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The Old World silversides are a
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 ...
, Atherinidae, of
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
in the order
Atheriniformes The Atheriniformes, also known as the silversides, are an order of ray-finned fishes that includes the Old & New World silversides, the rainbowfishes, and several less-familiar families, including the unusual Phallostethidae. The order incl ...
. Atherinidae are abundant and considered bony fish (teleost) that are widespread globally, living in rivers, estuaries, and coastal waters. They occur worldwide in
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and temperate waters. About two-thirds of the
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), ...
are marine, and the remainder live in fresh water. The 74 species are in 13
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
. The
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 ...
''Craterocephalus'' is the most diverse with 25 species. Four genera are
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
. Silversides are relatively small with most being less than in length, with several not attaining lengths of more than . The body is generally elongated. Distinctive characters include two
dorsal fin A dorsal fin is a fin on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...
s widely separated, with the first consisting of flexible spines and the second having one spine followed by soft rays, while the
anal fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported o ...
has one spine on the leading edge followed by soft rays. The
pectoral fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish aquatic locomotion, swim. Apart from the tail or caudal fin, fish fins have no direct connection with the vertebral column ...
s tend to be high, and there is no
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelia ...
. On the flanks is a broad, silvery band. The scales are relatively large. They feed on
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 ...
. Some species, such as the hardyhead silverside, ''Atherinomorus lacunosus'', are commercially fished. The family Atherinopsidae (
Neotropical silverside The neotropical silversides are a family, Atherinopsidae, of fishes in the order Atheriniformes. About 112 species in 13 genera are distributed throughout the tropical and temperate waters of the New World, including both marine and freshwater ha ...
s) is closely related, while the genus ''
Atherion ''Atherion'' is a small genus of old world silverside, silversides, known as the pricklenose silversides. It is the only genus in the Family (biology), family Atherionidae. Other authorities classify this as a monogeneric subfamily, Atherioninae, ...
'' has been given family status in the Atherionidae by some authorities.


Subfamilies and genera

With reference to the 5th edition of ''
Fishes of the World ''Fishes of the World'' is a standard reference for the systematics of fishes. It was first written in 1976 by the American ichthyologist Joseph S. Nelson (1937–2011). Now in its fifth edition (2016), the work is a comprehensive overview of t ...
'' and the 2020 introduction of the genus ''Doboatherina'', the family should be classified as follows: * Subfamily Atherininae Risso, 1827 ** '' Atherina''
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, 1758
(7 species) ** '' Atherinason'' Whitley, 1934 (monotypic) ** '' Atherinosoma'' Castelnau, 1872 (2 species) ** '' Kestratherina'' A. Pavlov, Ivantsoff,
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts come in many styles and sizes, depending on the exact job they are designed for. Common variations ...
& Crowley, 1988
(2 species) ** '' Leptatherina'' Pavlov, Ivantsoff, Last & Crowley, 1988 (2 species) * Subfamily Atherinomorinae Dyer & Chernoff, 1996 ** '' Alepidomus'' C. L. Hubbs, 1944 (monotypic) ** '' Atherinomorus'' Fowler, 1903 (9 species) ** '' Doboatherina'' Sasaki & Kimura, 2019 (10 species) ** '' Hypoatherina'' Schultz, 1948 (15 species) ** '' Stenatherina'' Schultz, 1948 (monotypic) ** '' Teramulus''
J.L.B. Smith James Leonard Brierley Smith (26 September 1897 – 8 January 1968) was a South African ichthyologist, organic chemist, and university professor. He was the first to identify a taxidermied fish as a coelacanth, at the time thought to be long ext ...
, 1965
(2 species) * Subfamily Bleheratherininae Aarn & Ivantsoff, 2009 ** '' Bleheratherina'' Aarn & Ivanstoff, 2009 (monotypic) * Subfamily Craterocephalinae Dyer & Chernoff, 1996 ** ''
Craterocephalus ''Craterocephalus'' is a genus of small and slender brackish or freshwater old world silverside, silversides from Australia and New Guinea. It is the most diverse genus in the family Atherinidae, containing 25 of the 71 species. Species The curr ...
'' McCulloch, 1912 (26 species) ** '' Sashatherina'' Ivantsoff &
Allen Allen, Allen's or Allens may refer to: Buildings * Allen Arena, an indoor arena at Lipscomb University in Nashville, Tennessee * Allen Center, a skyscraper complex in downtown Houston, Texas * Allen Fieldhouse, an indoor sports arena on the Univ ...
, 2011
(monotypic) The oldest known Old World silverside is "''Atherina''" ''macrocephala''
Agassiz Jean Louis Rodolphe Agassiz ( ; ) FRS (For) FRSE (May 28, 1807 – December 14, 1873) was a Swiss-born American biologist and geologist who is recognized as a scholar of Earth's natural history. Spending his early life in Switzerland, he recei ...
, 1835
from the
Early Eocene In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
-aged Monte Bolca site of Italy.


Subfamily Craterocephalinae


Origin and characteristics of genus ''Craterocephalus''

In the silverside family Atherinidae, the genus ''Craterocephalus'' is the only one containing primarily freshwater species and is the most speciose. It is also the only genus that experienced high levels of radiation in freshwater ecosystems. Overall, there are twenty-five species, of which twenty are freshwater organisms and five are marine fish found only in Australian coastal waters. Within the genus, numerous species live in Australia’s freshwater ecosystems, specifically, 18 species live in Australia, 6 in New Guinea, and 1 in East Timor. How ''Craterocephalus'' diverged from being marine water organisms is linked to the ancestors of Craterocephalinae and Atherininae, where both subfamilies lived in similar environments but were separated by a North-South disjunction around Australia. The separation caused one lineage to remain in Northern Australia, forming the genus ''Craterocephalus,'' which developed distinct characteristics from the other subfamily Atherininae. Since all of Atherinidae are primarily marine, ''Craterocephalus'' is most likely also of marine origin. Freshwater fish in the genus ''Craterocephalus'' are found in various ecosystems, such as rivers, rainforest streams, creeks, and hot desert springs. Characteristics of the fish include having short life spans that are 2–3 years long, and their body length ranges from 5–10 cm. Maturity is reached when fish are 3–5 cm in length, and most species spawn shortly after one year of being born. Spawning rates increase in the spring and continue to be high in the summer — typically, one fish releases 85 eggs daily. Species in this genus are tolerant of environments differing in salt concentrations and temperature, especially ''C. eyresii'' and ''C. cuneiceps''. Both these species can tolerate 70–100 ppm of salt, while ''C. dalhousiensis'' can withstand high water temperatures up to 42 °C. The genus is split into five groups: "eyresii", "stercusmuscarum", "new honoriae", "capreoli", and "stramineus". Two groups, "eyresii" and "stercusmuscarum", are mainly found in Australian freshwaters. The majority of the fish in the "new honoriae" are marine organisms found in the Northern area of Australia, and their habitat slightly overlaps with the habitat of fish in the subfamily Atherininae. After the formation of the ''Craterocephalus'', the "new honoriae" group established itself in Northern Australia’s marine environments, while the other groups invaded freshwater habitats.


Species within ''Craterocephalus''

''Craterocephalus stercusmuscarum'' is an Australian freshwater fish of marine origin that has moderate dispersal potential. The species is abundant in the Wet Tropics of Northern Australia and travels long distances upland to areas near waterfalls.


Subfamily Atherininae


Origin of Atherininae

The separation between the ancestors of ''Craterocephalinae'' and Atherininae caused ''Atherininae'' lineage to form in Southern Australia, which has cooler habitats than Northern Australia. The subfamily Atherininae has five genera. One is Atherina, which is found near the Eastern Atlantic and Mediterranean oceans; however, the remaining four are restricted to Australia’s southern coastal waters. Since Atherina’s biogeographic location is significantly different, it is likely that Atherina and the four genera are not monophyletic or do not descend from a common ancestor. Instead, Atherina is hypothesized to have a sister lineage with the ancestors of Atherininae and the ''Craterocephalinae''.


Species within genus ''Atherina''

''Atherina boyeri'' is found in estuaries, coastal waters, lagoons, salt marshes, and brackish water ecosystems; therefore, this species is euryhaline or can tolerate different levels of salinity. The fish is typically located in the Mediterranean seas but is also found in the northeast and northwest Atlantic. ''A. boyeri'' is a commercial fish in Greece, where the fishing period begins in February but only lasts a few weeks. In Greece, this species is found in the Mesolongi and Etolikon lagoons. The maximum length observed for females and males that were three years old was 103 mm and 83.1 mm, respectively. Rapid growth is seen during the first year of life, specifically, fish reach about 66% of their adult size after one year. However, the fish experience a 20% decrease in length after one year, and a 15% decrease in length in their third year. Their lifespan typically ranges from 1-to 3 years. Both males and females have a single gland for their gonads. Young females have white ovaries, but as they mature, their ovaries are covered with peritoneum and darken to black. Males testis do not experience any changes and are white. Sexual maturity is reached two or three months after their birth, and the GSI value of the fish begins to increase early in the year and reaches its maximum ratio in May, indicating that the reproductive period begins in March and ends in June. Females have larger GSI values than males. The reproductive period is extensive and lasts about five months. In terms of the sex ratio, there are slightly more females than males in the population. The sex ratio between males and females is approximately 1:1.2. ''Atherina hepsetus'' is found in marine and brackish waters across the Eastern Atlantic coasts and can swim between 5 and 20 meters in depth. Their diet includes zooplankton and benthic crustaceans. In the Mediterranean, this species is an important commercial fish that is also a food source for other commercial fish. The approximate length for females males is 12.07 cm and 11.83 cm, respectively. The average weight of both sexes is 6 grams, while the maximum age is about four years. Additionally, the ratio between females and males is about 1.3:1. In terms of growth, the fish significantly increase in length and weight during their first year of life, with females reaching a length of 11.3 cm and males reaching 11.15 cm. Maturation also begins after one year; however, after sexual maturation, both sexes' growth rate decreases. Spawning rates increase in March and fall down after April. During the spawning season, the GSI value of males is larger than females, but the mass of the female gonads is slightly greater than their male counterparts.


Species within genus ''Atherinosoma''

''Atherinosoma elongata'' is found in Australia’s south-western estuaries and ranges from 40 to 78 mm in length. Fish are usually bigger in size during the summer months than winter months. The lifespan of this species is about one year, so the fish grow significantly during their first months of life. When sexual maturity is reached, females are about 68.9 mm in length while males are 54 mm. The gonadosomatic index (GSI) for both males and females increases after August and remains high until December, indicating their breeding period is around these months.


Species within genus ''Leptatherina''

Two species of ''Leptatherina (L. presbyteroides and L. wallacei)'' are located in South-Western Australia. ''L. presbyteroides'' is a primarily marine species that can also live in estuaries and ''L. wallacei'' is found in both estuary and inland waters. ''Leptatherina presbyteroides'' have a lifespan of one year, and its length ranges from 25 to 75 mm. Specifically, females are about 69.1 mm while males are 59.1 mm in length when they reach sexual maturity. Smaller fish are present throughout the year, which means that breeding occurs over a long period of time. This species primarily lives in estuaries, so their breeding occurs there, but as the fish mature, they travel to the ocean and spawn there. Spawning begins from August to November due to the GSI of the fish being significantly high in both males and females. However, younger fish have their GSI relatively high during April and May, indicating that spawning may occur in the Spring. ''Leptatherina wallacei'' has a lifespan of one year, and its length ranges from 25 to 50 mm; females are longer than males. When sexual maturity is reached, females have an average length of 54.8 mm, and males have an approximate length of 44.7 mm. Additionally, the number of larger fish decreases during November and December while the number of smaller fish increases, indicating that the spawning period of this species begins in the Spring and mortality begins later in the year.


References

{{Taxonbar, from=Q446802 * Extant Eocene first appearances Ypresian first appearances