Zootaxa 2525: 19–50 (2010)www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

ArticleCopyright © 2010 · Magnolia PressZOOTAXAISSN 1175-5334 (online edition)

Marine fish diversity: history of knowledge and discovery (Pisces)
WILLIAM N. ESCHMEYER1, 5, RONALD FRICKE2, JON D. FONG3 & DENNIS A. POLACK4
1
Curator emeritus, California Academy of Sciences, San Francisco, California, U.S.A. 94118 and Research Associate, Florida
Museum of Natural History, Gainesville, Florida, U.S.A. 32611. E-mail: weschmeyer@calacademy.org
2
Ichthyology, Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany. E-mail: Ronald.fricke@smns-bw.de3
California Academy of Sciences, San Francisco, California, U.S.A. 94118. E-mail: jfong@calacademy.org
4
P.O. Box 518, Halfway House 1685, South Africa. E-mail: dennis@fishwise.cc
5
Corresponding Author. E-mail: weschmeyer@calacademy.org

Abstract
The increase in knowledge of marine fish biodiversity over the last 250 years is assessed. The Catalog of Fishes database(http://research.calacademy.org/ichthyology/catalog) on which this study is based, has been maintained for 25 years andincludes information on more than 50,000 available species names of fishes, with more than 31,000 of them currently
regarded as valid species. New marine species are being described at a rate of about 100–150 per year, with freshwater
numbers slightly higher. In addition, over 10,000 generic names are available ones of which 3,118 are deemed valid formarine fishes (as of Feb. 19, 2010).
This report concentrates on fishes with at least some stage of their life cycle in the sea. The number of valid marine
species, about 16,764 (Feb. 19, 2010), is about equal to that of freshwater fishes (15,170). Valid species of fishes
apparently restricted to brackish water number only 108. The sum (32,042) is more than the current total number of
31,362, valid species of fishes because some species occur in more than one habitat.
Presented is information on the description of species and genera over historic time, the authors describing taxa, andthe deemed validity of described species and genera. We characterize families and also geographic areas where marine
fishes are relatively well known and those where much discovery appears to remain. Endemism is also discussed. As
examples, the marine fish faunas of the Mediterranean Sea, the Red Sea and the Mascarene Islands are shown to be wellknown. Little new discovery has been found recently at the family level, and new discoveries of species and genera aremostly limited to certain families and geographic areas and habitats. Specialized collecting techniques are discussed.
Overall success rates for valid species discovery through time has only been about 50%, or two species described foreach valid one recognized; however this percentage has been improving over time. Because of recent improvements intechnology, literature availability, quality of analysis, better communication, and other factors, the current success rate
for validity of species is well over 90% (with a small lag time as status is confirmed or rejected by the ichthyologicalcommunity).
Two habitats where most new marine taxa will likely be found are deep-reefs and deep-slopes, areas poorly sampled
and studied. Some deep-sea areas, particularly in the Southern Hemisphere and throughout the Indian Ocean and inIndonesia, should reveal many new taxa from increased collecting efforts. Molecular genetic studies are proving valuable
in phylogenetic and phylogeographic analyses as well as in species’ population analyses, but these relatively newtechniques are not uncovering large numbers of new or cryptic taxa.
An estimate of marine fish species yet to be sampled and described is about 5,000, or twice the number described in
the last 19 years, for a projected total of approximately 21,800 valid marine species of fishes.
Key words: Marine fishes (all), history of discovery, taxonomy, valid taxa, statistics of discovery, biogeography,
families, genera, species

Introduction
Taxonomy is an on-going endeavor. The basic knowledge of diversity through species discovery anddescription is mostly complete for some areas of the world and for many families of fishes, but important gaps
remain. The present study is an analysis of the scientific discovery and description of marine fish taxa overAccepted by L. Page: 12 Jun. 2010; published: 2 Jul. 201019

time. Fishes constitute more than half of all vertebrates, with over 31,000 valid species, and of these over half
are marine fishes.
The Catalog of Fishes database was begun in the 1980s at the California Academy of Sciences where it ismaintained and updated continuously. Two large publications have resulted from it: the “Genera of Fishes”(Eschmeyer & Bailey 1990) and the “Catalog of Fishes” (Eschmeyer 1998a); online versions have been
updated several times each year since 1997. The database is available online at http://
research.calacademy.org/research/ichthyology/catalog/fishcatmain.asp.
The Catalog of Fishes database was designed especially for ichthyologists and other biologists, andincludes a large audience seeking accurate information on fish taxonomy, including evolutionary andenvironmental biologists, fisheries workers and ecologists. It is an on-going summary of current knowledge ofthe kinds of fishes. Special efforts were made to provide accurate spellings of taxonomic names, informationon the location of type specimens, and reliable information on dates of publication, correct authorship andother technical details. Nearly all original descriptions of species and genera were examined. Information in
the database has been incorporated into the independent Fishbase database and several other initiatives inrecent years. From the Catalog of Fishes database, we are able to analyze a number of trends, and provide anumber of graphs and tables to document the history of discovery of marine fishes.
Mora et al. (2008) used records from the Ocean Biogeographical Information System (OBIS) database(http://www.obis.org) to estimate the completeness of the global fish inventory through the extrapolation of
species discovery curves. They estimated that the global inventory is about 79% complete, or that 21% ofmarine fishes remain to be discovered, based on about 15,700 currently known. In the present study, adifferent approach is used, involving analysis by families of fishes and their distribution and biology, analysis
of geographic areas and basins, and descriptions of new species through time, and from various habitats.
Compared to Mora et al. (2008), our estimates are somewhat higher, reflecting more species from habitats
under-represented in their data, different assumptions, and consideration of biological factors.

Methods
The printed version of the Catalog of Fishes (Eschmeyer 1998a) provides information on the database and its
preparation and content. In this paper we include information on all taxa described by the end of 2009. Figures
and tables were prepared from information taken from the online version of 19 February 2010 (for data
through 2009). The database uses MS SQL Server 2005 and MS Access. The figures were prepared in MS
Excel. Some graphs and tables are presented in 5- or 10-year periods, except the first time period includes theextra year 1758 making it a 6- or 11-year period.
Definitions important to this paper are: Marine fishes: Species that spend at least a part of their life cyclein the sea. This includes anadromous species, such as salmon, as well as catadromous and amphidromous
species. Available name: A scientific name that meets the criteria of publication, authorship and otherrestrictions of the international Code of Zoological Nomenclature; it is a name that may be used in zoologicalnomenclature. Generally, we are concerned only with available names. Valid name: A species, subspecies orgenus that is considered to be a legitimate and recognizable “good” taxon. Synonym: A taxon considered to
be a junior synonym (duplicate) of a taxon describe earlier. Primary type locality: The geographic locality ofthe primary name-bearing type specimens (holotype, lectotype, neotype or syntypes). Species-group name:
The scientific name of a species or subspecies; for this analysis, specific and subspecific names are grouped
together as species-group taxa; a subspecies can be raised to a species or a species reduced to a subspecies
with no change in availability, spelling or authorship. Genus-group name: The scientific name of a genus orsubgenus; for this analysis, generic and subgeneric names are grouped together as generic-group taxa.
Endemism (endemic): Restricted in distribution to a specific geographic area (e.g., a species endemic to the
Hawaiian Islands). Status of taxa: Three categories are used—valid, uncertain, synonym. Species habitats:
Where a species lives; for this analysis a species may occur in one to three habitats—fresh, brackish or marine
waters.

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ESCHMEYER ET AL.

Results and discussion
Species over time. Descriptions of new fish species over time varies widely (Table 1, Fig. 1) for a variety ofreasons. The great exploring expeditions of the mid-1800’s provided high peaks in new marine species
descriptions. And, in the late 1800s and early 1900s, American marine expeditions produced anothersignificant peak. During the 1940s, a number of new workers entered the discipline. The advent of SCUBA
and use of ichthyocides (mainly rotenone) resulted in the discovery and description of many new marine
species since 1950, and there were significantly more researchers in subsequent years. The total number ofauthors of marine fish species was about 2,300 (as of the end of 2009), and the number of authors of names for
all habitats was about 4,300.
TABLE 1. Number of new marine species-level taxa (species and subspecies), number of names now considered to bevalid, and the total number of authors describing those taxa, per 5-year periods.
Years

New Taxa Valid

Authors

Years

New Taxa Valid

Authors

1758–1763

321

263

2

1884–1888

652

248

61

1764–1768

98

48

11

1889–1893

585

381

53

1769–1773

60

28

10

1894–1898

618

334

69

1774–1778

168

100

14

1899–1903

932

469

67

1779–1783

64

35

13

1904–1908

1217

582

88

1784–1788

161

73

18

1909–1913

1008

530

89

1789–1793

471

151

17

1914–1918

619

292

64

1794–1798

87

40

14

1919–1923

466

223

64

1799–1803

1009

267

17

1924–1928

733

324

95

1804–1808

96

20

12

1929–1933

952

398

113

1809–1813

367

93

15

1934–1938

1058

444

121

1814–1818

377

104

15

1939–1943

551

259

88

1819–1823

213

72

31

1944–1948

486

168

68

1824–1828

621

189

32

1949–1953

599

327

110

1829–1833

1524

578

52

1954–1958

592

305

141

1834–1838

746

304

53

1959–1963

647

403

152

1839–1843

1005

335

54

1964–1968

512

350

185

1844–1848

933

360

44

1969–1973

495

401

225

1849–1853

841

325

48

1974–1978

663

574

273

1854–1858

1232

397

43

1979–1983

832

759

320

1859–1863

1161

421

46

1984–1988

740

677

330

1864–1868

772

287

45

1989–1993

608

578

276

1869–1873

694

243

43

1994–1998

550

524

250

1874–1878

836

358

58

1999–2003

689

681

292

1879–1883

1059

475

64

2004–2008

859

846

382

Periods of high activity can be summarized as follows (numbered in Fig. 1). Stage 1, early discovery:
This covers the period after Linnaeus in 1758 to about 1825. During this time, very little new material became
available. Many species that had been described earlier were given new names in works authored by B. G. E.
Lacepède, M. E. Bloch and J. G. Schneider, C. S. Rafinesque, W. Swainson and G. Shaw. The percentage ofvalid species is low and authors few. There was no “rule book” comparable to today’s Code of Zoological
Nomenclature (International Commission on Zoological Nomenclature, 1999). Stage 2, worldwide

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explorations: This covers the period from about 1825 to about 1895, especially the mid-1800s. This was atime of the great marine expeditions by Europeans. There was much new material collected, including many
surprising discoveries. A code of nomenclature was established in 1842 [for an interpretation and history see
Eschmeyer (1998b, Appendix A)]. Stage 3, focused exploration: The period from about 1890 to 1920, saw
extensive American marine expeditions (e.g., Hawaiian Islands, Alaska, Bering Sea, Galapagos), andincreased deep-sea exploration by explorers from several countries. Many monographs on many groups offishes and geographical areas were published by David Starr Jordan and his co-workers. The number ofprofessional ichthyologists remained low, however. Stage 4, increased workers: This stage reached a peakduring the 1920s–1950s. There was an increase in taxonomic workers, and in the publication of regional
studies on marine fishes by such ichthyologists as H.W. Fowler, G. P. Whitley, Carl L. Hubbs and J.L.B.
Smith. Stage 5, new exploration, new techniques: After 1950, there was less exploration of offshore anddeep-sea areas, but much concentration in search for fishable commercial stocks. However, some significant
exploration of deep-sea and mid-water depths was undertaken, especially by German and Russian scientists.
Some geographic areas and many taxonomic groups (especially commercially important ones) became
relatively well studied. Technological advances contributed to ease of study. SCUBA was introduced, there
were many more workers, and new geographical areas were studied.

FIGURE 1. Number of new marine species-level taxa (species and subspecies), number of names considered to be valid(as species), and total number of authors, per 5-year periods.

After 1950 and increasing to the present, ichthyological studies became more efficient with better tools,
such as SCUBA (and just recently re-breathers), use of rotenone and other ichthyocides, manned and robotic
submersibles, molecular genetic studies, better communication, the Internet, computers, photocopiers

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followed by scanners, portable document format (PDFs), team approach, digital photography, online
literature, the Catalog of Fishes and Fishbase, electronic journals, and more publication outlets. The
sophistication and accuracy of description of new taxa increased substantially.
Use of subspecies rank in marine fishes. Subspecies are species-group names, and a subspecies may beelevated in rank to a full species with no change in authorship and date. On the other hand, a taxon described
as a new species can be demoted later to subspecies rank. Subspecies were much more commonly used inearlier literature than now. Table 2 illustrates the increase in description of (full) species and the decline of theuse of subspecies since 1940 in marine taxa.
TABLE 2. Average number of marine species and subspecies per 10-year period since 1940, and percent of total
described as subspecies.
PeriodSpecies

Subspecies% Subsp.

Total

1940–1949

85

9

11.52

94

1950–1959

118

11

9.63

129

1960–1969

101

5

5.59

106

1970–1979

114

5

4.63

119

1980–1989

153

3

2.15

156

1990–1999

117

1

1.53

118

2000–2009

150

1

0.67

151

The concept of subspecies has changed through time. Some authors used subspecies to identify closely
related populations that were not overlapping in distribution and usually showed differences that wereconsidered minor. More recent studies suggest that if populations show consistent differences and lack zones
of integration, they can be regarded as full species, especially with molecular confirmation. In freshwater, theuse of subspecies has similarly declined—if populations are in long-separated basins, they are usually now
called full species.
The proportion of subspecies described has declined in every decade since 1940 (Table 2). For the period
1940–1949, the average number of species described annually was 85, while an average of nine subspecies
were described per year. In 2000–2009, an average of 150 new marine species and only one marine subspecies
were described per year. Subspecies are now recognized mostly in only a few oceanic families withpopulations occurring in two or more ocean basins. Examples of currently recognized subspecies are found intropical marine families such as flyingfishes (Exocoetidae), halfbeaks (Hemiramphidae), and a few others.
Authors of new species. The number of authors of names of new marine fish species also has changed
substantially through time (Table 1, Fig. 1). Until about 1880, fewer than 60 authors per each 5-year period
were responsible for describing all new species, and nearly all of them were men associated with major
natural history museums. In general, the specimens went to the museums, having been collected byprofessional collectors and ichthyologists working for government agencies, universities, or museums. Since
1950, there has been an increase in number of authors, often including technicians and collectors.
A major change is that the describers of new marine species have “gone to obtain the specimens” rather
than the specimens coming to them. This includes many ichthyologists employing SCUBA to observe andcollect, such as John Randall, Gerald Allen, Richard Pyle and Ross Robertson among many others, and otherbiologists participating directly in trawling operations and in field collecting. They see the specimens in fresh
condition, can use digital photography to document live colors, usually employ global positioning systems(GPS), and often use specialized techniques. This contrasts to the early years when the specimens werebrought to the museum specialist, at most accompanied by a color drawing or painting. The early workers
tended to publish alone or with one co-author and without peer review, whereas more recent authors often
publish with one to often several co-authors and most journals require rigorous peer review of theirmanuscripts.

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Zootaxa 2525 © 2010 Magnolia Press · 23

TABLE 3. Number of described and valid marine species-level taxa (species and subspecies) described by the 50 mostprolific authors, from 1758 through 2009, including their success rate (% valid as species) and year of first publication.
Rank

AuthorDescribed

Valid% Valid

First Publication

1

Bleeker P.

1373

571

41.59

1846

2

Valenciennes A.

1247

459

36.81

1821

3

Jordan D. S.

1068

575

53.84

1878

4

Günther A.

988

551

55.77

1859

5

Cuvier G.

950

383

40.32

1798

6

Fowler H. W.

814

284

34.89

1900

7

Gilbert C. H.

743

579

77.93

1880

8

Randall J. E.

705

680

96.45

1955

9

Bloch M. E.

577

223

38.65

1779

10

Regan C. T.

518

281

54.25

1902

11

Lacepède B. G. E.

490

113

23.06

1798

12

Whitley G. P.

451

157

34.81

1926

13

Steindachner F.

442

200

45.25

1860

14

Richardson J.

431

182

42.23

1823

15

Poey F.

406

103

25.37

1851

16

Smith J. L. B.

386

189

48.96

1931

17

Linnaeus C.

379

300

79.16

1758

18

Schneider J. G.

372

114

30.65

1788

19

Seale A.

315

86

27.3

1896

20

Castelnau F. L.

294

52

17.69

1855

21

Ogilby J. D.

272

118

43.38

1885

22

Allen G. R.

271

256

94.46

1970

23

Garman S.

265

160

60.38

1875

24

Bean T. H.

261

158

60.54

1877

25

Evermann B. W.

251

97

38.65

1887

26

Risso A.

244

51

20.9

1810

27

Herre A. W. C. T.

234

88

37.61

1903

28

Snyder J. O.

230

122

53.04

1896

29

Rafinesque C. S.

230

30

13.04

1810

30

Macleay W.

227

44

19.38

1877

31

Gill T. N.

223

104

46.64

1859

32

Walbaum J. J.

216

40

18.52

1784

33

Kaup J. J.

210

61

29.05

1826

34

Schultz L. P.

196

107

54.59

1931

35

Schlegel H.

195

142

72.82

1839

36

Goode G. B.

192

129

67.19

1874

37

Rüppell W. P. E. S.

191

99

51.83

1828

38

Starks E. C.

189

103

54.5

1895

39

Temminck C. J.

185

136

73.51

1842

40

Trewavas E.

181

86

47.51

1929
Continued next page

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TABLE 3. (continued)
Rank

AuthorDescribed

Valid% Valid

First Publication

41

Nielsen J. G.

169

162

95.86

1961

42

De Vis C. W.

167

17

10.18

1882

43

Hildebrand S. F.

166

89

53.61

1912

44

Weber M.

164

99

60.37

1905

45

Alcock A. W.

162

131

80.86

1889

46

Last P. R.

161

160

99.38

1978

47

Andriashev A. P.

160

138

86.25

1934

48

Tanaka S.

160

67

41.88

1905

49

Shaw G.

159

24

15.09

1790

50

Gronow L. T.

158

1

0.633

1772

The authors who described the most marine species are listed in Table 3. Early workers (see date of firstpublication) dominated the description of new marine species. Of the top 20, all are deceased except the“living legend” John E. Randall of the Bishop Museum (Honolulu), who in 2009 ranked as number 8, andcontinues to publish new taxa. The top 50 includes only three other living persons (G. R. Allen at #22, J. G.
Nielsen at #41, and P. R. Last at #46). Randall has described the most marine species that are currently
considered valid. In a tally of new species from freshwater and estuarine as well as marine habitats, the top 16
authors of described species at the end of 2009 were: Bleeker (1,926 species, 44% valid), Valenciennes(1,843, 37), Günther (1,633, 57), Fowler (1,445, 41), Jordan [D. S.] (1,351, 52), Boulenger (1,136, 69), Cuvier(1,073, 40), Steindachner (1,032, 57), Regan (934, 63), Gilbert [C. H.] (813, 76), Eigenmann [C. H.] (736,72), Randall [J. E.] (717, 96), Bloch (714, 38), Lacepède (612, 23), Richardson (526, 40) and Linnaeus (499,78).
The number of new species per publication (Table 4) has decreased to an average of between one and two
species per publication since the period 1964–1968. This reflects new methods and fewer new marine species
being found. Major revisions with several new species are usually the result of the work by one or two or three
co-workers that have studied the group for many years, but these publications have become rather rare (see forexample Schwarzhans et al. 2005).
Genera of marine fishes. Most new genus-level taxa in early years were described by museum-basedtaxonomists who had available to them the necessary comparative specimens and literature. Description of
new taxa shows a decline after about 1929 until the most recent years. The numbers of new genera andsubgenera of marine fishes described since 1930, in 10-year periods are shown in Figure 2. The top 25 authorsof marine fish genera are listed in Table 5; all are deceased. David Starr Jordan (with his co-authors included)
tops the list, followed by T. N. Gill. “Success” rates (i.e., taxa now considered valid) vary widely. For marine
fishes, A. W. C. T. Herre has the worst “batting average”—only 22.6% of marine genera and subgenera hedescribed are now considered valid, followed by Henry Fowler with 24%. Peter Bleeker was a famous fieldoriented researcher, but many of his new marine genera came from southern Indonesia, which forms part ofthe large tropical Indo-Pacific faunal realm, thus some of them already had been described from other areas byearlier authors. Linnaeus was first to describe genera, so his success rate based on status of species at the endof 2009 is high. Other workers, such as C. T. Regan and C. H. Gilbert, had early access to major theretofore
relatively unknown faunas, thus their success rates also are relatively high.
For comparison, the top 15 authors of fish genera and subgenera from all habitats through 2009 are:Jordan (604 described, 212 valid, 35% valid), Bleeker (523, 299, 57), Fowler (479, 117, 24), Gill [T. N.] (473,
246, 52), Whitley (450, 127, 28). Günther (301, 186, 62), Eigenmann (209, 140, 67), Cuvier (206, 112, 54),
Rafinesque (190, 70, 37), Evermann [C. H.] (189, 50, 26), Regan (187, 110, 59), Smith [J. L. B.] (169, 71, 42),
Swainson (166, 59, 35), Hubbs [Carl L.] (140, 49, 35), Lacepède (137, 89, 65). No living fish taxonomist
ranks higher than 64th in this analysis. The top five authors of valid genera from all habitats (through 2009)
are: Bleeker (299 valid), Gill [T. N.] (246), Jordan [D. S.] (212), Günther (186) and Eigenmann (140).
MARINE FISH DIVERSITY

Zootaxa 2525 © 2010 Magnolia Press · 25

TABLE 4. Yearly average number of new marine species-level taxa (species and subspecies) described per publication,
per 5-year periods.
Years

Avg. Taxa

Years

Avg. Taxa

1758–1763

64.2

1884–1888

3.9

1764–1768

7.0

1889–1893

4.6

1769–1773

4.0

1894–1898

5.0

1774–1778

12.9

1899–1903

5.4

1779–1783

3.6

1904–1908

5.2

1784–1788

6.0

1909–1913

5.0

1789–1793

14.3

1914–1918

3.5

1794–1798

4.0

1919–1923

3.6

1799–1803

59.4

1924–1928

3.9

1804–1808

7.4

1929–1933

3.6

1809–1813

17.5

1934–1938

3.6

1814–1818

10.8

1939–1943

2.9

1819–1823

5.2

1944–1948

3.8

1824–1828

12.2

1949–1953

2.5

1829–1833

17.9

1954–1958

2.1

1834–1838

8.3

1959–1963

2.1

1839–1843

10.9

1964–1968

1.6

1844–1848

10.5

1969–1973

1.6

1849–1853

6.9

1974–1978

1.8

1854–1858

7.8

1979–1983

1.9

1859–1863

5.9

1984–1988

1.8

1864–1868

4.4

1989–1993

1.6

1869–1873

6.1

1994–1998

1.8

1874–1878

5.1

1999–2003

1.8

1879–1883

4.0

2004–2008

1.7

Described genera of fishes, as in other taxonomic disciplines, generally experienced a break-up of the(early) Linnaeus-period inclusive genera as more species diversity was discovered. New discovery anddescription of marine genera now have leveled off (Fig. 3). Recently described deep-reef species are mostly
relatives of species in shallow-living genera. Molecular genetic studies have resulted in some refinement ofgeneric limits, but in many cases as new genera are named, available names rather than new names can beused. This decline in description of new genera indicates that marine fish biodiversity is becoming wellknown, at least at the generic level.
Different taxonomic disciplines include very different “sized” genera—with, for example, some insect
and mollusk genera containing thousands of named species; concepts of generic limits also vary. More than
3,118 valid marine genera of fishes were recognized as of the end of 2009. Surprisingly, about 1,139 (about 36%) are monotypic (with a single valid included species) as of Feb. 2010. Marine genera with the mostcurrently deemed valid species are Apogon (163 species), Coelorinchus (121), Gymnothorax (120),
Paraliparis (118), Eustomias (117), Careproctus (116), Callionymus (109), Sebastes (109), Chromis (95) andEpinephelus (90). Nineteen species of Apogon were recently removed to Apogonichthyoides (Fraser & Allen
2010), reducing to 144 the number of species in Apogon.

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ESCHMEYER ET AL.

FIGURE 2. Number of new marine genus-level taxa (genera and subgenera) of marine fishes, number of names
considered to be valid (as genera), and number of authors, per 5-year periods.

The average genus of marine fishes contains 5.4 species recognized as valid. [For comparison, the average
number of species in freshwater fish genera is 6.7, where the largest genera are Barbus (314 species),
Haplochromis (205), Schistura (177), Corydoras (153), Etheostoma (149), Puntius (133), Trichomycterus(133), Hypostomus (129), Rivulus (129) and Synodontis (128).] Some of these genera probably will be split
into additional genera, and splitting for some has already been suggested, especially as a result of molecular
genetic analyses. The “size” of genera presumably represents evolutionary history and success, but also isdetermined by historical consensus among fish taxonomists. However, the success in defining monophyletic
groups for most marine taxa has not yet been tested.
Does the current description of new genera reflect a splitting of existing taxa or the discovery ofcompletely new biodiversity? Molecular and morphological studies using new techniques result in somesplitting, and less so the lumping of existing genera as an effort is made to identify monophyletic groups. At
the same time, description of new genera representing new biodiversity discoveries continues. One way to
distinguish between the two is to examine the type species of the new genera—is the type species of the
newly-named genus a new species, indicating new generic diversity discovery? Or was the new genus
established for a previously described species that has been moved to the new genus? This is somewhat of ageneralization because a new genus may contain both a previously described species and one or more new
species, and the type species selected may represent one of the previously described species.

MARINE FISH DIVERSITY

Zootaxa 2525 © 2010 Magnolia Press · 27

TABLE 5. Number of described and valid marine genus-level taxa (genera and subgenera) published by the 25 mostprolific authors, from 1758 through 2009, with the percentage valid (as genera) per author and the year of their firstpublication.
Rank

AuthorDescribed

Valid% Valid

First Publication

1

Jordan D. S.

496

177

35.69

1877

2

Gill T. N.

420

214

50.95

1858

3

Whitley G. P.

388

112

28.87

1927

4

Fowler H. W.

329

79

24.01

1901

5

Bleeker P.

294

161

54.76

1845

6

Günther A.

204

131

64.22

1859

7

Cuvier G.

168

89

52.98

1798

8

Evermann B. W.

166

45

27.11

1887

9

Smith J. L. B.

165

70

42.42

1931

10

Swainson W.

134

49

36.57

1838

11

Rafinesque C. S.

122

46

37.7

1810

12

Gilbert C. H.

117

62

52.99

1879

13

Lacepède B. G. E.

111

68

61.26

1798

14

Regan C. T.

110

58

52.73

1903

15

Kaup J. J.

107

45

42.06

1826

16

Hubbs C. L.

90

25

27.78

1915

17

Bean T. H.

72

47

65.28

1879

18

Valenciennes A.

72

34

47.22

1830

19

Ogilby J. D.

72

24

33.33

1885

20

Herre A. W. C. T.

62

14

22.58

1923

21

Goode G. B.

61

37

60.66

1879

22

Starks E. C.

60

32

53.33

1895

23

Castelnau F. L.

58

16

27.59

1855

24

Snyder J. O.

57

20

35.09

1900

25

Linnaeus C.

56

55

98.21

1758

Numbers of new marine genera described in combination with new species descriptions for the last 51
years are shown in Table 6. Note that the number of genera described for 10-year periods shows a decrease in
the description of new marine genera with a new species as the type species of the genus.
Marine species by selected family. Marine fish families fall into several groups, although some fishes ofsome families are in more than one category: commercial families with large species, commercial families
with small to medium-sized species, and families of no commercial value and/or with small species, or
species which are difficult to catch. Some families have large species (usually widespread) that have been
described numerous times (note percent valid in the table), especially ones with large changes with growth(such as ocean sunfishes, with 47 names for four valid species). We can distinguish ‘mature families’ (thosewith few new species in the last 10 years) from ‘active families’, those with new species now being found anddescribed. A list of all fish families/subfamilies through Feb. 19, 2010 (showing available names, valid
names, and new species in the last 10 years) is presented at http://www.calacademy.org/research/ichthyology/
catalog/SpeciesByFamily.asp. Table 7 shows data for selected marine fish families. The first four families
contain moderate to large species, most of which have wide ranges and are commercially important. The nextnine families/subfamilies are small and/or hard to collect. Gobiidae and Cottidae contain some freshwater
species which have been excluded.

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ESCHMEYER ET AL.

FIGURE 3. Number of currently valid marine genus-group taxa (genera and subgenera) described in combination withnew species from 1929 through 2009, by 10-year periods.

We can assess the depth of capture of new species of selected coral/rocky reef families in the last 20 years,
although the discovery of deeper-living species is just beginning. This shows (Table 8) that new species are
generally being collected in greater depths, thereby reflecting the more complete sampling of shallow-reef
species (except for species in some families containing small species).
New marine families. New macro-biodiversity discoveries can be defined as new families that are basedon discovery of a unique and unusual new genus. These are distinguishable from new families that result
from splitting or reorganizing of existing genera and species through phylogenetic studies. Most of thefamilies of fishes and much of the major biodiversity were discovered and named before 1900. There is nodatabase for fish families. Are we currently finding new and unusual taxa or have we discovered the unusual
fish evolutionary innovations? To our knowledge, in the last 50 years, there have been only eight recognized
new marine families described based on new biodiversity discoveries (about 1.6% of total fish families
currently recognized).

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Zootaxa 2525 © 2010 Magnolia Press · 29

TABLE 6. Number of new marine genus-level taxa (genera and subgenera) described in combination with descriptionsof new species as the type species from 1959 through 2009 by year, and at right by 10-year periods from 1930 through
2009.
Year

New

Year

New

Year

New

Years

New

1959

19

1976

12

1993

8

1960

11

1977

12

1994

6

1961

14

1978

18

1995

4

1962

6

1979

10

1996

3

1963

9

1980

8

1997

4

1930–1939

253

1964

10

1981

8

1998

3

1940–1949

135

1965

15

1982

9

1999

4

1950–1959

141

1966

23

1983

14

2000

1

1960–1969

115

1967

14

1984

10

2001

4

1970–1979

91

1968

4

1985

8

2002

4

1980–1989

89

1969

9

1986

6

2003

3

1990–1999

53

1970

9

1987

9

2004

8

2000–2009

52

1971

10

1988

7

2005

5

1972

7

1989

10

2006

5

1973

4

1990

8

2007

11

1974

3

1991

10

2008

2

1975

6

1992

3

2009

9

TABLE 7. Number of available species-level taxa (species and subspecies) for marine fishes, their current status, and thepercent valid at the species level for selected families and subfamilies for 1758-through 2009 and in the last 10 years.
Family/Subfamily

Available

Currently valid Synonym

Uncertain% valid

New last 10
years

Carangidae (Jacks and allies)

619

147

452

17

23.75

2

Istiophoridae (Billfishes)

61

10

49

2

16.39

0

Scombridae (Tunas and allies)

182

53

127

2

29.12

0

Carcharhinidae (Large sharks)

190

52

120

17

27.37

0

Gobiidae (Gobies)

1838

1165

644

24

63.38

180

Cottidae (Sculpins)

327

188

138

1

57.49

9

Symphurinae (Tongue soles)

92

77

13

2

83.7

8

Ogcocephalidae (Batfishes)

96

66

26

4

68.75

10

Callionymidae (Dragonets)

276

183

90

3

66.3

17

Bythitidae (Livebearing brotulas) 205

192

12

1

93.66

105

Tripterygiidae (Threefin blennies) 207

167

39

1

80.68

21

Syngnathidae (Pipefishes/
Seahorses)

557

300

250

6

53.86

29

Scorpaenidae (Scorpionfishes)

331

209

112

10

63.14

23

A surprising discovery was of the megamouth shark (Megachasma, family Megachasmidae Taylor et al.
1983), a large species that feeds by straining. It was unknown to scientists before that time; the first specimen
was caught tangled in a sea anchor. The discovery of deep gaseous hydrothermal vents revealed new families

30 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

and higher categories of invertebrates, but the only new fishes described was one species in a new genus
Thermarces of a known deep-sea family Zoarcidae.
TABLE 8. Average depth of capture (in meters) of new marine species-level taxa (species and subspecies) in selectedfamilies based on data published for primary types for 1985–2009. Numbers in parentheses are numbers of species
represented.
Family1985–1989

1990–1994

1995–1999

2000–2004

2005–2009

Apogonidae

12.38 (11)23.50 (19)26.63 (19)18.27 (18)30.30 (6)Chaetodontidae

28.25 (2)none

30.48 (1)140.00 (1)none

Gobiidae

26.08 (26)28.79 (45)51.53 (20)24.06 (50)36.17 (74)Labridae

25.51 (15)17.21 (12)26.42 (34)53.69 (21)33.56 (14)Pomacentridae

34.78 (15)

7.57 (23)9.74 (10)16.05 (19)45.14 (11)Pseudochromidae

22.95 (6)25.91 (7)17.34 (11)15.70 (17)30.33 (3)Serranidae

107.34 (17)93.50 (17)110.73 (14)61.31 (8)149.63 (8)At the same time, some very unusual bathypelagic (1000–4000 meters) families have been shown recently
to be members of existing families and genera, and they are no longer recognized—a decrease inbiodiversity! Kasidoroidae Robins & de Sylva (1965) was shown by de Sylva & Eschmeyer (1977) to be
based on unusual juveniles of the family Gibberichthyidae, and Johnson et al. (2009) synonymized two
families into a third. They studied transitional specimens, and based on morphological and mitogenomic
sequence data, showed that fishes assigned to Mirapinnidae, Megalomycteridae and Cetomimidae werelarvae, males and females, respectively, of the single family Cetomimidae.
TABLE 9. New marine families based on discoveries of new species from 1960–2009.
Year

FamilyComments

1965

Kasidoroidae

Robins & de Sylva (1965)—young of Gibberichthyidae (see de Sylva & Eschmeyer
1977)

1969

Amarsipidae

Haedrich (1969)—aberrant stromateoid, currently valid, one genus/one species

1980

Hexatrygonidae

Heemstra & Smith (1980)—stingray with 6 gill slits, now 5 species

1981

Hexatreatobatidae

Chu et al (1981)—synonym of earlier-named family Hexatrygonidae

1981

Hispidoberycidae

Kotlyar (1981)—stephanoberycoid, one genus/one species

1983

Megachasmidae

Taylor et al. (1983)—large shark, feeds by straining, now several specimens

1986

Dinopercidae

Heemstra & Hecht (1986)— percoid, valid, now two genera

2002

Paraulopidae

Sato & Nakabo (2002)— deep-sea aulopiform, two genera, now four species

Deep-reef and upper-slope areas would appear to be the major habitats where we can expect new unusual
discoveries above the species level; however, this is not supported by the data (Table 9). There are no new
families from deep-reef and upper-slope areas. At the generic level, Richard Pyle (pers. comm. 9/2009)
reports only two potentially new genera from deep coral reef depths were discovered in recent years. Deep
coral reef species are related to shallow water species occurring above them. We conclude that it is unlikelythat many new families of marine fishes will be discovered.
Mid-water fish biodiversity. Midwater fishes are pelagic and are often widespread, occurring in morethan one ocean. Most spend their entire life off the bottom. Many migrate each night to shallow water, such as
lanternfishes (Myctophidae). Most mid-water fishes are small, less than 15–20 cm in length. Many studies on
mid-water fishes were published in the 1960s–1970s, with fewer studies published in recent years. Mid-water
trawls, especially closing nets, are most effective. A few, mostly benthic families contain mid-water species

MARINE FISH DIVERSITY

Zootaxa 2525 © 2010 Magnolia Press · 31

(e.g., Macrouridae, Liparidae). Mid-water fishes by selected orders or suborders show that descriptions of
new taxa have declined based on comparing the last 30 years and the last 10 years (Table 10).
Deep-living mid-water species that do not migrate to shallower waters are less well known than those thatcan be caught near the surface at night. More new taxa are expected as deep-ocean trawling continues, but theincrease in new taxa is correlated with sampling effort. A recent example (Prokofiev & Kukuev 2007) is therevision of the swallowerfish genus Pseudoscopelus (Chiasmodontidae). The first species was described in
1892, with 7 more by 1971, and new species described in 2005 (1), 2006 (3), and 2007 (5) for a total of 17
species. Prokofiev & Kukuev (2007) state, “…. the fauna of [chiasmodontids] of the Indo-Pacific is still
poorly known.” Obviously, the number of new deep mid-water species to be found is mostly dependent on
trawling operations by staff employed by universities and governments.
TABLE 10. Number of available and valid species-level taxa (species and subspecies) (1758-through 2009), and numberof species and subspecies described in the last 30, and last 10, years in selected families of mid-water fishes.
Order

FamilyAvailable

Valid

Last 30
Years

Last 10Years

Saccopharyngiformes

Cyematidae

3

2

0

0

Saccopharyngidae

16

10

6

1

Eurypharyngidae

6

1

0

0

Monognathidae

15

15

9

0

Gonostomatidae

52

33

5

1

Sternoptychidae

91

75

23

2

Phosichthyidae

29

24

6

0

Stomiidae

413

289

94

28

Neoscopelidae

10

6

0

0

Myctophidae

395

250

31

10

Caulophrynidae

10

5

2

0

Neoceratiidae

1

1

0

0

Melanocetidae

25

6

2

0

Himantolophidae

27

19

11

0

Diceratiidae

7

6

2

1

Oneirodidae

120

66

17

4

Thaumatichthyidae

9

8

2

1

Centrophrynidae

1

1

0

0

Ceratiidae

22

4

0

0

Gigantactinidae

28

23

16

1

Linophrynidae

47

29

7

1

Stephanoberycidae

5

4

1

1

Hispidoberycidae

1

1

1

0

Melamphaidae

67

54

13

10

Gibberichthyidae

3

2

0

0

Stomiiformes

Myctophiformes
Lophiiformes

Stephanoberyciformes

Indo-Pacific fauna. The highest number of marine fish species occurs in tropical areas of the Indian andwestern Pacific oceans east to the central Pacific, an area known as the Indo-Pacific faunal region. IndoPacific shore fishes are the subject of a classic paper by J. E. Randall (1998). Randall discussed endemism,
disjunct distributions, and species versus subspecies. Another important paper, by Springer (1982), defined
the Pacific Plate bioregion. Halas & Winterbottom (2009) provided information on the origin of the East

32 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

Indies coral reef biota, and defined the “East Indies Triangle” as islands between Sumatra in the west, New
Guinea in the east, and Luzon, Philippines in the north. This area has the highest marine biodiversity,
including fishes, in the world. The number of marine fish species declines with distance from this area, with
622 species in the Hawaiian Islands (Randall 2007) and 139 shore species at Easter Island (Randall & Cea inpress). Authors differ as to whether the so-called center of origin of coral reef species is the Philippines(Carpenter & Springer 2005) or West Papua, Indonesia (Allen & Erdmann 2009). Randall (1998) and Halas &Winterbottom (2009) suggested causes for why the East Indian region contains the most marine species. Allen& Erdman (2009) reported 1,511 coral reef species in 451 genera, including 26 endemic species at Bird’s
Head Peninsula, West Papua, Indonesia. Halas & Winterbottom (2009) tested four mechanisms proposed forthe origin of the biodiversity in the East Indies coral triangle, without finding empirical support for any one
mechanism.
A sub-set of the fishes of the Indo-Pacific region is the tropical coral reef fauna. Discoveries of new coral
reef species have declined, and it is difficult to find a new shallow coral reef species (Randall, pers. comm.
Jan. 2010). This does not apply to a few groups containing small species, such as some families in thesuborder Gobioidea (gobies and allies).
Tomio Iwamoto, a world expert on deep-sea fishes and the first author (W.N.E.) have identified areas
where more work is needed, especially the Indian Ocean, deep-sea ridges and seamounts, and in deep water
generally (mostly in the Southern Hemisphere). A statement in a recent paper (Iwamoto et al. 2009: 47) noted,“The recent captures in more than 3,000 m off Japan, … are additional examples of this lack of adequate
collecting at greater ocean depths.” Grenadiers (Macrouridae) are Iwamoto’s specialty, and he and co-workers
have described 93 species in the last 39 years (through 2009); 390 species are currently recognized as valid.
Museum collections. Most type specimens and nearly all major collections of marine fishes are housed inscience museums, either independent or associated with universities or governments (Table 11). These
collections form the primary source of material for taxonomic studies. The museums established earliest
contain the most type specimens. Museums more recently established tend to contain more holotypes andfewer syntypes. Full names for collections and more information on collections can be found in Fricke &Eschmeyer (2010). An important historical summary of fish collections, with information on most majorcollections, is provided in Pietsch & Anderson (1997) under the title “Collection Building in Ichthyology andHerpetology.” Of importance to biodiversity studies, many museums have now prepared type catalogs, many
type localities are now geo-referenced, and complete holdings may be online. Note however, that many
specimens in large museums are misidentified or identified with an old name—now a synonym of anotherspecies. Type specimens that were presumed to be lost or missing are being found, especially at major
museums such as the Natural History Museum (London). Specimens of many new taxa are in small, mostly
new, regional museums. In the Catalog of Fishes online (under Ichthyological Collections) information isprovided on museums containing fish specimens. Ichthyology has benefited from the easy access to
specimens; there are few large private collections. Many museums reflect primarily the focus of the institution
and the specialties of the curators. Type catalogs also provide historical information on individual collections.
These collections contain an enormous amount of biodiversity information.
Museums serve other functions. As habitats continue to become degraded or destroyed, species become
extinct, and restrictions are placed on collecting, the specimens in museums may be the only ones available
for study. An important role is the housing of voucher specimens, especially for molecular genetic studies.
When vouchers are not available, many conclusions made in the literature can not be verified; this isespecially true for barcoding and data in GenBank.
Environmental factors and geographic areas. The most important factor in marine fish distribution istemperature, and this is why we can define families as cold-water and warm-water. After temperature, habitatis probably next—with soft-bottom continental species, reef-associated species, brackish-water species, etc.
The data support the hypothesis that the new marine species are fairly well known in some areas, but new
species are abundant in other areas (Table 12). For example, there has been only two new marine species
described from Canadian waters in the last 40 years, but 38 from Brazil and 90 from Japan in the last nine
years.

MARINE FISH DIVERSITY

Zootaxa 2525 © 2010 Magnolia Press · 33

TABLE 11. Number of marine species and subspecies represented by primary type specimens (as of the end of 2009) inmajor museum fish collections.
Rank

Abbreviation

Holotype, Lectotype, Neotype

Syntypes

Total

1

USNM (Washington)

3613

543

4156

2

BMNH (London)

1720

1245

2965

3

MNHN (Paris)

2025

731

2756

4

AMS (Sydney)

855

267

1122

5

RMNH (Leiden)

741

381

1122

6

CAS/SU (San Francisco)

939

145

1084

7

ZMB (Berlin)

474

454

928

8

ANSP (Philadelphia)

725

84

809

9

MCZ (Cambridge)

475

311

786

10

ZIN (Moscow)

578

157

735

11

BPBM (Honolulu)

704

10

714

12

ZMUC (Copenhagen)

473

119

592

13

NMW (Vienna)

185

290

475

14

SAIAB (Grahamstown)

452

0

452

15

ZSI (Calcutta)

229

150

379

16

SMF (Frankfurt)

240

116

356

17

QM (Brisbane)

233

57

290

18

WAM (Perth)

259

3

262

19

FMNH (Chicago)

243

15

258

20

ZUMT (Tokyo)

209

18

227

21

ZMA (Amsterdam)

126

91

217

22

CSIRO (Hobart)

194

0

194

23

SIO (La Jolla)

194

0

194

24

LACM (Los Angeles)

186

0

186

25

SAM (Cape Town)

129

55

184

TABLE 12. Number of new marine species-level taxa (species and subspecies) described from selected geographic areasin 10-year periods. Number of species is based on their primary type locality.
Locality

1950–1959

1960–19691970–1979

1980–1989

1990–1999

2000–2009

North Pacific

12

6

22

24

8

24

Antarctic

2

15

11

25

11

12

North Atlantic

25

7

8

13

7

18

California

43

22

24

20

18

13

Australia

89

42

87

163

119

240

New Zealand

17

12

22

43

16

10

Canada

1

3

3

1

0

1

Brazil

12

20

18

12

10

38

China

11

35

29

78

23

26

Taiwan

12

26

16

23

26

43

Japan

132

50

65

101

53

90

34 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

Basin and area analyses. We also have examined two basins and one group of islands in more detail.
Significant effort was made by one of us (R.F.) to identify valid species and synonyms and endemics fromthese basins. Species described in the early years often had multiple and/or questionable localities. Many had
general localities. For example, a Linnaeus species might have six or more sources, and one of them may
have involved a Mediterranean Sea locality. If a primary type locality has never been established for the
species, it is impossible to include that species with certainty as one with a Mediterranean Sea type locality.
So those species are excluded from the analysis. If a species is a Mediterranean Sea endemic, and theMediterranean Sea was one of several original localities, then that species is included.
TABLE 13. Number of new marine species-level taxa (species and subspecies), number currently valid (as species), andnumber of authors, in 5-year periods, from the Mediterranean Sea.
Years

Species

Valid

Authors

Years

Species

Valid

Authors

1758–1763

88

81

1

1884–1888

26

7

7

1764–1768

17

5

3

1889–1893

18

8

6

1769–1773

101

1894–1898

3

02

1774–1778

8

02

1899–1903

10

1

1779–1783

101

1904–1908

7

3

5

1784–1788

20

9

2

1909–1913

11

1

6

1789–1793

43

8

3

1914–1918

4

2

3

1794–17983

03

1919–19235

1

5

1799–1803

81

3

6

1924–1928

14

5

10

1804–18086

3

2

1929–19334

0

4

1809–1813

307

69

6

1934–1938

9

1

6

1814–1818

24

11

6

1939–1943

1

0

1

1819–1823

39

8

8

1944–19482

02

1824–1828

164

14

7

1949–1953

7

4

6

1829–1833

97

16

15

1954–1958

5

0

6

1834–1838

63

17

11

1959–1963

7

4

6

1839–1843

69

6

12

1964–1968

5

2

3

1844–1848

52

9

11

1969–1973938

1849–1853

23

4

8

1974–1978

3

3

4

1854–1858

50

2

8

1979–1983

2

12

1859–1863

32

3

9

1984–1988

4

3

3

1864–1868

15

4

8

1989–1993

2

2

3

1869–1873

62

5

1994–19982

2

3

1874–187892

6

1999–2003

2

2

4

1879–1883

45

8

9

2004–2008

1

1

3

The Mediterranean Sea. The Mediterranean Sea is an extension of the northeastern Atlantic Ocean,connected through the Strait of Gibraltar. In the northeast, there is an extension of the Mediterranean Sea—theBlack Sea with the Sea of Azov, connected through the Dardanelles, the Sea of Marmara and the Bosporus.
The present treatment deals only with the Mediterranean Sea proper. The Mediterranean Sea is about 3,730
km long, at its widest point 1,770 km wide, and has a surface area of approximately 2.5 million km². Its
deepest point is at 5,267 meters.
The Mediterranean Sea has a warm temperate fauna, reaching from 45°45’N (in the Gulf of Trieste) to
30°16’N (Gulf of Sidra, Libya). The climate does not support the growth of reef-building corals, so the shore

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habitats are mainly dominated by rocks, sand, and seagrass and algae beds. The large deep-sea basins of thewestern and eastern Mediterranean Sea contain a depauperate, in a small part endemic, deep-sea fish fauna.
Due to considerable evaporation and limited freshwater inflow, the salinity is higher than in the adjacentnortheastern Atlantic Ocean, especially in the eastern parts. Geologically, the Mediterranean Sea wasconnected to the Indian Ocean as part of the Tethys Sea in the past, so it has in part a relict fauna of Indo-West
Pacific origin. The connection closed ca. 18 million years ago. Later, during the Messinian salinity crisis of 5–
6 million years ago, the entrance through the Strait of Gibraltar was blocked by a collision of continental
plates; the sea water evaporated, and the Mediterranean was reduced to large hyper saline lakes which were
uninhabited by fishes. The present-day fish fauna survived in the adjacent Atlantic or the Black Sea, andimmigrated after the end of that event. The temporary isolation of fish populations resulted in a relatively highpercentage of endemic species (41) with an endemism rate of 7.3% (of 563 species); the total number of
species is 646 with the 82 Red Sea immigrants included.

FIGURE 4. Number of new marine species-level taxa (species and subspecies), number currently valid (as species), andnumber of authors, in 5-year periods, from the Mediterranean Sea.

In 1869, the man-made Suez Canal was opened, connecting the Mediterranean Sea with the Red Sea as asea-level waterway. It extends from Port Said (southeastern Mediterranean Sea) to Suez (northern Gulf ofSuez, Red Sea), and its depth does not exceed 200 meters. The dominant current through the Suez Canal issouthbound, but only a few Mediterranean fish species (perhaps 15) migrated to the Red Sea, while a larger
number of species (about 80) occurring in the Red Sea immigrated against the current to the Mediterranean
Sea (Golani 1998, Galil 2008, Mavruk & Avsar 2008).

36 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

The fish fauna of the Mediterranean Sea has been known for a long time. Aristotle (384–322 B.C.) was
the first to study fishes at Lesbos Island (Greece), and published a natural history (Aristoteles 1454, Latintranslation). Through the Middle Ages there was little research, but in the Renaissance period two importantworks dealing with marine fishes were published, Rondelet (1554, southern France) and Salviani (1558,
Rome, Italy). The history of knowledge on Mediterranean Sea fishes was summarized by Artedi (1738);
Artedi’s monograph was used as a major source for formally naming fish species by Linnaeus (1758). Duringa Danish Expedition to the Mediterranean Sea and Red Sea, Forsskål (1775) described fish species from
France, Malta, Turkey and Egypt observed in 1761–1762 (Fricke 2008). In the early 19th century, several
important works dealt with marine fishes of the Mediterranean (Rafinesque 1810, Risso 1810, 1827). Later,
Mediterranean fishes were included in the worldwide fish catalogs by Cuvier & Valenciennes (1828–1850)
and Günther (1859–1870). The first comprehensive modern treatment of the ichthyofauna of theMediterranean was by Hureau & Monod (1973), later published as an illustrated catalogue with distribution
maps by Whitehead et al. (1984–1986). In a checklist of the fishes of Turkey, 367 native marine species werereported from the Mediterranean (plus 41 Red Sea immigrants) (Fricke et al. 2007). Iglesias (2009a, 2009b)
recently started to establish a preliminary catalogue on Northeastern Atlantic and Mediterranean Sea fishes,
which is available only as an electronic publication.
New species described from the Mediterranean Sea over time are shown in Table 13 and Figure 4. Very
few new species have been discovered in the Mediterranean since about 1900, with an average of one to three
new species per year since 1974. Major peaks reflect the collections described by expeditions and researchers
discussed above.
The Red Sea. The Red Sea is a marine water body, an extension of the northwestern Indian Ocean,connected through the Bab el Mandeb. In the north, there are two branches, the relatively shallow Gulf ofSuez and the deep Gulf of Aqaba. The Red Sea is part of the Great Rift Valley; it is about 2,200 km long, at its
widest point 355 km wide, and has a surface area of approximately 438,000 km².
The Red Sea has a tropical fauna; as it reaches north to 30°N, it is the world’s northernmost tropical sea.
The shore fish fauna is limited to narrow fringing reefs along most of the shorelines, except for extensive sandand mud bottoms in the north (Gulf of Suez) and south. The main Red Sea and the Gulf of Aqaba contain adepauperate fish fauna, including endemic deep-sea species. Due to heavy evaporation and nearly no
freshwater inflow, the salinity is higher than in the Indian Ocean, especially in the northern parts. The Red Sea
was probably temporarily isolated from the Indian Ocean during glacial periods. This resulted in a highpercentage of endemic species (159 species, endemism rate 13.6%). Isolation of the fauna probably occurredat least twice (Winterbottom 1985) so there appear to be two periods of endemism. The total number of Red
Sea species of fishes numbers 1,188, including 15 immigrants from the Mediterranean Sea.
The peaks in Figure 5, based on data in Table 14, reflect collecting expeditions in the Red Sea (for history
see Fricke 2008). The marine fish fauna was mostly unknown before a Danish expedition with Peter Simon
Forsskål as naturalist (1732–1763). He collected fish specimens mainly in Saudi Arabia and Yemen, whichwere posthumously described by Forsskål (1775). In the 19th century, personnel from three German museums
explored the fish fauna of the Red Sea. The Berlin museum (ZMB) sent Friedrich Wilhelm Hemprich (1796–1825) and Christian Gottfried Ehrenberg (1795–1876) who collected specimens which were later described by
Cuvier & Valenciennes (1828–1850). The Senkenberg Museum in Frankfurt (SMF) received specimens
collected by Eduard Rüppell (1794–1884) which were described by Rüppell (1828–1830, 1835–1838). Later,
the Stuttgart museum (SMNS) sent Martin Theodor von Heuglin (1824–1876) and Carl Benjamin Klunzinger(1834–1914) to the Red Sea; Klunzinger described several new fish species and published the first catalogueof Red Sea fishes (Klunzinger 1870, 1871, 1884). More recently, Red Sea fish checklists were published by
Dor (1984) and Goren & Dor (1994), the latter listing 1,248 fish species as valid. Subsequently, several
additional species were recorded (e.g. Randall 1994, Khalaf & Zajonz 2007), or described as new. A recent
updated checklist is Golani & Bogorodsky 2010.
The Mascarene Islands. The Mascarenes are a group of volcanic and coralline islands in thesouthwestern Indian Ocean, east of Madagascar. The three main islands are Réunion (an overseas
Département of France), Mauritius and Rodrigues (both part of the Republic of Mauritius). The Mascarene

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fish fauna was listed by Fricke (1999; for history see p. 4–6). The Mascarenes were first discovered by
Portuguese in search of fresh water and food supplies on the route to India, and then ruled by France since1638 (Mauritius was given to Britain in 1814). The most important early explorer of the islands was Philibert
Commerson (1727–1773), who participated in Bougainville's expedition around the world in 1766–1768, andthen stayed in Mauritius, where he died in 1773. Commerson described and illustrated numerous Mascarene
fishes; the manuscripts were sent to Paris, where the fish species were posthumously described by Lacepède(1798–1803) and Cuvier & Valenciennes (1828–1850). Regional checklists were published by Heemstra et al.
(2004, Rodrigues) and Fricke et al. (2009, Réunion). Fish species occurring at the Mascarenes have both
continental and insular Western Indian Ocean relations. A total of 1,183 species occur in the Mascarenes; 45
species are endemic (3.4 %). New species described from the Mascarene Islands over time are shown in Table
15 and Figure 6. The peaks in Figure 6 mainly represent (from left to right) descriptions by Lacepède ('1800
peak'), Bennett and Cuvier & Valenciennes ('1833 peak'), Günther ('1863 peak'), Bleeker, Sauvage and Bliss('1883 peak'). The '1963 peak' mainly represents French explorations in Réunion. Recent descriptions are due
to SCUBA diving explorations and rotenone collections.
TABLE 14. Number of new marine species-level taxa (species and subspecies), number currently valid (as species), andnumber of authors, in 5-year periods, from the Red Sea.
Years

Species

Valid

Authors

Years

Species

Valid

Authors

1758–1763

2

2

1

1884–1888

20

7

3

1764–1768000

1889–1893212

1769–1773

000

1894–1898

5

4

1

1774–1778

128

89

2

1899–1903

52

4

1779–1783

101

1904–1908

101

1784–1788

601

1909–1913000

1789–1793

17

4

2

1914–1918

4

4

2

1794–17982

1

1

1919–1923

1

1

1

1799–1803

25

0

4

1924–1928

15

5

1

1804–1808000

1929–1933521

1809–1813000

1934–1938

4

2

6

1814–1818402

1939–1943

3

2

2

1819–1823

000

1944–19480

0

0

1824–1828

21

11

3

1949–19538

5

7

1829–1833

139

64

6

1954–1958

16

4

9

1834–1838

124

56

3

1959–19638

7

4

1839–1843

30

4

4

1964–1968

14

9

10

1844–18488

1

4

1969–1973

20

13

19

1849–18532

03

1974–1978

34

25

18

1854–1858622

1979–1983

39

32

25

1859–18634

02

1984–1988

25

25

21

1864–18684

0

3

1989–1993

12

12

16

1869–1873

69

27

4

1994–1998

12

11

10

1874–1878

20

4

4

1999–2003

8

8

9

1879–18834

02

2004–2008

6

6

10

38 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

FIGURE 5. Number of new marine species-level taxa (species and subspecies), number currently valid (as species), andnumber of authors, in 5-year periods, from the Red Sea.

Endemism. The concept of endemism as used here is a traditional use in descriptive biodiversity—ageographical area with taxa (typically species, or genera and species) that occur only in that area. It is
generally assumed that the taxa evolved there. Considerations of ancestral species, of extinctions, range
contractions and expansions that may have occurred are not addressed. Phylogenetic hypotheses for numerous
clades usually are not available to corroborate congruent patterns of distribution to test for vicariance ordispersal. A more rigorous cladistic approach is suggested by Harold & Mooi (1994).
In marine fishes, endemic genera have received less attention than other taxonomic groups, and they are
typically not mentioned in faunal studies. On the Biodiversity Hotpots website of Conservation International
under "endemic genera" (http://www.biodiversityhotspots.org/, accessed 25 Jan. 2010), it is stated that "Whenassessing biodiversity data, it is important to consider not only species distributions but also distribution ofhigher taxa." On that site they provide a table of endemic genera from selected geographic areas for plants,
mammals, birds, reptiles, amphibians and freshwater fishes. They concluded that, “Unsurprisingly though,regions with high species endemism also have greater levels of endemic genera.” That conclusion is notsupported by marine fish insular faunas, such as those of Easter Island, the Galapagos Archipelago, theMascarenes and the Hawaiian Islands, but it is supported by continental areas, such as southern Africa andsouthern Australia. Typically, island groups or small coastal areas are used for assessing endemism in marine
species. The presence of endemic genera (and species) involves many complex issues, such as geological ageof the island group or coastal area, its isolation and location, arrival of ancestral progenitors, competition,
suitable habitat, and environmental stability over time.

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TABLE 15. Number of new marine species-level taxa (species and subspecies), number currently valid (as species), andnumber of authors, in 5-year periods, from the Mascarene Islands.
Years

Species

Valid

Authors

Years

Species

Valid

Authors

1758–1763

000

1884–18883

2

1

1764–1768000

1889–1893

22

1

3

1769–1773

1

1

1

1894–1898

1

0

1

1774–1778000

1899–1903

4

1

2

1779–1783000

1904–1908201

1784–1788

3

1

1

1909–1913

101

1789–1793

101

1914–1918

5

12

1794–1798

3

3

1

1919–19232

21

1799–1803

72

31

4

1924–19281

0

2

1804–18085

03

1929–19331

02

1809–1813000

1934–1938

2

1

2

1814–18183

12

1939–1943

0

0

0

1819–1823

2

2

2

1944–1948

1

0

1

1824–1828

21

9

6

1949–1953

3

1

2

1829–1833

118

51

8

1954–1958

2

0

1

1834–1838

46

16

9

1959–1963

10

3

5

1839–1843

45

14

6

1964–1968

3

2

2

1844–18488

2

3

1969–19731

1

1

1849–18533

1

3

1974–1978

12

12

10

1854–1858

11

4

2

1979–198388

9

1859–1863

33

10

4

1984–1988

5

5

6

1864–18688

3

5

1989–1993

4

4

6

1869–18738

24

1994–1998

7

7

8

1874–1878

16

5

8

1999–2003

11

10

10

1879–1883

27

9

4

2004–2008

7

7

6

For the coastal area of southern Australia (Victoria to southern Western Australia), it is suggested here
(W.N.E.) that this area of reasonable size has the highest number of endemic marine fish genera anywhere(over 50)—this constitutes a marine hotspot; it is also rich in endemic species (see Gomon, et al. 2008). In the
broad Indo-West Pacific tropical fauna, there are relatively few restricted endemic genera because mostgenera are now widespread. For marine fishes, we have counts of endemic genera of selected island groups(authority in brackets): Hawaiian Islands [Randall, Mundy] (1 genus), Fiji [Greenfield] (0), Mediterranean
Sea basin [this study] (5), Red Sea basin [this study] (4), Easter Island [Randall] (1), Mascarenes [this study](1), Galapagos Archipelago [McCosker] (1), southern Africa [Heemstra] (36). Our count ofsouthern Australian endemic genera is over 50, with additional "non-endemic" genera having representatives
in New Zealand or northern areas of Australia (counted from Gomon et al. 2008). The area extends from
Victoria to southern Western Australia. South Africa is more complex with important considerations of ocean
currents and water temperature, but marine endemic genera in South Africa (Namibia around to Beira
Mozambique) number 36 (P. Heemstra, pers. comm., 26 Jan. 2010, and see Heemstra & Heemstra 2004).
New Zealand is another area with many endemic genera and species.
A very detailed analysis of the Australian fish fauna was published in 2006 (Hoese & Paxton 2006). The
total fish fauna in 2006 was about 4,500 described species (all habitats), and overall, 24% of the Australianfish fauna was endemic, with substantial new discoveries still anticipated.

40 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

FIGURE 6. Number of new marine species-level taxa (species and subspecies), number currently valid (as species), andnumber of authors, in 5-year periods, from the Mascarene Islands.

For endemic marine species, there have been estimates for certain island groups and areas, but the criteria
used vary—such as distance from shore, depth of capture, or the exclusion of widespread pelagic species
or deep-sea species that may be found near shore. An approximation for shallow water species is about 12 to
20% endemic species in such isolated areas as the Galapagos Archipelago, Hawaiian Islands and Easter
Island. The number of endemic species is low in less isolated areas, such as Fiji with 15 (Greenfield, pers.
comm., 1/27/2010) and a large majority of marine Indo-Pacific tropical reef species are widespread. Species
living on mud bottom areas, for example, are often not as widespread.
There is a temperate Southern Ocean fauna of mostly mobile species, with many occurring in Southern
Africa, southern Australia and the southern end of South America and southern islands. The fauna off Chile is
not well known, but is not extensive, partly because of its extreme southern location and lack of suitablecurrents, and it also reflects insufficient ichthyological exploration. Eschmeyer & Hureau (1971) showed thatthe eastern Pacific tropical area was a major barrier (nearly 100%) for movement of temperate faunas acrossthe eastern Pacific tropics, however they noted that one to three species of the North Pacific genus Sebastes
and a poacher (family Agonidae) crossed the tropics to Peru and Chile, and their closest relatives appeared tobe the most southern-living species in the North Pacific off California and Mexico. It was suggested that thiswas a fairly recent extreme anomaly of short duration, allowing minimal crossing by some fishes andinvertebrates. Movement of the Southern Ocean fauna in an easterly or westerly direction is facilitated by astable temperature regime and strong currents. Eschmeyer & Poss (1976, fig. 10) showed that if the 20 degree
isotherm shifted to the present 25 degree isotherm during a glacial period, a significant number of islands andnear-surface seamounts were available as “stepping stones” across the south Pacific.
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Contribution of molecular studies to biodiversity knowledge of marine fishes. Most fishes have many
external characters useful in taxonomy. It appears that most marine species (especially larger ones and adults)
are defined accurately, based on different researchers reaching the same conclusions and with consensus overtime. DNA studies are used in marine fish taxonomy for two primary purposes: defining higher categories
and relationships of taxa and defining species and populations. Of particular interest here are studies thatreveal biodiversity at the species level.
Molecular studies can be useful to evaluate color differences between populations that are otherwise
indistinguishable morphologically. Molecular studies clearly are useful in discovering species that, afterinvestigation, may have demonstrable morphological differences. Colborn et al. (2001) showed theworldwide bonefish (genus Albula) actually to be composed of several species. There are other warm-water
widespread non-reef species that are good candidates for molecular study, such as carangids. See also Zemlak
et al. (2009).
Many molecular studies could be mentioned, but Baldwin et al. (2009b) is an excellent representative
study. They reviewed the western Atlantic species of marine Coryphopterus, a genus containing small
species, many of which are similar to one another. The study was conducted in Belize, were nearly all the
species occurred. They collected fresh specimens, and took digital color photographs of young and adultspecimens. DNA barcoding (mitrochondrial cytochrome c oxidase I [COI] sequences) revealed distinct
genetic lineages. They used neighbor-joining trees and genetic distance matrices. Specimens were depositedin the U. S. National Museum, and DNA samples were placed in GenBank. They stated (p. 134), "Resolving
the identities of many Caribbean Coryphopterus in the absence of the DNA data would have been extremely
difficult." Morphological characters and color differences were used to define the species. Two of the authors
had classical training in systematic ichthyology. A carefully-prepared key to the western Atlantic species wasprovided. Using these techniques, they synonymized one recently-described species and provided justification
for recognizing up to three others. They did not discover any new species, but defined 12 valid species.
Often, there are names that are available for species that were regarded as synonyms but are now regarded asvalid species; that knowledge increases biodiversity valid species counts but not new species taxa. Moreinformation on barcoding and the “Barcode of Life” project can be found at www.fishbol.org/
www.barcodinglife.org.
A second use of the barcode approach in marine fishes is to confirm species identification, especially of
early life history stages (see Baldwin et al. 2000a). In the six species studied, identification of larvae andjuveniles was problematic because characters used to distinguish similar adults were absent or incomplete inyoung stages. Using barcode techniques along with chromatophore patterns recorded by digital photography,
they were able to link young with adults and provide new characters to distinguish the species at all life stages.
Barcoding techniques are still being evaluated, and Dasmahapatra et al. (2010) suggest that DNA barcoding
detects some “species” that are real and some that are not.
A recent example of a phylogenetic study at higher taxonomic levels using DNA sequence data is that ofThacker (2009); new ideas were supplemented by morphological character analysis.
Unusual fish families/subfamilies. The discovery of biodiversity also depends on the biology of fishes,and some families will show many new species. Family Liparidae: Fishes of this family occur in shallow
water and in the deep sea. They are small in size (most less than 15 cm long). Deep-living species place theireggs inside other animals (such as bivalve mollusks). They have a few large eggs and no larval stage, so they
are not limited by surface temperatures and are not transported by surface currents. Consequently, they areable to move freely in the deep sea. They have few “morphological characters” but it is assumed they are
distinguished accurately. Many species are known only from type specimens. The deep-living species
apparently have small ranges, but this environment is poorly collected. Many new discoveries will continue,
and two of the largest genera of marine fishes are in this family. The total species in the family Liparidae thatare considered as valid as of 2009 is 370, and 202 have been described since 1980. Family Ogcocephalidae(batfishes): H. Ho (pers. comm. 15 Nov. 2009) indicates that in his Ph.D. dissertation on ogcocephalids of the
Indo-Pacific, valid species will number about 65, with about half being new. Ogcocephalids are peculiar,
small bottom-living species occurring both in shallow and deep depths. Many tend to live on deep-sea rough42 · Zootaxa 2525 © 2010 Magnolia Press

ESCHMEYER ET AL.

bottom habitats that are difficult to sample. Subfamily Symphurinae (tonguesoles): The smallest and flattestof the flatfishes. They are difficult to sample and occur widely in the deepsea. Twenty species have been
described in the last 20 years, and all in the genus Symphurus. Many new species are expected.
Deep-reef fishes (50–500 m). Deep-reef species are defined as those fishes which are associated withcoral reefs and live below about 50 meters to a depth of about 500 meters; the fauna changes at about 500
meters to more deeper-living families. Richard Pyle is the most experienced person collecting fishes at these
depths, currently using re-breather SCUBA technology. He suggests (pers. comm.) that assemblages on deep
reefs show comparatively low distributional overlap (between different island groups and between sites withineach island group).
There are many deep reef genera of fishes than can be expected to contain many new species. For
example, a recent paper on species of Plectranthias (Serranidae) by Heemstra & Randall (2009) in thewestern Indian Ocean, describes one new species, but it contains the surprising statement that "…. most
Plectranthias species are rare in fish collections, and twenty of the 45 valid species are known from only one
or two specimens.” The authors note that species found in depths of 20–300 meters are solitary and sedentaryand live on rugged deep-reef habitats in 50 to 300 meters.
These deep-reef areas will be a significant source of new marine fish species. We do not know at this time
how widespread these deep-reef species are, but many new taxa will be found, especially when newtechnology is available (such as collecting from one-person submersible vehicles). Also deep trapping andlong-lining should be successful. Fishes from deep reefs are probably less well known than are the fishes inthe deep sea.
Deep-slope fishes. Deep slope includes depths from about 50 to 1000 or more meters, but not coralassociated species. This is another area where we expect many new species to be discovered.
An example of an isolated insular fauna with deep-slope species is the Galapagos Archipelago. The
Galapagos Islands are in the eastern Pacific Ocean. They are volcanic, and lie about 1000 km from themainland. The two closest islands are Malpelo (450 km ENE) and Cocos (630 km NE). To the west lies theEastern Pacific Ocean barrier. Sea temperatures and habitats vary widely. The fauna reflects primarily the
species that arrived on currents. The fishes of the Galapagos Islands have been analyzed in great detailrecently (McCosker & Rosenblatt, in press), and see also Grove & Lavenberg (1997). The addition of new
deepwater species gave a total of 75 near-shore and deep-slope species that were unique to the Galapagos.
They calculated that the rate of endemism was 13.6%, with a total of 550 species (not including mesopelagic
species). They state that the shorefishes to a depth of 60 m are fairly well known. Nearly half of the Galapagos
fish species are shared with the Panamic fauna to the east, 15.2% are found worldwide in the tropics and 15.9
% are Indo-Pacific shore fishes. Some are recorded as vagrants. There is one putatively endemic genus. They
predict that deep-slope and benthic collections below 500 m will make numerous remarkable discoveries.
Diving and collecting in deep-sea submersibles resulted in the discovery of 32 undescribed fish species
and many new records occurring on deep slope habitats (McCosker & Rosenblatt, in press). They collectedfrom rocky, nearly vertical, habitats and others that are difficult to sample. They were able to collect, film, andobserve from 60–1000 meters, and they adapted new techniques for poisoning, trapping and underwater hookand-line fishing. Use of submersibles has wide potential for future research, and use of this technology is
discussed in some detail by McCosker & Rosenblatt (in press).
The important unanswered question is how widespread are deep-slope species? We do not have thisinformation, an important unknown for estimating undiscovered marine fish biodiversity.
Collecting methods. Most collecting methods have changed little over time. New methods include use ofchemicals, scuba, re-breathing scuba and submersibles. Shallow water: Reef species are taken by hand,spear-guns, hand nets, etc., often with the use of immobilizing chemicals such as rotenone. Rotenone has long
been the choice for reef fish collecting (Smith-Vaniz et al. 2006) but has become somewhat controversial (seeRobertson & Smith-Vaniz 2008); clove oil has recently come into use (see Robertson & Smith-Vaniz 2010).
Dynamite was sometimes used in early years. Shore collections are made by hand, from specimens washedashore, use of small nets in association with chemicals, traps, and other methods. A variety of seines, dredges,
and trawls are used. In early years, specimens were often purchased in fish markets or directly fromMARINE FISH DIVERSITY

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fishermen. Fish market specimens still provide new species, most recently from Taiwan. Lights suspended in
the water at night attract specimens that can be caught with dip-nets. SCUBA: The invention of the demand
regulator by Emile Gagnan and Jacques Cousteau in 1942 was the beginning of a new era for underwater
exploration. Sixty-five years later, sales of SCUBA equipment in the U.S.A. totaled $804 million dollars. Thisnew technology enabled marine biologists to spend long periods of time underwater collecting and observing
fish in their natural habitat. Pure air easily allows divers to spend about 45 to 60 minutes at a depth of about 60
feet. “Rebreathers,” using different gas mixes allow technical divers to dive very much deeper and spend
longer times underwater. A closed circuit rebreather uses 100% of each breath and completely re-circulates
the breathing gases. It removes the carbon dioxide that has been generated and replenishes the oxygen.
Because there is no exhausting of gases there are no bubbles and virtually no noise. This is a non-invasive
method of collecting fishes. Submersibles: The use of submersibles in deep slope areas is discussed in
McCosker & Rosenblatt (in press) and is treated above. Use of this gear resulted in discovery of over 30 new
deep slope species at the Galapagos Islands (about 5% of the fauna). The submersibles allow use ofichthyocides (chemicals to stun fish), moveable collecting arms, and the ability to bring specimens to thesurface for study. Progress is being made on developing one-person submersibles. Traps, long-lines: Baited
traps can be used effectively at deep depths; two new genera were captured recently in about 1000 m by thismethod (Matallanas 2009). Depth is apparently limited when floatation/release mechanisms are used. Longlines have long been used to collect fishes, especially wide-ranging pelagic ones for many years, and the useof deep-set near-bottom long-lines is effective for finding new species, especially sharks and rays. Trawling:
Bottom and mid-water trawls are the typical way of collecting offshore species. They can be used at any
depth. Mid-water nets that can be closed before retrieval provide accurate bracketed depth ranges of captures.
Specimen illustration and photography. The recording of images of new specimens in publications haschanged dramatically through the years. In earliest times, especially before the early 1800s, types were notillustrated at all or they were only shown by simple line drawings. Artists accompanied the great exploring
expeditions in the mid-1800s, and paintings were often made of fresh specimens. Laboratory line drawings ofhigh quality were the norm for expeditions and collections into the 1900s and forward. Some publications
that were illustrated in color were produced by artists painting each color illustration by hand. They used one
reference original so each plate differed slightly from the others. Most of these original illustrations are stillavailable in natural history museums.
Pen-and-ink and pencil drawings, some involving special rendering surfaces, were typical from the
middle 1800s to late 1900s. The recent use of digital cameras to illustrate specimens in lateral view has
virtually obviated the need for drawings and has replaced traditional film photography for illustrating
specimens. Pen-and-ink drawings are still used to show special features.
Specimen photography. Specimen photography of “live colors” was perfected by John Randall, using atechnique of pinning erect the fins of specimens, usually on the evening of capture. The median fins were
"painted" with formalin to “fix” them in an erect position; the pins were removed and color photographs werethen taken in lateral view. Type specimens for hundreds of publications of new marine taxa in the last half ofthe 1900s were prepared in this way. Smaller specimens were often photographed in water-filled tanks withthe specimens positioned by glass plates. Use of antioxidants to preserve color (see Waller & Eschmeyer1965) was successful for preserving only some colors, and freezing and reducing exposure to light are shortterm solutions only. Digital photography has virtually replaced whole specimen drawings, and current
publications tend to have both posed and in situ photographs. The resolution of digital photography allows
for close-up photographs of specific parts or areas of specimens.
Underwater photography. Underwater photography has simplified the work of ichthyologists andhelped in the discovery of new species. Underwater photography has come far since the early Calypso camera
was introduced in 1960. This camera was the first 35-mm self-contained underwater camera. The Calypso
was conceived by Jacques Cousteau, designed by Jean de Wouters and manufactured by Atoms in France.
Production was taken over by Nikon, and the camera was sold as the Nikonos. The Nikonos became a wellknown and accepted series of cameras. The Nikonos was not an SLR camera, and it had to be manuallyfocused. The quality of pictures taken using these cameras was generally very poor. Housed 35-mm SLR

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cameras were a major breakthrough and enabled photographers to consistently take excellent pictures
underwater. They came with the disadvantage that only 36 pictures could be taken on each dive. Processing
the film was costly and normally not possible in remote locations. The advent of digital cameras has
revolutionized all photography and has made underwater photography available to the masses. Digital SLR
cameras in underwater housings overcome the disadvantages of film cameras. It is now possible to take inexcess of 400 pictures during one dive, the number being limited more by the dive time than the technology.
The results can be viewed immediately and adjustments made if necessary. Today many new species are being
photographed first, identified as a potential new species and then collected. The posting of photographs on theinternet by amateur divers is now an important source of information on possible new taxa, live color, and forenlarging the known ranges of species.
Preservation of specimens. Preservation of specimens has changed through time. One of the earliest
methods was to prepare dried skins. Specimens were typically mounted and stuffed, often for subsequentmuseum display. Sometimes "half-skins" (one side of a specimen) were prepared. The use of "spirits" forpreserving fishes was also used in the 1800s. The early museums (e.g., London, Vienna, Paris, Leiden) haveboth dried collections and spirit collections. Nearly all of the dried specimens are still available today. By the
1900s and thereafter, specimens were first placed in formaldehyde [termed formalin when diluted with water]
to stop all decay. Specimens were subsequently rinsed and then placed in ethyl or isopropyl alcohol for longterm storage. Additional techniques to preserve and study specimens became available more recently.
Skeletons were prepared by placing a fish carcass in a dermestid colony; the insects consume the soft tissues
leaving the bones free of flesh for study. Radiographs (x-rays) were useful to study internal parts. Staining
bones, cartilage and nerves in different colors was possible, and this was then accompanied by digesting themuscle tissue in whole specimens (termed clearing and staining); this was initiated by Taylor (1967), and asummary of these techniques is given by Song & Parenti (1995).
How many marine species are there? One of the goals of this analysis was to make a prediction of themarine fish biodiversity that remains to be discovered and documented. It is by analysis of the tables andgraphs—and informed conjecture—that we arrive at a value. We do have fairly complete knowledge ofcommercial food fishes, as well as others used in commerce, such as in the aquarium hobbyist trade. We know
most of the dominant shore species, and we have a good understanding of many deep sea families. We also
have shown that new marine fish families will be very few, and new genera based on unique discovery of new
species will be modest.
As discussed above, we assume that nearly all species in the Northern Hemisphere, including theMediterranean Sea and Red Sea basins, have been found. Some additional species are expected from thenorthwestern Pacific, some deep-sea areas in the Middle Atlantic Ridge and the Bering Sea area. The number
we expect to be discovered is perhaps only 200–300.
We anticipate some new shallow-reef species, but mainly in families with small-sized species, such as
gobies and blennies, but most shallow-reef species have been found. We place that estimate at about 300 to
500. Deep reef species, however, are poorly known. Estimates as high as 1,500 yet-to-be-discovered species
have been made (Pyle, 2005, abstract, 7th Indo Pacific Fish Conference, based on 30% above about 5,000
known shallow-reef species). Similarly, deep slope species are poorly known. More than 80 new marine
species of sharks and rays have been described from Australian waters since 2005 (Last & Stevens 2009), andmore new species are being discovered in the Indo-Australian Archipelago. If the work in the Galapagos
Archipelago is representative, then the increase of deep-slope species will be on the order of 10–15% newtaxa—perhaps another 2,000 or more, however it is unknown how widespread deep slope species are.
Deep-sea species will continue to be found, especially in the Indian Ocean and the southern Pacific.
Collecting in the deep sea is expensive, and this endeavor will continue for decades. We suggest that 800
deep-sea species remain to be described. It is anticipated that certain geographical areas will be most fruitful,such as southern Australia, South Pacific island areas and Indonesia.
Add to this the new taxa that may be revealed by molecular studies. A recent publication by Zemlak et al.
(2009) shows interesting results for populations sharing a presence in South Africa and Australia. Theysuggest that sequence divergence patterns identify species-level differences between populations for a large

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number of species. They only studied populations at these two geographic extremes, and did not consider
populations of these species in other areas. One of their pairs is known to involve misidentified specimens (W.
Smith-Vaniz, pers. comm., 8 Mar. 2010 after communication with the persons making the identifications).
Mora et al. (2008) estimate that 21% of marine species remain to be discovered. This seems too low for
the same reasons that limit our ability to make a good prediction using our approach. We are in agreement in
the stage of completeness for some areas, such as shallow-reef-associated species and pelagic species. We
suspect that their estimate for deep-sea species is a little too high; the deep-sea has been poorly sampled asthey suggest, but many species are widespread. However, besides the deep sea, they do not mention the two
habitats that we (W.N.E.) believe will provide the most new species in the next 20 or 30 years—deep-reef
tropical areas and deep-slope habitats. In spatial terms, the shallow areas above these habitats have beensampled adequately, but deeper collections in the same area have not been sufficiently sampled. On ageographically-based special resolutions system as used by Mora et al., it would seem to be somewhat
difficult to separate adjacent habitats.
A reasonable estimate would be at least 5,000 new species awaiting discovery in the next four decades, oran increase of about 30%. We can assume a success rate of at least 90 to 95% of described versus valid
species. In the last 20 years (1990–2009), 2,701 new species of marine fishes have been described based on
Catalog of Fishes entries. Of these, over 97 % are recognized as valid. The number of new marine species
actually shows an increase (average 119 per year for 1990–1999 and 151 for 2000–2008). There is no reason
to doubt that 2,500 new species will be described in the next 20 years, and 2,500 more in the 20 years
thereafter.

Acknowledgments
The basic text was primarily prepared by Eschmeyer, with basin descriptions and analysis by Fricke. Fong andPolack assembled and proofed information for the tables and graphs; Fong prepared the tables and graphs andPolack checked for data accuracy and prepared some text sections. Fong also served as the programmer for
the Catalog of Fishes project for the last 10 years.
We thank Jesse Ausubel and the Alfred P. Sloan Foundation for a grant to assist in preparation andpublication. This is a contribution to the Census of Marine Life and its Ocean Biogeographical Information
System (OBIS). Many persons have participated in the Catalog of Fishes database, and we thank them all.
Rainer Froese (Fishbase) co-prepared a presentation by W.N.E. in 1999 of similar information for all fishes,and Froese was responsible for the design of some graphs that were modified for use in this work. Stan Blum,as the California Academy of Sciences’ informatics specialist helped with technical details, as did librarians,
administrators, the computer department, and other staff at the California Academy of Sciences.
The development of the Catalog of Fishes database was supported in many ways through the years by
funding from granting agencies. Initial progress was mainly with the help of funds from the U.S. National
Science Foundation. The California Academy of Sciences has supported the project for over 25 years.
Discussions and email exchanges with ichthyologists were helpful in the preparation of this paper. Many
sent specific information, and we thank Gerry Allen, Diane Bray, Carole Baldwin, George Burgess, Tom
Fraser, David Greenfield, Martin Gomon, Phil Heemstra, Hans Ho, Doug Hoese, Tomio Iwamoto, John
McCosker, Bruce Mundy, John Randall and Richard Winterbottom. Many other assisted, included Edward
Clark, Jill Johnson, Alan Leviton, Rob Robins and Alfred Thompson.
We are grateful for comments and suggestions on an early draft by Richard Winterbottom, Bill SmithVaniz and Derek Tittensor. Robert Lea generously reviewed all, and Lloyd Findley part of the manuscript, andthey made many useful suggestions. Larry Page carefully edited the manuscript, and we thank him for his
valuable assistance.

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