A Century of Copepods: The U.S. Fisheries Steamer Albatross

DAVID M. DAMKAER

“You were kind enough to say something very complimentary about the reflected the experiences of other pre­ work that has been done upon the collection of parasitic copepods. It does decessors, the Coast Survey’s Bache not seem to me as if the work was as worthy of credit as is the collection (1872) and Blake (1874). itself. This collection is the natural outcome of the activities of the Bureau The making and equipping of the Al­ of Fisheries extended over a long period of years. Beginning away back in batross has been told by others (Tanner, the sixties of the last century someone has been collecting fish parasites 1885a,b; Hedgpeth, 1945), and will be pretty steadily ever since. The result is the present Museum collection the stuff of many more fine stories to . . . which is by far the largest and richest collection in the world.” come. Likewise, Albatross journeys and accomplishments in far-flung geograph­ — C. B. Wilson letter to W. L. Schmitt, 9 March 1922 ic and academic fields will be rever­ ently spoken of as long as men admire the sea (Agassiz, 1913; Andrews, 1929; Introduction means (Hobart, 1995). This fortunate Dunn, 1996a,b). convergence soon created the Albatross Tanner (1885a) described in incred­ The marine invertebrates of North (1882), a classic vessel at the center of ible detail the equipment of the Alba­ America received little attention before the golden age of oceanography. For tross and how it was used. The men of the arrival of Louis Agassiz in 1846. length of service, areas explored, and the Albatross were as proud of her then Agassiz and his students, particularly volumes written, the Albatross record as any of today’s engineers would be of Addison E. Verrill and Richard Rath- is likely unsurpassed. a satellite space station. A particularly bun, and Agassiz’s colleague Spencer The Albatross (Fig. 1) was not the engaging passage related to the com­ Fullerton Baird, provided the concept first vessel for the U.S. Fish Commis­ mencement of trawling, in some cases and stimulus for expanded investiga- sion (Galtsoff, 1962; Nelson, 1971). to 1,000 fathoms: tions. Baird, through the establishment The Fish Hawk (157 feet), built for of the U.S. Commission of Fish and near-shore investigations, was earlier “When the vessel reaches the in­ Fisheries in 1871, provided a principal by 2 years, and she worked longer by tended station the officer of the 6 years. The Fish Hawk, however, did deck stops her with her stern to the not visit such exotic climes as the Alba- David M. Damkaer, now retired, was with the wind, has the patent log hauled in, Northwest Fisheries Science Center, National tross (234 feet), nor did she have such and then takes his station on the Marine Fisheries Service, NOAA, Seattle, Wash. an army of distinguished investigators grating at the sounding machine, His current address is 21318—195th Avenue, associated with her. The Albatross also S.E., Monroe, WA 98272-9481. where he superintends the sound­ ing, and maneuvers the vessel to keep the wire vertical during the descent. Having satisfied himself ABSTRACT—The marine invertebrates tion. Other collections were sent to the Nor­ that the specimen cup is properly of North America received little attention wegian Georg Ossian Sars. The American bent to the stray line, the sinker ad­ before the arrival of Louis Agassiz in 1846. Charles Branch Wilson eventually added justed, the thermometer and water Agassiz and his students, particularly Addi- planktonic copepods to his extensive pub­ son E. Verrill and Richard Rathbun, and lished works on the parasitic copepods from bottle clamped, the friction rope Agassiz’s colleague Spencer F. Baird, pro- the Albatross. The Albatross copepods from properly attended by a careful man vided the concept and stimulus for expanded San Francisco Bay were reported upon by detailed for the purpose, a man investigations. Baird’s U.S. Commission of Calvin Olin Esterly in 1924. forward of the machine at the Fish and Fisheries (1871) provided a princi- Henry Bryant Bigelow accompanied the last brake, one abaft it with the crank pal means, especially through the U.S. Fish- scientific cruise of the Albatross in 1920. Big­ eries Steamer Albatross (1882). Rathbun elow incorporated the 1920 copepods into his shipped, and another on the grating participated in the first and third Albatross definitive study of the plankton of the Gulf to attend the guide pulley, he will scientific cruises in 1883–84 and published of Maine. The late Otohiko Tanaka, in 1969, lower away gently until the ap­ the first accounts of Albatross parasitic published two reviews of Albatross copepods. paratus is under water, then seize copepods. Albatross copepods will long be worked and The first report of Albatross planktonic reworked. This great ship and her shipmates the small lead to the stray line, copepods was published in 1895 by Wilhelm were mutually inspiring, and they inspire us caution the record keeper to look Giesbrecht of the Naples Zoological Sta- still. out, have the pawl thrown back

61(4), 1999 69 and the crank unshipped, and order ‘Lower away!’”

With respect to the copepod crusta­ ceans collected by the Albatross, it is essential to recognize the two princi­ pal, although artificial, copepod groups. These groups, the free-living and the parasitic copepods, are generally caught with different methods. On the Al­ batross, the parasitic copepods were mostly taken from fish, although cope­ pods could be parasitic on most any marine animal, even whales. The free­ living copepods were taken primarily by plankton nets, which would catch pelagic as well as occasional bottom­ dwelling copepods. In some plankton Figure 1.—U.S. Fisheries Steamer Albatross under partial sail in Alaska (from Towns­ samples, free-swimming stages of par­ end, 1901). asitic copepods were encountered. I am pleased to record that copepod investigators were present on the first was a major advance in ecology of investigate the environment of the com­ and the last Albatross cruises, and that our seas. Although there was consider­ mercial fishes, Verrill began compre­ considerable and praiseworthy attention able new information on many inverte­ hensive surveys for the Fish Commis­ was given to copepods throughout and brate groups, especially mollusks (Ver­ sion in 1871 (Coe, 1930). beyond the active life of this famous rill) and large crustaceans (Smith), one Verrill was at the center of America’s oceanographic ship. Most of the prom­ found little on copepods: major growth in zoology from Agas­ ising young men who sailed with the Al­ siz’s prime through the experimental batross had brilliant careers. This great “The . . . minute Copeopoda [sic] era of the first quarter of the 20th cen­ ship and her shipmates were mutually of our coast have not yet been suf­ tury. Verrill participated in the general inspiring, and they inspire us still. ficiently studied by any one for trend of zoological fashion from tax­ us to attempt to enumerate even onomy, to evolutionary adaptations, to A. E. Verrill and S. I. Smith the more common species” (Ver­ embryology, and to experimental phys­ The Albatross was launched in an rill and Smith, 1873). iology and genetics. However, taking a undercurrent of anticipation of an im­ long look back at his work, the single mense harvest of little-known marine Only 20 copepod species were includ­ item that probably still brings the great­ life. Those who would go down to the ed, and all but one of these were parasit­ est recognition is the 1873 report on the sea for invertebrates were not the prime ic. This was the basis on which the Al­ northeastern invertebrates. movers of the Albatross, for its essen­ batross copepod collections began. [In In this monumental study, Verrill tial task was always toward vertebrate Verrill and Smith’s taxonomy, the old was ably assisted by his brother-in-law fisheries research. However, persons of lower-crustacean group Entomostraca Sydney Irving Smith (1843–1926). influence recognized direct and indirect included the “Copeopoda” (apparently Smith (Fig. 2) had followed Verrill to links between fish and invertebrates, only the planktonic species, and here Yale as a student; in the following year, such as the copepod crustaceans, and only Sapphirina sp.) and the Siphono­ Verrill married Smith’s sister. In 1867, their study was encouraged in those stoma, the parasitic species (19 listed, Smith became Verrill’s assistant, co­ early days. The Fish Commission had all from fishes). In early references, the worker, and eventually coauthor on the no staff for this work, so it was obliged advanced parasitic forms were some­ Fish Commission’s invertebrate report to accept outside help. times also referred to as “lernaeids.”] (Coe, 1929). In 1875, Smith was ap­ Indeed, the very first annual report of Addison Emery Verrill (1839–1926) pointed Professor of ComparativeAnat­ the Commissioner of Fisheries summa­ had been a student of and assistant to omy at Yale. He was also one of the rized the essential knowledge of Amer­ Louis Agassiz at Harvard’s Museum of founders of the Marine Biological Lab­ ican marine invertebrates to that time Comparative Zoology. In 1864, Agas­ oratory at Woods Hole. (Verrill and Smith, 1873). There had siz recommended Verrill to Yale, where been nothing approaching this in the he remained for 43 years as Professor R. Rathbun generation since Gould’s 1841 pioneer of Zoology. Verrill continued his stud­ Richard Rathbun (1852–1918) (Fig. 3) report on the invertebrates of Massachu­ ies of invertebrates along the northeast was linked to the Albatross even before setts. Verrill and Smith’s large volume coast. Responding to Baird’s desire to it was afloat. That he became the first

70 Marine Fisheries Review Figure 3.— Richard Rathbun (1852–1918) (from Benjamin, 1918).

Figure 2.—Sydney Irving Smith assistant. Rathbun remained at Cam­ retary until 1888. In 1880 and 1882, (1843–1926) (USNM photograph). bridge until 1875. During his summers Rathbun was on the Fish Hawk, sur­ there, he began his long association veying fishery resources out to the Gulf with Spencer Baird, at first as a volun­ Stream: scientist to report on the enormous teer scientific assistant. numbers of copepods collected from In 1875, Rathbun joined Hartt’s staff, “At each dredging station, collec­ this celebrated ship gives him a promi­ in Brazil, where Hartt was conducting tions were made with the towing nent part in the present narrative. He a geological survey. Rathbun’s interests net [Fig. 4], which is designed to was born in Buffalo, N.Y., and at age were particularly in coral reefs and fossils. scoop in the free-swimming forms, 15 began a 4-year apprenticeship as a When Hartt died of yellow fever in March living at the surface and at in­ clerk for his father’s contracting and 1878, Baird offered Rathbun a position as termediate depths. . . . The an­ stone business. At this early age, Rath­ a paid Scientific Assistant in his new U.S. imals obtained [included] cope­ bun had a fascination with fossils of Fish Commission. Rathbun was detailed pods, the latter frequently occur­ western New York, finding the first to Verrill at Yale where, although Verrill ring in countless numbers. They in his father’s own quarries (Rathbun, was in charge of the work, Rathbun had serve as food for the surface-swim­ 1969). His collections and studies of day-to-day responsibility, making many ming fish, such as menhaden and these, and his obvious knowledge and invertebrate collections over the follow­ mackerel” (Tanner, 1884). enthusiasm, led him even then to the ing 2 years. Rathbun, although based at honorary post of curator of paleontolo­ various offices, retained this formal posi­ As both the USNM and the Fish Com­ gy at the museum of the Buffalo Soci­ tion with the Fish Commission through mission grew, Baird relied more and ety of Natural History. At the urging of 1896 (Benjamin, 1918). more on Rathbun for administrative Charles F. Hartt (an Agassiz pupil and Rathbun was therefore a protégé of duties, until Baird’s death in 1887. This Professor of Geology at Cornell), Rath­ both Agassiz and Baird. Through Baird, turned Rathbun into an administrator of bun began science studies at Cornell Rathbun also spanned both the Fish remarkable ability. University in 1871 (Benjamin, 1918; Commission and the growing U.S. Na­ One of the essential housekeeping Coe, 1918). tional Museum (USNM), and continued chores was the review and recording of Rathbun’s first publications, on fos­ the close relations between the two in­ the earliest lists of stations and activ­ sils, began at this time. His research stitutions. In 1880, with a new National ities of the Fish Commission, copub­ soon brought him to Cambridge, Mass., Museum building, Rathbun transferred lished by Rathbun (Smith and Rath­ where he took classes from Louis to Washington, D.C., and was assigned bun, 1882). This was followed by a Agassiz at the Museum of Compar­ as Curator of the Department of Marine number of summaries of North Atlan­ ative Zoology in 1872–73, Agassiz’s Invertebrates. Rathbun thus joined an tic fisheries, including accounts of the last years. In September 1873, Rath­ eager community of biologists in the natural history of crustaceans, worms, bun was on the Bache, dredging inver­ Nation’s capital, in time to be one of echinoderms, and sponges, comprising tebrates off the coast of Maine with ten founders of the Biological Society altogether more than 500 pages (Coe, A. S. Packard, at that time an Agassiz of Washington, for which he was sec­ 1918). This was his best zoological

61(4), 1999 71 work, and it established Rathbun as an authority on the investigation and eco­ nomics of marine zoology. Rathbun’s first obvious connection with the Albatross was his role in the preparation of the extensive catalog accompanying the U.S. exhibit at the “Great International Fisheries Exhibi­ tion” in London in 1883. Rathbun au­ thored a section on the economic crusta­ ceans, worms, echinoderms, and spong­ es. In a second section, he reviewed the “apparatus of scientific research” (Rath­ bun, 1883). The U.S. exhibit outlined fisheries work by American scientists, particularly from U.S. Coast Survey and Fish Commission vessels, but the Albatross, her equipment, and expected scientific harvest, was the main feature of the presentation. Baird had planned to move Rathbun wholly into the USNM, but after his Figure 4.—Surface net and dip nets in use on the Albatross (from Townsend, 1901). death, George Brown Goode (1857– 1896), the interim Commissioner of Fish­ eries, persuaded Rathbun to retain his “Absorbed in the details of his var­ first Albatross scientific cruise conclud­ Fish Commission position. During this ious activities, all of which had to ed thus: time, Rathbun assisted the international do with the institution to which fur seal commission and thereby contin­ he gave his life, he had but little “You will give to the naturalist of ued with the results of Albatross inves­ time for other interests” (Benja­ the expedition all possible facili­ tigations through 1896. In 1892, he was min, 1918). ties for collecting and preserving the U.S. representative for fisheries on the such specimens as you may meet Boundary Commission and visited both One of Rathbun’s great accomplish­ during the cruise. coasts and the Great Lakes, from the Gulf ments, from 1913, was the present Nat­ of St. Lawrence to Cape Flattery. Rath­ ural History Museum building, where “P. S.—The operations of dredg­ bun’s extensive reviews of the Nation’s I was employed for several years, and ing and trawling should be car­ fisheries (e.g. Rathbun, 1884, 1899) are where Rathbun was still remembered ried on as frequently as opportu­ classics of their kind. Altogether, Rath­ with fondness. (Rathbun’s attention to nity offers. . . .” (Tanner, 1885b). bun published about 100 papers. In 1894, every detail of the planning and con­ Rathbun was given an Honorary Doctor­ struction of this building ensured him The Albatross then put to sea on 24 ate by Bowdoin College, and he includ­ the repetitive duty with respect to the April, and the first “official” sampling ed among his many affiliations, member­ building of the National Gallery of Art.) was with a beam trawl at Station 2007 ship in the American Fisheries Society. Waldo L. Schmitt (1887–1977), who on 27 April between Washington and Rathbun did not leave the Fish Com­ had been Naturalist on the Albatross, Norfolk, Va. mission until after Goode’s successor, was the protégé of Rathbun’s sister, The earlier parts of Tanner’s (1885b) Marshall McDonald, died in 1895. Mary Jane Rathbun (1860–1943), crus­ report of the first Albatross year spoke Rathbun began working solely for the tacean specialist at the USNM. Through mainly of fish, but soon the response to USNM on New Year’s Day 1897. His their close relationship, Richard Rath­ Baird’s orders “to determine . . . the bi­ previous work there ensured an extraor­ bun extended every courtesy to Schmitt, ological peculiarities” of the New Eng­ dinarily rapid rise. Within a month, who spoke of him often and with great land seas became evident as detailed Rathbun was Assistant Secretary of the respect. I was privileged to consider observations and the concurrent excite­ Smithsonian Institution, the parent or­ Schmitt a friend in the last decade of ment of Verrill’s crew reached the Com­ ganization of the USNM, in which, as his life; I was that close to the Alba­ mander. With “the naturalists finding an old habit, he retained his other of­ tross. no difficulty in picking over the con­ ficial title. In mid 1898, he was named The Albatross was taken on a prelim­ tents of the trawl,” log-book entries in­ Director of the USNM. He held this po­ inary cruise in March 1883, when much dicated the examination of the surface sition until 1914, and the Smithsonian’s of the equipment was tested. Baird’s and gills of fish for parasites. By May, until his death in 1918: orders, dated 10 April 1883, for the with “many new and interesting forms

72 Marine Fisheries Review having been brought up,” Commander ist, James E. Benedict (Tanner, 1885b). batross during 1883. Verrill acknowl­ Z. L. Tanner (1835–1906) was making Underwater lights were often used, typ­ edged that “very interesting additions familiar entries of salps, foraminifera, ically just beneath the surface, aiding in to our collections were made in nearly globigerina, large red crabs, starfish, the capture of several surprising forms. every class.” His particular interest was squid, “coral growth,” barnacles, and Among these were “early stages of var­ in the mollusks. Smith studied the crus­ lobsters, reflecting the growing inter­ ious Copepods,” passed to Professor taceans, principally the decapods, while est of all in that portion of the harvest. Sydney Smith. Rathbun dealt with the copepods. The Some of these notes mentioned genera; Richard Rathbun participated on var­ only copepod specifically mentioned by “minute crustaceans” sometimes made ious legs of the first scientific cruise Verrill was found on a red sea anemone, their way into the ship’s logs. “A number of the Albatross, in 1883, dredging and of unpleasant if not dangerous stinging of naturalists” (July) became “a large trawling from Woods Hole to the Gulf abilities, common below 150 fathoms: party of naturalists” (September), and Stream until 14 November, when the Al­ produced “many valuable specimens.” batross tied up in the Washington Navy “A very singular, large, soft, pink­ In October, “all the fish taken were Yard. Among the invertebrates which ish Lernean crustacean (Antheach­ carefully examined for parasites.” attracted Rathbun most were parasitic eres duebenii [Michael] Sars, fig. In spite of the careful handling of copepods found on fish. Rathbun also 167–8) lives parasitically in the ship and equipment, “the trawl was was on the third Albatross cruise, in stomach of this Actinian, with lost” was an all-too-common entry in the same area, in 1884, and from 1885 which it agrees in color. It is not the early reports. In one case, the Com­ to 1887 he published the first three ac­ uncommon” (Verrill, 1885). mander stated that counts of Albatross copepods (Rathbun, 1885, 1886, 1887). In another section, it was acknowl­ “. . . the accident was the result Rathbun published pre-Albatross lists edged that of kinking and the fault rests be­ of invertebrates from the collections of tween myself and the dredge rope. the USNM (Rathbun, 1882a, b); these “. . . the Copepoda . . . are very I have not yet been able to judge mentioned copepods, but only from the abundant, both in the lots obtained satisfactorily which is responsible Fish Hawk or from the Commission’s in the trawl-wings and in the sur­ for the frequent losses during our work along the northeast coast. The first face collections. Very many fine present trip” (Tanner, 1885b). Albatross copepods, parasites of fish, species were noticed, but they have were listed and described by Rathbun in not yet been reported upon by Mr. One mystery for which the Albatross 1885. At that date, in the museum col­ Rathbun, who has charge of [this was charged was to determine what At­ lections, there were 22 identified spe­ group]. He has studied a number lantic menhaden, Brevoortia tyrannus, cies of copepods (not counting 4 species of interesting and novel forms of eat. The skipper of a fishing boat met at of argulids, then classified as copepods), Lerneans found parasitic on sever­ sea in August said that “their food is a of which 9 were from Albatross Cruis­ al of the deep-sea fishes” (Verrill, very small marine insect, which appears es 1 and 3 off the northeastern United 1885). under the microscope to be a species States. The various legs of these first of crab” (Tanner, 1885b). This is not a two Atlantic cruises were under Verrill’s The trawl wings (Fig. 5) were muslin bad description of copepods, which, of direction, but Verrill himself apparently nets attached so as to be just above the course, make up a large part of the food never shipped on the Albatross. bottom at the ends of the trawl frame. of the filter-feeding menhaden. There­ Rathbun (1886) followed soon with These were first used by Verrill on the fore, it soon was apparent that menha­ a second publication on parasitic co­ Fish Hawk. Verrill noted that many pe­ den, unlike Atlantic cod, Gadus morhua, pepods. This included figures for three lagic species which were not caught at the were not “ground grubbers,” which was new species of Chondracanthus taken surface were collected in the trawl wings, commonly held, nor did menhaden eat during the Albatross first cruise, in but that the precise depth of collection mud. Another old fisherman, speaking 1883, as well as a second Albatross could, of course, not be determined. of the food of mackerel, Scomber spp., record (1885) for a previously reported For the Gulf Stream, said that it “consists in part of what species. is known among fishermen as ‘cay­ In a third and final report, Rathbun “Copepod crustaceans are usually enne,’ only seen when the water is very (1887) described three new species of the most abundant forms of small smooth. It then appears to skip out of parasitic copepods from Albatross col­ surface animals, occurring in great the water.” This is an early American lections, taken from sharks, menhaden, quantities and of many genera reference to the surface swarming of the and bluefish, Pomatomus saltatrix, in and species. Various species of copepod Calanus (and others), a phe­ Vineyard Sound or off Florida. the genus Calanus are the most nomenon by then well known to Scan­ Before leaving Rathbun’s direct par­ common. Several species of the dinavian biologists (Gunnerus, 1770). ticipation, it must be noted that Verrill genus Sapphirina were taken, Copepods also surfaced briefly in the (1885) published a large report on gen­ some of them very brilliant in report of the Albatross’s first Natural­ eral invertebrates collected by the Al­ colors” (Verrill, 1885).

61(4), 1999 73 Verrill’s report only hinted at what Nelson, 1971). In describing the de­ favorable auspices. The ship was were to become overwhelming numbers parture of the Albatross for the Pacific well equipped and thoroughly sea­ of pelagic copepods in the Albatross Ocean on 20 November, Commander worthy in every respect. She had collections. Tanner wrote that on board an efficient corps of of­ The second, the longest, and the most ficers and scientists, and her crew important period of the Albatross began “It is seldom the fortune of men to could not be excelled” (Tanner, at the end of 1887 (Hedgpeth, 1945; start on a long voyage under more 1891).

The Albatross sampled around South America, via the Strait of Magellan, and reached San Francisco 11 May 1888, 4 days ahead of schedule. W. Giesbrecht The Albatross collected vast num­ bers of planktonic copepods, especial­ ly in the Pacific Ocean. The first report of these was published in 1895 by Wil­ helm Giesbrecht (1854–1913), the vir­ tuoso German copepodologist on the staff of the Naples Zoological Station (Damkaer, 1995b). Giesbrecht (Fig. 6) was born in Dan­ zig and earned a doctorate at the Uni­ versity of Kiel. An appointment as guest researcher at Naples in 1881 re­ sulted in a lifetime position. Giesbre­ cht’s major effort was the unexcelled 1892 monograph on pelagic copepods. Giesbrecht was the first to clearly show that parasitism arose independently at several phases of copepod evolution, and through consideration of morpho­ logical development, he removed the prevailing distinctions between free-liv­ ing and parasitic copepods. Through his writing and artistic skills, he was a re­ spected editor at Naples and for other publishers. The Giesbrecht (1895) copepod col­ lection was from the 1891 Albatross cruise through the warm Pacific from California to the Galapagos. This was the first of three cruises under the direc­ tion and patronage of Alexander Agas­ siz, esteemed son and student of Louis Agassiz. Ironically, this exotic region had been traversed by the U.S. Explor­ ing Expedition (1838–1842), the Chal­ lenger (Great Britain, 1872–1876), and the Vettor Pisani (Italy, 1882–1885), with the copepods incorporated into classic publications. Giesbrecht him­ self published several reports on cope­ pods from the latter cruise, and he sum­ Figure 5.—The improved beam trawl, with trawl wings, as used on the Albatross marized these in his well-known 1892 (from Tanner, 1885a). monograph.

74 Marine Fisheries Review Although the number of individual Figure 6.— copepods was small, this Albatross col­ Wilhelm Giesbrecht lection, only 32 samples, contained a (1854–1913) large number of species; this now well­ (Naples known feature of tropical waters was Zoological not so obvious in the early days. As a Station supplement to earlier work, Giesbrecht photograph). had a keen interest in the collection. Most samples were from the surface, but there were some reaching 200 to 600 m, and one to 3,000 m. Three sam­ ples had been obtained with a closing net (Fig. 7). From the deep-sea sam­ ples, Giesbrecht suggested that some surface forms from cold-water regions were living at depth in the warm-water regions. This is another notion that has firmed with time. Giesbrecht also found special interest in the species which were common to both the Atlantic and Pacific. Giesbrecht discussed 48 Alba­ tross copepod species, describing and illustrating 10 new species and 3 new genera. Giesbrecht honored both Alexander Agassiz and Lt. Commander Z. L. Tanner ogist. Ossian Sars’s international fame his already extensive published works through the new copepods Pontella agas­ was based on his revelation that most on the parasitic copepods from the Al­ sizi and Heterochaeta tanneri. of the commercial fishes had plankton­ batross. A measure of the task may be ic eggs and larval stages. His interests suggested by the fact that Wilson, too, G. O. Sars then moved toward the lower crusta­ died with the work far from complete. Other Albatross collections of pelag­ ceans, and during the next 60 years, ic copepods were sent to the Norwegian his production of definitive and well­ C. B. Wilson Georg Ossian Sars (1837–1927), who illustrated monographs made him the Charles Branch Wilson (1861–1941) died with his report less than half com­ best-known Norwegian zoologist. He probably never looked seriously at a pleted. The material comprised seven became a professor at the University of copepod until he was 35 years old, distinct collections: 1) around South Christiania. He was a planner, partici­ when he became a professor, and later America and through the eastern Pacific pant, and reporter for the Norwegian head of the science department, at the (1887–88), 2) across the tropical Pacific North Atlantic Expedition (1876–78). State Normal School in Westfield, Mass. to Japan and Kamchatka (1899–1901), Another acclaimed publication was Before that time, he had experimented 3) Hawaiian Islands (1902), 4) Cali­ Sars’s account of the copepods col­ with embryonic development of am­ fornia and south (1904), 5) Alaskan lected by his brother-in-law, Fridtjof phibians. Opportunities for field work salmon (1903, 1905), 6) Northwest Pa­ Nansen, during the Norwegian North in Jamaica and in California in 1897 cific (1906), and 7) the 3-year cruise Polar Expedition (1893–96). Sars, as and 1899, respectively, turned most of to the Philippines (1907–10), with the many of his efforts proved, was not his energy toward parasitic marine co­ most specimens. Sars sorted and la­ averse to field work. Indeed, his inge­ pepods. In fact, by expressing a solid in­ beled about 40% of the copepods (those nious applications are still appreciated terest, Wilson received the mantle from collected before 1905) in this large as­ by those who would again collect ma­ Rathbun and became his protégé: sortment, and he made drawings of new terial revealed by him. However, the species, at which time he felt an obli­ enormous numbers of specimens sent “I am pleased to know that some gation to an earlier study on the cope­ to Sars from all over the world ensured one has again taken up the sub­ pods collected by the Prince of Monaco. that he would remain near his labora­ ject of parasitic crustaceans in this Sars devoted the rest of his life to fin­ tory (Damkaer, 1993). country, and trust that you may ishing that exquisite monograph and After Sars’s death, the Albatross col­ be able to make good progress in never looked at the Albatross copepods lection, with Sars’s notes and drawings, the work. Some years ago I made again. was returned to the USNM, and the quite a collection in that line at Sars (Fig. 8) was the son of Michael work was taken up by C. B. Wilson, Woods Hole and elsewhere along Sars, a priest turned world-class zool- who added the plankton copepods to the coast, but lack of opportunity

61(4), 1999 75 to work them up accounts for the little I have published. The spec­ imens are now here in the Na­ tional Museum in good condition. Would you care to study and report upon them to the Museum? As I recall it, there are still several un­ described species and much else of interest in the material. I shall be glad to hear from you on the subject at your convenience. Yours very truly, Richard Rathbun, As­ sistant Secretary” (24 September 1900).

Wilson (Fig. 9) joined the circle of in­ dependent Fish Commission research­ ers in 1901 by spending the first of many summers at the Woods Hole Laborato­ ry. He took up the parasitic copepods with a vengeance, and for a generation no one in this country, and few abroad, came even close to his output and quali­ ty (Damkaer, 1995a). Like Sars, Wilson was the recipient of many specimens from around the world, as his long list of publications clearly shows. Most of Wilson’s copepod papers were published by the USNM. Wilson’s papers can be grouped somewhat ar­ bitrarily as systematic reviews (source not stressed) or as summaries of geo­ graphically limited collections. Not sur­ prisingly, the collections available to him included many specimens from the Albatross. In the first category are eight reports describing Albatross copepods. The first, Wilson (1905), was based mainly on Fish Commission collections off the U.S. east coast. Rathbun had turned over to Wilson all of his lists and notes from preliminary work on this collec­ tion. Although some Albatross cope­ pods were mentioned, including some taken on the trip around South America and in the Pacific, the source was not given for all specimens; USNM records would have to be revisited to determine the exact number of Albatross cope­ pods from this study. The emphasis in the report was on hosts, descriptions, and redescriptions of species. Recall that many “good” parasitic copepods had been described from hosts obtained at markets, and that the precise location Figure 7.—Tanner’s first closing net, as used on the Albatross (from Tanner, 1893). was less important in those days than a

76 Marine Fisheries Review description of the species. Among the Figure 8.— accounts of 36 caligids were 10 from Georg Ossian Sars the Albatross, including a new genus (1837–1927) (photograph and 6 new species, one of which had ca. 1925 by been attracted by the electric light. Typ- Hjalmar Broch, ical for Wilson’s papers, there was an University of Oslo), excellent review of the literature. The next two papers, Wilson (1907 a,b), reported three Albatross species, one of which, from 1887–88, was new, and from various Atlantic and Pacific localities, about 20 species from the Al­ batross, one of which was new. A large and well-known monograph covering marine and freshwater para­ sitic copepods of Wilson’s (1915) Le­ rnaeopodidae included ten Albatross species, seven of which were new. One of the latter was Brachiella nitida, found in the mouth of the fish Albatrossia pec­ toralis from a 1906 North Pacific local­ ity. A companion report on the Lernae­ idae (Wilson, 1917) described 13 At­ lantic and Pacific Albatross species, of which 9 were new species. Three new lernaeid genera from the Albatross col­ lections were introduced. A following report (Wilson, 1919) dealt with four Atlantic and Pacific Albatross species, three of which were new. A new genus Figure 9.— was described from Albatross collec- Charles tions off New Jersey in 1883. Wilson Branch Wilson (1924) reported two Albatross cope- (1861–1941) pods, one from British Columbia and (photograph ca. 1925 by one from the Strait of Magellan. The Marine Biological date of the latter collection, stated as Laboratory, 1908, cannot be correct; conversely, Woods Hole). perhaps the locality is incorrect. The final paper in this group of sys­ tematic reports is Wilson (1944), com­ pleted only a few weeks before he died. Therefore, this was the last paper of Wilson’s own doing. Only one cope­ pod species was clearly from Albatross collections, and this represented a new genus and species from the New Jersey coast. In Wilson’s papers based on geo­ graphically limited collections are six with Albatross material. The first is Wilson (1908) on parasitic copepods found on fishes of the Pacific coast, with descriptions of new genera and species; “some of them are the result of work done upon different expeditions of the steamer Albatross.” Except for two lots sent directly to Wilson (one from

61(4), 1999 77 Ritter collected in 1904 and another duction of any agitation in refer­ and dredging in the Pacific Ocean (i.e. from Kofoid), all Albatross specimens ence to an 8-hour working-day. . . . plankton copepods), essentially the were sent to Wilson via the USNM. Evidently in the interests of science same samples that had been sent to Of 13 Albatross species, 4 were new; a 17-hour day was not deemed im­ Sars, was covered in Wilson (1950). Wilson also described one new genus. possible by those enthusiasts. This collection comprised thousands Two new copepods, one from an echi­ “If all this wealth of material be of mostly surface samples; in amount noderm in Japan and the other from a considered in connection with the and coverage of plankton, these com­ Hawaiian mollusk, were reported from several papers that Doctor Rath­ pared with the greatest collections to Albatross collections in Wilson (1921). bun published, it can easily be seen that time: U.S. Exploring Expedition, Wilson (1923) described a new genus that he would have become one Challenger, Prince of Monaco cruises and species of parasitic copepod from a of the foremost authorities upon (1885–1914), Siboga (1899–1900), and Lower Californian annelid dredged by copepods, had not his executive the Carnegie (Wilson, 1942). Altogeth­ the Albatross in 1911. duties compelled him to give up er, from the Albatross, there were 473 Wilson’s (1932) “Copepods of the research work. It is, therefore, em­ copepod species (including one larval Woods Hole region Massachusetts” is inently appropriate that the present Pennella, a parasitic copepod), of which likely the most-used American publica­ paper, which embodies so much 29 were new. Males of pelagic cope­ tion on copepods—the copepod “bible” of his work, be dedicated to his pods are frequently scarce, and this col­ for a generation, and still a ready com­ memory” (Wilson, 1932). lection provided previously unknown panion for those working in the North males for 26 species. From similar areas Atlantic. Wilson defined this region as The Albatross was in the Atlantic for traversed by other expeditions, 93 spe­ ca. 200 miles wide and 150 miles long: only a relatively short time, although cies were found only by the Albatross virtually all marine copepods reported (including, of course, the 29 new spe­ “Any reduction of the area would in Wilson (1932) could have been col­ cies); only 12 species were on all of the of necessity eliminate some of the lected from that ship. Most of the cope­ lists. excellent material gathered during pod species were from Grampus collec­ The methods employed on the Alba­ the earliest cruises of the . . . Alba­ tions, which may have had a higher pri­ tross were not quantitative, and there tross. As these were all included ority. Noted from the Albatross were 57 were few comparisons of numbers. The in the notes and drawings made by calanoid species and 11 species from nets were 1) surface ring nets, 12 in to Dr. Richard Rathbun, it has been other groups, including a new genus 51/2 ft; an electric light added to num­ deemed best to keep the collection and species. This represents 18% of the bers of copepods captured; 2) paired intact.” 373 copepods reported and described in ring nets for horizontal and vertical this monograph. Many of the other co­ tows, sizes to 10-ft diameter; 3) ca. 3-ft Rathbun spent his summers and early pepods were from fresh water or brack­ diameter closing nets at intermediate autumns during 1880–85 at Woods ish ponds and beaches. Wilson included depths: Hole, and his investigations formed the a key to all the known copepod genera, basis of this classic work. He made co­ worldwide, ensuring the broad and last­ “A good start has been made pious notes regarding living forms and ing value of this work. toward a knowledge of the sur­ their colors. For marine copepods in Wilson’s (1935) report on parasitic face plankton, but it is only a start, this area, northern and southern faunas copepods from the Pacific coast sup­ and a broad field is still left for overlap, so diversity is high, and a plemented Wilson (1908) by adding future investigation and discov­ three species from the Albatross, one of ery. Many species have been ob­ “ . . . remarkable feature of the pres­ which was new and named for Ed Rick­ tained in vertical hauls from con­ ent collection is the large number etts of “Cannery Row” fame (Hedg­ siderable depths, but we have ab­ of species that have never before peth, 1978a, b). Other specimens came solutely no definitive knowledge been reported from the Woods Hole from Stanford University and Univer­ as to where they entered the net” region . . . [or] anywhere along the sity of California collections. (Wilson, 1950). Atlantic shores of North America. In Wilson’s publications covering Al­ . . . How does it happen that so batross copepods, only the last, Wilson Each species was described, and many many species escaped the atten­ (1950), dealt exclusively with this col­ were illustrated. Each station is listed tion of those investigators . . . al­ lection. Wilson died in 1941, and his separately (location, depth, date) with ready mentioned? The first answer colleagues at the USNM, especially the species found there. Wilson was is suggested by a perusal of the Waldo L. Schmitt, completed the refer­ lavish in his praise for Sars and the notes and records made by Doctor ences, edited the records, and brought preliminary work on this collection; Rathbun and his associates. They this report to publication. if not for the rules of nomenclature, were exceptionally thorough in The Albatross Atlantic forms were Sars would have been credited with their search, and their task was ac­ addressed in “Copepods of Woods many new species. Wilson (and his col­ complished long before the intro­ Hole”—the remainder, from tow nets leagues) included an exceptionally good

78 Marine Fisheries Review literature list, which has been my con­ Giesbrecht, Sars, and Thomas and stant companion for 40 years. Andrew Scott, died August 18, Therefore, the Albatross copepods re­ 1941” (J. A. Fleming of the Carn­ ported by Wilson in 14 publications egie Institution in Wilson, 1942). comprised 623 pelagic and parasitic spe­ cies, including 66 new species and 9 new C. O. Esterly genera. The numbers would be larger if In 1912 and 1913, the Albatross sur­ the Albatross argulids, no longer held as veyed the biology and hydrography of copepods, were included. San Francisco Bay in a cooperative Wilson retired in 1932 but retained effort by the U.S. Bureau of Fisheries his laboratory at Westfield, Mass., and (the U.S. Fish Commission was re­ returned in summers to Woods Hole. named in 1903) and the University of Toward the end of his life, through the California. Francis B. Sumner (1874– urgings of Schmitt, Wilson was per­ 1945) was Naturalist for the Bureau, and suaded to take on the planktonic cope­ Charles A. Kofoid (1865–1947) repre­ pods from the Albatross. Schmitt had sented the University. Waldo Schmitt more than one regret over this; first, be­ was a Scientific Assistant. cause he felt that the task contributed to Nearly 200 stations were occupied Wilson’s death, and second, that Wilson throughout San Francisco Bay, from was never able to give the planktonic January 1912 until July 1913. Plankton Figure 10.—Calvin Olin Esterly (1879– copepods the deliberation that he might samples were collected approximately 1928) (photograph ca. 1926, Occiden­ have as a younger man. The day before every 2 months; often there were mul­ tal College, Los Angeles). Wilson’s death, Schmitt visited him at tiple plankton samples at a particular Westfield, where Wilson went over the station. Although he did not participate Albatross records and showed Schmitt in the Albatross cruises, the separated dissertation on copepods. Copepods re­ some of the recent additions to his copepods from these samples were sent mained the principal research field to C. O. Esterly (Fig. 10) for his identi­ “ . . . incomparable library of co­ fications and report. “. . . to which Esterly set himself, pepod literature that he had built Calvin Olin Esterly (1879–1928) had and pursued for more than two de­ up in the course of a busy lifetime. already distinguished himself in this cades without a moment of doubt . . . [Wilson was] one of the most area by publishing the first compre­ about its worth, of swerving in valued scientific collaborators on hensive accounts of copepods from the purpose relative to it, or relaxation the rolls of the Museum. He be­ U.S. west coast. The San Francisco in pursuing it. . . . The history of queathed to the Museum his li­ Bay report (Esterly, 1924) supplement­ marine biology does not furnish brary of copepod literature, which ed Esterly’s long series of papers on the many instances of such a well-bal­ is perhaps the most complete of systematics and distribution of pelagic anced combination of field, statis­ its kind in the world” (Schmitt, copepods from the San Diego region, tical work and laboratory experi­ 1941). beginning in 1905. From the San Fran­ mentation as is furnished by the cisco Bay samples, Esterly described researches of Esterly on the Co­ Wilson’s library has been kept up to 11 species and their marked seasonal pepoda of the ‘San Diego area’ of date as a distinct world-class resource. abundances. The bay-wide distribution the Pacific” (Ritter, 1928). Many of Wilson’s classic papers on of four species was related to salinity. parasitic copepods, systematically deal­ Esterly’s characteristically practical il­ Returning from Harvard, Esterly was ing with groups worldwide, included lustrations make this report useful still. appointed Professor of Zoology at Oc­ annotated literature reviews that have Esterly was born in Texas and earned cidental College in Los Angeles, a post never been surpassed. In contrast, his a B.A. from the University of Cali­ he held for the rest of his short life. He papers on planktonic copepods from fornia in 1902. His interests turned was named to the “Non-Resident Staff” the Albatross (Wilson, 1950) and the early toward marine invertebrates, and of Scripps Institution of Oceanography Carnegie (Wilson, 1942) were essen­ he came to the notice of William E. in 1910. Esterly only worked one full tially limited to the specific collections, Ritter, the first Director of Scripps In­ year at Scripps, but he was a virtual no doubt due to the necessities of post­ stitution of Oceanography. Esterly was volunteer every summer otherwise. He humous editing: with Ritter at the institution’s begin­ died at La Jolla, Calif., in 1928. ning, as the San Diego Marine Bio­ “Dr. Charles BranchWilson, the last logical Laboratory, in 1903 (Raitt and H. B. Bigelow of that outstanding group of great Moulton, 1967). It was fitting that copepods would monographers of the marine cope­ Esterly studied at Harvard under E. L. have been collected by the multifacet­ pods which included Brady, Dana, Mark, obtaining a Ph.D. in 1907, with a ed Henry Bryant Bigelow (1879–1967)

61(4), 1999 79 Some copepods from the Albatross collections: Euchaeta concinna (a) from Wilson (1950); Paralabidocera amphitrites (b) from Esterly (1924); Gaetanus recticornis (c) from Wilson (1950); Scottocalanus infrequens (d) from Tanaka (1969); Pseudomolgus hawaiiensis (e) from Wilson (1921); continued on next page

80 Marine Fisheries Review (Cont. from facing page) Pandarus cranchi (f) from Rathbun (1886); Kroyerina elongata (g) from Wilson (1944); Lernanthropus pomatomi (h) from Rathbun (1887); Lernaeopoda bicauliculata (i) from Wilson (1908); Phrixocephalus cincinnatus (j) from Wilson (1908). Figures a–d are of free-living, planktonic species; the others are of parasitic copepods.

61(4), 1999 81 on the last scientific cruise of the Al­ the problems of quantitative investiga­ up Albatross, we would, I suppose, have batross, from February into May 1920. tions for copepods and acknowledged been required to consider those cope­ The last Albatross station was occu­ the primary deficiencies in this study. pods as a continuation of the Albatross pied 19 May 1920, and Bigelow was There are better quantitative methods tradition. However, the report from this there. No one was more entitled to write today. In Bigelow’s survey, the sam­ work (Fish, 1925), except for a few about these copepods than Bigelow. A pling and the counting were standard­ new observations in Vineyard Sound, 1906 Harvard Ph.D., he was a protégé ized, so that at least relative abundanc­ was based on samples collected entire­ of Alexander Agassiz, and had been es could be considered. ly from the Woods Hole dock. During on Agassiz’s third Albatross cruise in From the 1920 Albatross cruise, there the Albatross years, the complementary 1905. Bigelow’s earliest publications were about 75 vertical and about 50 sur­ reports on copepods by Wheeler (1900) were on the systematics and distribu­ face plankton hauls. Data tables showed and Sharpe (1911) should also be men­ tion of planktonic coelenterates, which stations, dates, depths, numbers of each tioned here only to record that they did quickly gave him an international rep­ species, and numbers of each species not include Albatross material. utation. He contributed to this field per square meter sea-surface. for more than 30 years. To these stud­ Bigelow’s (1926) report, “being eco­ O. Tanaka ies, he soon added fishes, and ichthy­ logic and not systematic,” included The copepods from the Albatross ology became his major field between copepods as one of “the more impor­ continued to be worked and reworked. 1925 and 1955. Remarkably, Bigelow tant groups.” Indeed, the specific cope­ Otohiko Tanaka (1902–1990), in 1969, added a third scientific discipline, that pod section comprised more than one­ published two reviews of Albatross of oceanography and general plankton fourth of the total. Bigelow’s summary copepods (Tanaka, 1969; Tanaka and studies between 1914 and 1940. No on the food of copepods, and copepods Omori, 1969). Tanaka was a 1928 grad­ one could accuse him of claiming har­ as food for other organisms, is a classic uate of the Department of Fisheries of vests gathered by others; throughout in the field. Nearly 100 plankton cope­ the Imperial University of Tokyo. In his career, Bigelow was a preeminent pod species are listed, with those from 1931, he joined the university as an as­ field man. Associated with the Museum Albatross collections being the most di­ sistant, already with his life-long inter­ of Comparative Zoology, he became a verse. Detailed information is provided est in copepods. In 1934, Tanaka joined principal spokesman for the develop­ for more than 50 species. Figures are the staff of a private biological labora­ ment of oceanography in the United given for the distributions and abun­ tory on the coast of the Izu Peninsula, States, and he was the first director dances of many 1920 Albatross species. where he began a long series of excep­ (1931–40) of the Woods Hole Oceano­ There are good photographs of selected tional papers on the marine copepods of graphic Institution (Anonymous, 1955). plankton samples, including many co­ that region. This work was interrupted After that time, Bigelow was a Profes­ pepods from the Albatross collections. by World War II, when Tanaka served sor at Harvard until his death. Bigelow compared his information as an engineer officer in Manchuria. Bigelow incorporated the 1920 Alba­ with what other studies there had been After the war, Tanaka was appoint­ tross copepods into his definitive study in the North Atlantic, by Canadian and ed to the faculty of Kyushu University, of the plankton of the Gulf of Maine, European researchers. All in all, his where he renewed his study of the Izu begun as an “oceanographic and bio­ report filled a large and detailed piece copepods. He retired in 1965, whereup­ logic survey” in 1912 (Bigelow, 1926). of the emerging picture of the biology on he joined the Ocean Research Insti­ Most of the plankton for this study was of many important groups. For such a tute of the University of Tokyo. Tanaka collected from the Fish Hawk and the large area, this was probably the most (Fig. 11) was widely regarded for his Grampus (90 feet), a Fish Commission detailed study of its kind, stressing re­ nearly 50 papers on copepods, and kept schooner used from 1886 until 1918. petitive sampling at the same stations, up a lively correspondence and person­ Bigelow’s tour de force was a review and it remains extremely useful. al interest until his death (Nishida and of a tremendous quantity of data on the After her active use, the Albatross Omori, 1991). horizontal and vertical distribution and was tied to the Bureau of Fisheries dock In a review of the genus Euchirella seasonal abundance of phyto- and zoo­ at Woods Hole until June 1924, when (Tanaka and Omori, 1969), advantage plankton, as well as chemistry and hy­ it was sold to a Boston firm (Hedg­ was taken of the large number of spec­ drography of the region. The plankton peth, 1945). During this period, Paul S. imens available from Wilson’s Alba­ study complemented Bigelow’s survey Galtsoff was assigned to the now-dry­ tross collections. A new species was de­ of the fishes of the same region. land but funded position of “Albatross scribed from samples off the west coast The copepods were identified and Naturalist.” Charles Fish continued the of South America, and 13 other species enumerated by C. B. Wilson, and they work on the seasonal distributions of were redescribed. Seven of these had were so acknowledged in the full title as Woods Hole copepods under Galtsoff’s not been reported by Wilson (1950). well as the text. Copepods were the only direction (Galtsoff, 1962). Fish’s title In the same year, Tanaka (1969) de­ group counted routinely in the samples was “General Assistant, U.S.F.S. Al­ scribed or redescribed 13 other interest­ from vertical hauls (including some Al­ batross,” another now-dry-land assign­ ing copepod species that he had encoun­ batross collections). Bigelow discussed ment. If Fish had sampled from the tied­ tered in the review of Euchirella species

82 Marine Fisheries Review Figure 11.— Chauncey Thomas, Jr. (1850–1919), Com­ Otohiko Tanaka manding: conflict aboard the U.S. Fish Com­ mission Steamer Albatross in 1902. Mar. Fish. (1902–1990) Rev. 58(1–2):3–16. (photograph 1965, ______. 1996b. Charles Henry Gilbert Kyushu (1859–1928), Naturalist-in-Charge: the 1906 University). North Pacific Expedition of the Steamer Alba­ tross. Mar. Fish. Rev. 58(1–2):17–28. Esterly, C. O. 1905. The pelagic Copepoda of the San Diego region. Univ. Calif. Publ. Zool. 2(4):113–233. ______. 1924. The free-swimming Copepoda of San Francisco Bay. Univ. Calif. Publ. Zool. 26(5):81–129. Fish, C. J. 1925. Seasonal distribution of the plankton of the Woods Hole region. U.S. Bur. Fish., Bull. 41(975):91–179. Galtsoff, P. S. 1962. The story of the Bureau of Commercial Fisheries Biological Laboratory Woods Hole, Massachusetts. U.S. Dep. Inter., Fish. Wildl. Serv., Bur. Comm. Fish., Circ. 145, 121 p. Giesbrecht, W. 1895. Reports on the dredging operations off the west coast of Central Amer­ ica to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission Steamer “Alba­ tross,” during 1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding. XVI. Die pela­ gischen Copepoden. Bull. Mus. Comp. Zool. 25(12):243–263, 4 pl. Gould, A. A. 1841. Report on the Invertebrata of Massachusetts, Comprising the Mollusca, Crustacea, Annelida, and Radiata. Folsom, Wells, and Thurston, Camb., xiii, 373 p. Gunnerus, J. E. 1770. Nogle smaa rare og meestendeelen nye norske soedyr. Skrifter, sum udi det Kiobenhavnske Selskab af Laer­ doms og Videnskabers Elskere, 1765–1769, 10:166–176. Hedgpeth, J. W. 1945. The United States Fish Commission Steamer Albatross. Am. Neptune 5(1):5–26. ______. 1978a. The outer shores. Part 1. Ed Ricketts and John Steinbeck explore the collected by the Albatross. Again, seven Aldrich, J. W. 1980. The Biological Society of Pacific coast. Mad River Press, Eureka, Calif., Washington: a centennial history 1880–1980. xii, 128 p. of the species had not been reported Bull. Biolog. Soc. Washington 4:1–40. ______. 1978b. The outer shores. Part 2. by Wilson (1950). These included three Andrews, R. C. 1929. Ends of the earth. National Breaking through. Mad River Press, Eureka, new species, one of which, Wilsonidius Travel Club, N.Y., 355 p. Calif., x, 182 p. Anonymous. 1955. Foreword. Papers in marine Hobart, W. L. (Editor). 1995. Baird’s legacy: alaskaensis, represented a new deep­ biology and oceanography. Deep-Sea Res. the history and accomplishments of NOAA’s water genus named for Charles Branch Suppl. 3:xi–xx, portrait. National Marine Fisheries Service, 1871– Benjamin, M. 1918. Richard Rathbun. Science 1996. U.S. Dep. Commer., NOAA Tech. Wilson. 48(1236):231–235, portrait. Memo. NMFS-F/SPO-18, 46 p. With marine copepods, pelagic or Bigelow, H. B. 1926. Plankton of the offshore Nelson, S. B. 1971. Oceanographic ships fore and parasitic, whether you are studying new waters of the Gulf of Maine. With tables of aft. Gov. Print. Off., Wash., D.C., 240 p. copepods by C. B. Wilson and tables of dia­ Nishida, S., and M. Omori. 1991. Otohiko Tanaka or old genera or species, descriptions, toms by Albert Mann. U.S. Bur. Fish., Bull. (1902–1990). J. Crustacean Biol. 11(2):336– or distributions, sooner or later you will 40(2):1–509. 338, portrait. encounter the Albatross. About 1,600 Coe, W. R. 1918. Richard Rathbun and his con­ Raitt, H., and B. Moulton. 1967. Scripps Institu­ tributions to zoology. Am. J. Sci., Ser. 4, tion of Oceanography; first fifty years. Ward virtually untouched Albatross samples 46(276):757–763. Ritchie Press, Los Angeles, 217 p. remain on the shelves of the Smithso­ ______. 1929. Biographical memoir of Sidney Rathbun, H. W. 1969. Richard Rathbun Irving Smith 1843–1926. Natl. Acad. Sci. (1852–1918). 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