by John C. Briggs (1960)
In these days of concentration on fishes of relatively restricted areas, by means of regional checklists, circumscribed faunal works, and geographically limited revisions, one is apt to forget that many fishes are relatively unrestricted in distribution, and therefore cannot be said to belong to any one zoogeographic area. Although they may tend to interfere with attempts to give precise definition to the various zoogeographic regions, such species must not be overlooked, as they form an undeniable and interesting part of the general pattern of the dispersal of marine fishes.
The purpose of this contribution is to call attention to those forms which are apparently worldwide in distribution, in the hope that they may receive additional consideration. It should be emphasized that the majority of the species mentioned in this work have not been subjected to recent systematic appraisal. Consequently, it is expected that this list will undergo considerable change in the near future.
The following annotated list of 107 species includes only those which, according to the present state of our knowledge, form homogeneous species populations extending entirely around the world in tropical or temperate waters. Fishes that seem to be worldwide at the species level but that apparently break up into subspecific populations in parts of their range are not included. Under each species are listed only those recent references that seem to give a reasonably good indication of circumglobal distribution. In many instances, these references are reinforced by the citation of a definite record from the Eastern Pacific area. Several species have been listed for which no Indian Ocean captures are yet known. Presumably, this is a reflection of the paucity of collections from that region rather than an actual discontinuity in the range of the species.
Worldwide fishes were found to occupy four different marine habitats. The "shore" species are those found in waters that overlie the continental shelf where the depth is less than 200 meters (109 fathoms). "Pelagic" fishes generally inhabit the surface layers of water--at depths of less than 200 meters--in the offshore regions, usually beyond the limits of the continental shelf. Species inhabiting the offshore waters below 200 meters, but not including the sea bottom, are in the "bathypelagic" zone. The "benthic" forms are bottom fishes found at depths greater than 200 meters. In many cases the designation of habitat has been quite arbitrary, especially in regard to the pelagic versus the shore species. For example, many of the carangids are fast swimmers and exhibit a body shape and coloration that is apparently well adapted to a pelagic existence yet they may be most often caught in relatively shallow water. Perhaps such fishes should be placed in a special "shore-pelagic" category.
Although relatively few of the total number of recent fish families are represented on this list, it can be seen that worldwide species have cropped up at many different evolutionary stages. At the primitive extreme, it may be noted that there are not only many cosmopolitan elasmobranchs but that two lancelets, Asymmetron lucayanum Andrews and the larval Amphioxides pelagicus (Günther), have almost achieved a similar distribution, since they are missing only from the Eastern Pacific. At the other end of the scale, we find several plectognaths and at least not ceratioid species with circumglobal distributions.
As one might suspect, there is a high percentage of wide-ranging species in the families that have exploited the pelagic and bathypelagic environments most successfully.
Despite the fact that the species of marine shore fishes are exceedingly numerous, it is still surprising to find that 14 stray from their littoral habitat, as larvae or adults, often enough to establish worldwide populations.
Ekman (1953: 73), as well as a number of others, has emphasized that the broad expanse of open ocean between Polynesia and America is responsible for the most pronounced break in the circumtropical shore fauna. Since we have noted that 14 shore fishes have been able to cross this "East Pacific Barrier" in establish worldwide ranges, the question may arise as to just how [[p. 172]] effective is the barrier in limiting the distribution of shore species that have shown the ability to become widely dispersed in other parts of the world? A search of the ichthyological literature reveals that apparently only five or six shore species are well established in the warm waters of all oceans but the Eastern Pacific. These are the southern lancelet, Asymmetron lucayanum Andrews, the snakefish Trachinocephalus myops (Forster), the ruby snapper Etelis carbunculus (Cuvier)--usually known as E. oculatus (Cuvier) in the Western Atlantic, the great barracuda Sphyraena barracuda (Walbaum), and the bristle filefish Stephanolepis setifer (Bennett). To these five may be added the flat needlefish Ablennes hians hians (Valenciennes) if it should be properly considered a shore instead of a pelagic species. The black triggerfish Melichthys radula (Solander) and the smooth triggerfish Canthidermis maculatus (Bloch) are not included because these forms have apparently succeeded in crossing the major portion of the barrier in becoming established around certain offshore islands of the Eastern Pacific.
These data make possible a generalization that seems justified in the light of our present knowledge about the species concerned: Of those tropical shore fishes that have a broad distribution elsewhere (on both sides of the Atlantic, the Indian Ocean, and the Western Pacific), the majority have been able to cross the East Pacific zoogeographic barrier.
The foregoing statement is applicable only to the wide-ranging shore fishes. For other shore species, which constitute an overwhelming majority, the East Pacific barrier is very effective indeed. Its efficiency depends not only on distance, as has been generally assumed, but also on ecological factors.
BATHYPELAGIC AND PELAGIC FISHES
As would be expected, by far the majority of fishes showing a worldwide distribution can be considered as typical of the open ocean waters. The 42 bathypelagic and 48 pelagic species comprise 84.1 percent of the entire list. During the process of compiling distributional records, it was noted that an impressive number of these species are apparently affected by the same East Pacific Barrier that operates so noticeably on the shallow-water marine fauna.
No fewer than 28 species of bathypelagic fishes, which show evidence of a broad distribution in the other oceans, have failed to become established in the Eastern Pacific. A similar pattern is shown by at least 15 of the pelagic forms. Many of these bathypelagic species are infrequently captured and are difficult to properly identify and the same can be said for at least a few of the pelagic fishes. For this reason, the absence of so many from the Eastern Pacific may be more apparent than real. With regard to the present distributional picture, however, it can be stated that more than one-third of the wide-ranging bathypelagic and pelagic fishes are evidently prevented from achieving a worldwide distribution by their inability to cross the East Pacific Barrier.
If in the light of future investigations the East Pacific Barrier (previously considered to be operative only in the case of the littoral fauna) continues to appear effective for bathypelagic and pelagic fishes, it may be concluded that many species, usually considered to be well adapted to life in the open ocean, may actually be dependent on a certain degree of proximity to land masses or to relatively shallow water areas.
Rapid progress in systematic work on the bathypelagic fishes is yielding increasing evidence of the wide distribution of a large percentage of the species. At present only a few of these forms are known from waters deeper than 2000 meters (1000 fathoms). Most of the 17 species from such depths listed by Grey (1956: 311-2) are already held to be very widely dispersed: seven are worldwide and two more have been taken from almost all areas except the Eastern Pacific.
Although our knowledge of the benthic fishes is still very limited, it may be said that, in general, their dispersal is quite restricted. Grey (1956: 804) gives a distributional list of benthic species that have been taken below a depth of 2000 meters.
As may be noted by reference to the list herein presented, only three benthic fishes were found to have a circumtropical distribution. As matters now stand, there is, among the wide-ranging members of this group also, some indication of an unexpected scarcity in the Eastern Pacific. Such species as Heptranchias perlo (Bonnaterre), Dalatias licha (Bonnaterre), Bathysaurus ferox Günther, Ipnops murrayi Günther, Synaphobranchus kaupi Johnson, Venefica procera, (Goode and Bean), Chascanopsetta lugubris Alcock, Nematonurus armatus (Hector) seem to have a broad distribution elsewhere but, so far, there is no [[p. 173]] indication of their presence in the depths of the Eastern Pacific area.
1. Hexanchus griseus (Bonnaterre)--sixgill cowshark. Benthic. A species from moderately deep water, usually 75-300 fathoms. Good evidence for a cosmopolitan distribution is given by Bigelow and Schroeder (1948: 80-7).
2. Carcharodon carcharias (Linnaeus)--white shark. Pelagic. Good evidence for a cosmopolitan distribution is given by Bigelow and Schroeder (1948: 134-45).
3. Cetorhinus maximus (Gunner)--basking shark. Pelagic. Although this species is almost always taken in cold or temperate waters, it is found in both northern and southern hemispheres indicating some movement across the equatorial regions. Range from Bigelow and Schroeder (1948: 147-60).
4. Alopias vulpinus (Bonnaterre)--common thresher. Pelagic. Considered to be worldwide, although the Western Pacific species may be distinct from the A. vulpinus of the other oceans, according to Bigelow and Schroeder (1948: 167-78).
5. Rhineodon typhus Smith--whale shark. Pelagic. Reliable reports from the tropical waters of all oceans are listed by Bigelow and Schroeder (1948: 193-5).
6. Galeocerdo cuvieri (Peron and Lesueur)--tiger shark. Shore.
Well substantiated records from the tropical and subtropical portions
of all the oceans are summarized by Bigelow and Schroeder (1948: 271-5).
10. Sphyrna tiburo (Linnaeus)--bonnethead. Shore. Although Bigelow
and Schroeder (1948: 425) thought that this species was not likely to
be found in Indian or Western Pacific waters, Herre (1953: 27-8) includes
it on his Philippine list and gives additional records.
13. Isistius brasiliensis (Quoy and Gaimard). Pelagic. As Bigelow and Schroeder (1948: 509-14) show, there is little doubt that this distinctive species is indeed worldwide.
14. Echinorhinus brucus (Bonnaterre)--bramble shark. Shore. This rare species is at present to be considered to be cosmopolitan, as by Bigelow and Schroeder (1948: 527-32), but more than one form may be included.
15. Squalus fernandinus Molina. Pelagic. The recent description of three Western Atlantic specimens by Bigelow, Schroeder, and Springer (1953: 221) established a circumtropical range for this species. Chilean range is given by Mann (1954: 108).
[[p. 174]] Family Mylobatidae
16. Aetobatus narinari (Euphrasen)--spotted duckbill ray. Shore. The circumtropical range of this ray is well established. The most complete modern account is that of Bigelow and Schroeder (1954: 453-65).
17. Manta birostris (Walbaum)--manta. Pelagic. Considered to be circumtropical although, according to Bigelow and Schroeder (1954: 511), there is some doubt about the specific relationship of the Atlantic and Pacific populations.
18. Albula vulpes (Linnaeus)--bonefish. Shore. Circumtropical range is shown by Smith (1949: 436). Eastern Pacific records are listed by Fowler (1944: 481).
According to the literature on the genus Argyropelecus, there are three species (A. affinis Garman, A. olfersi (Cuvier), and A. sladeni Regan) which could be considered worldwide in distribution. However, Dr. Leonard P. Schultz has in press a review of this group and he has written (personal communication) that he has no evidence of such broad distribution for any of the species.
19. Sternoptyx diophana Hermann. Bathypelagic. As was mentioned by Haig (1955: 319), this species has been reported from many parts of the Atlantic, Indian, and Pacific oceans.
20. Cyclothone microdon (Günther)--small-tooth bristlemouth.
Bathypelagic. Listed by Smith (1949: 437) as occurring in all oceans but
the Eastern Pacific. Recorded from California by Hubbs and Follett (1953:
Although Ege (1934: 9) in his revision of the genus Stomias indicates a worldwide distribution for S. affinis Günther, Dr. Carl L. Hubbs (personal communication) states that neither of the two Eastern Pacific species is referable to this form.
A circumtropical distribution for Idiacanthus fasciola Peters is indicated by Smith (1949: 437) but Dr. Carl L. Hubbs writes (pers. comm.) that it has not been taken in the Eastern Pacific.
26. Neoscopelus macrolepidotus Johnson. Bathypelagic. The general occurrence is given by Fraser-Brunner (1949: 1039) as "temperate and warm seas."
27. Ceratoscopelus townsendi (Eigenmann and Eigenmann). Bathypelagic.
Bolin (1959: 37) states that this species occurs commonly in a broad circumglobal
belt between the latitudes of approximately 35° N. and S.
45. Scopelarchus analis Brauer--common pearleye. Bathypelagic. Records for the Eastern Atlantic, Indian, and Antarctic oceans are cited by Fowler (1936: 354). Listed for Bahama Islands by Fowler (1944: 438) and for California by Hubbs and Follett (1953: 17).
46. Eupharynx pelecanoides Vaillant--gulper. Bathypelagic. A summary of the distributional records for this species is given by Grey (1956: 139).
47. Nemichthys scolopaceus Richardson--common snipe-eel. Bathypelagic.
Occurrences in the North Atlantic, Mediterranean, Indian Ocean, and Western
Pacific listed by Beebe and Crane (1937: 353), and it was recently recorded
from California by Hubbs and Follett (1953: 18).
49. Regalecus glesne (Ascanius)--oarfish. Pelagic. Presence in all oceans but the Eastern Pacific was indicated by Smith (1949: 440). Recorded from California by Hubbs and Follett (1953: 20).
50. Trachipterus cristatus Bonelli--scalloped ribbonfish. Pelagic.
Circumtropical range based on a personal communication from Dr. Vladimir
Walters who with Mr. John E. Fitch and Dr. Richard H. Backus is revising
52. Lophotus capellei Temminck and Schlegel--crestfish. Pelagic. Worldwide range indicated by Briggs (1952: 206).
53. Lampris regius (Bonnaterre)--opah. Pelagic. Smith (1949: 440) gives a circumtropical range. Presence in the Eastern Pacific is verified by Hubbs and Follett (1953: 19).
54. Gadomus longifilis (Goode and Bean)--rattail. Benthic. Parr (1946: 10) accepts this species as cosmopolitan but Grey (1956: 166) remarks that it may be confined to the Atlantic.
55. Antimora rostrata Günther. Benthic. Grey (1956: 160) lists many records that show this to be a rather common, worldwide species.
56. Anoplogaster cornuta Valenciennes--fangtooth. Bathypelagic. Norman (1930: 347) gives records (listed as Caulolepis longidens) from the Atlantic, California coast, and Hawaiian Islands. Mrs. Marion Grey has further evidence (unpublished) that suggests a worldwide distribution.
According to the published literature on this family, both Melamphaes mizolepis (Günther) and M. nigrescens Brauer should be considered circumtropical. However, Dr. Alfred W. Ebeling is completing a study on this group and advises (pers. comm.) that the above species are replaced, in the Eastern Pacific, by related forms.
57. Priacanthus cruentatus (Lacépède)--bigeye. Shore. Herre (1953: 375) and others indicate the range as Atlantic (both sides) to the Indo-Pacific. However, there are specimens in both the University of British Columbia and the University of California at Los Angeles collections from San Lucas Bay, Baja California, and Dr. Boyd Walker has recently provided information (via personal communication) about a U.C.L.A. specimen from Panama Bay.
58. Naucrates ductor (Linnaeus)--pilotfish. Pelagic. The records
for this common, widespread species are most recently compiled by Herre
[[p. 177]] Family Coryphaenidae
64. Coryphaena hippurus Linnaeus--dolphin. Pelagic. The distribution
of this common, worldwide species is most recently summarized by Gibbs
and Collette (1959: 136).
66. Ruvettus pretiosus Cocco--escolar. Pelagic. Most recent
account is that of Herre (1953: 253), who considers this to be a circumtropical
species. Presence in Eastern Pacific is verified by Hubbs and Follett
69. Brama brama (Bonnaterre)--common pomfret. Pelagic. Smith
(1949: 455) indicates a scattered, worldwide distribution (under the designation
of Brama raii). Presence in the Eastern Pacific is verified by
Hubbs and Follett (1953: 24). It is a common offshore species in British
71. Thunnus alalunga (Gmelin)--albacore. Pelagic. Smith (1949:
454) shows a distribution in all oceans but the Eastern Pacific. Hubbs
and Follett (1953: 25) note its occurrence in California waters. Rivas
(1951: 222) mentions a lack of authentic Western Atlantic records but
Bullis and Mather (1956: 8) supply these.
76. Xiphias gladius Linnaeus--swordfish. Pelagic. Herre (1953: 256) and almost all previous authors list this species for all tropical and temperate seas.
77. Makaira nigricans Lacépède--blue marlin. Pelagic. As the result of a comparison of Atlantic and Pacific specimens, Dr. C. Richard Robins (mimeographed Progress Rept. No. 6. The Charles F. Johnson Oceanic Gamefish Investigations, 1960) concludes that the blue marlin is a single, worldwide species.
78. Tetragonurus atlanticus Lowe--square-tail. Pelagic. A cosmopolitan distribution for this species is shown by Grey (1955: 33) in her revision of the genus.
[[p. 178]] Family Mugilidae
79. Mugil cephalus Linnaeus--striped mullet. Shore. Although most authors consider this to be a single, worldwide species, some recent evidence indicates that detailed morphological investigation will disclose a complex of species or subspecies (see de Sylva, Stearns, and Tabb, 1956: 1-45).
80. Remora remora (Linnaeus)--remora. Pelagic. Herre (1953:
775) states that this species is found in all warm seas. Presence in the
Eastern Pacific is verified by Hubbs and Follett (1953: 24).
86. Rachycentron canadus (Linnaeus)--crabeater. Shore. Range given by Herre (1953: 287) as "all warm seas but Eastern Pacific." However, it is recorded from the coast of Chile (Iquique) by Fowler (1944: 502).
87. Pomatomus saltatrix (Linnaeus)--bluefish. Pelagic. Shown from all oceans but the Eastern Pacific by Smith (1949: 448). Recorded from the coast of Chile (Valparaiso) by Fowler (1944: 502).
88. Macrorhamphosus gracilis (Lowe)--slender snipefish. Pelagic.
In her fine revision of the family, Mohr (1937: 61) shows a worldwide
distribution for this species.
90. Chiasmodon niger Johnson--black swallower. Bathypelagic. This interesting species is recorded by Norman (1930: 349) from the Caribbean Sea and the Atlantic and Indian oceans. Listed from California by Hubbs and Follett (1953: 34).
91. Diodon holacanthus Linnaeus--balloon fish. Shore. Smith
(1949: 463) shows a distribution to all oceans except the Eastern Pacific.
Recorded from a number of Eastern Pacific localities by Fowler (1944:
93. Mola mola (Linnaeus)--headfish. Pelagic. In his recent revision
of the family, Fraser-Brunner (1951: 113), gives an extensive synonymy
which demonstrates a worldwide distribution.
[[p. 179]] Family Balistidae
95. Xanthichthys ringens (Linnaeus)--redtail triggerfish. Shore. Clark (1949: 10) indicates that X. lineopunctatus (Hollard), a widespread Indo-Pacific species, is the same as X. ringens (Linnaeus) of the Atlantic. It is reported, under the former name, from California by Hubbs and Follett (1953: 38).
96. Alutera monoceros (Osbeck)--unicorn filefish. Shore. The
only Eastern Pacific record for this distinctive species seems to be that
of Hiyama in the book by Kumada (1937: 54, pl. 86) from somewhere along
the Mexican coast. However, there is a specimen in the U. B. C. collection
(BC60-31) taken by Mr. Tony Pletcher 2.5 miles south of Mazatlán,
Sinaloa, Mexico, on July 11, 1959. There are many records in the literature
for both sides of the Atlantic, the Indian Ocean, and the Western Pacific.
98. Cryptopsaras couesi Gill. Bathypelagic. This and the following
species of this family are listed from the distributional table provided
by Bertelsen (1951: 222) in his excellent treatise on the family based
on the material from the various "Dana" expeditions.
Acknowledgements.--I wish to express my appreciation, especially, to Rolf L. Bolin of the Hopkins Marine Station for much helpful information about the distribution of the Myctophidae and to Marion Grey of the Chicago Natural History Museum for sending data about several other deepwater fishes. I am also indebted to Carl L. Hubbs and Alfred W. Ebeling of the Scripps Institution of Oceanography, Vladimir Walters of the American Museum of Natural History, Leonard P. Schultz and Ernest A. Lachner of the U. S. National Museum, Giles W. Mead of the Museum of Comparative Zoology at Harvard, and Boyd W. Walker of the University of California at Los Angeles for providing useful information upon certain species groups or individual specimens. Finally, my thanks to Carl L. Hubbs and to Peter A. Larkin and Casimir C. Lindsey of the University of British Columbia for their helpful criticism of the manuscript.
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INSTITUTE OF FISHERIES, UNIVERSITY OF BRITISH COLUMBIA, VANCOUVER, CANADA.