Overview

Comprehensive Description

Biology

Commonly found close inshore. Forms large schools. Feeds on small bony fishes and cephalopods. Viviparous (with a yolk-sac placenta), 1 to 6 young per litter. Size at birth 51 to 64 cm. Presumably eaten fresh and dried salted.
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Distribution

Range Description

Northwest, western central and southwest Atlantic: USA (Possibly north as a vagrant to New York, rare off North Carolina, and normally from off South Carolina, Georgia, the Atlantic coast of Florida and the Gulf of Mexico coast of Florida, Alabama, Mississippi and Texas). A record from Cuba is apparently erroneous (Compagno in prep, Castro 1993). However, Wiley and Simpfendorfer (2007) recently extended the US range to approximately 25°N which increases the likelihood of exchange between the US and Cuba. In South America, it is known with certainty from São Paulo to Santa Catarina States in southern Brazil (Soto 2001, Compagno et al. 2005). It is also reported from Trinidad and Guyana and may be more wide-ranging than currently documented (Garrick 1985, Castro 1993, Compagno in prep).

Records of the species from the eastern central Atlantic, from Senegal and Guinea-Bissau, are unconfirmed and probably incorrect (Compagno in prep, M. Ducrocq pers. comm. 2006).
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Distribution

N.Y. to Cuba and n. Gulf of Mexico; also in e. Atlantic
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Distribution

European waters (ERMS scope), Gulf of Mexico, North West Atlantic
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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Western Atlantic: North Carolina, USA to Cuba, the Gulf of Mexico, and southern Brazil to Uruguay (Ref. 58839). Occurrence in the eastern Atlantic, specifically in Senegal and Guinea-Bissau have not been confirmed, and may be based on Carcharhinus brevipinna.
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Physical Description

Size

Maximum size: 1900 mm TL
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Max. size

190 cm TL (male/unsexed; (Ref. 10731))
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
A coastal shark, occurring in shallow waters from the intertidal to depths of ~20 m (Compagno et al. 2005), which is common off beaches, in bays, estuaries and off river mouths. They often form large schools.

Throughout the southeastern USA and Gulf of Mexico, finetooth sharks use coastal bays and estuaries as nursery grounds (Carlson and Brusher 1999, McCandless et al. 2002). Adults and juveniles are common in shallow coastal waters off South Carolina, Georgia, northern states along the Gulf of Mexico during the warm summer months and migrate south when surface water temperatures drop below 68°F (20°C). In the Atlantic Ocean off the USA, they spend the winter months in the waters off the coast of Florida.

Estimates of size and age-at-maturity for male and female sharks from the Gulf of Mexico were different from those in the US Atlantic and northern Gulf of Mexico (Carlson et al. 2003, Drymon et al. 2006). Fork length at which 50% of the population reached maturity is 1,022 mm in the US South Atlantic and 990 mm in the Gulf of Mexico for females and was found to be significantly different. Median fork length at maturity for males is 988 mm and 935 mm for the US South Atlantic and Gulf of Mexico, respectively. Median age-at-maturity was 6.2 and 4.2 years for females, and 4.9 and 3.5 years for males for sharks in the US Atlantic and Gulf of Mexico, respectively (Carlson et al. 2003, Drymon et al. 2006).

Size at birth is 55?58 cm TL and fecundity is 4.0 (SD=0.79) pups for sharks in the US South Atlantic (Castro 1993). Off Louisiana, Neer and Thompson (2004) observed three gravid females with litters of three, four and five pups, respectively. The reproductive cycle is currently assumed to be biennial (Castro 1993).

Significant differences between von Bertalanffy growth curves were found between sharks in the US Atlantic and northern Gulf of Mexico for females but not males (Carlson et al. 2003, Drymon et al. 2006). In the US Atlantic, growth coefficients (K) were 0.19 yr-1 for females and 0.33 yr-1 for males whereas female growth coefficient were 0.24 yr-1 and males were 0.41 yr-1 in the Gulf of Mexico. The maximum observed ages, based on vertebral band counts, were 8.2 and 10.3 years for male sharks from the Gulf of Mexico and US Atlantic, respectively (Carlson et al. 2003).

Castro (1993) published descriptive diet data for the finetooth shark in the northwest Atlantic off the coasts of South Carolina and Daytona Beach, FL, and found teleosts (Atlantic Menhaden (Brevoortia tyrannus), Spot Croaker (Leiostomus xanthurus), Spanish Mackerel (Scomberomorus maculatus), and Mugil species) to be the primary prey. In northwest Florida, Finetooth Sharks fed almost entirely on menhaden and showed no ontogenetic shift in diet (Bethea et al. 2004).

Systems
  • Marine
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Habitat

benthic
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Depth range based on 3 specimens in 1 taxon.

Environmental ranges
  Depth range (m): 6 - 9

Graphical representation

Depth range (m): 6 - 9
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.
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Habitat Type: Marine

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Environment

demersal; marine; depth range ? - 10 m (Ref. 55182)
  • Florida Museum of Natural History 2005 Biological profiles: finetooth shark. Retrieved on 26 August 2005, from www.flmnh.ufl.edu/fish/Gallery/Descript/finetoothshark/finetoothshark.html. Ichthyology at the Florida Museum of Natural History: Education-Biological Profiles. FLMNH, University of Florida. (Ref. 55182)   http://www.fishbase.org/references/FBRefSummary.php?id=55182&speccode=872 External link.
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Migration

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

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Life History and Behavior

Life Cycle

Distinct pairing with embrace (Ref. 205). Viviparous, placental (Ref. 50449).
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Life Expectancy

Lifespan, longevity, and ageing

Maximum longevity: 8 years (wild)
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Carcharhinus isodon

The following is a representative barcode sequence, the centroid of all available sequences for this species. 

 
There are 12 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
 
ESHKC115-07|07-GHRI-0666|Carcharhinus isodon| ------------------------------------------CTTTACCTAATTTTTGGTGCATGAGCAGGTATAGTAGGAACAGCCCTA---AGTCTCCTAATTCGAGCTGAACTTGGGCAACCTGGATCACTTTTAGGAGAT---GATCAGATCTACAATGTAATCGTAACCGCCCACGCTTTTGTAATAATCTTTTTCATGGTTATACCAATTATAATTGGTGGTTTCGGAAATTGACTAGTTCCCTTAATA---ATTGGTGCGCCGGATATAGCCTTCCCACGAATAAATAACATAAGTTTCTGACTTCTTCCACCGTCATTTCTTCTTCTCCTCGCCTCTGCTGGAGTAGAAGCTGGAGCAGGTACTGGCTGAACAGTCTATCCTCCATTAGCTAGCAACCTAGCACATGCTGGACCATCTGTTGACTTA---GCTATCTTCTCTCTTCACTTAGCTGGTGTTTCATCAATTTTAGCTTCAATTAATTTTATCACAACTATTATTAATATAAAACCACCAGCCATTTCCCAATATCAAACACCATTATTTGTTTGATCTATTCTTGTAACCACTATTCTTCTTCTCCTTTCACTTCCAGTTCTTGCAGCA---GGGATTACAATACTACTTACAGATCGCAACCTTAATACTACATTCTTTGACCCTGCAGGTGGAGGAGACCCAATTCTTTATCAACACCTA------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------- 
-- end --

Download FASTA File
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Statistics of barcoding coverage: Carcharhinus isodon

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 12
Species: 22
Species With Barcodes: 1

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2009

Assessor/s
Carlson, J., Kyne, P.M. & Valenti, S.V.

Reviewer/s
Stevens, J.D., Dudley, S., Pollard, S., Soldo, A. & Kulka, D.W. (Shark Red List Authority)

Contributor/s

Justification
The Finetooth Shark (Carcharhinus isodon) is a coastal species occurring in shallow waters from the intertidal to depths of ~20 m. It is predominantly distributed in US waters in the western Atlantic and the northern Gulf of México where it is locally common. In South America, it is reported from São Paulo to Santa Catarina States in southern Brazil, Trinidad and Guyana. Stock assessments for the finetooth shark in US Atlantic waters and Gulf of Mexico indicate that the current status of the population is above maximum sustainable yield and no overfishing is occurring, therefore this population is assessed as Least Concern. This species is apparently rare throughout its reported range off South America, and virtually no information is available from this region. This rarity, together with its shallow inshore occurrence in a region which faces heavy coastal fishing pressure raises concern for the species? threat status in South America, particularly as significant declines have been documented in other elasmobranchs that occur in coastal waters of Brazil. However, due to infrequent records and a total lack of information on the species? full distribution and catches in fisheries, it is not possible to assess this population beyond Data Deficient at present. Further research is urgently required into the species? full distribution, catch levels, impact of fisheries and population trends off South America. Available information suggests that the majority of total global population occurs in the US Atlantic and Gulf of Mexico; thus the species is currently listed as Least Concern globally. With further information on the South American population of this species, this assessment may need to be revisited.
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National NatureServe Conservation Status

United States

Rounded National Status Rank: NNR - Unranked

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NatureServe Conservation Status

Rounded Global Status Rank: GNR - Not Yet Ranked

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Population

Population
The population structure of this species is poorly known throughout its range, but it is reasonable to assume that little to no exchange occurs between the population of this highly coastal species off the US Atlantic and Gulf of Mexico and the population off South America.

US Atlantic and Gulf of Mexico
Common in inshore waters off the eastern coast of the USA and the Gulf of Mexico. It is abundant along the coast of the southeastern United States and the Gulf of Mexico (Carlson et al. 2003, Wiley and Simpfendorfer 2007). Stock assessments for the US population of finetooth shark indicate that the current status of the population is above maximum sustainable yield and no overfishing is occurring (Cortés 2007).

South America
This species is poorly documented off South America, but is apparently rare throughout its range there (Castro 1993, Soto 2001, Compagno et al. 2005).

Population Trend
Unknown
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Threats

Threats

Major Threats
US Atlantic and Gulf of Mexico
Targeted by recreational fishers, small gillnet fisheries and occasionally taken as bycatch in demersal shrimp trawls that occur along the southeast coast of the USA and Gulf of Mexico. Estimated commercial landings were 69,258 lbs dressed weight in 2000 while recreational catches of small coastal sharks were 86,167 lbs (Cortés 2002). Finetooth Shark makes up a small proportion of these landings but the actual number is unknown. The group ?small coastal sharks? includes Atlantic Sharpnose Shark (Rhizoprionodon terraenovae), Blacknose Shark (Carcharhinus acronotus) and Bonnethead Shark (Sphyrna tiburo), as well as Finetooth Shark (Carcharhinus isodon).

The Finetooth Shark is vulnerable to overfishing and depletion because of targeted fishing in gillnet fisheries in the southeastern United States (Carlson and Bethea 2007). Despite this, recent stock assessments have shown that the US population of this species is above maximum sustainable yield, with current fishing levels not threatening its viability (Cortés 2007). This species is also restricted by its limited distribution (both bathymetric and geographic), with the majority of individuals living in shallow inshore waters adjacent to built-up areas, with extensive habitat degradation and some fisheries. However, the coastal nature of this species affords it some protection because many gillnet fisheries in US state waters have been banned (NMFS 2007).

South America
Shark fisheries have increased dramatically in the southern part of the Finetooth Shark?s range during the past half-century (L.J.V. Compagno pers. comm. 2008). The species? apparent rarity is of concern in the southern part of its range because rare species captured as bycatch of coastal shark fisheries may become even rarer to the point of local extirpation as the fisheries continue, supported by the more common species (L.J.V. Compagno pers. comm. 2008).

Coastal species are the most important commercial elasmobranchs in the Southwest Atlantic, and inshore fisheries are generally very intense off southern Brazil (Bonfil et al. 2005). Intensive fishing by pair trawl, shrimp trawl, gillnet and beach seine in near-shore waters is documented off Rio Grande do Sul State (Vooren and Klippel 2005). Although this species? range does not extend to Rio Grande do Sul, similar fisheries are likely to operate throughout much of the coast of Santa Catarina to São Paulo States, from which it is known.

Fisheries in Guyana and Trinidad and Tobago are generally inshore and artisanal (Chan A Shing 1999). Intensive trawling occurs in Trinidad and Tobago's waters (Mohammed and Chan A Shing 2003). Bycatch of the shrimp trawl fleet is considerably higher than the target catches. Total bycatch in these fisheries declined from 13,712 t in 1987 to 4,099 t in 2001 although the species composition of this bycatch has not yet been examined (Mohammed and Chan A Shing 2003). An artisanal gillnet and line fishery targets mackerel in coastal waters off Trinidad and Tobago and takes sharks as bycatch. This is the most widespread fishing method, accounting for over 85% of artisanal shark landings (Chan A Shing 1999).

In Guyana, there is a partially directed gillnet fishery for sharks (~600 vessels) which captures both demersal and pelagic inshore species. This fishery operates in estuarine and shallow coastal waters (at depths of <40 m) (Chan A Shing 1999). About 100 industrial shrimp trawlers also take sharks and finfish as bycatch, but no details are available on the composition of the bycatch. An artisanal demersal longline fishery targets sharks and catfish at depths of 9?20 m (Chan A Shing 1999).
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Least Concern (LC)
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Management

Conservation Actions

Conservation Actions
US Atlantic and Gulf of Mexico
Finetooth sharks are managed under the US Fishery Management Plan for Atlantic tunas, swordfish, and sharks and management actions are underway to limit potential increases in fishing. Moreover, gillnet bans in state waters of Florida, Georgia, South Carolina, and Texas will give added protection to this species, as it primarily occurs in state waters. See http://www.nmfs.noaa.gov/sfa/hms/hmsdocument_files/sharks.htm for up-to-date information and the regulations that apply to the small coastal shark complex.

South America
No measures in place. Recommended: The status of this species in South American waters should be investigated. Research is needed into distribution, catch levels, impact of fisheries and population trends.
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Relevance to Humans and Ecosystems

Benefits

Importance

fisheries: minor commercial
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Wikipedia

Finetooth shark

The finetooth shark (Carcharhinus isodon) is a species of requiem shark, family Carcharhinidae, found in the western Atlantic Ocean from North Carolina to Brazil. It forms large schools in shallow, coastal waters and migrates seasonally following warm water. A relatively small, slender-bodied shark, the finetooth shark can be identified by its needle-like teeth, dark blue-gray dorsal coloration, and long gill slits. It attains a maximum length of 1.9 m (6.2 ft). The diet of this species consists primarily of small bony fishes, in particular menhaden. Like other members of its family, it is viviparous with females giving birth to 2–6 pups in estuarine nursery areas every other year.

Valued for its meat, the finetooth shark forms an important component of the commercial gillnet shark fishery operating off the southeastern United States. Population assessments suggest that this fishery does not currently pose a threat to U.S. populations of the species. The International Union for Conservation of Nature (IUCN) has therefore listed the finetooth shark under Least Concern, though there is no fishery data available for this species off South America. This shark is not known to pose a danger to humans, though it snaps vigorously when captured and should be handled with caution.

Contents

Taxonomy and phylogeny

The finetooth shark was originally described as Carcharias (Aprion) isodon by French zoologist Achille Valenciennes, in Müller and Henle's 1839 Systematische Beschreibung der Plagiostomen. The type specimen is a 65 cm (26 in) juvenile male, possibly caught off the U.S. state of New York. This species was later moved to the genus Carcharhinus.[2] The specific epithet isodon means "equal teeth" in Greek, and refers to the similar number of teeth in the upper and lower jaws.[3] This species may also be referred to as the eventooth shark, smoothtooth shark, or night shark (usually used for C. signatus).[4]

As is the case for most Carcharhinus species, attempts to analyze the finetooth shark's phylogenetic relationships have yielded variable results. In 1988, Leonard Compagno grouped this species with the spinner shark (C. brevipinna), blacktip shark (C. limbatus), graceful shark (C. amblyrhynchoides), and the smooth tooth blacktip shark (C. leiodon), on the basis of morphological characters. Gavin Naylor's 1992 allozyme analysis found that the finetooth shark is the second-most basal member of the genus next to the blacknose shark (C. acronotus).[5] Mine Dosay-Akbulut's 2008 study, based on ribosomal DNA, suggested that the closest relative of the finetooth shark is the smalltail shark (C. porosus), and that the two species form a clade apart from other Carcharhinus species.[6]

Distribution and habitat

In North American waters, the finetooth shark is common and found from North Carolina to the northern Gulf of Mexico, and very occasionally straying as far north as New York. In Central and South American waters, it is rare but may occur more widely than presently known, having been reported off Trinidad and Guyana, infrequently from the Caribbean Sea, and off southern Brazil from São Paulo to Santa Catarina. The northwestern Atlantic, Gulf of Mexico, and South America populations are distinct, with little interchange between them.[1][7] There are old records of this species in the eastern Atlantic off Senegal and Guinea-Bissau, but these likely represent misidentifications of spinner sharks (C. brevipinna).[2]

An inhabitant of coastal habitats, the finetooth shark is often found near beaches and in bays and estuaries. It ranges no deeper than 10 m (33 ft) deep in the summer and 20 m (66 ft) deep in the winter.[7] Historically it was known to venture into rivers in the Gulf Coastal Plain of Texas, though most of paths into this area are now blocked by dams.[8] The northwestern Atlantic population of this species is strongly migratory: juveniles, followed by adults, arrive off South Carolina from late March to early May, when the water temperature rises above 20 °C (68 °F). They remain until September to mid-October, at which point the water temperature drops and they move south to Florida. The movements of other populations are unknown.[7]

Description

Drawing of a finetooth shark and its tooth

The body of the finetooth shark is slender and streamlined. The snout is long and pointed, with the nares preceded by short, broadly triangular flaps of skin. The eyes are large and round, with nictitating membranes (protective third eyelids). The mouth is broad with well-defined furrows at the corners. There are 12–15 tooth rows on either side of the upper jaw and 13–14 tooth rows on either side of the lower jaw. Each tooth is small and needle-like, with a narrow central cusp and smooth to minutely serrated edges. The five pairs of gill slits are long, measuring about half the length of the dorsal fin base.[2]

The first dorsal fin is high and triangular with a pointed apex, originating forward of the free rear tips of the pectoral fins. The second dorsal fin is relatively large and originates over the anal fin. There is no ridge running between the dorsal fins. The pectoral fins are small and falcate (sickle-shaped), with pointed tips.[2] The dermal denticles are small and overlapping, each bearing three horizontal ridges leading to marginal teeth. Living finetooth sharks are a distinctive dark bluish-gray above and white below, with a faint pale stripe on the flanks and no prominent fin markings.[3] Some individuals from Florida have green eyes.[9] Males average 1.6 m (5.2 ft) in length and females 1.7 m (5.4 ft); the largest shark on record was 1.9 m (6.2 ft) long.[3]

Biology and ecology

A male finetooth shark caught in the Gulf of Mexico

Both adults and juvenile finetooth sharks form large schools.[2] This energetic, fast-moving predator feeds mainly on small bony fishes, often entering the surf zone during the day to hunt.[10] The most important prey of this species in the northwestern Atlantic is the Atlantic menhaden (Brevoortia tyrannus), with sharks of all ages off northwestern Florida eat almost nothing else. The menhaden are swallowed whole after the head has been removed. Other known prey species include spot croaker (Leiostomus xanthurus), Spanish mackerel (Scomberomorus maculatus), mullet (Mugil spp.), shrimp, and in one case a juvenile Atlantic sharpnose shark (Rhizoprionodon terraenovae), which may have been scavenged from the bycatch discard of a shrimp trawler.[7] The finetooth shark may be preyed upon by larger sharks.[3]

Life history

Like other requiem sharks, the finetooth shark is viviparous: the developing embryos are nourished by yolk for the first 15 weeks, after which the depleted yolk sac develops into a placental connection to the mother. Females produce litters of 2–6 pups every other year. In the northwestern Atlantic, mating occurs from early May to early June and the young are born at around the same time the following year after a 12-month gestation period. The males bite at the female to hold her for copulation. The semen exuded by the male congeals into a large spongy mass inside the female's uterus, in which the individual spermatozoa are embedded. Called a "spermatozeugma", the function of this short-lived structure is unknown. Newborn finetooth sharks measure 48–64 cm (19–25 in) long.[2][7] Shallow bays and estuaries, such as Bull's Bay in South Carolina, serve as critical nursery areas for newborns and juveniles.[11]

Female finetooth sharks grow much more slowly and to a larger ultimate size than males.[12] Individuals in the northwestern Atlantic population have a larger body size and reach sexual maturation later than those in the Gulf of Mexico population; males in the two populations grow at similar rates, but northwestern Atlantic females grow more slowly than Gulf of Mexico females. In the northwestern Atlantic, males mature at a fork length (length from snout tip to caudal fin fork) of 99 cm (39 in) and females at 102 cm (40 in), corresponding to ages of 5 and 6 years respectively. In the Gulf of Mexico, males mature at a fork length of 94 cm (37 in) and females at 99 cm (39 in), corresponding to ages of 4 and 5 years respectively.[1][13] The maximum lifespan has been estimated to be at least 9 years for males and 14 years for females.[12]

Human interactions

A finetooth shark tagged by the NOAA Fisheries Shark Tagging Program

The finetooth shark has never been implicated in an attack on humans.[3] However, when caught this shark will thrash and snap at anything within range, and people have been bitten attempting to handle it.[14] Finetooth sharks are utilized for human consumption fresh or dried and salted. Other than off the southeastern United States, this species is of little commercial importance: it is small and occurs in water too shallow for most commercial and recreational fisheries, and is generally too fast-swimming to be caught by shrimp trawlers. Small numbers are taken incidentally by floating longlines and on hook-and-line.[7] This species is susceptible to overfishing due to its low reproductive rate, and to habitat degradation due to its inshore habits.[1]

Substantial numbers of finetooth sharks are caught in drift gillnets operated by the shark fishery off southeastern United States, which peaked in 1999 at about 117 tons landed.[12] Stock assessments conducted in 2002 suggested that U.S. populations had not yet been overfished but that the catch rate was unsustainable,[11] whereas assessments conducted in 2007 concluded that catch rates did not exceed sustainable levels and that populations were stable.[1] Fishing for this species in U.S. waters is regulated by the National Marine Fisheries Service (NMFS) 1993 Fisheries Management Plan (FMP) for Atlantic and Gulf of Mexico sharks; it is classified as a "Small Coastal Shark" (SCS) for the purposes of commercial quotas and recreational bag limits.[12] As a result, the International Union for Conservation of Nature (IUCN) has assessed the finetooth shark as of Least Concern overall and off the U.S. and Mexico. There is concern for this species in South America, where its numbers seem naturally low and it is potentially under heavy pressure by widespread, intensive coastal fisheries. However, at present the IUCN does not have sufficient data to assess its status in the region beyond Data Deficient.

References

  1. ^ a b c d e Carlson, J., P.M. Kyne and S.V. Valenti (2008). "Carcharhinus isodon". IUCN Red List of Threatened Species. Version 2010.1. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/161524. Retrieved April 24, 2010. 
  2. ^ a b c d e f Compagno, L.J.V. (1984). Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. Rome: Food and Agricultural Organization. pp. 477–478. ISBN 92-5-101384-5. 
  3. ^ a b c d e Bester, C. Biological Profiles: Finetooth Shark. Florida Museum of Natural History Ichthyology Department. Retrieved on May 5, 2009.
  4. ^ Froese, Rainer, and Daniel Pauly, eds. (2009). "Carcharhinus isodon" in FishBase. May 2009 version.
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  13. ^ Carlson, J.K., Drymon, J.M. and Neer, J.A. (2007). "Life history parameters for finetooth sharks, Carcharhinus isodon, from the United States South Atlantic Ocean and northern Gulf of Mexico". SEDAR 13 Small Coastal Sharks Data Workshop, Working Document SEDAR 13-DW-11.
  14. ^ Parsons, G.R. (2006). Sharks, Skates, and Rays of the Gulf of Mexico: A Field Guide. University Press of Mississippi. p. 64. ISBN 1-57806-827-4. 
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