The plates in holbrook's ichthyology of south carolina, 1860.

Abstract

Best known for North American Herpetology, John Edwards Holbrook (1794-1871) also published several ichthyological works, the most important of which was his Ichthyology of South Carolina (18556, 1860). Because of their rarity, reproductions of the plates from the 1860 edition are presented herein, along with comments on the fishes illustrated, Holbrook's artists, his other ichthyological work, and his herpetological contributions. In order to put his efforts in perspective, I also include a biographical sketch of Holbrook.

Zusammenfassung

John Edwards Holbrook (1794-1871), der hauptsachlich durch seine North American Herpetology (Nordamerikanische Reptilienkunde) bekannt wurde, hat auch mehrere ichthyol-ogische Werke geschrieben, vor allem eine aber die Ichthyology ofSouth Carolina (1855b, 1860) (Fischlcunde Sad-Carolinas). Weil das Buch selten zu bekommen ist, werden hier die Tafeln der Ausgabe von 1860 wiedergegeben, verbunden mit Erlauterungen zu den abgebildeten Fischen, den beteiligten Kansdern, Holbrooks anderen fischkundlichen Arbeiten und seinen reptilienkundlichen Beitragen. In Wardigung seiner Lebensleistung wird auSerdem eine biografische Slcizze Holbrooks angefagt.

Resume

Surtout connu pour son North American Herpetology, John Edwards Holbrook (1794-1871) a aussi publie plusieurs ouvrages ichtyologiques dont le plus important est son Ichthyology of South Carolina (1855b, 1860). A cause de leur rarete, des reproductions des illustrations de l'edition de 1860 sont presentees ici, avec des commen-taires sur les poissons representes, les artistes d'Holbrook, le reste de son travail ichtyologique et ses contributions herpetologiques. Min de mettre son oeuvre en perspective, j'ai aussi indus une esquisse biographique d'Holbrook.

Sommario

Meglio conosciuto nell'erpetologia del Nord America, John Edwards Holbrook (1794-1871) pubblico anche diversi lavori nel campo dell'ittiologia, il piu importante dei quali fu Ichthyology of South Carolina (1855b, 1860). A causa della rarita di questo testo, sono proposte in questo articolo le riproduzioni delle tavole dell'edizione del 1860, corredate dei commenti sui pesci illustrati, opera di Holbrook, e dagli altri contributi ittiologici e erpetologici dell'autore. Al fine di inquadrare il suo impegno in una dimensione storica, si include anche un profilo biografico di Holbrook.

INTRODUCTION

Most of the early work on the cold-blooded vertebrates of North American was done by Europeans or in some cases by Americans who studied relatively small geographic areas, but this would change. A notable exception to such investigations by Americans was William Bartram's Travels (1791), which was based on field work in a large part of what is now the southeastern United States. More Americans became interested in the scientific study of the biota of their homeland in the years between 1790 and 1830, resulting in more published works on various aspects of natural history. John Edwards Holbrook's efforts in the middle third of the nineteenth century were among the most important contributions made by anyone in those years to an understanding and appreciation of the enormous variety of native plants and animals living on the continent of North America. The publication of Holbrook's North American Herpetology (1836-1840, 1842) signaled the beginning of a new day in the study of amphibians and reptiles in the United States, and his detailed descriptions and excellent illustrations in the Ichthyology of South Carolina, though geographically limited in scope, were harbingers of things to come in the world of fishes (Worthington & Worthington, 1976).

HOLBROOK

John Edwards Holbrook (Fig. 1) was born in Beaufort, South Carolina, his mother's home, on 30 December 1794. Holbrook's father, Silas (17681800), left his home in Wrentham, Massachusetts, to teach school in Beaufort, a coastal town 50 miles southwest of Charleston, where he met his future wife, Mary Edwards (?-1826), daughter of John Edwards, an immigrant from Wales. A short time after the birth of their second son, Silas (17961834), the Holbrooks moved to Wrentham, where John Edwards' mother remained after her husband's death in 1800. Holbrook received his early education from a local preparatory school, graduated from Brown University in 1815, and received his M.D. degree from the University of Pennsylvania in 1818. Soon thereafter he moved to Boston and studied with a well-known physician for a few weeks, but, with bigger goals in sight, he left for Scotland in June 1818 to continue his medical studies at the University of Edinburgh and to travel extensively in the British Isles. Later he spent some time at the Jardin des Plantes in Paris where he became acquainted with a number of the leading French naturalists at the Museum d'Histoire na-turelle, including Gabriel Bibron (1805-1848), Jean Leopold Nicholas Frederic Cuvier (better known as Georges Cuvier, 1769-1832), Andre Marie Constant Dumeril (1774-1860), and Achille Valenciennes (1794-1865) (Worthington & Worthington 1976; Stephens 1997, 2000; Sanders 8c Anderson 1999).

[FIGURE 1 OMITTED]

Holbrook established a medical practice in Charleston, South Carolina, in 1822, and two years later became Professor of Anatomy at the Medical College of South Carolina, an institution that he helped to found. In 1834, Holbrook joined the faculty of the newly established Medical College of the State of South Carolina and maintained an association with that institution for the remainder of his active life. In 1827, he married Harriott Pinckney Rutledge (1802-1863), a union that bore no children. Harriott Holbrook's ancestors introduced the culture of indigo to South Carolina and were signatories to both the American Declaration of Independence and to the fledgling country's constitution (Worthington & Worthington 1976; Stephens 1997, 2000; Sanders & Anderson 1999; Anderson & Stephens 2002).

During the American Civil War (1861-1865), Holbrook was chairman of the Examining Board of Surgeons for South Carolina, and served in the field, at the age of 70, as medical officer with the Confederate Army, enduring hardship and exposure, sleeping under an army wagon, and sharing the very modest fare of the ordinary soldier. Better recognized for his contributions to herpetology than for activities in ichthyology, Holbrook's efforts in ichthyology were far from negligible. He was held in great esteem by his contemporaries, who derived from his name numerous patronymics for poikilotherms; several of those names are currently considered valid (e. g., Diplodus holbrooki, Spottail Pinfish [Sparidae]; Holbrookia, a genus of earless lizards [Phrynosomatidae]). In 1868 Holbrook was elected to the United States National Academy of Sciences, the first southern scientist so honored, a signal acknowledgement of his accomplishments in zoology. (1) During the last six years of his life, Holbrook made frequent trips to Massachusetts, dying on 8 September 1871 in North Wrentham (now Norfolk), Massachusetts, the town of his early years, but he was buried in Charleston (Agassiz 1872; Gill 1905; Worthington & Worthington 1976; Adler 1989; Stephens 1997, 2000; Sanders & Anderson 1999; Anderson & Stephens 2002).

HOLBROOK'S ARTISTS

The high quality of the illustrations in his publications is a distinctive feature of Holbrook's work. With the goal of obtaining drawings that were both lifelike and scientifically accurate, he went to great lengths to find artists who could satisfy his needs. At least 17 artists contributed to his Herpetology, drawing from living specimens when possible. Prominent among those artists were J. Sera (?-1837) and John H. Richard (ca. 1807-1881). Holbrook even recruited the Right Reverend John Henry Hopkins (1792-1868), Bishop of the Episcopal Diocese of Vermont, to make a drawing for Herpetology. In Charleston, an immigrant Italian artist, J. Sera, was an important provider of artwork for Herpetology. Sera had received great praise for his efforts in preparing theater paintings and decorations. "For Holbrook, Sera produced watercolor drawings almost like miniature portraits" (Blum, 1993: 147). After Sera's death, Richard, also an outstanding artist, was hired. Later Richard, whose origins are obscure (perhaps German, perhaps Alsatian French), was hired as an illustrator by Spencer Fullerton Baird (1823-1887) at the Smithsonian Institution. Other artists contributing to Herpetology include Maria Martin (1796-1863), Albert Newsarn (1809-1864), and James Fuller Queen (ca. 1820-1886, Blum 1993: 150; also reported as 1824-ca. 1877, Falk 1999: 2683). Maria Martin was the second wife of John Bachman (1790-1874), minister for many years at St. John's Lutheran Church in Charleston and coauthor with John James Audubon (1785-1851) of extensive works on the quadrupeds of North America (Audubon & Bachman 1845-1854, 1849-1854). Newsam and Queen, highly skilled employees of a Philadelphia lithographer, produced a considerable number of the plates for Herpetology (Rutledge 1949: 217; Worthington & Worthington 1976: xix-md; Adler 1989: 33; Blum 1993: 147, 150, 167, 363, 364; Falk 1999: 2683; Sanders & Anderson 1999: 73, 74; Stephens 2000: 87, 90, 95, 99).

Richard rendered the plates for Southern Ichthyology (Holbrook 1847, 1848), and most of them for the first edition of Holbrook's (1855b) Ichthyology of South Carolina (Stephens 1997, 2000; Sanders tic Anderson 1999; Anderson 86 Stephens, 2002). In the Preface to the second edition of Ichthyology of South Carolina, Holbrook (1860) wrote "The new drawings are from nature, and have been made by the best artists,--as A. J. Ibbotson and A. Sonrel. The colour of the fish has been, in almost every instance, taken from living specimens, by J. Burkhardt, an artist of great merit." Indeed, Jacques Burkhardt (ca. 1808-1867),2 was "an artist of great merit," serving as Louis Agassiz's principal artist for many years. Among other activities, Burkhardt accompanied the Thayer Expedition (1865-1866) to Brazil "where he made nearly 2,000 watercolor drawings of fresh fishes and local habitats." (http://www.mcz.harvard.edu/Departments/Ichthyology/expeditions_thayer_hassler.html)

Antoine Sonrel (2) (ca. 1805-1879) and Burkhardt worked for Louis Agassiz in Neuchatel and both moved to the United States some time after Agassiz became established there, maintaining their associations with him for many years (Irmscher 2013: 98, 154, 155). Sonrel, an extraordinarily talented man, made very important contributions in assisting Agassiz in the preparation of Contributions to the Natural History of the United States of America (Agassiz 1857-1862). Agassiz was particularly impressed with the skill displayed by Sonrel in illustrating Cyanea capillata (Linnaeus), the Lion's-Mane Jellyfish (Irmscher 2013: 155). A. J. (presumably Arvah J.) Ibbotson was a lithographer in Philadelphia in the mid-nineteenth century (Falk 1999: 1680).

HOLBROOK AND HERPETOLOGY

Holbrook is best known for North American Herpetology (Holbrook 1836-1840, 1842). This monumental effort laid the groundwork for studies of amphibians and reptiles of North America. Agassiz (1872: 348) wrote that Holbrook "first compelled European recognition of American science by the accuracy and originality of his investigations" and that his "elaborate history of American Herpetology was far above any previous work on the same subject. In that branch of investigation Europe had at that time nothing which could compare with it." Earlier Girard (3) (1851: 200) had dubbed Holbrook "the father of American herpetology." In his magnum opus, which he began in the mid-1820s, Holbrook described and illustrated every species of amphibian and reptile then known to occur in the United States (at that time limited largely to the area east of the Mississippi River) (Adler 1989). His Herpetology is noteworthy in being "the first great synthesis of information on the topic, and

Table I. Species in Holbrook's Ichthyology of South Carolina, 1860.

Name in              Valid name           Family      Page  Plate
Holbrook, 1860                                              (Fig.)

Perca            Perca flavescens      Percidae          2  I(1)
fluvescens

Pomotis          Lepomis gibbosus      Centrarchidae     8  I(2)
vulgaris

Ichthelis        Lepomis nacrochirus   Centrarchidae    12  II(1)
incisor

Icluthelis       Lepomis auritus       Centrarchidae    15  II(2)
rubricanda

Centrarchus      Centrarchus           Centrarchidae    18  III(1)
irideus          macropterus--juveile

Labrax           Morone americana      Moronidae        20  III(2)
americanus

Labrax lineatus  Morone saxatilis      Moronidae        24  IV(1)

Grystes          Micropterus           Centrarchidae    28  IV(2)
salmoides        salmoides

Diplectrum       Diplectrum formosum   Serranidae       35  V(1)
fasciculare

Serranus         Epinephelus morio     Epinephelidae    32  V(2)
erythrogaster

Pomoxis          Pomoxis               Centrarchidae    39  VI(1)
hexacanthus      nigromaculatus

Rhypticus        Rypticus maculatus    Serranidae       42  VI(2)
maculatus

Centropristes    Ccntropristis         Serranidae       49  VII(1)
trifurca         pliiladclphica

Centropristes    Ceniroprislis         Serranidae       45  VII(2)
atrariits        siriata

Lagodon          Lagodon rhomhoides    Sparidae         59  VIII(1)
rhomboides

Sargus ovis*     Archosargns           Sparidae         54  VIII(2)
                 probatoccphalus

Cybium           Scomberomorus         Scom bridae      68  IX(1)
maculatum        maculatus

Temnodon         Pontatomus saltatrix  Pomatomidae      64  IX(2)
saliator

Seriola zonata   Seriola               Carangidae       75  X(1)
                 zonara--juvenile

Scriola          Seriola               Carangidae       72  X( 2)
carolinensis     zonata--adult

Seriola chloris  Chloroscombrus        Carangidae       79  XI(1)
                 chrysurus

Bothrolacmus     Trctchinotus          Carangidae       83  XI(2)
pampanus*        carolinus

Caranx defensor  Caranx hippos         Carangidae       87  XII(1)

Caronx hippos    Caranx crysos         Carangidae       90  XII(2)

Caronx richardi  Caranx latus          Carangidae       96  XIII(1)

Caranx falcatus  Hemicaranx            Carangidae       94  XIII2)
                 amblyrhynchus

Echeneis         Echcneis naucrates    Echeneidae      102  XIV(1)
lineata

Elecate Canada   Rachyccntron Canadum  Rachycentridae   97  XIV(2)

Ephippus faber   Chactodiptcrus        Ephippidae      110  XV(1)
                 fabcr--juvenile

Ephippus gigas   Chaetodipterus        Ephippidae      107  XV(2)
                 faher--adult

Pogonias         Pogonias              Sciaenidae      119  XVI(1)
fasciatus        cromis--juvenile

Pogonias cromis  Pogonias              Sciaenidae      114  XVI(2)
                 cromis--adult

Haemulon         Hacmulon              Haemulidae      121  XVII(1)
chrysopteron     aitrolincainm

Haemulon         Hacnndon plumierii    Haemulidae      124  XVII(2)
arcnatum

Otolithus        Cynoscion regalis     Sciaenidae      129  XVIII(1)
regalis

Otolithus        Cynoscion regalis     Sciaenidae      133  XVIII(2)
thalassinus

Otolithus        Cynoscion nothus      Sciaenidae      134  XIX(1)
nothus

Otolithus        Cynoscion nebuldosus  Sciaenidae      136  XIX(2)
carolinensis

Umbrina          Menlicirrluts         Sciaenidae      137  XX(1)
alburnus*        americanus

Umbrina          Menticirrhus          Sciaenidae      144  XX(2)
littoralis       littoralis

Micropogon       Micropogonias         Sciaenidae      146  XXI(1)
undulatus        undulatus

Corvina          Sciaenops ocellatus   Sciaenidae      150  XXI(2)
ocellata

Larimus          Larimus fasciatus     Sciaenidae      154  XXII(1)
fasciatus

Pristapoma       Orthopristis          Haemulidae      157  XXII(2)
fulvo-maculatum  chrysoptera

Leiastomus       Leiostomus xanthurus  Sciaenidae      160  XXIII(1)
obliquus

Homoprion        Bairdiella chrysoura  Sciaenidae      164  XXIII(2)
xanthurus**

Homoprion        Stellifer             Sciaenidae      167  XXIV(1)
lanceolatus      lanceolatus

Lobotes          Lobotes surinamensis  Lobotidae       169  XXIV(2)
surinamensis

Pagrus argyrops  Slenotomus chrysops   Sparidae        174  XXV(1)

Serranus         Hyporthodus nigritus  Epinephelidae   177  XXV(2)
nigritus

Saurux foetens   Synodus foetens       Synodontidae    187  XXVI(1)

Mops saurus      Mops saurus or E.     Elopidae        180  XXVI(2)
                 smithi

Esox affinis*    Esox niger            Esocidae        198  XXVII(1)

Esox ravenelii*  Esox americanus       Esocidae        201  XXVII(2)

Trachinotus      Trachinotus goodei    Carangidae      192  XXVIII(1)
glaucus*

Huenudon         ?Haemdon striatum     Haemulidae      195  XXVIII(2)
quadrilineatum*

Not in text      Alosa mediocris       Clupeidae            XXIX(1)

Not in text      AIosa sapidissima     Clupeidae            XXX(2)

Not in text      Brevoortia lyrannus   Clupeidae            XXX(1)

Not in text      Dorosoma cepedianum   Clupeidae            XXX(2)

*Plate number in text does not agree with number on plate.
**Description in text appears to be in part of Leiostomus xanthurus
and in part of Bairdiella clirysoura: illustration is of Bairdiella
chrysoura.

Holbrook carefully researched the literature to determine the correct scientific names to be used, thus making a major contribution to stabilizing the nomenclature" (Adler 1989: 33). Holbrook's great desire for accuracy can best be appreciated in his own words (Holbrook 1842, I: ix):

With an immense mass of materials, without libraries to refer to, and only defective museums for comparison, I have constantly been in fear of describing animals as new that have long been known to European Naturalists. In no department of American Zoology is there so much confusion as in Herpetology. This is to be traced partly to the earlier Naturalists, partly to the practice of describing from specimens preserved in alcohol, or from prepared skins. I have endeavoured to avoid error in this respect, by describing in almost every instance from the living animal, and often after a comparison of many individuals.

In the last sentence of the quotation above, Holbrook showed an appreciation, rare in his day, for individual variation. Of the 37 herpetological taxa described as new by Holbrook (1836-1840, 1842), 24 (three genera and 21 species) are considered valid today (Adler 1976, 1989; Sanders and Anderson 1999).

HOLBROOK AND ICHTHYOLOGY

Holbrook seems to have developed an interest in fishes as early as the late 1820s, as shown by his donations of several specimens to the Museum di-Es-toire naturelle in Paris by the early 1830s (Stephens, 1997, 2000); that interest eventually led to an association with Louis Agassiz. The noted Swiss naturalist Jean Louis Rodolphe Agassiz (1807-1873) arrived in Charleston in 1847 for an extended visit. He became acquainted with Holbrook, and probably in large part because of a mutual interest in ichthyology, they became good friends. Over a period of several years, Agassiz made a number of trips to Charleston, usually bringing with him his outstanding artist Jacques Burkhardt. Agassiz and Holbrook conducted fieldwork together, and some of the fishes they collected in South Carolina were deposited in Harvard's Museum of Comparative Zoology: 103 lots by Agassiz, 26 by Holbrook and Agassiz, 19 by Jacques Burkhardt, and five by Agas-siz, Holbrook, and Burkhardt. In addition, Holbrook placed 64 lots in Agassiz's new museum at Harvard (4) (Stephens 1997, 2000; Sanders & Anderson 1999: 71).

Table II. Fish taxa described by Holbrook. Holbrook's names in
current use are in boldface.

Holbrook's name                  Valid name              Family

Diplectrum             Diplectrum                     Serranidae

Bothrolaemus           Trachinotus                    Carangidae

Lagodon                Lagodon                        Sparidae

Homoprion              Stellifer                      Sciaenidae

Pimelodus marmoratus   Ameiurus nebulosus marmoratus  Ictaluridae

Esox affinis           Esox niger                     Esocidae

Esox ravenelii         Esox americanus                Esocidae

Serranus nigritus      Hyporthodus nigritus           Epinephelidae

Rypticus maculatus     Rypticus maculatus             Serranidae

Bryttus fasciatus      Enneacanthus obesus            Centrarchidae

Bryttus gloriosus      Enneacanthus gloriosus         Centrarchidae

Calliurus floridensis  Lepomis gulosus                Centrarchidae

Pomotis elongatus      Lepomis auritus                Centrarchidae

Pomotis niarginatus    Lepomis marginatus             Centrarchidae

Pomotis speciosus      Lepomis macrochirus            Centrarchidae

Boleosoma barratti     Etheostoma fusiforme barratti  Percidae

Caranx falcatus        Hemicaranx amblyrhynchus       Carangidae

Caranx richardi        Caranx latus                   Carangidae

Seriola carolinensis   Seriola zonata                 Carangidae

Otolithus nothus       Cynoscion nothus               Sciaenidae

Otolithus thalassinus  Cynoscion regalis              Sciaenidae

Otolithus thallapinus  Cynoscion regalis              Sciaenidae

Larimus fasciatus      Larimus fasciatus              Sciaenidae

Urnbrina littoralis    Menticirrhus littoralis        Sciaenidae

Homoprion lanceolatus  Stellifer lanceolatus          Sciaenidae

Eventually Holbrook's interest in fishes led him to write Southern Ichthyology (1847, 1848), in which he provided 14 species accounts and 17 species illustrations (one text figure and eight plates with two figures each); two of the species, members of the drum family (Sciaenidae), were new to science. Only a small part of what Holbrook apparently intended to present in Southern Ichthyology ever reached print. Ultimately, he must have concluded that his original plans were beyond his resources of time, energy, and money. Consequently, he narrowed his scope and decided to concentrate on fishes of South Carolina, eventually publishing Ichthyology of South Carolina (Stephens 1997, 2000; Anderson & Stephens 2002).

Nevertheless, despite narrowing his focus, Holbrook did not ignore the biota of other southern states, as shown by his lists of fishes, amphibians, and reptiles of Georgia (Holbrook 1849) and a short paper on fishes mainly collected in Florida and Georgia (Holbrook 1855a). In the latter, he contributed accounts of eight species, describing all of them as new. Of those eight, two were given specific names considered valid today, and two others are sometimes recognized at the subspecific level (Anderson 8c Stephens 2002: 319). Altogether, Holbrook described 25 fish taxa, 12 of the names he proposed are considered valid today.

ICHTHYOLOGY OF SOUTH CAROLINA

The first edition of Ichthyology of South Carolina (Holbrook 1855b), with plates much improved over those in Southern Ichthyology, was followed five years later by the second edition (Holbrook 1860), which for the most part has even better plates (Gill 1864: 89). Two genera and four species covered in the first edition were new to science. The first edition with 27 color plates ends at the bottom of page 184, following the words "The palate-..." The second edition, containing 205 pages of text and 28 color plates (each with two figures; one copy of this edition has 30 plates, the last two of which are uncol-ored) (5), is quite similar to the first, as noted by Gill (1864: 89). The 1860 edition includes accounts of 102 taxa (nine families, 37 genera, 56 species).

Both editions of Holbrooks' Ichthyology are quite rare, only present in a small number of libraries and in an unknown, presumably small, number of private collections. However, the 1855 edition is available online at the website for the Biodiversity Heritage Library (http://wwvv.biodiversitylibrary.org/bibliography/6911), and the 1860 edition can be found at the Hathi Trust Digital Library. Since few libraries own the second edition, it seems appropriate to make it available in part, at least, for individuals studying North American fishes and for others with interests in the history of biology. Accordingly, the plates from the second edition are reproduced herein.

PLATES IN THE SECOND EDITION OF ICHTHYOLOGY OF SOUTH CAROLINA

The sequence of plates in the second edition of Holbrook's Ichthyology differs very little from the sequence in the first edition. Not surprisingly, neither arrangement is consistent with current concepts of systematic relationships. Herein the plates (each with two figures) are presented in the same order as that in the second edition and are listed in the same manner in Table I. I am aware of one copy of the 1860 edition that has two additional plates (Plates XXIX and XXX, each with two uncolored figures); the species in those plates are included in the notes below. (5) In the commentary on the species represented in the plates, I follow the systematic sequence used by Page et al. (2013).

Comments on the Species Illustrated: The species are grouped within families, as presently understood. For each species, following the name used by Holbrook (1860), are the valid scientific name, if different, common name(s) (which for the most part are those given by Page et al. 2013), references to the pagination and plate in the 1860 edition, and comments on the biology. In preparing the remarks on the biology of the species included in this section I have consulted the following without giving individual attribution under each species: Robins & Ray (1986), Smith (1997), Smith-Vaniz et al. (1999), Carpenter (2003), Boschung & Mayden (2004), Marcy et al. (2005), Rohde et al. (2009), Kells & Carpenter (2011), Page & Burr (2011), and Murdy & Musick (2013).

ELOPIDAE

Elops saurus: Ladyfish, Tenpounder; p. 180, Plate XXVI (Fig. 2). McBride et al. (2010) described a new Tenpo under, Elops smithi, distinguishable morphologically only by myomere (larvae) or vertebral (adults) counts from Elops saurus; E. smithi is sympatric with E. saurus along the east coast of the United States (including South Carolina and Georgia) (McBride & Horodysky 2004). Also, McBride et al. (2010: 29) noted that "the sequence divergence in mtDNA cytochrome b (d = 0.023-0.029) between E. smithi and E. saurus is similar to or greater than that measured between recognized species of Flops in different ocean basins." McBride and Horodyski (2004) and McBride et al. (2010) concluded that as a result of stressful environmental conditions (severe thermal, salinity, and predatory regimes) few E. smithi survive for more than a few years in the waters off the eastern United States. Although it is impossible to say with certainty which species Holbrook described and illustrated, he most likely had Elops saurus.

Elops saurus ranges along the east coast of the United States from southern New England (uncommon north of North Carolina) to southern Florida and in the Gulf of Mexico, while E. smithi occurs from the northern coast of South America through the Caribbean basin and the Bahamas and c'sympatrically with E. saurus in the Gulf of Mexico and along the eastern seaboard of North America" (McBride et al. 2010: 37). There are also two records of Elops from Bermuda, at least one of which appears to be of E. smithi, but no resident population has been found there (Smith-Vaniz et al. 1999: 126). Elops is found in estuaries and coastal waters, being capable of living in a wide range of salinities but appearing infrequently in fresh water.

CLUPEIDAE

Alosa mediocris (Mitchill): Hickory Shad; no account in text, Plate XXIX (Fig. 1). Anadromous in fresh, brackish, and coastal marine waters; most of adult life spent at sea, in spring ascending coastal streams to spawn. Commonly occurring from Cape Cod, Massachusetts, to northern Florida, also reported from as far north as New Brunswick.

Alosa sapidissima (Wilson): American Shad; no account in text, Plate XXIX (Fig. 2). Anadromous in fresh, brackish, and coastal marine waters, spending most of adult life at sea in large schools, in spring ascending coastal streams to spawn. Nova Scotia to Florida.

Brevoortia tyrannus (Latrobe): Atlantic Menhaden; no account in text, Plate Mc (Fig. 1). Marine and brackish inshore waters, being most abundant near estuaries; migratory, forming large tight schools near surface. Nova Scotia to Florida.

Dorosoma cepedianum (Lesueur): Gizzard Shad; no account in text, Plate XXX (Fig. 2). Anadro-mous in rivers, bays, and estuaries. St. Lawrence River to Florida; in Gulf of Mexico from Florida to Mexico.

ESOCIDAE

Esox affinis = Esox niger Lesueur: Chain Pickerel, Jackfish; p. 198, Plate XXVII (Fig. 1). Freshwater, occurring in vegetated lakes, swamps, and backwaters and quiet pools of streams. Atlantic and Gulf of Mexico slopes, Nova Scotia (where introduced) to south Florida and west to Red River drainage, Oklahoma, and Sabine River drainage, Texas; introduced elsewhere.

Esox ravenelii = Esox americanus Gmelin: Redfin Pickerel; p. 201, Plate XXVII (Fig. 2).Freshwater, living in lakes, swamps, backwaters, and sluggish pools of streams, most commonly in clear water among vegetation. Esox americanus americanus ranges on the Atlantic Slope from the St. Lawrence River drainage, Quebec, to south Georgia; the species has been introduced elsewhere.

SYNODONTIDAE

Saurus foetens = Synodus foetens (Linnaeus): Inshore Lizardfish; p. 187, Plate XXVI (Fig. 1). Found along beaches and in bays, inlets, estuaries, and lagoons to depths of about 200 meters, more commonly in shallow water; burrows into bottom sediments from which prey can be ambushed. Bermuda and Massachusetts to southern Florida, Gulf of Mexico, Bahamas, Antilles, and northern Caribbean Sea. As a result of resurrecting S. bondi from the synonymy of S. foetens, Frable et al. (2012) were able to clarify the range of the latter species.

MORONIDAE

Labrax americanus = Marone americana (Gmelin): White Perch; p. 20, Plate III (Fig. 2).

Anadromous in fresh, brackish, and coastal marine waters; usually over mud bottoms in quiet areas in medium to large rivers. Atlantic Slope from Saint Lawrence drainage, Quebec, southward to Savannah River drainage, Georgia; introduced elsewhere.

Labrax lineatus = Marone saxatilis (Walbaum): Striped Bass, Rockfish; p. 24, Plate IV (Fig. 1).

Anadromous, seasonally migratory; occurs in rivers, freshwater impoundments, estuaries, and nearshore waters; some populations landlocked in fresh waters. Atlantic and Gulf of Mexico slope drainages from Saint Lawrence River drainage, Quebec, to north Florida, and from west Florida to east Texas; widely introduced.

EPINEPHELIDAE

Serranus erythrogaster = Epinephelus mono (Valenciennes): Red Grouper; p. 32, Plate V (Fig. 2). Protogynous hermaphrodite; fished commercially and for sport, although it is listed as "Near Threatened" by the International Union for Conservation of Nature and Natural Resources in its Red list of Threatened Species (IUCN, 2014). Found over hard bottoms (around reefs, in crevices, under ledges), mud, and sand from near shore to about 290 meters. Bermuda and North Carolina to southern Brazil, including the Bahamas, Gulf of Mexico, Antilles, and Caribbean Sea; strays have been reported from as far north as Massachusetts.

Serranus nigritus = Hyporthodus nigritus (Holbrook): Warsaw Grouper; p. 177, Plate XXV (Fig. 2). Listed as "Critically Endangered" by the IUCN (2014). Juveniles found around reefs and jetties, adults over rocky bottoms from 55 to 520 meters. Massachusetts to Florida, Gulf of Mexico, Cuba, Haiti, Trinidad, and from Venezuela to Rio de Janeiro, Brazil; unsubstantiated reports from Bermuda.

SERRANIDAE

Centropristes trifurca = Centropristis philadelph-ica (Linnaeus): Rock Sea Bass; p. 49, Plate VII (Fig. 1). Protogynous hermaphrodite. Occurs over sand and mud bottoms in depths of about 10 to 170 meters. Virginia to Palm Beach, Florida, and the northern Gulf of Mexico.

Centropristes atrarius = Centropristis striata (Linnaeus): Black Sea Bass, Blackfish; p. 45, Plate VII (Fig. 2). Protogynous hermaphrodite. Most commonly found on rocky bottoms and around pilings, seawalls, and jetties; near shore to about 75 meters. Massachusetts to central Florida, occasionally to the Florida Keys; in Gulf of Mexico along coast of Florida from Pensacola to Placida.

Diplectrum fasciculare = Diplectrum formosum (Linnaeus): Sand Perch; p. 35, Plate V (Fig. 1).

Over bottoms of sand, shell, mud, or rocky rubble; found from inshore grass flats to offshore depth of about 75 meters, most commonly between 7 and 50 meters. Bermuda; Virginia to Florida, throughout Gulf of Mexico, and from Colombia to Sao Paulo, Brazil; rare in West Indies.

Rhypticus maculatus = Rypticus maculatus Holbrook: Whitespotted Soapfish; p. 42, Plate VI (Fig. 2). Near bottom around rocky areas, coral reefs, jetties, and pilings from near shore to about 90 meters. North Carolina to Florida and in the Gulf of Mexico.

CENT PARCHIDAE

Centrachus irideus = Centrarchus macropterus (Lacepede): Flier (juvenile); p. 18, Plate III (Fig. 1). Usually over mud in swamps, vegetated lakes, ponds, sloughs and backwaters and in pools in creeks and small rivers. Coastal Plain drainages from Potomac River, Maryland, to central Florida, west to Trinity River, Texas, and north in former Mississippi Embayment to southern Illinois and southern Indiana.

Ichthelis rubricauda = Lepomis auritus (Linnaeus): Redbreast, Redbreast Sunfish; p. 15, Plate II (Fig. 2). Rocky and sandy pools of creeks and small to medium size rivers, also margins of lakes in rocks and vegetation. Atlantic slope drainages from New Brunswick to central Florida, Gulf of Mexico drainages of Apalachicola and Chocta-whatchee (possibly) rivers in Georgia, Alabama, and Florida; introduced elsewhere.

Pomotis vulgaris = Lepomis gibbosus (Linnaeus): Pumpkinseed; p. 8, Plate I (Fig. 2).

Vegetated lakes, ponds, and quiet pools of creeks and small rivers. Atlantic drainages from New Brunswick to Savannah River, Georgia; Great Lakes, Hudson Bay (Red River), and upper Mississippi River basins; introduced elsewhere.

Ichthelis incisor = Lepomis macrochirus Rafinesque: Bluegill, Bream; p. 12, Plate II (Fig. 1). Vegetated shores of impoundments, oxbow lakes, ponds, swamps, and pools of creeks and small to large rivers. Native range includes much of the fresh waters of eastern United States, widely introduced elsewhere; two or three subspecies recognized, but exact limits of their individual distributions uncertain.

Grystes salmoides = Micropterus salmoides (La-cepede): Largemouth Bass; p. 28, Plate IV (Fig. 2). Usually over mud or sand in lakes, ponds, backwaters, swamps, and in pools in creeks and small to large rivers. Widely distributed in eastern and central United States and southernmost eastern and central Canada; in Atlantic and Gulf of Mexico drainages from North Carolina to northern Mexico; introduced into much of the United States and southern Canada and also into Africa and Eurasia.

Pomoxis hexacan thus = Pomoxis nigromaculatus (Lesueur): Black Crappie; p. 39, Plate VI (Fig. 1). Usually in clear water among vegetation over mud or sandy bottoms in lakes, ponds, sloughs, backwaters, and stream pools. Native range much of eastern and central United States and southernmost eastern and central Canada; widely introduced in other parts of the United States.

PERCIDAE

Perca flavescens (Mitchill): Yellow Perch; p. 2, Plate I (Fig. 1). Lakes, ponds, and pools in creeks and rivers, usually in clear water near vegetation. Native to Atlantic, Arctic, Great Lakes, and Mississippi River basins in much of Canada and northern United States east of the Continental Divide; ranges on Atlantic slope south to Savannah River, Georgia, and occurring on Gulf slope in Apalachicola River basin in Georgia, Alabama, and Florida; also native disjunct populations present in Choctawhatchie River drainage and in a very limited area around Mobile Bay, Alabama; widely introduced elsewhere in the United States.

POMATOMIDAE

Temnodon saltator = Pomatomus saltatrix (Linnaeus): Bluefish; P. 64, Plate IX (Fig. 2). Moving in large schools in coastal waters, juveniles in bays and estuaries. In coastal subtropical and temperate waters in eight major isolated populations, except absent from the eastern Pacific and the Indo-west Pacific north of the equator; in western Atlantic from Bermuda, Nova Scotia to Gulf of Mexico and Colombia to Argentina but absent from Bahamas, Antilles (except northern coast of Cuba), and Caribbean coast of Central America.

CARANGIDAE

Caranx hippos = Caranx crysos (Mitchill): Blue Runner; p. 90, Plate XII (Fig. 2). A schooling species found mainly inshore (uncommon around reefs), off southeastern United States probably spawns offshore, young frequently occur with sar-gassum. Both sides of the Atlantic including western Mediterranaean; in western Atlantic from Nova Scotia to Argentina, including Bermuda, Gulf of Mexico, Bahamas, Caribbean Sea, and Antilles.

Caranx defensor = Caranx hippos (Linnaeus): Crevalle Jack, Jack Crevalle; p. 87, Plate XII (Fig. 1). Common on shallow flats, larger individuals may be found in offshore waters, common in brackish waters, may enter rivers. Both sides of Atlantic; in western Atlantic from Nova Scotia to Uruguay, including Gulf of Mexico and Caribbean Sea, rare with patchy distribution in Bahamas and Antilles; occasionally found at Bermuda (Smith-Vaniz & Collette 2013: 173).

Caranx richardi = Caranx latus Agassiz: Horseeye Jack; p. 96, Plate XIII (Fig. 1). Mostly in small schools around islands, offshore, and off sandy beaches in the tropics; may enter brackish waters and rivers. Both sides of the Atlantic, including Ascension Island, St. Helena, and the Gulf of Guinea; western Atlantic from New Jersey to Rio de Janeiro, Brazil, including Bermuda, Gulf of Mexico, and Caribbean Sea.

Seriola chloris = Chloroscombrus chrysurus (Linnaeus): Atlantic Bumper; p. 79, Plate XI (Fig. 1). Mostly in shallow marine and estuarine waters and mangrove-lined lagoons. Both sides of the Atlantic; western Atlantic from Massachusetts to Uruguay (including Gulf of Mexico and Caribbean Sea), rare at Bermuda, possibly present throughout the Antilles.

Caranx fidcatus = Hemicaranx amblyrhynchus (Cuvier): Bluntnose Jack; p. 94, Plate XIII (Fig. 2). Inshore species, enters brackish water. Western Atlantic, historical records from the Carolinas, but very rare along east coast of United States; known from Gulf of Mexico to Florianopolis, Brazil, few records from West Indies except for Cuba and Trinidad.

Seriola zonata (Mitchill): Banded Rudderfish (juvenile); p. 75, Plate X (Fig. 1). Juveniles with six dark bars on sides and dark band through eye to origin of first dorsal fin. Pelagic and epibenthic over continental shelf. Maine (possibly Nova Scotia) to Santos, Brazil (including Gulf of Mexico and Greater Antilles).

Seriola carolinensis = Seriola zonata (Mitchill): Banded Rudderfish (adult); p. 72, Plate X (Fig. 2). See preceding account.

Bothrolaemus pampanus = Trachinotus carolinus (Linnaeus): Pompano, Florida Pompano; p. 83, Plate XI (Fig. 2). Sandy beaches, inlets, and brackish bays. Massachusetts to Brazil (including Gulf of Mexico and Caribbean Sea), occurring irregularly at Jamaica, Puerto Rico, Tobago, and Trinidad.

Trachinotus glaucus = Thachinotus goodei Jordan & Evermann: Palometa; p. 192, Plate XXVIII (Fig. 1). Usually in large schools in surf zone of sandy beaches, also around reefs and rocky areas, most often in high salinity waters. Massachusetts and Bermuda to Argentina (including Gulf of Mexico and Caribbean Sea), and throughout the West Indies.

RACHYCENTRIDAE

Elecate canada = Rachycentron canadum (Linnaeus): Cobia; p. 97, Plate XIV (Fig. 2). Coastal, pelagic; to depths of 50 meters over waters as deep as 1,200 meters, also on shallow coral reefs and off rocky shores, occasionally in estuaries. Almost worldwide in tropical and subtropical waters, except absent from eastern Pacific and Pacific Plate; in western Atlantic from Massachusetts and Bermuda to Argentina, including Gulf of Mexico, West Indies, and Caribbean Sea.

ECHENEIDAE

Echeneis lineata = Echeneis naucrates Linnaeus: Sharksucker; p. 102, Plate XIV (Fig. 1). Attaches temporarily to a wide variety of living hosts, especially sharks, but also rays, some teleosts, and sea tur-des, and to ships and buoys. Worldwide in tropical and temperate seas except for the eastern Pacific.

LOBOTIDAE

Lobotes surinamensis (Bloch): Atlantic Tripletail; p. 169, Plate XXIV (Fig. 2). Sluggish offshore species, often found floating on its side near surface associated with flotsam, occasionally drifting into shallow water; young often drift with sargassum and on occasion mimic mangrove leaves. Lobotes occurs in all warm seas, except on Pacific Plate; Lobotes surinamensis in western Atlantic ranges from Nova Scotia and Bermuda to Argentina, including Gulf of Mexico, West Indies, and Caribbean Sea.

HAEMULIDAE

Haemulon chrysopteron = Haemulon aurolina-turn Cuvier: Tomtate; p. 121, Plate XVII (Fig. 1). Found in a variety of natural and artificial habitats, from near shore to about 40 meters. Bermuda and Chesapeake Bay to Florida, Bahamas, Gulf of Mexico, Antilles, and Caribbean Sea south to Brazil.

Haemulon arcuatum = Haemulon plumierii (La-cepede): White Grunt; p. 124, Plate XVII (Fig. 2). From shore to outer reefs (depths of about 40 meters) over a variety of bottom types. Chesapeake Bay to south Florida, Bahamas, Gulf of Mexico, Antilles, and Caribbean Sea to Brazil.

Haemulon quadrilineatum of Holbrook, 1860 = ? Haemulon striatum (Linnaeus): Striped Grunt, La Gorette a Quatre Lignes, Cricri; p. 195, Plate XXVIII (Fig. 2). Although I am not aware of any references in the current literature of Haemulon striatum occurring off South Carolina, there are specimens of this species in the collections of the Grice Marine Biological Laboratory obtained from two localities off South Carolina. Some credence can be given to Holbrook's report, because he wrote that this fish "is rarely found in our waters, though it is abundant in the Caribbean Sea" (Holbrook, 1860: 197), and the counts of dorsal and anal fin rays (except for dorsal soft rays) on the specimen depicted in Plate XXVIII, Figure 2, fall within the ranges given for this species by Courtenay (1961: 126, 134). Despite this, the overall body shape (particularly anteriorly) of the illustrated specimen does not resemble very closely individuals of Haemulon striatum depicted in other publications examined. Haemulon striatum is found over outer reefs in depths of 12 to 100 meters and is reported in the recent literature from southern Florida, Bahamas, southern Gulf of Mexico, Caribbean Sea (including the Antilles) southward to Brazil. Although reported from Bermuda, Smith-Vaniz et al. (1999: 265) deleted it from their list of Bermuda fishes.

Pristipoma fulvo-maculatum = Orthopristis chrysoptera (Linnaeus): Pigfish; p. 157, Plate XXII (Fig. 2). Shallow waters near shore over soft bottoms, also in bays and other estuaries. Bermuda, coastal waters from New Jersey to Florida Keys, Cuba, and Gulf of Mexico to Yucatan.

SPARIDAE

Sargus ovis = Archosargus probatocephalus (Wal-baum): Sheepshead; p. 54, Plate VIII (Fig. 2). Along coast and in estuaries, over mud, rocks, and other hard substrates. Nova Scotia to Florida and Gulf of Mexico, scattered reports from Honduras to Rio de Janeiro; absent from Bermuda and West Indies.

Lagodon rhomboides (Linnaeus): Pinfish; p. 59, Plate VIII (Fig. 1). Shallow-water species usually over vegetated bottoms, sometimes over rocks and in mangrove areas, enters brackish waters, may enter fresh water. Bermuda and from Cape Cod, Massachusetts, to Campeche Banks off Yucatan, including entire coast of the Gulf of Mexico and northern coast of Cuba.

Pagrus argyrops = Stenotomus chrysops (Linnaeus): Scup; p. 174, Plate XXV (Fig. 1). Coastal waters mainly over hard bottoms. Nova Scotia to Florida, rare south of North Carolina.

SCIAENIDAE

Homoprion xanthurus: p. 164, Plate XXIII (Fig. 2). Description in text appears to be in part of Leiostomus xanthurus Lacepede and in part of Bairdiella chrysoura (Lacepede); the illustration is of B. chrysoura. Bairdiella chrysoura: coastal waters over sandy and muddy bottoms, in estuaries in summer. Cape Cod, Massachusetts, to south Florida, in Gulf of Mexico from Florida to northeastern Mexico.

Otolithus carolinensis = Cynoscion nebulosus (Cuvier): Spotted Seatrout, Spotted Weakfish; p. 136, Plate XIX (Fig. 2). Usually in estuaries and shallow coastal waters over sand bottoms, also associated with seagrass beds (nursery areas for young), salt marshes, and high-salinity tidal pools. Atlantic coast from Long Island, New York, to Florida, and in Gulf of Mexico from Florida to Laguna Madre, Mexico.

Otolithus nothus = Cynoscion nothus (Holbrook): Silver Seatrout; p. 134, Plate XIX (Fig. 1). Usually over sandy bottoms along beaches, in inlets, and in mouths of rivers. Chesapeake Bay to southern Florida and eastern and northern Gulf of Mexico to Texas.

Otolithus regalis = Cynoscion regalis (Bloch & Schneider): Weakfish; p. 129, Plate XVIII (Fig. 1). In shallow coastal waters over sand and sandy mud bottoms, relatively common in sounds and along beaches; in estuaries in summer. Nova Scotia to south Florida to southwest Florida (where infrequent), recent record from Bermuda (Smith-Vaniz 8c Collette, 2013: 179).

Otolithus thalassinus = Cynoscion regalis (Bloch 8z Schneider): Weakfish; p. 133, Plate XVIII (Fig. 2). See preceding account.

Larimus fasciatus Holbrook: Banded Drum: p. 154, Plate XXII (Fig. 1). Over mud and sandy bottoms in coastal waters to about 60 meters, uncommon in estuaries. Massachusetts to south Florida and Gulf of Mexico from Florida to Veracruz, Mexico.

Leiostomus obliquus = Leiostomus xanthurus La-cepede: Spot; p. 160, Plate XXIII (Fig. 1). Over sandy and muddy bottoms in coastal waters to about 60 meters. Cape Cod, Massachusetts, to Florida and in Gulf of Mexico from Florida to Veracruz, Mexico.

Umbrina alburnus = Menticirrhus americanus (Linnaeus): Southern Kingfish; p. 137, Plate XX (Fig. 1). Over sandy mud to hard sand bottoms in shallow coastal waters, also in surf and estuaries, juveniles often in brackish waters. Cape Cod, Massachusetts, to south Florida; Gulf of Mexico from Cape Sable, Florida, to Bay of Campeche, Mexico; western Caribbean Sea to southern Brazil; few records from the Antilles; rare from south Florida and Venezuela.

Umbrina littoralis = Menticirrhus littoralis (Holbrook): Gulf Kingfish; p. 144, Plate XX (Fig. 2). Coastal waters over sandy and sandy mud bottoms, most abundant in surf zone; occasionally in estuaries. Chesapeake Bay to south Florida, Gulf of Mexico, western Caribbean Sea to southern Brazil.

Micropogon undulatus = Micropogonias undula-tus (Linnaeus): Atlantic Croaker; p. 146, Plate )0U (Fig. 1). Coastal waters over sandy and muddy bottoms from near shore to about 100 meters and in estuaries. Cape Cod, Massachusetts, to Florida, in Gulf of Mexico from Florida to Bay of Campeche, Mexico.

Pogonias fasciatus = Pogonias cromis (Linnaeus): Black Drum (juvenile); p. 119, Plate XVI (Fig. 1). Juveniles with four or five black bars on sides. Over sand and sandy mud bottoms in surf, near mouths of rivers, and juveniles in estuaries. Nova Scotia to south Florida and in Gulf of Mexico from Florida to Texas, uncommon in Antilles and off southern Caribbean coast, also occurs off Atlantic coast of South America from Orinoco Delta to Argentina (unknown from northeastern Brazil).

Pogonias cromis (Linnaeus): Black Drum (adult); p. 114, Plate XVI (Fig. 2). See preceding account.

Corvina ocellata = Sciaenops ocellatus (Linnaeus): Red Drum, Redfish, Channel Bass, Spottail Bass; p. 150, Plate )0U (Fig. 2). Over sand and sandy mud bottoms in coastal waters, juveniles often in estuaries. Massachusetts to Florida and in Gulf of Mexico from west coast of Florida at least to the Laguna Madre in northeastern Mexico.

Homoprion lanceolatus = Stellifer lanceolatus (Holbrook): Star Drum; p. 167, Plate XXIV (Fig. 1). Over sandy-mud bottoms in coastal waters from near shore to about 20 meters. Chesapeake Bay to south Florida and in Gulf of Mexico from Florida to Bay of Campeche, Mexico, also reported from coast of Belize.

EPHIPPIDAE

Ephippus faber = Chaetodipterus faber (Brous-sonet): Adantic Spadefsh (juvenile); p. 110, Plate XV (Fig. 1). Shallow coastal waters over reefs and wrecks, sandy beaches, around buoys and pilings, in mangroves and harbors, and under bridges. Massachusetts to south Florida, Gulf of Mexico, Bahamas, Antilles, Caribbean Sea to southeastern Brazil.

Ephippus gigas = Chaetodipterus faber (Brous-sonet): Atlantic Spadefsh (adult); p. 107, Plate XV (Fig. 2). See preceding account.

SCOMBRIDAE

Cybium maculatum = Scomberomorus maculatus (Mitchill): Spanish Mackerel; p. 68, Plate IX (Fig. 1). Forms schools, migratory over continental shelf and slope, also found in estuaries. Nova Scotia to south Florida, in Gulf of Mexico from Florida to Yucatan.

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ADDITIONAL FISHES DRAWN FOR HOL-BROOK

Years ago while searching for copies of Holbrook's Southern Ichthyology, I learned that the Senckenber-gische Bibliotek in Frankfurt am Main has six color plates (each with two figures) of North American fishes that were received either directly or indirectly from Holbrook by the Senckenbeigische Natur-forschende Gesellschaft (Anderson, 2003) and that the Bibliotheque centrale, Museum national d'His-toire naturelle, Paris, has eight Holbrook plates (each with two figures) depicting fishes (Anderson, 2012). The fishes illustrated in the plates in Southern Ichthyology and those in the Senckenberg and Paris plates are listed by Anderson (2012: 4, table 1)

In November 2013, I visited the Natural-History Rare-Books section of the library at the National Museum of Natural History (Smithsonian Institution) and examined 23 loose plates (cataloged as qQL628 .S6H72 1855 c.2 SCNHRB) associated with one of that library's two copies of the 1855 edition of Holbrook's Ichthyology of South Carolina. Each of those plates bears figures of two fishes and, with the exception of one, is not colored, and all but two have the initials JHR (6) (= John H. Richard) printed at the right beneath the lower figure on the plate. The single color plate has a Roman numeral W at the far upper right that may have been handwritten; it depicts the same species (Pomoxis nigro-maculatus and Rypticus maculatus), as does Plate VI in the 1855 edition, but it lacks the initials JHR, and in view of the major differences in the two plates was probably drawn by another artist. Five of the plates have numbers printed at the right, just above the posterior end of the upper fish figured on the plate. One of those plates, bearing "Pl. XXVIII," has drawings of two herrings (Clupei-dae), viz., Alosa mediocris (Mitchill), Hickory Shad, and Alosa sapidissima (Wilson), American Shad, and is the only one of the loose plates that does not have a counterpart in Holbrook's Ichthyology (1855 or 1860, except one copy of the 1860 edition has a plate, Pl. XXIX, that is quite similar to loose Pl. XXVIII). The drawings in the other loose plates are very similar to those in the color plates of the 1855 edition, and in many cases the loose plates appear almost identical to those of 1855, except for being uncolored. The plate labeled "Pl. XX" has handwritten at the top: "Holbrook's Ichthyology of S. Carolina 1855." Based on the preceding, it is reasonable to conclude that at least some of the loose plates at the Smithsonian are extra proofs for the 1855 edition. (7)

CONCLUSION

Although well liked and highly regarded as a physician, lecturer in anatomy, and researcher, Holbrook was apparently not without his foibles. One of his friends described him "as a careless man who never took care of anything," stating that "he was a type of the poco curante," nevertheless "liked by every one and regarded as very able in many departments of research" (Gill, 1905: 53). It is difficult to reconcile Holbrook's alleged indifference with his considerable accomplishments.

Holbrook contributed significantly not only to the herpetology of North America but also to the ichthyology of southeastern United States. In total, Holbrook described as new 36 vertebrate taxa that are considered valid today: 12 fish taxa (two genera, eight species, two subspecies; see Table II) and 24 amphibian and reptilian taxa. In view of the fact that he worked under less than ideal conditions, this is truly an impressive array. In order to pursue his interests in zoology, Holbrook had to overcome the impediments of inadequate library (8) and museum resources, absence of local ichthyological and herpetological colleagues with whom to confer, and remoteness from the great centers of natural history investigations. Viewed in that context, John Edwards Holbrook was a most remarkable man, one who has not received the full recognition he deserves as a student of cold-blooded vertebrates (Anderson & Stephens, 2002: 328).

ACKNOWLEDGEMENTS

William A. Roumillat allowed me to borrow his copy of the 1860 edition of Holbrook's Ichthyology for an extended period of time, and Dany E. Burgess made digital photographs of the plates in Roumillat's copy. Ted Steinbock sent me photographs of the extra plates (Nos. XXIX and XXX) that are in his copy of Holbrook's 1860 Ichthyology. Leslie K. Overstreet was extremely helpful, answering a number of queries about the Holbrook material in the Natural-History Rare-Books section of the library at the National Museum of Natural History (Smithsonian Institution). Mary R. Bull, a reference librarian at the Thomas Cooper Library, University of South Carolina, assisted in finding material on Holbrook's artists--as did Helen A. Ivy, Librarian at the Marine Resources Library, College of Charleston. William N. Eschmeyer provided a list of the fish taxa described by Holbrook. Kraig Adler, Benjamin W. Frable, Carter R. Gilbert, Karsten E. Hartel, Theodore W. Pietsch, William A. Roumillat, Mark Sabaj Perez, William F. Smith-Vaniz, and Lester D. Stephens furnished information. G. David Johnson, using Photoshop, improved three of the images in the manuscript. Presubmission drafts of the manuscript were read by Kraig Adler, Theodore W. Pietsch, William F. Smith-Vaniz, and Lester D. Stephens. This is contribution number 424 of the Grice Marine Biological Laboratory, College of Charleston.

Received: 21 August 2014--Accepted: 09 June 2014

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ADLER, K. Ed. 1989. Contributions to the history of herpetology. Society for the Study of Amphibians and Reptiles, Contributions to Herpetology, No. 5. Oxford, Ohio. Pp. 1-202.

AGASSIZ, L. 1857-1862. Contributions to the natural history of the United States ofAmerica. Four vols. Little, Brown & Company, Boston.

AGASSIZ, L. 1872. Eulogy on Dr. J. E. Holbrook. Proceedings of the Boston Society of Natural History 14: 347-351.

ANDERSON, W. D., Jr. 2003. John Edwards Holbrook's Senckenberg plates and the fishes they portray. Archives of Natural History 30 (1): 1-12, 7 figs.

ANDERSON, W. D., Jr. 2012. John Edwards Holbrook's illustrations of fishes in the Bibliotheque centrale, Museum national d'Histoire naturelle, Paris. Archives of Natural History 39 (1): 1-6, 4 figs.

ANDERSON, W. D., Jr. & STEPHENS, L. D. 2002. John Edwards Holbrook (1794-1871) and his Southern Ichthyology (1847-1848). Archives of Natural History 29 (3): 317332, 5 figs.

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CARPENTER, K. E. Ed. 2003. The living marine resources of the western central Atlantic. Vols. 2 and 3: Bony fishes. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5. Food and Agriculture Organization of the United Nations, Rome. [Dated 2002, but published in 2003.]

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COURTENAY, W R., Jr. 1961. Western Atlantic fishes of the genus Haemulon (Pomadasyidae): Systematic status and juvenile pigmentation. Bulletin of Marine Science of the Gulf and Caribbean11 (1): 66-149.

FALK, P. H. Ed. 1999. Who was who in American Art, 1564-1975: 400 years of artists in America. Three vols. Sound View Press, Madison, Connecticut.

FRABLE, B. W., BALDWIN, C. C., LUTHER, B. M. & WEIGHT, L. A. 2013. A new species of western Atlantic lizard fish (Teleostei: Synodontidae: Synodus) and resurrection of Synodus bondi Fowler, 1939, as a valid species from the Caribbean with redescriptions of S. bondi, S. fietens (Linnaeus, 1766), and S. intermedius (Agassiz, 1829). Fishery Bulletin 111 (2): 122-146.

GILL, T. 1864. Review of Holbrook's ichthyology of South Carolina. The American Journal of Science and Arts, 2'd series, 37: 89-94.

GILL, T. 1905. Biographical memoir of John Edwards Holbrook, 1794-1871. National Academy of Sciences Biographical Memoirs 5: 47-77.

GIRARD, C., 1851. On a new American saurian reptile. Proceedings of the American Association for the Advancement of Science 4: 200-202.

HOLBROOK, J. E. 1836-1840. North American herpetology. Four vols. J. Dobson, Philadelphia.

HOLBROOK, J. E. 1842. North American herpetology. Second edition. Five vols. J. Dobson, Philadelphia. (Facsimile reprint edition, 1976. Society for the Study of Amphibians and Reptiles, Athens, Ohio.)

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HOLBROOK, J. E. 1848. Southern ichthyology: or, a description of the fishes inhabiting the waters of South Carolina, Georgia, and Florida. No. III. Pp. 33-60. Wiley & Putnam, New York & London.

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ERRATA:

aqua Vol. 20 (3), 29 July 2014: Hypsolebias trifasciatus, a new species of annual fish (Cyprinodontiformes: Rivulidae) from the rio Preto, rio Sao Francisco basin, northeastern Brazil

Page 131: Wrong author name: Amer Faour not Four

NOTES

(1.) Gill (1905:65) wrote: "Had he [Holbrook] been in the North at the time of the formation of the Academy (1863), he would probably have been one of its founders; but then he was widely separated by distance as well as war and sympathies."

(2.) Jacques Burkhardt was also known as Jacques Burghardt and Jacob Burkhardt; Antoine Sonrel was also called Auguste Sonrel (Kraig Adler, in litt., 16 February 2014).

(3.) Charles Frederic Girard (1822-1895) was educated at Neuchatel, where he became associated with Louis Agassiz, first as a student and later as an assistant. He followed Agassiz to the United States and during the 1850s worked with Spencer F. Baird at the Smithsonian Institution. He spent the summer of 1851 in Charleston helping Holbrook prepare his Ichthyology. In the decade before the American Civil War, Girard's efforts yielded more than 170 notices, papers, and reports. His main interests were fishes and reptiles, but he also wrote on a variety of other organisms (Clark, 1960:319; Hook, 1962:19; Jackson and Kimler, 1999:518).

(4.) From a list of holdings of ichthyological specimens from South Carolina in the Museum of Comparative Zoology, Harvard University, furnished by Karsten E. Hartel, 8 May 1996.

(5.) Kraig Adler informed me (in litt.12 April 2014) that he knew a natural-history book dealer who had on consignment, for a time, a copy of the 1860 edition of Holbrook's Ichthyology that has 30 plates, but that the copy was never sold. Every other copy that I am aware of has only 28 plates. I telephoned the book dealer who told me how to get in touch with the owner of the book. The owner, Ted Steinbock of Louisville, Kentucky, sent me photographs of the two additional plates (Nos. XXIX & XXX). Those plates, reproduced herein, are uncolored and each contains two figures.

(6.) Richard's initials (JHR) were formed on the loose plates and on the plates in the 1855 edition of Holbrook's Ichthy-ologylv upper-case letters "j" and "R" joined by a crossbar.

(7.) Gill (1905:77) reported that Holbrook had three engraved plates ready for issue, of which proofi were obtained by the Smithsonian Institution, and have been bound at the end of the imperfect copy of the first edition. Two of these were numbered ("Pl. XXVIII" and 'PI XXIX") and apparently had been drawn by Richard. ... The other was unlettered and much more finely executed; the artist probably was Sonreh and the plate had probably been drawn and engraved much later than the others [see HOLBROOK'S ARTISTS, above]. On "Pl. XXVIII" are represented the Tailor Herring (Pomolobus mediocris [. Alosa mediocris] and Shad (Alosa sapidissi-ma). On "Pl. XXXIX" are delineated the Menhaden (Brevoor-tia tyrannus) and Gizzard Shad (Dorosoma cepedianum). On the unnumbered plate are ten figures representing various species of Cyprinodonts or Poeciliids, viz., . ...

During my visit in November 2013 to the Natural-History Rare-Books section of the library at the National Museum of Natural History (Smithsonian Institution), I did not find "PI. XXDC" or the unnumbered plate mentioned by Gill, but I did find "Pl. XXVIII" (see ADDITIONAL FISHES DRAWN FOR HOLBROOK, above).

(8.) Although the Charleston Library Society was established in 1748, its holdings of books, journals, and magazines were small compared with those in the libraries of Boston, New York, and Philadelphia (Stephens, 2000: 8).

Grice Marine Biological Laboratory, College of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412-9110, USA. E-mail: andersonwd@cofc.edu

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