Chaetogaster Limnaei
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''Chaetogaster'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
belonging to the segmented worms ( Annelida). It is classified in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Naididae The Naididae (including the former family Tubificidae) are a family of clitellate oligochaete worms like the sludge worm, '' Tubifex tubifex''. They are key components of the benthic communities of many freshwater and marine ecosystems. In fres ...
(subfamily Naidinae) in the order
Oligochaeta Oligochaeta () is a subclass of 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 megadrile earthworm ...
. These are ca. 2–25 mm long transparent
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 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 of clitellate oligochaete worms like the sludge worm, '' Tubifex tubifex''. They are key components of the benthic communities of many freshwater and marine ecosystems. In fres ...
, however, in distinction from the other
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, they bear
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
(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. Although the term specifically excludes seawater and brackish water, it does include ...
, 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 estua ...
. 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 biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
(''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 the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John H ...
s,
water flea The Diplostraca or Cladocera, commonly known as water fleas, are a superorder of small crustaceans that feed on microscopic chunks of organic matter (excluding some predatory forms). Over 1000 species have been recognised so far, with many more ...
s etc.), ''Ch. limnaei'' e.g. selects organisms in the order of 0.03-0.3 mm, which are mainly
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
(
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s and
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
) or very small
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 ...
. 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 single ...
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 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 chordate ...
s with acid stomach content, just like
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
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, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gastro ...
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, 179 ...
'', ''
Physa ''Physa'' is a genus of small, left-handed or sinistral, air-breathing freshwater snails, aquatic pulmonate gastropod mollusks in the subfamily Physinae of the family Physidae.MolluscaBase eds. (2021). MolluscaBase. Physa Draparnaud, 1801. Ac ...
'', ''
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. (2 ...
'', '' 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 fro ...
ectosymbiosis Ectosymbiosis is a form of symbiotic behavior in which a parasite lives on the body surface of the host, including internal surfaces such as the lining of the digestive tube and the ducts of glands.  The parasitic species is generally an im ...
). 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 Trematodes are parasitic flatworms of the class ''Trematoda'', specifically parasitic flukes with two suckers: one ventral and the other oral. Trematodes are covered by a tegument, that protects the organism from the environment by providing secr ...
and miracidiae (two types of
trematode Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive 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 know ...
. 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 developi ...
), 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 mitochondria of eukaryotes. It is the last enzyme in the respiratory electr ...
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 Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' 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 ''Chaetogaster'' is a genus belonging to the segmented worms ( Annelida). It is classified in the family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' Sporka 1983; uncommon (gelegentlich). *''
Chaetogaster krasnopolskiae ''Chaetogaster'' is a genus belonging to the segmented worms ( Annelida). It is classified in the family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' Lastockin 1937; uncommon (gelegentlich). *''
Chaetogaster langi ''Chaetogaster'' is a genus belonging to the segmented worms ( Annelida). It is classified in the family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' Bretscher 1896; widely distributed. *''
Chaetogaster limnaei ''Chaetogaster'' is a genus belonging to the segmented worms ( Annelida). It is classified in the family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' Baer 1827; widely distributed. **''Ch. l. limnaei'' Baer, 1827; epizootic on snails. **''Ch. l. vaghini'' Gruffydd 1965; parasitic in snails. *''
Chaetogaster parvus ''Chaetogaster'' is a genus belonging to the segmented worms ( Annelida). It is classified in the family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' Pointner 1914; widely distributed. *''
Chaetogaster setosus ''Chaetogaster'' is a genus belonging to the segmented worms ( Annelida). It is classified in the family Naididae (subfamily Naidinae) in the order Oligochaeta. These are ca. 2–25 mm long transparent worms that are very common in fresh wat ...
'' Svetlov 1925; widely distributed.


References


External links

{{Taxonbar, from=Q1058727 Haplotaxida