology), class of flatworms known as trematodes, and commonly as flukes. They are obligate parasite, obligate Endoparasites, internal parasites with a complex biological life cycle, life cycle requiring at least two Host ( ...
larvae) are also among its food items. The ecological relationships in this biological control of parasites have been studied by various groups. It seems to play a particularly important role, but can apparently not be used in any practical way. The subspecies ''Ch. l. limnaei'' is also often found on freshwater
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...
s, especially of the genus ''
Sphaerium ''Sphaerium'' is a genus of very small freshwater clams, aquatic bivalve molluscs in the family Sphaeriidae, known as the fingernail clams. The small clams in this genus are unusual in that many of them, such as ''Sphaerium corneum'', can climb ...
''.


Reproduction

Like almost all members of the Naidinae, ''Chaeogaster'' species normally reproduce asexually. They thereby form characteristic chains of two or three worms that come into being through a form of
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
. These chains give rise to new individuals through division (
paratomy Paratomy is a form of asexual reproduction in animals where the organism splits in a plane perpendicular to the antero-posterior axis and the split is preceded by the "pregeneration" of the anterior structures in the posterior portion. The develop ...
), which have, because of the asexual nature of the multiplication, the same genetic blueprint as the original worm. However, the worms switch to sexual reproduction during difficult times like the winter. The subspecies ''Ch. l. limnaei'' then leaves its host in late summer or autumn, lives and sexually reproduces freely in the water for some time and then in spring, attaches itself to new snail hosts.


Systematics

The membership of the genus ''Chaetogaster'' to the family Tubificidae and the subfamily Naidinae has meanwhile been confirmed not only on morphological but also on molecular-genetic grounds. According to
cytochrome oxidase The enzyme cytochrome c oxidase or Complex IV (was , now reclassified as a translocasEC 7.1.1.9 is a large transmembrane protein complex found in bacteria, archaea, and the mitochondria of eukaryotes. It is the last enzyme in the respiratory e ...
I sequence studies, its closest relationship is with the genus '' Amphichaeta'' of the same subfamily.Bely, A.E., Wray, G.A. (2003): Molecular phylogeny of naidid worms (Annelida: Clitellata) based on cytochrome oxidase I. Mol. Phyl. Evol. 2004: 50-63. Nowadays, nine valid species are placed in the genus ''Chaetogaster'' Von Baer 1827: *''
Chaetogaster cristallinus ''Chaetogaster'' is a genus belonging to the segmented worms (Annelida). It is classified in the family (biology), family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very co ...
'' Vejdovsky 1883, often written as ''Chaetogaster crystallinus'' Vejdovsky; worm chains up to 7 mm in length; common. *'' Chaetogaster diaphanus'' (Gruithuisen 1828); usually from 10–15 mm in length, with extremes down to 2.5 mm (single worms) and up to 25 mm (worm chains); common. *'' Chaetogaster diversisetosus'' Sporka 1983; uncommon (gelegentlich). *'' Chaetogaster krasnopolskiae'' Lastockin 1937; uncommon (gelegentlich). *'' Chaetogaster langi'' Bretscher 1896; widely distributed. *''
Chaetogaster limnaei ''Chaetogaster'' is a genus belonging to the segmented worms (Annelida). It is classified in the family (biology), family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very co ...
'' Baer 1827; widely distributed. **''Ch. l. limnaei'' Baer, 1827; epizootic on snails. **''Ch. l. vaghini'' Gruffydd 1965; parasitic in snails. *'' Chaetogaster parvus'' Pointner 1914; widely distributed. *'' Chaetogaster setosus'' Svetlov 1925; widely distributed.


References


External links

{{Taxonbar, from=Q1058727 Haplotaxida