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Cladus: Eukaryota
Supergroup: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Classis: Chondrichthyes
Subclassis: Elasmobranchii
Superordo: Selachimorpha
Ordo: Carcharhiniformes
Familia: Scyliorhinidae
Genus: Galeus
Species: G. antillensi - G. arae - G. atlanticus - G. boardmani - G. cadenati - G. eastmani - G. melastomus - G. mincaronei - G. murinus - G. nipponensis - G. piperatus - G. polli - G. sauteri - G. schultzi - G. springeri -

Name

Galeus Rafinesque, 1810

Galeus is a genus of catshark, family Scyliorhinidae, commonly known as sawtail catsharks in reference to a distinctive saw-toothed crest of enlarged dermal denticles found along the upper edges of their caudal fins.[3] They are found in the Atlantic, the western and central Pacific, and the Gulf of California, inhabiting deep water on or close to the bottom. Members of this genus are rather small, slim sharks with firm bodies and thick, rough skin. Their heads are usually fairly long and pointed, and have large mouths with well-developed furrows at the corners. They have large pectoral and anal fins, and two similar dorsal fins placed well back. Many species are ornately patterned with dark saddles and/or blotches. Sawtail catsharks feed on various invertebrates and fishes, and may be either egg-laying or live-bearing. These harmless sharks are sometimes caught as bycatch but are of minimal commercial value.

Taxonomy
The illustration that accompanied the 1904 description of the mouse catshark, as Pristurus murinus.

Galeus, derived from the Greek galeos meaning "shark", is one of the oldest carcharhiniform generic names. It was first used in a binomial by Constantine Samuel Rafinesque in his 1810 Caratteri di alcuni nuovi generi e nuove specie di animali (principalmente di pesci) e piante della Sicilia: con varie osservazioni sopra i medesimi.[4] Rafinesque listed G. melastomus, G. vulpecula (=Alopias vulpinus), G. mustelus (=Mustelus mustelus), and G. catulus (=Scyliorhinus canicula). Subsequently, in 1816 Georges Cuvier used Galeus to refer to the genus presently known as Galeorhinus, and in 1818 William Elford Leach used Galeus to refer to genus presently known as Mustelus. As a result, 19th century authors generally used Galeus for the tope sharks, and Pristiurus, coined by Charles Lucien Bonaparte in 1834, for the sawtail catsharks.[3]

Rafinesque likely intended G. mustelus to be the type species for Galeus, but of his listed species he furnished a description only for G. melastomus. Therefore, in 1908 Henry Weed Fowler designated G. melastomus as the type species of Galeus, establishing the genus to contain the sawtail catsharks.[5] Pristiurus became a junior synonym, though it continued to appear in scientific literature for some time after. Fowler's definition of Galeus gained widespread acceptance after Henry Bryant Bigelow and William Charles Schroeder's 1948 taxonomic review.[3][6] In 1952, Philip Orkin advocated that Pristiurus take precedence over Galeus, based on David Starr Jordan and Barton Warren Evermann's (possibly questionable) designation of G. mustelus as a type species for Galeus in 1896.[7] Leonard Compagno and most other recent authors have not upheld his proposal, in the interests of taxonomic stability.[3]

Species

* Antilles catshark, Galeus antillensis Springer, 1979
* Roughtail catshark, Galeus arae (Nichols, 1927)
* Atlantic sawtail catshark, Galeus atlanticus (Vaillant, 1888)
* Longfin sawtail catshark, Galeus cadenati Springer, 1966
* Gecko catshark, Galeus eastmani (Jordan & Snyder, 1904)
* Slender sawtail catshark, Galeus gracilis Compagno & Stevens, 1993
* Longnose sawtail catshark, Galeus longirostris Tachikawa & Taniuchi, 1987
* Blackmouth catshark, Galeus melastomus Rafinesque, 1810
* Southern sawtail catshark, Galeus mincaronei Soto, 2001
* Mouse catshark, Galeus murinus (Collett, 1904)
* Broadfin sawtail catshark, Galeus nipponensis Nakaya, 1975
* Peppered catshark, Galeus piperatus Springer & Wagner, 1966
* African sawtail catshark, Galeus polli Cadenat, 1959
* Galeus priapus Séret & Last, 2008
* Blacktip sawtail catshark, Galeus sauteri (Jordan & Richardson, 1909)
* Dwarf sawtail catshark, Galeus schultzi Springer, 1979
* Springer's sawtail catshark, Galeus springeri Konstantinou & Cozzi, 1998


Phylogeny and evolution

Most taxonomic studies have concluded the closest relatives of Galeus to be Apristurus, Asymbolus, Parmaturus, and/or Cephalurus.[8] Leonard Compagno has placed Galeus with Apristurus, Bythaelurus, Cephalurus, Parmaturus, and Pentanchus in the tribe Pentanchini of the subfamily Pentanchinae, based on morphological characters.[9] Galeus was suggested to be the sister group of Apristurus in a 2005 phylogenetic study based on mitochondrial and nuclear DNA gene sequences, by Samuel Iglésias and colleagues.[8] The affinity between Galeus and Apristurus was also upheld, albeit weakly, in a 2006 phylogenetic analysis based on three mitochondrial DNA genes, by Brett Human and colleagues.[9] Within the genus, though the G. arae species complex, G. atlanticus, G. eastmani, G. melastomus, G. piperatus, G. polli, and G. sauteri definitely form a monophyletic group, the assignment of other species (e.g. G. murinus) within the genus is more problematic.[3] In the aforementioned study by Iglésias and colleagues, which included five Galeus species, G. eastmani, G. gracilis, and G. sauteri were grouped into one clade and G. melastomus and G. murinus grouped into another.[8] Galeus fossils, dating to the Burdigalian (20.43–15.97 Ma) and Langhian (15.97–13.65 Ma) stages of the early Miocene, have been recovered from France.[1]

Distribution and habitat
A roughtail catshark (G. arae) in its natural habitat; members of this genus occur close to the bottom in deep water.

The centers of biodiversity for Galeus are the North Atlantic (8 species) and the northwestern Pacific (4 species). A few outlying species are found in the South Atlantic (G. mincaronei and G. polli), Oceania (G. gracilis and G. priapus), and the Gulf of California (G. piperatus). In the western Indian Ocean, this genus appears to be replaced by the ecologically similar genus Holohalaelurus.[3][10][11] Sawtail catsharks are demersal in habits and occur in deep water over outer continental and insular shelves and upper slopes.[10][12]

Description


Sawtail catsharks reach maximum lengths of between 25 and 90 cm (9.8 and 35 in). They have slender, firm bodies and narrow, slightly flattened heads with short to long, pointed snouts. The nostrils are divided into incurrent and excurrent openings by triangular flaps of skin on their anterior rims. The horizontally oval eyes are placed mostly on the sides of the head and equipped with rudimentary nictitating membranes (protective third eyelids); below each eye is a subtle ridge, and behind is a small spiracle (auxiliary respiratory opening). The mouth is rather large and wide, and when closed the upper teeth are exposed. There are short to long furrows around the corners of the jaws. The teeth are small and number 47–78 rows in the upper jaw and 48–82 rows in the lower jaw; each tooth has a narrow central cusp flanked by one or more smaller cusplets on either side. There are five pairs of gill slits.[3][12]
Close-up from above of the caudal fin denticle crest of the longfin sawtail catshark (G. cadenati); such crests are shared by all Galeus species.

The two dorsal fins are nearly equal in size and shape, and are placed well back on the body, behind the origins of the pelvic fins. The pectoral fins are fairly large and broad, with angular to rounded corners. The pelvic fins are much smaller, and bear claspers in males; in G. murinus and G. nipponensis, the pelvic fin inner margins are partially fused to form an "apron" over the base of the claspers. The anal fin is elongated and much larger than the pelvic and dorsal fins; its position relative to the pelvic and caudal fins varies from very close to well-spaced. The caudal peduncle can be nearly cylindrical to laterally compressed, depending on species. The caudal fin comprises more or less a quarter of the total length, and is low with a small lower lobe and a ventral notch near the tip of the upper lobe.[3][12][13]

The skin is thick and densely covered by small, overlapping, well-calcified dermal denticles; each denticle has a leaf-shaped crown with a horizontal ridge and three teeth on the posterior margin. There is a prominent, saw-like crest of enlarged denticles along the dorsal margin of the caudal fin. G. murinus and G. springeri also have a similar crest along the ventral margin of the caudal fin. Galeus species are typically grayish or brownish above and lighter below, and most have a pattern of darker saddles and/or blotches along the back and tail. The interior of the mouth may be light or dark.[3][12][14]

Biology and ecology

Natural history data is scant for most Galeus species. They feed on various types of invertebrates and fishes on or near the bottom.[10] In Suruga Bay, the dietary compositions of G. eastmani and juvenile G. nipponensis differ significantly, suggesting there is reduced interspecific competition between co-occurring Galeus species.[15] Reproductive modes within the genus are notably diverse: while most species are oviparous and lay encapsulated eggs on the sea floor, there is a single aplacental viviparous species (G. polli) that retains eggs internally and gives live birth. Among the oviparous species, most (e.g. G. murinus, G. nipponensis) exhibit single oviparity, in which only a single egg matures within each of the female's oviducts at a time. In contrast, a few species such as G. atlanticus and G. melastomus exhibit multiple oviparity, in which several eggs can mature within each oviduct simultaneously. Single oviparity is considered to be the basal condition, while multiple oviparity and aplacental viviparity are thought to be more derived.[16]

Human interactions

Sawtail catsharks pose no danger to humans and have little economic value, though varying numbers are caught incidentally by deepwater commercial fisheries. Some of the larger species, such as G. melastomus and G. polli, are occasionally utilized for meat, fishmeal, and/or leather.[12] The International Union for Conservation of Nature (IUCN) has listed G. atlanticus and G. mincaronei, both of which have very restricted distributions, as Near Threatened and Vulnerable respectively.[17][18]

References

1. ^ a b Marsili, S. (2007). "A new bathyal shark fauna from the Pleistocene sediments of Fiumefreddo (Sicily, Italy)". Geodiversitas 29 (2): 229–247. http://www.mnhn.fr/museum/front/medias/publication/10535_g07n2a1.pdf.
2. ^ Rafinesque, C.S. (1810). Caratteri di alcuni nuovi generi e nuove specie di animali e piante della Sicilia, con varie osservazioni sopra i medisimi. (Part 1 involves fishes, pp. [i-iv] 3-69, Part 2 with slightly different title, pp. ia-iva + 71-105). Pls. 1-20.
3. ^ a b c d e f g h i Compagno, L.J.V. (1988). Sharks of the Order Carcharhiniformes. Blackburn Press. pp. 134–142. ISBN 1930665768.
4. ^ "Characters of various new genera and species of animals (mostly fishes) and plants, with various observations about them"
5. ^ Fowler, H.W. (1908). "Notes on sharks". Proceedings of the Academy of Natural Sciences, Philadelphia 60: 52–70.
6. ^ Bigelow, H.W. and W.C. Schroeder. "Sharks". Memoirs of the Sears Foundation of Marine Research 1: 53–576.
7. ^ Orkin, P.A. (December 1952). "Galeus Rafinesque, 1810 (Chondrichthyes, Triakidae), an invalid generic name". Annals and Magazine of Natural History 5 (60): 1112.
8. ^ a b c Iglésias, S.P., G. Lecointre and D.Y. Sellos (2005). "Extensive paraphylies within sharks of the order Carcharhiniformes inferred from nuclear and mitochondrial genes". Molecular Phylogenetics and Evolution 34 (3): 569–583. doi:10.1016/j.ympev.2004.10.022. PMID 15683930. http://linkinghub.elsevier.com/retrieve/pii/S1055790304003409.
9. ^ a b Human, B.A., E.P. Owen, L.J.V. Compagno and E.H. Harley (May 2006). "Testing morphologically based phylogenetic theories within the cartilaginous fishes with molecular data, with special reference to the catshark family (Chondrichthyes; Scyliorhinidae) and the interrelationships within them". Molecular Phylogenetics and Evolution 39 (2): 384–391. doi:10.1016/j.ympev.2005.09.009. PMID 16293425. http://linkinghub.elsevier.com/retrieve/pii/S1055790305002964.
10. ^ a b c Compagno, L.J.V., M. Dando and S. Fowler (2005). Sharks of the World. Princeton University Press. pp. 223–232. ISBN 9780691120720.
11. ^ Séret, B. and P.R. Last (June 30, 2008). "Galeus priapus sp. nov., a new species of sawtail catsharks (Charcharhiniformes: Scyliorhinidae) from New Caledonia". Zootaxa 1813: 19–28.
12. ^ a b c d e Compagno, L.J.V. (1984). Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. Food and Agricultural Organization. pp. 306–318. ISBN 9251013845.
13. ^ Nakaya, K. (1975). "Taxonomy, comparative anatomy and phylogeny of Japanese catsharks, Scyliorhinidae". Memoirs of the Faculty of Fisheries, Hokkaido University 23: 1–94.
14. ^ Konstantinou, H. and J.R. Cozzi (1998). "Galeus springeri, a new species of sawtail catshark from the Caribbean Sea (Chondrichthys, Scyliorhinidae)". Copeia 1998 (1): 151–158. doi:10.2307/1447711. http://jstor.org/stable/1447711.
15. ^ Horie, T. and S. Tanaka (2000). "Reproduction and food habits of two species of sawtail catsharks, Galeus eastmani and G. nipponensis, in Suruga Bay, Japan". Fisheries Science 6: 812–825.
16. ^ Iglésias, S.P., M.H. du Buit, and K. Nakaya (2002). "Egg capsules of deep-sea catsharks from eastern North Atlantic, with first descriptions of the capsule of Galeus murinus and Apristurus aphyodes (Chondrichthyes: Scyliorhinidae)". Cybium 26 (1): 59–63.
17. ^ Coelho, R., J. Rey, F. Serena, and C. Mancusi (2007). Galeus atlanticus. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. Downloaded on October 29, 2010.
18. ^ Rincon, G. (2004). Galeus mincaronei. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. Downloaded on October 29, 2010.

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