I iilij! i iSffif! |: i Mb w mmmmm I ;ii#iti !: ■i! i ill: l!!!i!l)i m DEPARTMENT OF TERRESTRIAL MAGNETISM J. A. Fleming, Director Scientific Results of Cruise VII of the Carnegie during 1928-1929 under Command of Captain J. P. Ault BIOLOGY- IV BIOLOGICAL RESULTS OF THE LAST CRUISE OF THE CARNEGIE I THE PHYTOPLANKTON II MARINE ALGAE III POLYCHAETOUS ANNELIDS IV THE MYSIDS • V THE ISOPODS VI THE HALOBATES Herbert W. Graham William Albert Setchell Aaron L. Tread well W. M. Tattersall James O. Maloney Harry G. Barber Alexander Wetmore VII LIST OF BIRDS VIII MISCELLANEOUS DETERMINATIONS THE SPONGE M. W. de Laubenfels THE ECHINODERMS Austin H. Clark THE INSECTS AND MITES E. A. Chapin and others Hoyt S. Hopkins THE PYROSOMIDS THE LIZARD Doris M. Cochran CARNEGIE INSTITUTION OF WASHINGTON PUBLICATION 555 WASHINGTON, D. C. 1943 This book first issued July 15, 1943 PREFACE Of the 110,000 nautical miles planned for the seventh cruise of the nonmagnetic ship Carnegie of the Carnegie Institution of Washington, nearly one-half had been com- pleted on her arrival at Apia, November 28, 1929. The extensive program of observation in terrestrial magnet- ism, terrestrial electricity, chemical oceanography, physical oceanography, marine biology, and marine me- teorology was being carried out in virtually every detail. Practical techniques and instrumental appliances for oceanographic work on a sailing vessel had been most successfully developed by Captain J. P. Ault, master and chief of the scientific personnel, and his colleagues. The high standards established under the energetic and re- sourceful leadership of Dr. Louis A. Bauer and his co- workers were maintained, and the achievements which had marked the previous work of the Carnegie extended. But this cruise was tragically the last of the seven great adventures represented by the world cruises of the vessel. Early in the afternoon of November 29, 1929, while she was in the harbor at Apia completing the storage of 2000 gallons of gasoline, there was an explosion as a result of which Captain Ault and cabin boy Anthony Kolar lost their lives, five officers and seamen were injured, and the vessel with all her equipment was destroyed. In 376 days at sea nearly 45,000 nautical miles had been covered (see map, p. iv). In addition to the exten- sive magnetic and atmospheric-electric observations, a great number of data and marine collections had been obtained in the field of chemistry, physics, and biology, including bottom samples and depth determinations. These observations were made at 162 stations, at an av- erage distance apart of 300 nautical miles. The distri- bution of these stations is shown in the map, which de- lineates also the course followed by the vessel from Washington, May 1, 1928, to Apia, November 28, 1929. At each station, salinities and temperatures were ob- tained at depths of 0, 5, 25, 50, 75, 100, 200, 300, 400, 500, 700, 1000, 1500, etc., meters, down to the bottom or to a maximum of 6000 meters, and complete physical and chemical determinations were made. Biological sam- ples to the number of 1014 were obtained both by net and by pump, usually at 0, 50, and 100 meters. Numerous physical and chemical data were obtained at the surface. Sonic depths were determined at 1500 points and bottom samples were obtained at 87 points. Since, in accord- ance with the established policy of the Department of Terrestrial Magnetism, all observational data and ma- terials were forwarded regularly to Washington from each port of call, the records of only one observation were lost with the ship, namely, a depth determination on the short leg between Pago Pago and Apia. The compilations of, and reports on, the scientific results obtained during this last cruise of the Carnegie are being published under the classifications Physical Oceanography, Chemical Oceanography, Meteorology, and Biology, in a series numbered, under each subject, I, n, and m, etc. A general account of the expedition has been prepared and published by J. Harland Paul, ship's surgeon and ob- server, under the title The last cruise of the Carnegie. and contains a brief chapter on the previous cruises of the Carnegie, a description of the vessel and her equip- ment, and a full narrative of the cruise (Baltimore, Wil- liams and Wilkins Company, 1932; xiii + 331 pages with 198 illustrations). Thepreparationsfor, and the realization of, the pro- gram would have been impossible without the generous cooperation, expert advice, and contributions of special equipment and books received on all sides from inter- ested organizations and investigators beth in America and in Europe. Among these, the Carnegie Institution of Washington is indebted to the following: the United States Navy Department, including particularly its Hydrographic Office and Naval Research Laboratory; the Signal Corps and the Air Corps of the War Department; the National Museum, the Bureau of Fisheries, the Weather Bureau, the Coast Guard, and the Coast and Geodetic Survey; the Scripps Institution of Oceanography of the University of California; the Museum of Comparative Zoology of Har- vard University; the School of Geography of Clark Uni- versity; the American Radio Relay League; the Geophys- ical Institute, Bergen, Norway; the Marine Biological Association of the United Kingdom, Plymouth, England; the German Atlantic Expedition of the Meteor , Institut fur Meereskunde, Berlin, Germany; the British Admiral- ty, London, England; the Carlsberg Laboratorim, Bu- reau International pour I'Exploration de la Mer, and Laboratoire Hydrographique, Copenhagen, Denmark; and many others. Dr. H. U. Sverdrup, now Director of the Scripps Institution of Oceanography of the University of California, at La Jolla, California, who was then a Re- search Associate of the Carnegie Institution of Washing- ton at the Geophysical Institute at Bergen, Norway, was consulting oceanographer and physicist. In summarizing an enterprise such as the magnetic, electric, and oceanographic surveys of the Carnegie and of her predecessor the Galilee, which covered a quar- ter of a century, and which required cooperative effort and unselfish interest on the part of many skilled scien- tists, it is impossible to allocate full and appropriate credit. Captain W. J. Peters laid the broad foundation of the work during the early cruises of both vessels, and Captain J. P. Ault, who had had the good fortune to serve under him, continued and developed that which Captain Peters had so well begun. The original plan of the work was envisioned by L. A. Bauer, the first Director of the Department of Terrestrial Magnetism, Carnegie Institu- tion of Washington; the development of suitable methods and apparatus was the result of the painstaking efforts of his co-workers at Washington. Truly, as was stated by Captain Ault in an address during the commemorative exercises held on board the Carnegi e in San Francisco, August 26, 1929, "The story of individual endeavor and enterprise, of invention and accomplishment, cannot be told." Dr. H. W. Graham, who succeeded H. R. Seiwell as chemist and biologist, had charge of the biological work on board the Carnegie from August 1929 until the loss of the vessel at Apia, Samoa. After his return to this coun- try. Dr. Graham was placed in charge of the biological collections, attending to their subsequent care, segrega- tion, and distribution to various specialists for examina- tion and report, he, himself, undertaking the reportingof the Peridinials. His memoir, "Studies in the morphol- ogy, taxonomy, and ecology of the Peridiniales," is Bi- ology III of this series. He also examined and prepared a report on the "Phytoplankton" which is the first of twelve biological reports grouped in the present volume. The macroscopic algae, gathered from the North Atlantic and South Pacific oceans, were examined by us a o 13 u <7 IS CO c 5 rt a o o Z c u •§ W o a- M 111 ^ 6 S3 K > > w W ^ I 100 cells <100 cells >100 cells <100 cells >100 cells <100 cells Northern Tropical Southern Northern Tropical Southern 50 100 50 100 50 100 50 100 50 100 50 100 Temperature, °C 14.23 (17) 11.92 (10) 10.45 (2) 23.24 (12) 20.55 (2) 13.90 (1) 16.90 (1) 23.00 (22) 17.87 (30 14.34 (31) 26.72 (42) 24.07 (46 22.20 (40) 20.31 (18) 17.45 (16 15.38 (14) 33.65 (17) 33.83 (10 33.71 (2) 35.00 (12) 34.74 (2) 34.99 (1) 34.05 (1) Salinity, o/oo 34.64 34.29 22) 30) 31) 42) 46) 34.24 35.03 35.33 35.42 (40) 34.91 (18) 34.73 (16 34.65 (14) Hydrogen-ion, pH values 8.14 (16) 8.30 (22) 8.11 (9) 8.15 (2) 8.15 (12) 8.04 (2 7.84 (1) 8.05 (1) Phosphate, mg/m^ 11 (22 31 (30 59 (30 22 (42 44 45 69 (15) 92 (8 84 (1) 39 (12) 30 (2) 159 (1) 46 (1) 56 (39 21 (18 27 (16 34 (14 Oxygen saturation, o/o 98 (5) 98 (15) 99 (4 103 (16) 100 (1) 96 (15) 83 (1) 95 (5) 15 (1) 91 4) 51 (6) 93 (2) ;!;;!!!!!!! "W'd) Silicate, mg/m^ 0.26 (2) 0.57 (13) 0.30 (1) 0.57 (15) 0.60 (12) 0.37 (3) 1.50 (1) 0.40 (2 0.88 (3) at from a study of the Carnegie data alone. They fur- ther indicate that the abundance of diatoms bears little relation to the silicate content of the water or to the percentage saturation with oxygen during the summer at least. Conditions at La Jolla (Allen, 1928) on the coast of southern California during the second quarter of the year when diatom production is the greatest, are simi- lar to conditions at Monterey in July, so that region will not be discussed. Lewis (1927) in studying surface catches of phyto- plankton off the coast of Oregon concludes that the colder water is more favorable for the production of diatoms. Water with temperatures below 11° C main- tained a greater production than water above that tem- perature. Table 4. Comparison of surface conditions at Carnegie northern stations with those at Monterey Bay Carnegie stations Means at Element (cells per liter) Monterey Bay for 100 100 July 1928 Temperature, °C 23.00 14.23 13.40 Salinity, o/oo 34.64 33.65 33.87 pH 8.30 8.14 8.14a PO4, mg/m3 11 69 48 Si02, gm/m'S 0.57 0.26 0.40 O2, per cent total 98 98 99 a Since Bigelow and Leslie did not include pH de- terminations in their analyses, this value is the ap- proximate mean of the second quarter of four years at La Jolla, California, taken from Moberg (14). VERTICAL CIRCULATION AND REPLENISHMENT OF NUTRIENTS The data presented here show that certain condi- tions usually accompany a growth of phytoplankton. In any study of the relation between the abundance of phytoplankton and environmental factors, the charac- teristics of a phytoplankton pulse must be kept in mind. Atkins (1928) and others have found that when favor- able conditions occur, such as an abundance of nutri- ent salts and sufficient illumination, a rapid growth of phytoplankton is initiated which continues until the nu- trients are exhausted. If one were to examine the plankton and the existing conditions of the environ- ment at different times during the progress of the pulse, different correlations would be found at different stages. Just before the rapid growth begins there is a sparse population in the presence of favorable conditions, such as high concentrations of nutrients. In other words. there is a negative relation between abundance of plank- ton and abundance of nutrients. After rapid growth has started there is a dense population occurring with com- paratively high concentrations of nutrients, or a positive relation between abundance of phytoplankton and of nu- trients. Near the close of the pulse, when the nutrients are practically depleted, there is a dense population in the presence of low concentrations of nutrients which presents a negative relation again. Still further, after the close of the pulse, the decay of the organic detritus may result in a regeneration of nutrients in the upper levels which creates another minor pulse, but before this pulse begins there occur together again high con- centrations of nutrients and a sparse population ofphyto- planktonic organisms, a negative relation again. It is obvious, therefore, that any correlations be- BIOLOGICAL RESULTS OF LAST CRUISE OF CARNEGIE tween the abundance of phytoplankton and the abundance of nutrients, which are drawn from observations made at any particular instant during a phytoplankton pulse, are meaningless unless the stage in the development of the pulse is considered, for, during a single pulse, every possible combination of the two factors occurs. Such a phytoplankton pulse as described, however, can run its course only when there is a single supply of nutrients which is not augmented during the pulse. Such a situa- tion is characteristic of cold-temperate regions. In these latitudes a supply of nutrients is brought to the up- per levels during the winter when the absence of strong thermal stratification permits considerable vertical cir- culation. In the early spring a phytoplankton pulse prob- ably is initiated by an increased amount of light. At the same time the temperature of the upper strata begins to rise and the body of water attains a more stable condi- tion so that nutrients are no longer supplied in great quantities from lower depths. It must also be remem- bered that the abundance of plankton in high latitudes is of a higher order of magnitude than that in lower lati- tudes so that the phytoplankton still existing after the close of a typical northern pulse or "bloom" may be large as compared with that ordinarily found in other regions. The vertical stability of the water probably has more to do with the abundance of plankton than any other factor. Whatever is the chemical substance limiting growth at any particular time, it probably occurs in abimdance in the deep water where organic remains are decomposing and any vertical circulation of the water that brings the deep water to the photosynthetic zone will increase the fertility of that zone. Various inves- tigators, viz.. Gran (1912 and 1928), Ruud (1932), Mar- shall and Orr (1927), and Atkins (1928) hive shown that the development of the phytoplankton is dependent on a replenishment of the supply of such nutrient salts as nitrates and phosphates in the photosynthetic zone, and that such a replenishment can usually take place only by some type of vertical circulation which carries quanti- ties of these salts from the rich stores in the deeper water. At Carnegie stations the richer plankton samples were obtained in regions where the surface water had characteristics of subsurface water, whereas the poor- er samples were obtained mostly in regions where a strong stratification of the water caused the conditions in the surface water to be considerably different from those in the deep water. Figure 2 shows the vertical distribution of temperature, salinity, density, phosphate, and pH in the upper 1000 m at station 137 in the North Pacific where there was little circulation of any nature. Conditions here were typical of regions in which the plankton population was extremely scanty. It is evident in fig\ire 2 that any vertical circulation which would re- new the phosphate at the surface would have to extend to more than 200 m, for there was practically no phosphate in the water to that depth. It is equally evident from the temperature and density curves that there was a very pronounced thermocline and stratification of the water above 700 m so that there would be considerable resist- ance to any such vertical mixing. We may now inspect conditions in two regions where the phytoplankton was relatively abundant, namely, south of the Aleutian Islands and off the coast of Peru. Condi- tions at the former region are represented by station 122 in figure 3 and at the latter by station 70 shown in figure 4. At station 122 there was an admixture of cold water of low salinity from the Bering Sea where vertical mixing is common. At station 70 there was an upwelling of water from a depth of about 500 to 700 m. At these two stations the difference in density between the sur- face water and the deep water is relatively not so great and the most rapid gradient is in the upper 100 m. Con- sequently, there is less resistance to vertical mixing and this fact is reflected in the higher concentrations of phosphate at the surface and in the correspondingly hieh concentration in the upper 500 m. At station 70, how- ever, this is owing partly to the upwelling of subsurface water. Any vertical circulation which extended to the 100-m level at these stations would carry great quanti- ties of phosphate to the surface layer for there were high concentrations at that depth. These observations were made in the summer when the most stable conditions oc- cur. In the winter there would be an even greater re- plenishment of the nutrients in the upper levels. Al- though phytoplankton pulses may occur in these regions, it is probable that there is always a greater production here than in such regions as represented by station 137. To return to the correlations discussed above, the richer phytoplankton samples were found in water of lower temperature, salinity, and pH, and of higher phos- phate content. A glance at figures 2, 3, and 4 will show that any vertical circulation that carries phosphate-rich water to the surface and thus increases the fertility of the surface water will also lower the temperature and pH of that water. Under such conditions the salinity of the surface water may be increased or decreased. For example, in figures 2 and 3 there is an increase in salin- ity with depth, but in figure 4 there is a decrease. It is also evident in figure 2 that a transport of water from lower levels should increase the silicate content of the surface water, and it is probable that a correlation be- tween silicate content and abundance of phytoplankton similar to that between phosphate content and abundance of phytoplankton would be found if the silicate data were as extensive as those for phosphate. It is well known that the cold waters of high lati- tudes support the greatest production of plankton. It is probable that low temperature, per se, is not particu- larly favorable for plankton growth except as it affects the particular species which are now adapted to low temperatures. The more abundant life of the colder seas is probably only indirectly related to low tempera- tures. Lower surface temperatures cause less stable conditions and thus permit a more rapid renewal of the fertility of the photosynthetic zone. DISSOLVED OXYGEN IN THE PHYTOSYNTHETIC ZONE A study of the dissolved oxygen in the photosynthet- ic zone reveals an interesting situation. Of the twenty stations (stations 130 to 149) in the northern region at which oxygen determinations were made, all but two showed supersaturation in the upper 100 m. Figure 5 shows the vertical distribution curve of percentage sat- uration of oxygen at station 145 which is characteristic of the region. Supersaturation occurred here between THE PHYTOPLANKTON 9 20 and 80 m. Now the question arises as to the source of this oxygen. It cannot have been brought up from low- er levels because the adjoining water was deficient in oxygen. It is very unlikely that a thin horizontal layer could move in from some region rich in oxygen along this high level where vertical mixing so often occurs. Furthermore, the temperature and salinity data do not permit such a conclusion. The only other possible source of oxygen is the photosynthetic organisms inhab- iting the upper strata of water. Of the twenty oxygen stations, only six produced samples with more than 100 diatom and dinoflagellate cells per liter, and at only one of these were samples collected giving over 1000 cells per liter. Zero counts were obtained at a subsurface level at four of these stations. It does not seem possi- ble that such a low population of diatoms and dinoflagel- lates could produce such an excess of oxygen. There are two possible explanations for this condition. Either there is present in the upper 100 m a large population of photosynthetic organisms which pass through the ordi- nary filter net, or else it is possible for a layer of water to exist in the ocean at this level and maintain its super - saturation with respect to oxygen after the phytoplankton, which has produced the oxygen, has disappeared. The latter explanation is hardly tenable except for regions where the water strata are in a very stable condition. Gran (1912) found in the Atlantic that the Coccolitho- phoridae, which readily passed through the finest net, far outnumbered the diatoms and dinoflagellates which were retained in the net. It was doubted, however, if the photosynthetic activity of such small organisms, even in such large numbers, could surpass that of the larger forms occurring in smaller numbers. The Car- negfie data lead one to suspect very strongly that in some instances the photosynthetic activity in the sea is con- fined principally to such minute organisms. Much more work must be done in the Pacific before such problems as this can be satisfactorily solved. At the fifteen oxygen stations in the tropical region only one observation showed complete saturation. This is in strong contrast with the northern group of stations among which were only two that did not show saturation at some level. Moberg and Graham (1930) have pointed out that this "can be accounted for only by assuming that at these stations at least part of the water had re- cently come from depths at which oxygen content is normally low." Unfortunately the quantitative plankton samples for most of these stations were lost, but the samples from six of them and from other tropical sta- tions indicate that the plankton flora in this region is much scantier than in the north, which condition also contributes to the low oxygen content of the upper strata. SUMMARY 1. There was a very sparse population of diatoms and dinoflagellates in the open waters of the Pacific in 1928 and 1929 at 0-, 50-, and 100-m levels. 2. Although dinoflagellates never reached the large numbers attained by the diatoms at some stations, they outnumbered the diatoms in 25 per cent of the samples. 3. When a rich sample is defined as one with more than 100 combined diatom and dinoflagellate cells per liter, the richer samples occurred in waters of lower temperature, lower salinity, higher phosphates, and higher hydrogen-ion concentration. No definite corre- lations were found between either dissolved oxygen or silicate and number of plant cells. The richer areas occurred in regions where subsurface water is brought to the surface by vertical circulation. 4. A comparison of the Carnegie data with results obtained by Bigelow and Leslie in Monterey Bay shows the same correlation between the above factors and quantity of phytoplankton. 5. Supersaturation of oxygen in the upper 100 m in regions where the diatom and dinoflagellate populations were very low suggested the presence of photosynthetic organisms not captured in the ordinary filter net. 6. The Carnegie investigations have shown the ne- cessity of making quantitative studies of the phytoplank- ton that pass through the ordinary filter net and of se- curing more complete vertical series over the whole ocean. LITERATURE CITED Allen, W. E. 1928. Review of five years of studies on phytoplankton at southern California piers, 1920- 1924 inclusive. Bull. Scripps Inst., Tech. Ser., vol. 1, no. 16. and R. Lewis. 1927. Surface-catches of ma- rine diatoms and dinoflagellates from Pacific high seas in 1925 and 1926. Bull. Scripps Inst., Tech. Ser., vol. 1, no. 12, pp. 197-200. Atkins, W. R. G. 1923. The silica-content of some nat- ural waters and of culture media. Jour. Marine Biol. Assoc, vol. 13, pp. 151-159. 1925. Seasonal changes in the phosphate- content of sea-water in relation to the growth of the algal plankton during 1923 and 1924. Jour. Marine Biol. Assoc, vol. 13, pp. 700-720. 1928. Seasonal variation in the phosphate and silicate content of sea water during 1926 and 1927 in relation to the phytoplankton crop. Jour. Marine Biol. Associ, vol. 15, pp. 191-205. Bigelow, H. B., and M. Leslie. 1930. Reconnaissance of the waters and plankton of Monterey Bay, July, 1928. Bull. Mus. Comp. Zool., vol. 70, no. 5. Gran, H. H. 1912. Pelagic plant-life in "The depths of the ocean" by Murray and Hjort, pp. 307-386. London. 1928. The conditions of life for plankton in the coastal waters of northern Europe. Rapp. Pro- ces-Verbaux. vol. 47, Conseil. Perm. Internat. Expl. Mer. pp. 196-204. Jacobsen, J. P. 1921. Manuel pratique de I'analyse de I'eau de mer --II. Dosage de I'oxyg^ne dans I'eau de mer par la methode de Winkler. Bull. Inst. Oceanogr. Monaco, no. 390, 16 pp. ______ 1925. Die Loslichkeit vonSauerstoff im Meer- wasser. Medd. Komm. Havundersog., Serie Hydro- grafi, vol. 1, no. 8. King, E. J., and C. C. Lucas. 1928. The use of picric acid as an artificial standard in the colorimetric estimation of silica. Jour. Amer. Chem. Soc, vol. 50, pp. 2395-2398. Lewis, R. 1927. Surface-catches of marine diatoms and dinoflagellates off the coast of Oregon by U.S.S. Guide in 1924. Bull. Scripps Inst., Tech. Ser., vol. 1, no. 11. Lohmann, H. 1912. Untersuchungen ueber das Pflanzen- und Tierleben der Hochsee im Atlantischen Ozean wahrend der Ausreise der Deutschland. SitzBer. Gesellsch. Naturfors. Preunde, Berlin, no. 2a, pp. 23-54. Marshall, S. M., and A. P. Orr. 1927. The relation of the plankton to some chemical and physical factors in the Clyde Sea area. Jour. Marine Biol. Assoc, vol. 14, pp. 837-868. , 1928. The photosynthesis of diatom cultures in the sea. Jour. Marine Biol. Assoc. United Kingdom, vol. 15, pp. 321-360. Moberg, E. G. 1926. The hydrogen-ion concentration of sea-water off the coast of southern California. Proc Third Pan-Pacific Sci. Cong., Tokyo, pp. 221-229. and H. W. Graham. 1930. The distribution of oxygen in the Pacific as an index of the circulation of the water. Rept. and Comm., Stockholm Assem- bly, Internat. Geod. Geophys. Union, Sect. Oceanogr., pp. 95-97. Pettersson, V. I. 1928. Apparatus for quantitative measurements of plankton in situ. Jour. Conseil. Internat. Expl. Mer., vol. 3, no. 13. Poole, H. H., and W. R. G. Atkins. 1926. On the pene- tration of light into sea water. Jour. Marine Biol. Assoc, vol. 14, pp. 177-198. Ruud, J. T. 1932. On the biology of southern Euphausi- idae. HvalrSdets Skrifter. Sci. Res. Marine Biol. Res. no. 2. Det Norske Videnskaps-akademi, pp. 1- 105. Wenner, F, E. H. Smith, and F. M. Soule. 1930. Appa- ratus for the determination aboard ship of the salin- ity of sea-water by the electrical conductivity meth- od. Bur. Stand. Jour. Res., vol. 5, pp. 711-732. Whipple, G. C. 1927. The microscopy of drinking wa- ter, pp. 95-98. 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Meyen (better known as Sargassum baccllerum [Turn.] C. Ag.) and S. fluitans Boerg. seem to become members of true self- perpetuating pleuston formations. Of such collections, some twenty-six bottles of various sizes were placed in the hands of the author for examination and report, all of which except one (bottle 1798) contained at least some minute fragment of the larger marine algae. The col- lections were gathered from the North Atlantic and the South Pacific oceans. The three samples (nos. 309, 317, and 318) from the neighborhood of Easter Island in the Pacific Ocean and bottle 8169, from Apia, Samoa repre- sent localities south of the equator. Sargassum fluitans Boerg. Station Latitude Longitude Sample Bottle Depth Date Quantity o 1 o / 1928 1 38 14 N 67 34 W 2 1976 70 m May 12 1 scrap 1- 2 37 45 N 53 26 W 6 3020 Surface May 16 1 vesicle 1- 2 37 53 N 52 46 W 9 3009 Surface May 16 1 scrap 1- 2 37 53 N 52 46 W 11 3868 Surface May 16 2 or 3 vesicles 34 11 18 N 78 34 W 200 4745 Surface Oct. 9 Abundant 34 11 18 N 78 34 W 202 4773 50 m Oct. 9 1 scrap c sargassum [latans (L.) Meyen Station Latitude Longitude Sample Bottle Depth Date Quantity ° / o / 1928 1 38 14 N 67 34 W 1 3805 Surface May 12 2 vesicles 1- 2 37 53 N 52 46 W 9 3009 Surface May 16 Abundant 2 39 06 N 45 41 W 15 3900 Surface May 18 1 vesicle 2- 3 40 00N* 44 00 W* 16 3023 Surface May 18 Scraps 13-14 42 ION 47 19 W 90 4232 Surface Aug. 9 Abundant 14 42 ION 47 19 W 96 4257 Surface Aug. 9 Scraps 15 38 48 N 48 48 W 100 4258 Surface Aug. 11 1 scrap 16 36 47 N 46 31 W 106 4298 Surface Aug. 13 Abundant 16 36 47 N 46 31 W 107 4300 50 m Aug. 13 1 scrap 17 33 42 N 42 41 W 112 4338 50 m Aug. 15 1 vesicle 18 29 47 N 40 36 W 117 3724 Surface Aug. 17 1 scrap 19 24 00 N 39 36 W 122 4383 Surface Aug. 20 1 scrap * Approximately ATLANTIC COLLECTIONS The majority of the samples show, as might be ex- pected, fragments of the two most common Sargasso weeds of the floating types, Sargassum natans (L.) Meyen and S. fluitans Boerg., both of which are usually desig- nated as Sargassum bacclferum (Turn.) C. Ag. Although scraps (at times a single vesicle) are not always cer- - tainly to be identified as to the one or the other species, most of these samples seem fairly clear as to distinc- tion. In the neighborhood of station 6, latitude 50° 22' north and longitude 13° 31' west, off the coast of Ireland, the dip net (sample 38, bottle 4011) brought up floating Ascophyllum nodosum (L.) Le Jolis, or "bottle wrack." North from this, on the way to station 7, off the Faroes, the 1/2 meter net caught more of this species (sample 51, bottle 3040, June 6, 1928) on which there was abun- dant epiphytic Scytosiphon lomentarius (Lyng.) J. Ag. There were also 2 or 3 large fragments of Fucus vesl- cnlosos L. and on this was considerable epiphytic Pylaiella litoralis (L.) Kjellm. Sample 51 also showed as epiphytes, excellently developed specimens of Entero- morpha crinita (Roth) J. Ag., a widely spread species closely related to, if not identical with, E. plumosa Kuetz. On June 4 (sample 49, bottle 4074) the 1/2 meter net caught from the surface a single bladder of Asco- phyllum. The Atlantic collections present nothing except what was to be expected. The data are given for reference as to floating conditions and possible depth relations. 17 18 BIOLOGICAL RESULTS OF LAST CRUISE OF CARNEGIE PACIFIC COLLECTIONS A collection was made in April 1929 on the reef at Apia, on Upolu of the Samoan group (bottle 8169). Sev- eral entire specimens of the bright green pom pom-like Chlorodesmls comosa Harv. et Bail, were found in the bottle: two fairly large specimens of the light green, areolate Dlctyosphaeria australis Setchell; and two spec- imens of a light red, flat, and laciniately much branched alga with the surfaces covered with low, blunt papillae, which seems to be referable to Meristotheca papulosa (Mont.) J. Ag., but with more the habit of Halymenia ceylonica Harv. Unfortunately it is sterile. Between stations 115 and 116, latitude 38° 06' north and longitude 146° 53' east (sample 699), the 1/2 meter net caught a couple of small fragments of what seems to be the Japanese brown alga, Cystophyllum hakodatense Yendo, a very probable find. Around Easter Island, south of the equator, three samples containing Sargassum prove to be the most in- teresting and novel catches of the whole plankton collec- tions, so far as the macroscopic marine algae are con- cerned. The samples (309, 317, and 318) represent dif- ferent phases of attached and floating forms, and seem also to have a definite bearing on the "Sargasso Sea" type of situation in the Indo-Pacific Ocean area. The specimens, which are excellent as such material goes, are closely related to a species group to which may be referred Sargassum stenophyllum J. Ag. (non Martius), S. lanceolatum J. Ag. (non Greville), and S. Skottsbergil Sjostedt. Sample 309 of the Carnegie collections was brought up from a depth of 15.4 fathoms by the snapper. It shows several plants. Two of these are species of Zonarla, probably to be referred to Z. variegata (Lam'x) Mert. and to Z. crenata J. Ag.; the former a prostrate speci- men, the latter represented by two upright specimens. Unfortunately, all the Zonarla plants are both young and sterile. There are, in addition, several specimens of Sargassum, two of which lack only the very holdfasts themselves of being complete. They show short (1 to 2 cm in these specimens), relatively stout, cylindrical, erect primary axes with sparse lateral tubercles, and with the primary branches grouped about the apex. The primary branches are compressed (even to flat below), and the leaves and secondary branches are arranged dis- tichously. All branches show marginal cryptostomata. The leaves are linear -lanceolate, sharply cuneate at the base, and attenuate above to a sharper or a somewhat obtuse apex. The margins are all sharply and rather coarsely dentate, with teeth one- to two-fifths the diam- eter of the leaf, projecting outward and sharply forward, and broadly or narrowly triangular in outline. The costa is stout and conspicuous below, broadening toward the upper part to the point of vanishing somewhat below the apex. The cryptostomata are smaller, less prominent, and scattered, in the lower leaves, becoming larger and more conspicuous above, arranged uniseriately on each side of the costa, frequently in pairs along it. The leaves are short-ly petiolate. The secondary branches are frequent (at intervals of approximately I cm) and short (2 to 4 cm long) as compared with the primary branch (up to 30 cm long), whence they arise. Only one of the plants shows receptacles. These are cymosely branched, about 1 cm in height. The individual branch- lets are arranged dichotomously below, but subdichoto- mously, almost racemosely, above, are compressed cy- lindrical, tapering above, and alternately torulose as the individual conceptacles mature and become protu- berant. All the conceptacles show antheridia only, thus suggesting a dioecious species. Sample 317 is made up of about sixteen fragments, evidently of the same species as those of sample 309. It was collected about two miles northeast of Easter Island, in latitude 27° 25' south and longitude 109° 25' west, from the surface. Axes and leaves of these tips of plants agree with those of sample 309. Evidently they had not been long detached, since in color, consist- ency, etc., they seem not at all modified. The recepta- cles, however, are varied. Both antheridial and oogoni- al receptacles are present, but on different fragments, the majority being antheridial. The antheridial recep- tacles of these plants are similar in shape and branch- ing to those of sample 309, except that some of them are more elongated (up to 1.5 and 1.75 cm) and very slender. The oogonial receptacles, though still of the Malaco- carpic, or unarmed (i.e. without teeth or spines) type, are very different from the long linear antheridial type. They are shorter (about 5 mm high) and less divided (only once or twice dichotomously cleft or lobed), the divisions being narrowly ellipsoidal fusiform, with the conceptacular openings broader and more gaping. In the Carnegie specimens, only the apical conceptacles of each receptacle show oogonia, but the lower concepta- cles show swollen structures such as are found in an- theridial conceptacles. Sample 318 was taken from floating masses (sur- face) about 8.5 miles southeast of Easter Island. The masses show the peculiar yellow of the Sargassa of the Sargasso Sea and might readily be taken for similar masses of Sargassum natans (L.) Meyen. The short in- dividual fragments (up to 10 cm long) seem likely to have arisen in the floating state. The stems, leaves, bladders, and receptacles, however, are practically identical in the principal details of their structure with those of samples 309 and 317. The costae (or midribs) of the leaves, however, seem stouter, the cryptostomata fewer and less conspicuous (at least in some of the frag- ments) and there are encrusting bryozoans, etc., on them similar to those occurring on the floating frag- ments of S. natans in the Sargasso Sea of the Atlantic Ocean. There does not seem to exist for the Pacific or In- dian oceans any real approximation to the Sargasso Sea of the North Atlantic. In the extreme south Pacific a pleuston, often very conspicuous and of large masses, is frequently met with, especially in the vicinity of Cape Horn and the west Patagonian coast. This has for its chief constituent, plants of Macrocystis. In the extreme northern Pacific Ocean, vast floating masses of Nereo- cystis or of Alaria fistulosa may be seen, particularly near Unimak Pass, in the Aleutian Islands. These masses, often of several acres in extent, are made up of stems and the huge terminal bladders of the Nereo- cystis or of the long leaves of the Alaria with their in- terruptedly swollen and inflated midribs. Both the Macrocystis formations (which may be true pleustoi^ and the floating masses of Nereocystis or of Alaria (not true pleustons) are characteristic of cold waters, but waifs from them seemingly stray into the edges of trop- MARINE ALGAE 19 ical waters. This is indicated by the attribution of Macrocystis and Lessonla, and even of Durvlllaea, spe- cies of the west South American coasts to such tropical localities as the coasts of Tahiti. Nevertheless, patches of floating seaweeds may be seen at times near the various islands or archipelagoes within the tropics (or warmer waters) and forms ap- proaching very closely to S. natans (L.) Meyen (S. bac- ciferum [Turn.] C. Ag.) are to be found in herbaria, des- ignated as from Indo-Pacific areas. The exact locali- ties are not always certain, but it may be presumed that at times the Atlantic species may pass around the Cape of Good Hope. Indigenous Indo-Pacific species of Sar- gassam may be suspected as possible of adopting, as well as adapting themselves to, a pleuston (or continued floating and self -propagating existence), but there is lit- tle certainty in this direction. Sample 318, taken in con- nection with samples 309 and 317, is a plausible sugges- tion along this line and may have a bearing on the ques- tion as to a limited type of Sargasso Sea in the south- eastern tropical Pacific. The Sargass«un of samples 309, 317, and 318 raise morphological, taxonomic, and distributional questions. Sample 309 is composed of attached specimens, living at a depth of 15.4 fathoms. As already stated, this species agrees most nearly with the description of Sargassum lanceolatum J. ^g. J. G. Agardh, however, does not indi- cate of what sex (or sexes?) his type specimens were, but Grunow states that the receptacles in this species (as he understands it) are androgynous. It seems that some dioecious species may have androgynous states (or possibly seasonal conditions). It has not been possi- ble to examine the type plants (from the west coast of Australia, preserved in Herb. Mus. Paris), so the deter- mination as to S. lanceolatum is not fully substantiated. The only species hitherto known with certainty from Easter Island is Sargassum Skottsbergii Sjostedt (in Skottsberg, The Natural History of Juan Fernandez and Easter Island, vol. II, part III, p. 311, 1924). The plants of this species are very close to those of sample 309, except that the leaves of the antheridial plant are entire or only sparingly and minutely denticulate, with few (at times no) and inconspicuous cryptostomata, whereas those of the oOgonial plant are more conspicuously and acutely denticulate, with the cryptostomata more numer- ous and conspicuous. The receptacles vary somewhat in both Sjostedt's plants, and the Carnegie samples 317 and 318 are fully as much swollen as those figured by SjO- stedt (loc. cit., fig. 3), and thoroughly merit his descrip- tion of "breviora, latiora, conica." Plants appearing to be identical with those of S. Skottsbergii occur common- ly on the shores of Tongatabu in the Tonga Archipelago. Both, however, seem to correspond in detail to the de- scription of S. stenophyllum J. Ag., from the western shores of Australia (type specimen in Herb. Mus. Paris, not seen). Grunow says that the receptacles of the type specimen of S. stenophyUum are androgynous, whereas all the Tongatabu and Easter Island specimens examined seem varyingly androgynous or polygamous (i.e. recep- tacles with exclusively male and female conceptacles, receptacles with^largely male or female conceptacles, and finally receptacles with conceptacles in which male and female are mixed). Therefore the three species represented by the binomials S. stenophyllum J. Ag., S. lanceolatum J. Ag., and S. Skottsbergii Sjost., seem very near to one another, sufficiently so to form a narrow spe- cies group, or even possibly a species cycle. Assuming that S. stenophyllum J. Ag. (1848) may be the same as S. Skottsbergii SjOstedt, the latter name has the preference since S. stenophyllum Martins (now S. cymosum C. Ag.) dates from much earlier (at least as early as 1827) than S. stenophyllum J. Ag. (1848). If all are considered to be only members of one form cycle about S. lanceolatum J. Ag. (1848), this name will apply, the S. lanceolatum Greville (1849) being different (now S. coriiiollum J. Ag.) from S. lanceolatum J. Ag. (1848), and of sufficiently probable later date of publication as to be rejected (not being an earlier homonym). A number of photographs are reproduced in connec- tion with the collection of samples 309, 317, and 318, in order to make more clear and more certain than may be accomplished by words, the variation of the receptacles in size, shape, and the coordination with sex segregation. Figure 1 shows two plants of diverse age and devel- opment of sample 309, from fourteen fathoms depth off Easter Island. The younger plant (above) shows the short main axis (characteristic of this species group) giving off several lateral axes. The older plant is only the upper part of a lateral axis, with branches, leaves, and well-developed receptacles which are antheridial. Figure 2 shows the receptacular branchlet enlarged 6 diameters. The coarse, compact, cymose receptacle is somewhat flattened and sufficiently condensed to be den- tate on the margins (because of suppressed branchlets). Because of this, the Agardhian arrangement would re- move this plant far from those of the other numbers (in- to the "Series" Acanthicarpicae, "Tribe" Biserrulae of J. G. Agardh). This seems unsuitable from other re- semblances of habit and structure between this and the other numbers of the Carnegie collections and also from plants collected in the Tonga Islands. Figure 3 shows a sheet of fragments of collection sample 317, collected floating about two miles northeast of Easter Island, but it seemed to be fairly recently torn away. The two lower fragments have elongated, slender antheridial receptacles (see figure 4,o*), whereas those of the upper row have shorter, more robust receptacles (see figure 4,?), antheridial below, oogonial toward the apex. Figure 5 shows receptacles from the fragments shown 6 diameters. The details of the long, slender an- theridial receptacles and the short, robust receptacles with antheridial conceptacles below and oogonial recep- tacles above are shown. The receptacles of sample 317 are from fragments, are neither definitely flattened nor toothed, and consequently are not ancipate. They differ in these respects from those of sample 309. Sample 309 would be placed under "Series" Malacocarpicae, "Tribe" Cymosae of J. G. Agardh. Figure 6 shows receptacular branchlets from two fragments of still other plants of sample 317, both oOgo- nial toward the apex, but antheridial below. The dispro- portion between the length -breadth indices of the two sets of receptacles is marked. Figure 7 shows four samples from the mass of float- ing fragments of the light yellowish color characteristic of the floating fragments from the Atlantic Sargasso Sea, under sample 318. The mass of sample 318 was floating about eight miles southeast of Easter Island. Figure 8 shows characteristic sterile fragments from this collection, whereas figures 9 and 10, enlarged 2 diameters, show characteristic fruiting fragments, Iwth with androgynous receptacles. In figure 9 these are shorter and more robust (as seen enlarged 6 diameters in figure 11). In figure 10 they are longer and more 20 BIOLOGICAL RESULTS OF LAST CRUISE OF CARNEGIE slender than some (enlarged 6 diameters in figure 12). The dark oogonia may readily be seen in the upper parts, and even fairly evenly distributed in the receptacles, of figure 11, whereas none are seen in figure 12. There is, however, one oogonium in an antheridial conceptacle of the receptacles represented on figure 12. In the type collection of Sargassum Skottsbergii Sjost. from Easter Island, described as dioecious, the author has seen an occasional oogonium in an antheridial receptacle and an occasional antheridial conceptacle in receptacles other- wise exclusively oOgonial. It seems, therefore, that in this naturally smaller assemblage of species passing under the names of S. stenophylliun J. Ag., S. lanceolatum J. Ag., and S. Skotts- bergii Sjost., the sexes may vary from partially even to completely segregated, but the receptacles are more properly polygamous, some plants having receptacles exclusively (or almost exclusively) of one sex or the other whereas other plants may have the sexes in sepa- rate conceptacles but both kinds of conceptacles in the same receptacle. According to the type of dominance the receptacles vary, elongated and slender for recep- tacles purely (or approximately pure) antheridial but shorter and more robust according to the admixture of, or purity of, oogonial conceptacles. In plants of this group from the Tonga Islands, seemingly otherwise of the same species cycle, there has been seen, chiefly, specimens in which oogonia and antheridia are found in the same receptacle, at least in the middle parts of the moderately long and moderately robust receptacles. To the same smaller group as the one being dis- cussed belongs also another cycle of species, viz., S. Grevlllei J. Ag., S. oligocystum Mont., and S.oligocy- stoides Grun. These species have broader leaves than those of the Easter Island group. Their receptacles are polygamous as are those of the Easter Island group, and with similar variation in length-breadth index, compres- sion, and absence or presence of denticulate or spinose margins. 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