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The Spanish toothcarp (''Aphanius iberus''), also known as the Spanish pupfish or Iberian
killifish A killifish is any of various oviparous (egg-laying) cyprinodontiform fish (including families Aplocheilidae, Cyprinodontidae, Fundulidae, Profundulidae and Valenciidae). All together, there are 1,270 species of killifish, the biggest family ...
, is a small,
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
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 ...
of fish 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 ...
Cyprinodontidae. Its risk of extinction is one of the greatest of any Iberian
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, ...
. Its limited range, coupled with the drastic population decline the species has suffered in the last two decades, has caused it to be placed on
endangered species An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
lists, both in
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
and internationally. In addition, habitat fragmentation, likely due to humans, has resulted in this species becoming increasingly stagnant and has led to increased genetic drift. Also found to be contributing to their endangerment was genetic diversity of mitochondrial DNA. The mitochondrial DNA coupled with A. iberus’s geographic distribution has been able to affect their population genetic structure gradually in different spaces. Its
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
in the south of the
Iberian peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
has notably worsened. A similar fish in the peninsula's south-west has been classified as an independent species, ''
Aphanius baeticus ''Aphanius baeticus'' is a species of fish in the family Cyprinodontidae. It is endemic to a small part of Southern Spain, between the Gulf of Cadiz and Huelva. Its natural habitats are rivers, estuarine waters, and coastal saline lagoons. It i ...
''. '' Aphanius saourensis'' is another similar species in
Algeria ) , image_map = Algeria (centered orthographic projection).svg , map_caption = , image_map2 = , capital = Algiers , coordinates = , largest_city = capital , relig ...
.


Description


Morphology

This is a small fish, rarely more than in length. Females are longer than males of the same age. It has an oblong body and rounded fins. Its
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through conv ...
is underdeveloped with respect to the fish's girth. It has large
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
, numbering 20 to 26 across the body at its widest point. It has obvious
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
. The males are marked with bluish to silver-colored vertical stripes on the flanks and dark bands on the
caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
. The females are generally greenish-brown with dark blotches distributed irregularly over their bodies; their fins are transparent and mostly unpatterned. Some populations, most notably those in
Murcia Murcia (, , ) is a city in south-eastern Spain, the capital and most populous city of the autonomous community of the Region of Murcia, and the seventh largest city in the country. It has a population of 460,349 inhabitants in 2021 (about one ...
, have short, dark stripes as opposed to blotches.


Reproduction

The lifespan of a Spanish toothcarp is short. They reach
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
at age three months. The females
spawn Spawn or spawning may refer to: * Spawn (biology), the eggs and sperm of aquatic animals Arts, entertainment, and media * Spawn (character), a fictional character in the comic series of the same name and in the associated franchise ** '' Spawn: Ar ...
repeatedly each season, producing from 100 to 900 eggs. Toothcarps in the Murcia region tend to spawn between April and August, while more northerly populations, such as those in the Delta del Ebro, lay their eggs between May and August. Spawning is usually done in areas with nearby
vegetation Vegetation is an assemblage of plant species and the ground cover they provide. It is a general term, without specific reference to particular taxa, life forms, structure, spatial extent, or any other specific botanical or geographic character ...
, which shelters the eggs.
Hatchling In oviparous biology, a hatchling is a newly hatched fish, amphibian, reptile, or bird. A group of mammals called monotremes lay eggs, and their young are hatchlings as well. Fish Fish hatchlings generally do not receive parental care, similar to ...
s emerge approximately eight days after the eggs are laid. Females that hatch in April are able to reproduce in June. While the females spawn, adult males set up small territories that they defend from other males in ritualized combats through which they
court A court is any person or institution, often as a government institution, with the authority to adjudicate legal disputes between parties and carry out the administration of justice in civil, criminal, and administrative matters in accordance ...
the females.


Behavior

The Spanish toothcarp is an omnivore, and eats
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
,
crustaceans Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
,
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 and
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 ...
. Generally, toothcarps travel in small groups, staying near underwater vegetation, where they normally go unnoticed. The biology of the Spanish toothcarp is characterized by a high growth rate, early maturity, a high reproductive rate, multiple periods of egg-laying and a short lifespan. From an
evolutionary ecology Evolutionary ecology lies at the intersection of ecology and evolutionary biology. It approaches the study of ecology in a way that explicitly considers the evolutionary histories of species and the interactions between them. Conversely, it can ...
point of view, this lifestyle is highly adaptive for fish that live in unstable environments, such as
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environment ...
, where unpredictable conditions increase mortality. This strategy allows the toothcarp to exploit environmentally favorable conditions during the short intervals in which they occur, thereby revitalizing the population.


Habitat

Spanish toothcarp inhabit shallow, slow-moving bodies of water such as river-mouths, coastal lakes, and
ponds A pond is an area filled with water, either natural or artificial, that is smaller than a lake. Defining them to be less than in area, less than deep, and with less than 30% emergent vegetation helps in distinguishing their ecology from th ...
. They occur in
saline Saline may refer to: * Saline (medicine), a liquid with salt content to match the human body * Saline water, non-medicinal salt water * Saline, a historical term (especially US) for a salt works or saltern Places * Saline, Calvados, a commune in ...
and
fresh Fresh or FRESH may refer to: People *DJ Fresh (born 1977), UK-based drum and bass artist *DJ Fresh (producer), US-based R&B producer born Marqus Brown Arts, entertainment, and media Films * ''Fresh'' (1994 film), a crime film * ''Fresh'' (200 ...
water alike, due to their ability to tolerate high levels of
salinity Salinity () is the saltiness or amount of salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensionless and equal ...
: they are
euryhaline Euryhaline organisms are able to adapt to a wide range of salinities. An example of a euryhaline fish is the molly (''Poecilia sphenops'') which can live in fresh water, brackish water, or salt water. The green crab (''Carcinus maenas'') is an e ...
. The Spanish toothcarp can tolerate temperatures of , and pH levels between 6.5 and 7.5. Despite their adaptability, they have been displaced by
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
, such as both species of
mosquitofish The western Mosquitofish (''Gambusia affinis'') is a North American freshwater fish, also known commonly, if ambiguously, as simply Mosquitofish or by its generic name, ''Gambusia'', or by the common name gambezi. Its sister species, the easte ...
, introduced from North America to Spain in 1921 by doctor Sadí de Buen Lozano in an attempt to control
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...
l mosquitoes. The toothcarp has retreated to high salinity areas where the mosquitofish cannot live.


Distribution

The Spanish toothcarp is characteristic of the Iberian peninsula, and extends from the Aigüamolls of
Alt Empordà Alt Empordà (, "Upper Empordà") is a comarca (county) in Girona, Catalonia, Spain, one of two into which Empordà Emporda (from the official name in ca, Empordà, , name in es, Ampurdán, ) is a natural and historical region of Catalonia ...
to Lake Adra in
Almería Almería (, , ) is a city and municipality of Spain, located in Andalusia. It is the capital of the province of the same name. It lies on southeastern Iberia on the Mediterranean Sea. Caliph Abd al-Rahman III founded the city in 955. The city gr ...
. In the last two decades, the Spanish toothcarp has suffered a severe decline, due in part to destruction of suitable habitats. Only a few dozen isolated populations are known. They have been found in the
alluvial plain An alluvial plain is a largely flat landform created by the deposition of sediment over a long period of time by one or more rivers coming from highland regions, from which alluvial soil forms. A floodplain is part of the process, being the sma ...
of the
Segura River Segura (, ; la, Thader; ar, شقورة, Shaqūrah, or ) is a medium-sized river in southeastern Spain. It has its source in the Sierra de Segura. Course The river begins at Santiago Pontones ( province of Jaén), passes Calasparra, Cie ...
, the Chicamo River in Abanilla, the wetlands surrounding the
Mar Menor Mar Menor (, "minor/smaller sea") is a coastal saltwater lagoon in the Iberian Peninsula located south-east of the Autonomous Community of Murcia, Spain, near Cartagena. Its name is the opposite of the Mediterranean, which is the (greater/larg ...
and
Valencian Community The Valencian Community ( ca-valencia, Comunitat Valenciana, es, Comunidad Valenciana) is an autonomous community of Spain. It is the fourth most populous Spanish autonomous community after Andalusia, Catalonia and the Community of Madrid with ...
(more specifically in the Parque Natural de El Hondo, and in the Parque Natural de la Albufera with its springs), the Parque Natural del Delta del Ebro in
Tarragona Tarragona (, ; Phoenician: ''Tarqon''; la, Tarraco) is a port city located in northeast Spain on the Costa Daurada by the Mediterranean Sea. Founded before the fifth century BC, it is the capital of the Province of Tarragona, and part of Tar ...
, and the Albufera de Adra and the Adra River.


Divergence between populations

Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
relationships and
genetic divergence Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes (mutations) through time, often leading to reproductive isolation and continued mutation even after the populations hav ...
have been observed in Iberian populations of ''Aphanius iberus'' by using
isozymes In biochemistry, isozymes (also known as isoenzymes or more generally as multiple forms of enzymes) are enzymes that differ in amino acid sequence but catalyze the same chemical reaction. Isozymes usually have different kinetic parameters (e.g. dif ...
along with the complete
genetic sequence A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are usua ...
of
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
B. The results for nuclear genes were consistent with those for
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
, and demonstrated that the Atlantic and Mediterranean toothcarp populations derive from separate lineages. The level of genetic divergence shows that they were isolated early and that
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
has not occurred. The divergence between these two
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...
is the same as that found between other species of
pupfish Pupfish are a group of small killifish belonging to ten genera of the family Cyprinodontidae of ray-finned fish. Pupfish are especially noted for being found in extreme and isolated situations. They are primarily found in North America, South Am ...
. This discovery makes it necessary to separate the Atlantic and Mediterranean populations in order to protect their
diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...
and genetic identities. Molecular analysis, together with
morphometric Morphometrics (from Greek μορϕή ''morphe'', "shape, form", and -μετρία ''metria'', "measurement") or morphometry refers to the quantitative analysis of ''form'', a concept that encompasses size and shape. Morphometric analyses are co ...
data, has taxonomic implications. In fact, the populations of the
Atlantic basin The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe and ...
are so different from those in Europe that they have been recognized as a separate species, ''Aphanius baeticus''. The species name ''Aphanius iberus'' is limited to populations that live in the Mediterranean.


Conservation status

Despite being distributed over a large area on the peninsula, the Spanish toothcarp is largely found in small, threatened areas. Because of this, the Spanish Ministero de Medio Ambiente lists it as an
endangered species An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
. Among the threats are:
* Water contamination by Agricultural waste, agricultural and urban
waste Waste (or wastes) are unwanted or unusable materials. Waste is any substance discarded after primary use, or is worthless, defective and of no use. A by-product, by contrast is a joint product of relatively minor economic value. A waste prod ...
* Habitat destruction, primarily from development *
Introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...
such as Louisiana crawfish, mosquitofish and
largemouth bass The largemouth bass (''Micropterus salmoides'') is a carnivorous freshwater gamefish in the Centrarchidae ( sunfish) family, a species of black bass native to the eastern and central United States, southeastern Canada and northern Mexico, but ...
* Disappearance of
channels Channel, channels, channeling, etc., may refer to: Geography * Channel (geography), in physical geography, a landform consisting of the outline (banks) of the path of a narrow body of water. Australia * Channel Country, region of outback Austral ...
and irrigation ponds *
Stream bed A stream bed or streambed is the bottom of a stream or river (bathymetry) or the physical confine of the normal water flow (Channel (geography), channel). The lateral confines or channel margins are known as the stream Bank (geography), banks ...
contamination by accidental spills * Unintended consequences of combating
eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytopla ...
from
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 ...
, using toxic chemicals such as
copper sulfate Copper sulfate may refer to: * Copper(II) sulfate, CuSO4, a common compound used as a fungicide and herbicide * Copper(I) sulfate Copper(I) sulfate, also known as cuprous sulfate, is an inorganic compound with the chemical formula Cu2 SO4. It ...
, or by covering them to prevent
sunlight Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when t ...
from reaching the water *
Eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytopla ...
from algae * Contamination of
albufera The Albufera, La Albufera or L'Albufera de València (, meaning "lagoon" in Valencian, from Arabic ''al-buhayra'', "small sea"), is a freshwater lagoon and estuary on the Gulf of Valencia coast of the Valencian Community in eastern Spain. It ...
s * Keeping them as pets once increased their decline, as they were highly valued by
aquarists Fishkeeping is a popular hobby, practiced by aquarists, concerned with keeping fish in a home aquarium or garden pond. There is also a piscicultural fishkeeping industry, serving as a branch of agriculture. Origins of fishkeeping Fish have ...
. This interest has recently shifted to more colorful
tropical fish Tropical fish are generally those fish found in aquatic tropical environments around the world. Fishkeepers often keep tropical fish in freshwater and saltwater aquariums. The term "tropical fish" is not a taxonomic group, but rather is a gene ...
.


Vernacular names

This fish is known as ''fartet'' or ''peixet de sequiol'' in
Spanish Spanish might refer to: * Items from or related to Spain: **Spaniards are a nation and ethnic group indigenous to Spain **Spanish language, spoken in Spain and many Latin American countries **Spanish cuisine Other places * Spanish, Ontario, Cana ...
and
Catalan Catalan may refer to: Catalonia From, or related to Catalonia: * Catalan language, a Romance language * Catalans, an ethnic group formed by the people from, or with origins in, Northern or southern Catalonia Places * 13178 Catalan, asteroid #1 ...
(the latter sometimes uses the
diminutive A diminutive is a root word that has been modified to convey a slighter degree of its root meaning, either to convey the smallness of the object or quality named, or to convey a sense of intimacy or endearment. A (abbreviated ) is a word-formati ...
''fartonet'').


See also

*''
Samaruc The ''Valencia hispanica'' or Valencia toothcarp ( ca-valencia, samaruc) is a species of freshwater fish in the family Valenciidae endemic to the south of Catalonia and the Valencian Community, Spain. There is at least one historical record of ...
''


References


External links

* {{Taxonbar, from=Q2010263 Cyprinodontidae Endemic fish of the Iberian Peninsula Fish described in 1846 Aphanius Taxobox binomials not recognized by IUCN