Occurrence of the Smallscale Codlet, Bregmaceros nectabanus in the Mediterranean Sea, previously misidentified as B. atlanticus in this region
© The Author(s) 2016
Received: 23 June 2016
Accepted: 24 June 2016
Published: 1 August 2016
We report the occurrence of Bregmaceros nectabanus as the only species of this genus occurring in the Mediterranean Sea.
All previously published records of B. atlanticus from the Mediterranean are based on misidentifications. Among the features that distinguish B. nectabanus are its nearly unpigmented abdomen, presence of a thin dorsolateral longitudinal stripe below the second dorsal fin, and a distally fimbriate opercular spine.
Since this species is known also from the Red Sea it is evidently a Lessepsian migrant that reached the Mediterranean via the Suez Canal.
The Bregmacerotidae or codlets, are small elongated fishes with a maximum length of 12 cm. Members of this family are characterized by two dorsal fins, the first being a long single ray on the back of the head, and the second dorsal fin being long-based with rudimentary rays in the middle portion of the fin. The anal fin is very similar in form to the second dorsal and is situated directly below it. The family has been reviewed by D’Ancona & Cavinato (1965) and Belyanina (1974) and contains the single genus, Bregmaceros Thompson, 1840, containing 14 valid species (Eschmeyer, 2015). Codlets occur in tropical and subtropical neritic environments, with habitats ranging from mesopelagic to inshore and even estuaries. The state of our knowledge of the biology and geographic distribution of these fishes is not particularly good, even though they may be commonly encountered in midwater tows through much of their range. Even in the Mediterranean Sea, it is not well known which species of Bregmaceros occur there. D’Ancona & Cavinato (1965) reported B. atlanticus Goode & Bean, 1886 from the Mediterranean based on a single questionable record, and to date that is the only species reported from the region (e.g., Turan et al., 2011; Aydin & Akyol, 2013).
In this paper we conclude that the previous records of B. atlanticus from the Mediterranean Sea were misidentifications of B. nectabanus Whitley, 1941, the only member of this genus we confirm that occurs there. We provide a brief account here of B. nectabanus, including comparisons with material from the Indo-Pacific (the region including the type locality of B. nectabanus). We also provide a discussion of the occurrence of the species in the Mediterranean Sea possibly through dispersal from the Indo-Pacific via the Suez Canal and the Red Sea.
Materials and methods
Morphometric, meristic and other characters were determined following the methods and terminology of Hubbs & Lagler (1958), with modifications by D’Ancona & Cavinato (1965) and Torii et al. (2003a, b). Counts of rays for the second dorsal fin, anal fin, pectoral fin, and pelvic fin are the total number of rays. Number of rays in the caudal fin are reported as principal caudal fin rays or total caudal fin rays (including segmented and procurrent rays). Fin rays and counts of other serially repeating structures were determined from alcohol preserved specimens or from radiographic images.
Morphological characters and associated terms: D1, occipital first dorsal fin; D2, second (main) dorsal fin; A, anal fin; Pc, pectoral fin; Pv, pelvic fin; longitudinal scale count is the number of scales in a midlateral longitudinal series from the pectoral girdle to the posterior termination of the vertebral column (posterior margin of hypural plate); transverse scale count is the number of horizontal scale rows between the origins of the second dorsal fin and the anal fin; predorsal scale count is the number of lateral line scales along the dorsum (anterior, elevated portion of the lateral line) between the occiput and the origin of the second dorsal fin.
Morphometric characters were measured point to point with a Mitutoyo digital caliper. Names and descriptions of morphometric characters are as follows: Standard length (SL), distance (mm) from snout (most anterior location on premaxilla) to posterior termination of hypural plate; Preanal length, distance from snout to anal fin origin; Predorsal length, distance from snout to origin of second dorsal fin; Body depth, distance from second dorsal fin origin to anal fin origin; Head length (HL), maximum distance between snout and bony posterior margin of gill cover; Snout length, minimum distance between snout and anterior bony margin of orbit; Eye diameter, maximum horizontal distance across the eye; Interorbital width, minimum distance across cranium between orbits; Values of morphometric characters are reported as %SL, except those taken from within the head (i.e., Snout length, Eye diameter, Interorbital width) which are %HL.
Infraclass teleostei Müller, 1845
Order gadiformes Goodrich, 1909
Family bregmacerotidae Gill, 1872
Genus Bregmaceros Thompson, 1840
Bregmaceros nectabanus Whitley, 1941(Fig. 1, Table 1)
Meristic and morphometric character values for Bregmaceros nectabanus from the Mediterranean Sea (HUJ 19751 and 19925) compared with Indo-Pacific material, including the holotype
Indian Ocean (Kotthaus, 1969)
Toyama Bay, Japan
D2 fin rays
A fin rays
P fin rays
Principal caudal fin rays
Total caudal fin rays
72– ~ 80
Head length %SL
Predorsal length %SL
Preanal length %SL
Body depth %SL
Snout length %HL
Eye diameter %HL
Interorbital width %HL
Bregmaceros nectabanus Whitley, 1941: 25, Figure 18 (type locality Darwin, Northern Territory, Australia [Arafura Sea, eastern Indian Ocean]). Munro, 1950: 41 (northwestern Australia); Shen, 1960: 68 (northern Australia); D’Ancona & Cavinato, 1965: 47 (Indian Ocean; central and western Pacific); Kotthaus, 1969: 39 (northwestern Indian Ocean; Red Sea); Belyanina, 1974: 11 [English translation] (Indian Ocean; western and central Pacific); Masuda & Ozawa, 1979 (type locality off Darwin, Northern Territory, Australia; Japan): 267; Dor, 1984: 58 (Red Sea); Okamura, 1984: 92 (Indian Ocean; western and southern Pacific); Masuda et al., 1986: 393 (type locality off Darwin, northern Australia; Japan); Smith, 1986: 330 (Indo-West Pacific); Paxton & Hanley, 1989: 306 (northern Australia); Rivaton, 1989: 147–148 (New Caledonia); Cohen, 1990: 525 (Indian Ocean; tropical western Pacific to Fiji); Mamhot et al., 1992: 50–53 (northern Australia; Japan); Goren & Dor, 1994: 15 (Red Sea); Iwamoto, 1999: 1997 (western central Pacific); Nakabo 2000: 415 (off Japan); Hutchins, 2001: 23 (off northwestern Australia); Nakabo, 2002: 415 (off Japan); Torii et al., 2003a: 30 (northwestern Australia); Torii et al., 2003b: 138 (northern Australia; Japan); Paxton et al., 2006: 622 (Indian Ocean; western Pacific); Shao et al., 2008: 242 (South China Sea off Taiwan; northwestern Australia); Fricke et al., 2011: 362 (New Caledonia); Shinohara et al., 2011: 41 (Sea of Japan; northern Australia); Larson et al., 2013: 52 (northern Australia); Shinohara et al., 2014: 242 (Sea of Japan; northern Australia); Bogorodsky et al., 2014: 414 (Red Sea; Indo-West Pacific).
Bregmaceros atlanticus (non Goode & Bean, 1886). Goren & Galil, 2006: 1 (off Palmahim, eastern Mediterranean Sea); Filiz et al., 2007: 108 (Aegean Sea); Turan et al., 2011: 189 (eastern Mediterranean Sea); Aydin & Akyol, 2013: 245 (Izmir Bay, northern Aegean Sea).
Type material examined
Holotype: AMS IA.1719 (30.8 mm), Darwin, Northern Territory, Australia, 12°27′S 130°50′E, 1923, digital image and radiograph only.
Non-type material examined
BMNH unreg. (9, 29.9–38.9 mm SL), R/V Discovery Sta. 54009 #1, Indian Ocean. BMNH unreg. (8, 25.3–45.1 mm SL), R/V Discovery Sta. 54009 #2, Indian Ocean. CAS 81511 (2, 25.8–52.3 mm), R/V Stranger Sta. 60–451, Gulf of Thailand, 13°21′45“N 100°32′43”E, 14 m, 14 Dec 1960. CAS uncat. (1, 34.0 mm), R/V Stranger Sta. 60–449, Gulf of Thailand, 12°19′15“N 100°43′40”E, 33 m, 13 Dec 1960. HUJ 19751 (6, 51.9–63.6 mm), off Israel. HUJ 19925 (4, 52.2–66.9 mm), off Israel. HUJ 19928 (1, 61.0 mm), trawled between Mikhmoret and Ga’ash off Israel.
Comparative material examined
Bregmaceros atlanticus Goode & Bean, 1886. Holotype, MCZ 32323 (44.1 mm), Caribbean Sea and Straits of Florida, USCSS Blake Exped., Stas. 045, 150 and 262 mixed, depth 90–500 fathoms (165–914 m), 1877. Non-types, Bregmaceros atlanticus, USNM 325140 (2, 66.0–71.3 mm), R/V Oregon Sta. 4417, off Venezuela, Caribbean Sea, 0–439 m, 04 Oct 1963.
A species of Bregmaceros with 47–55 dorsal and 47–57 anal fin rays, 15–19 pectoral fin rays, 13 principal caudal fin rays, 69–80 longitudinal scales, 15–17 transverse scales, unpigmented (or nearly so) abdomen, and a distinctive thin brown dorsolateral longitudinal stripe below the second dorsal fin. Opercular spine distally fimbriate.
Bregmaceros contains 14 valid nominal species (Eschmeyer, 2015) and at least four undescribed species (Harold, unpublished). Of the valid nominal species, B. nectabanus is distinguished from B. atlanticus, B. cayorum Nichols, 1952 , B. houdei Saksena & Richards, 1986 , B. japonicus Tanaka, 1908 , B. neonectabanus Masuda et al., 1986 , B. bathymaster Jordan & Bollman, 1890 , B. rarisquamosus Munro, 1950, and B. cantori Milliken & Houde, 1984, based on the presence of a distally fimbriate opercular spine in B. nectabanus, compared with a spine with a distally unornamented point in the other species (see Torii et al., 2003a, b). Species that share the presence of a fimbriate opercular spine with B. nectabanus are B. arabicus D’Ancona & Cavinato, 1965 , B. pescadorus Shen, 1960 , B. mcclellandi Thompson, 1840 , B. lanceolatus Shen, 1960, and B. pseudolanceolatus Torii et al., 2004. Of these species, B. nectabanus, B. arabicus, B. pescadorus, and B. mcclellandi all have an indented caudal fin posterior margin as compared with the two species, B. lanceolatus and B. pseudolanceolatus, with a lanceolate caudal fin. Bregmaceros nectabanus has a distinctive thin brown dorsolateral longitudinal stripe below the second dorsal fin, as compared with multiple longitudinal rows of minute spots in B. arabicus and B. pescadorus, or a darkly pigmented dorsum but lacking any longitudinal pigment stripes or rows of dots in B. mcclellandi. With respect to overall similarity, B. nectabanus is closest to B. neonectabanus, but the former differs in its lack of pigmentation on the abdomen (compared with the presence of dark pigment in that area), and the presence of 13 principal caudal fin rays in B. nectabanus, as compared with 14.
The only species of Bregmaceros to have been reported from the Mediterranean Sea to date is B. atlanticus (D’Ancona & Cavinato, 1965, and others cited above). Based on the original description and our examination of the type and other non-type specimens from the Atlantic ocean, B. atlanticus is very darkly pigmented over almost the entire body, has a relatively elongate body (body depth 12.4–14.4 % SL), and a simple opercular spine termination. On the other hand, B. nectabanus has the distinctive body pigmentation pattern including a thin dorsolateral stripe, a typically greater body depth (13.2–17.5 % SL in the B. nectabanus material reported herein from the Mediterranean, the Indian Ocean, and the type locality) and a fimbriate opercular spine termination. The minumum value of body depth for B. nectabanus from off Japan was reported by Masuda et al. (1986) to be 10.8 % SL (Table 1), a rather low value for the species, but we are unable to verify it.
The status of Bregmaceros nectabanus and associated type material was considered in detail by Masuda et al. (1986). The species had previously been known to have a broader distribution, including the southeastern Atlantic, the Indian and the western Pacific regions. Masuda et al. (1986) restricted the use of B. nectabanus and designated a new species, B. neonectabanus, based on a number of meristic and pigmentation characters. The summary of the distribution of B. nectabanus provided by Eschmeyer (2015) includes the Red Sea, the Indo-West Pacific region and the Atlantic, although Masuda et al. (1986: 392) excluded the Atlantic material and reassigned it to B. neonectabanus. Occurrences of B. nectabanus in the western Indian Ocean and the Red Sea are few (Kotthaus, 1969; Dor, 1984) but we have been able to verify the identity of material from the area. Other relevant material from the Red Sea reported by Bogorodsky et al. (2014) is similar to B. nectabanus, but may not be conspecific.
The first record of a member of the Bregmacerotidae from the Mediterranean was reported by D’Ancona & Cavinato (1965) off the coast of southern Italy, based on a questionable identification of a specimen identified as Bregmaceros atlanticus. It is not clear at all from the account of the species provided by D’Ancona & Cavinato (1965) what the source of the record was and, furthermore, they may not have examined the specimen. However, others have followed, indicating the questionable status of that record (Cohen, 1986; Goren & Galil, 2006). These authors were also unable to examine the specimen, nor is the specimen explicitly referred to or listed as material examined in D’Ancona & Cavinato (1965). Since the source of this initial record of B. atlanticus from the Mediterranean is unknown, and neither specimens nor descriptive data are known, we treat it as Bregmaceros sp. only. We also find that their account of B. atlanticus takes a rather broad view of variation in the species and may include representatives of at least one other undescribed species. It has been shown that D’Ancona & Cavinato (1965) often employed a rather expansive interpretation of intraspecific variation (Torii et al., 2003b). The first record of B. atlanticus from the Mediterranean for which specimens and data were reported was Yilmaz et al. (2004), who reported two specimens from the stomach contents of the Lessepsian (Red Sea) migrant, the Brushtooth Lizardfish Saurida undosquamis, in Antalya Bay, Turkey. Although the counts of fin rays for the second dorsal and anal fins fall within the range of B. atlanticus, the pattern of pigmentation is not consistent with published descriptions (“silver-grey on the belly with dense pigmentation along the dorsum”), as compared with darkly pigmented over the entire body and fins (D’Ancona & Cavinato, 1965; Belyanina, 1974). About 2 years later, Goren & Galil (2006) reported four specimens of B. atlanticus from a trawl catch from a depth of about 35 m off the coast of Israel. The records of Filiz et al. (2007) extended the range of B. atlanticus in the Mediterranean to the eastern Aegean Sea, and later to the northern region of that sea (Aydin & Akyol, 2013). Turan et al. (2011) reported the species from a depth of 120 m in Iskenderun Bay at the northeastern corner of the Mediterranean. However, based on the specimen data and photographs published in the foregoing accounts these records represent occurrences of B. nectabanus in the Mediterranean, not B. atlanticus. Apparently, B. nectabanus has established a sustainable population along the Israeli coast with dozens of specimens being collected, mainly between depths of 30 and 50 m, and why it was not collected in the eastern Mediterranean prior to D’Ancona & Cavinato’s (1965) report is unknown. It is possible it invaded the area or increased in population size significantly since the time of their report.
Regarding its distribution in the Atlantic, according to the revision of Masuda et al. (1986), B. nectabanus is not known to occur there, thus excluding it as a possible source of individuals dispersing into the Mediterranean Sea. We report here the first occurrence of B. nectabanus from the Mediterrnean Sea. Previous reports of B. atlanticus from the Mediterranean cannot be substantiated and the former may be the only species of the genus, and indeed the family Bregmacerotidae, to occur there.
Three species of Bregmaceros have been mentioned in the literature as possibly occurring in the Red Sea: B. nectabanus, B. arabicus, according to Aron & Goodyear (1969), and B. mcclellandi, however, according to D’Ancona & Cavinato (1965) the latter’s occurrence there was most likely a result of misidentification of B. arabicus (see Dor, 1984). Bregmaceros mcclellandi, in the restricted sense of Torii et al. (2003b), and as supported by observations of the first author, has not been recorded from the western Indian Ocean or the Red Sea. As for B. nectabanus, this species was reported by Kotthaus (1969) and Bogorodsky et al. (2014) from the Red Sea and by Belyanina (1974) from the Gulf of Oman at the southern entrance of the Red Sea. Our material from the Mediterranean is clearly conspecific with B. nectabanus, with little variation from specimens reported from across the range of the species (Table 1; Kotthaus, 1969; Masuda et al. 1986).
Goren & Galil (2006) wondered as to the source of Bregmaceros species in the Mediterranean. Given that numerous fish collections in the region over many years did not reveal Bregmaceros species, it seems likely that the population became established there following dispersal from elsewhere. The opening of the Suez Canal in 1869 resulted in the influx of over 600 species from the Red Sea into the Mediterranean (Galil, 2009; Zenetos, 2010), among these were 97 migrant fish species (Fricke et al., in press). Such species have been referred to as Lessepsian migrants (see Golani, 2010). Goren & Galil (2006) raised the possibility that Bregmaceros species entered the Mediterranean in ballast water or through dispersal via the Suez Canal. The present findings and the inferred rapid population growth, which is typical of Lessepsian migrant fish species (Golani, 2010), indicate that B. nectabanus most likely entered from the Red Sea via the Suez Canal. Kotthaus (1969) reported this species from the extreme southern end of the Red Sea as well as the Gulf of Oman and the Arabian Sea; additional records tentatively referred to this species from the Red Sea have been reported (Bogorodsky et al., 2014). There are no substantiated records of B. nectabanus from the Atlantic, thus excluding it as a possible source. Bregmaceros nectabanus had been recorded from the South Atlantic (D’Ancona & Cavinato, 1965) and it is widely reported to occur there but Masuda et al. (1986) reassigned those records to B. neonectabanus.
We have reviewed published records of Bregmaceros atlanticus from the Mediterranean Sea and examined type and non-type specimens of this and other species belonging to this genus. Morphometric and meristic data, in addition to qualitative observations were used as the basis of interspecific comparisons. We have determined that all previously published records of B. atlanticus from the Mediterranean Sea, the first being D’Ancona and Cavinato (1965), are based on misidentifications, and that B. nectabanus is the correct identity of all such verifiable records. Therefore, this is the first record of B. nectabanus from the Mediterranean and it is the only species of Bregmaceros known to occur in the region. Bregmaceros nectabanus was originally described from off Australia and is known to have a natural range including much of the Indo-Pacific region but not the Atlantic. We conclude that the species now occurs in the Mediterranean Sea as a result of dispersal from the Red Sea via the Suez Canal (Lessepsian migrant).
Material examined is listed by institutional acronym and catalog number or vessel and collecting station number; number of specimens examined and the size range in standard length of those specimens from which data were taken; institutional abbreviations are as follows:
AMS, (Australian Museum, Sydney); BMNH, (The Natural History Museum, London); CAS, (California Academy of Sciences); HUJ, (Hebrew University of Jerusalem); KAUMM, (King Abdulaziz University Marine Museum, Jeddah, Saudi Arabia); MCZ, (Museum of Comparative Zoology, Harvard University); SMF, (Senckenberg Forschungsinstitut und Naturmuseum); USNM, (National Museum of Natural History, Smithsonian Institution) (abbreviations as listed in Sabaj Perez, 2014).
We thank Dr. H. Yakovlev-Golani for the translation from Russian, Dr. D. Darom for the photograph, M. McGrouther (AMS), N.E. Merrit and O. Crimmen (BMNH), D. Catania and T. Iwamoto (CAS), Alistair Graham (CSIRO), K.E. Hartel (MCZ), H.J. Walker and C. Klepadlo (SIO), J. Williams (USNM) for loans of specimens, D.A. Baltzegar, S.K. Shore and R. Simpson for assistance in morphometric data acquisition, and the Grice Marine Laboratory, College of Charleston for space, funding and other resources.
This research was supported by Department of Biology, College of Charleston, Charleston, South Carolina, research and development funds provided to the corresponding author.
Availability of data and materials
Data available from the corresponding author, upon request. Material available through institutional loan, subject to approval by host institution.
The authors contributed approximately equally to all aspects of this paper. Both authors read and approved the final manuscript.
The authors declare that they have no competing interests.
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- Aron W, Goodyear WH. Fishes collected during a midwater trawling survey of the Gulf of Elat and the Red Sea. Isr J Zool. 1969;18:237–44.Google Scholar
- Aydin I, Akyol O. New record of the antenna codlet, Bregmaceros atlanticus Goode and Bean, 1886 (Gadiformes: Bregmacerotidae), from the northern Aegean Sea (Izmir Bay, Turkey). J Appl Ichthyol. 2013;29:245–6.View ArticleGoogle Scholar
- Belyanina T.N. (1974) Material on the development, systematics and distribution of fishes of the family Bregmacerotidae. Trudy Instituta Okeanologii 96, 143–188 [in Russian; English translation].Google Scholar
- Bogorodsky SV, Alpermann TJ, Mal AO, Gabr MH. Survey of demersal fishes from southern Saudi Arabia, with five new records for the Red Sea. Zootaxa. 2014;3852:401–37.View ArticleGoogle Scholar
- Cohen DM. Bregmacerotidae. In: Whitehead PJP, Bauchot ML, Hureau JC, Nielsen JG, Tortonese E, editors. Fishes of the North-eastern Atlantic and the Mediterranean. Paris: UNESCO; 1986. p. 711–2.Google Scholar
- Cohen DM. Bregmacerotidae. In: Quéro J-C, Hureau J-C, Karrer C, Post A, Saldanha L, editors. Check-list of the fishes of the eastern tropical Atlantic. CLOFETA. Paris: UNESCO; 1990. p. 16–7.Google Scholar
- D’Ancona U, Cavinato G. The fishes of the family Bregmacerotidae. 1965. Dana Report, no. 64, 91 pp.Google Scholar
- Dor M. (1984) Checklist of the fishes of the Red Sea. CLOFRES. Jerusalem: The Israel Academy of Sciences and Humanities, i–xxii, map + 437 pp.Google Scholar
- Eschmeyer W.N. (ed) (2015) Catalog of Fishes, electronic version. San Francisco (California Academy of Sciences). Available at: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. Accessed 09 Jan 2015.
- Filiz H, Akçinar SC, Ulutürk E, Bayhan B, Taşkavak E, Sever TM, et al. New records of Bregmaceros atlanticus (Bregmacerotidae), Echiodon dentatus (Carapidae) and Nemichthys scolopaceus (Nemichthyidae) from the Aegean Sea. Acta Ichthyol Piscat. 2007;37:107–12.Google Scholar
- Fricke, R., Golani, D. and Appelbaum-Golani, B. (in press). First record of the Indian anchovy Stolephorus indicus (Hasselt, 1823) in the Mediterranean Sea (Clupeiformes: Engraulidae). BioInvasions Records.Google Scholar
- Fricke R, Kulbicki M, Wantiez L. Checklist of the fishes of New Caledonia, and their distribution in the Southwest Pacific Ocean (Pisces). Stuttgarter Beiträge zur Naturkunde A, Neue Serie. 2011;4:341–463.Google Scholar
- Galil BS. Taking stock: inventory of alien species in the Mediterranean Sea. Biol Invasions. 2009;11:359–72.View ArticleGoogle Scholar
- Golani D. Colonization of the Mediterranean by Red Sea fishes via the Suez Canal - Lessepsian migration. In: Golani D, Appelbaum-Golani B, editors. Fish invasions of the Mediterranean Sea: change and renewal. Sofia: Pensoft; 2010. p. 145–88.Google Scholar
- Golani D, Bogorodsky SV. The fishes of the Red Sea – reappraisal and updated checklist. Zootaxa. 2010;2463:1–135.Google Scholar
- Goode GB, Bean TH. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877–78) and in the Caribbean Sea (1879–80), by the U.S. Coast Survey Steamer “Blake”, Lieut.-Commander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. XXVIII. Description of thirteen species and two genera of fishes from the Blake collection. Bull Mus Comp Zool Harvard College. 1886;12:153–70.Google Scholar
- Goren M, Dor M. An updated checklist of the fishes of the Red Sea, CLOFRES II. Jerusalem: The Israel Academy of Sciences and Humanities; 1994. 120 pp.Google Scholar
- Goren M, Galil BS. Additional records of Bregmaceros atlanticus in the eastern Mediterranean - an invasion through the Suez Canal or ballast water? Mar Biodivers Rec. 2006;1(e42):1–3.Google Scholar
- Hubbs CL, Lagler KF. Fishes of the Great Lakes region. Ann Arbor, Michigan: University of Michigan Press; 1958. 213 pp.Google Scholar
- Hutchins J.B. (2001) Checklist of the fishes of Western Australia. Records of the Western Australian Museum, Supplement, no. 63, 9–50.Google Scholar
- Iwamoto T. Bregmacerotidae. In: Carpenter KE, Niem VH, editors. Species identification guide for fisheries purposes. The living marine resources of the western central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae). Rome: FAO; 1999. p. 1997–8.Google Scholar
- Jordan D.S. and Bollman C.H. (1890) Scientific results of explorations by the U.S. Fish Commission Steamer Albatross, No. IV. Descriptions of new species of fishes collected at the Galapagos Islands and along the coast of the United States of Colombia: 1887–88. Proceedings of the United States National Museum 12 (no. 770), 149–183.Google Scholar
- Kotthaus A. Fisches des Indischen Ozean. Ergebnisse der ichthyologiiscen Untersuchungen während der Expedition des Forschungsschiffes “Meteor” in den Indischen Ozeean, Oktober 1964 bis Mai 1965. A. Systematischer Teil V: Solenichthyes und Anacanthini (1). Meteor Forschungsergebnisse Reihe D, Biologie. 1969;no. 4:35–46.Google Scholar
- Larson HK, Williams RS, Hammer MP. An annotated checklist of the fishes of the Northern Territory, Australia. Zootaxa. 2013;3696:1–293.View ArticleGoogle Scholar
- Mamhot JR, Ozawa T, Masuda Y. Occurrence and abundance of bregmacerotid larvae in Kagoshima Bay, southern Japan, with descriptions of ontogenetic larval characters. Jpn J Ichthyol. 1992;39:49–58.Google Scholar
- Masuda S, Ozawa T. Reexamination of the holotypes of Bregmaceros japonicus Tanaka and B. nectabanus Whitley. Jpn J Ichthyol. 1979;25:266–8.Google Scholar
- Masuda S, Ozawa T, Tabeta O. Bregmaceros neonectabanus, a new species of the family Bregmacerotidae, Gadiformes. Jpn J Ichthyol. 1986;32:392–9.Google Scholar
- Milliken DM, Houde ED. A new species of Bregmacerotidae (Pisces), Bregmaceros cantori, from the western Atlantic ocean. Bull Mar Sci. 1984;35:11–9.Google Scholar
- Munro ISR. Revision of Bregmaceros with descriptions of larval stages from Australasia. Proc R Soc Queensl. 1950;61:37–53.Google Scholar
- Nakabo T. (ed) (2000) Fishes of Japan with pictorial keys to the species. 2nd edition. Tokyo: Tokai University Press, vol. 1, i–lvi + 866 pp. [In Japanese.]Google Scholar
- Nakabo T. (ed) (2002) Fishes of Japan with pictorial keys to the species. English edition. Tokyo: Tokai University Press. vol. 1: i–lvi + 1–866.Google Scholar
- Nichols JT. A new fish of the genus Bregmaceros from the Straits of Florida. Am Mus Novit. 1952;1556:1–3.Google Scholar
- Okamura O. Order Gadiformes. In: Masuda H, Amaoka K, Araga C, Uyeno T, Yoshino T, editors. The fishes of the Japanese Archipelago. Tokyo: Tokai University Press; 1984. p. 92, pl. 344C.Google Scholar
- Paxton JR, Hanley JE. Bregmacerotidae (225). In: Paxton JR, Hoese DF, Allen GR, Hanley JE, editors. Zoological catalogue of Australia, vol. 7. Canberra: Australian Government Publishing Service; 1989. p. 305–6.Google Scholar
- Paxton J.R., Gates J.E. and Bray D.J. (2006) Zoological catalogue of Australia, fishes. Canberra: CSIRO Publishing, vol. 35, parts 1–3, pp. 2248.Google Scholar
- Rivaton J. Premières observations sur la faune ichtyologique des Iles Chesterfield (Mer du Corail). Cybium. 1989;13:139–64.Google Scholar
- Sabaj Perez M.H. (editor) (2014) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 5.0 (22 September 2014). Electronically accessible at http://www.asih.org/, American Society of Ichthyologists and Herpetologists, Washington, DC.
- Saksena VP, Richards WJ. A new species of gadiform fish, Bregmaceros houdei, from the western north Atlantic. Bull Mar Sci. 1986;38:285–92.Google Scholar
- Shao K-T, Ho H-C, Lin P-L, Lee P-F, Lee M-Y, Tsai C-Y, et al. A checklist of the fishes of southern Taiwan, northern South China Sea. Raff Bull Zool Suppl. 2008;no. 19:233–71.Google Scholar
- Shen S-C. Bregmaceros lanceolatus and Bregmaceros pescadorus, two new species of dwarf fishes from southern Taiwan and Pescadore Islands. Quart J Taiwan Mus (Taipei). 1960;13:67–74.Google Scholar
- Shinohara G, Nakae M, Ueda Y, Kojima S, Matsuura K. Annotated checklist of deep-sea fishes of the Sea of Japan. In: Fujita T, editor. Deep-sea fauna of the Sea of Japan, National Museum of Nature and Science Monographs, vol. 44. 2014. p. 225–91.Google Scholar
- Shinohara G, Shirai AM, Nazarkin MV, Yabe M. Preliminary list of the deep-sea fishes of the Sea of Japan. Bull Natl Mus Nat Sci (Ser A). 2011;37:35–62.Google Scholar
- Smith MM. Family No. 92: Bregmacerotidae. In: Smith MM, Heemstra PC, editors. Smith’s sea fishes. Johannesburg: Macmillan South Africa; 1986. p. 329–30.View ArticleGoogle Scholar
- Tanaka S. Descriptions of eight new species of fishes from Japan. Annotationes Zoologicae Japonenses 7 (pt. 1), 1908;27–47.Google Scholar
- Thompson W. (1840) On a new genus of fishes from India. Charlesworth's Magazine of Natural History, New Series 4 (art. 6), 184–187.Google Scholar
- Torii A, Harold AS, Ozawa T. Redescription of type specimens of three Bregmaceros species (Gadiformes: Bregmacerotidae): B. bathymaster, B. rarisquamosus, and B. cayorum. Memoirs of the Faculty of Fisheries. Kagoshima University. 2003a;52:23–32.Google Scholar
- Torii A, Harold AS, Ozawa T, Iwatsuki Y. Redescription of Bregmaceros mcclellandi Thompson, 1840 (Gadiformes; Bregmacerotidae). Ichthyol Res. 2003b;50:129–39.View ArticleGoogle Scholar
- Torii A, Javonillo R, Ozawa T. Reexamination of Bregmaceros lanceolatus Shen, 1960 with description of a new species Bregmaceros pseudolanceolatus (Gadiformes: Bregmacerotidae). Ichthyol Res. 2004;51:106–12.View ArticleGoogle Scholar
- Turan C, Yağlıoğlu D, Gürlek M, Ergüden D. A new record of Antenna Codlet Bregmaceros atlanticus Goode and Bean, 1886 (Bregmacerotidae: Gadiformes) from the northeastern Mediterranean coast of Turkey. J Black Sea/Mediterr Environ. 2011;17:186–92.Google Scholar
- Whitley G.P. (1941) Ichthyological notes and illustrations. Australian Zoologist 10, part 1, 1–50.Google Scholar
- Yilmaz R, Bilecenoğlu M, Hoşsucu B. First record of the Antenna Codlet, Bregmaceros atlanticus Goode and Bean, 1886 (Osteichthyes: Bregmacerotidae), from the eastern Mediterranean Sea. Zool Middle East. 2004;31:111–2.View ArticleGoogle Scholar
- Zenetos A. Trend in aliens species in the Mediterranean. An answer to Galil, 2009, “Taking stock: inventory of alien species in the Mediterranean Sea”. Biol Invasions. 2010;12:3379–81.View ArticleGoogle Scholar