CHAPTER VIII SPONGES, COELENTERATES, AND CTENOPHORES Blank page retained for pagination THE PORIFERA OF THE GULF OF MEXICO 1

By J. Q. TIERNEY. Marine Laboratory, University of Miami

Sponges are one of the dominant sessile inverte­ groups. The. floor of the Gulf between the bars brate groups in the Gulf of Mexico: they extend is sparsely populated. The majority of the ani­ from the intertidal zone down to the deepest mals and plants are concentrated on the rocky Parts of the basin, and almost all of the firm or ledges and outcroppings. rocky sections of the bottom provide attachment The most abundant sponges on these reefs are for them. of several genera representing most of the orders Members of the class Hyalospongea. (Hexacti­ of the class Demospongea. Several species of nellidea) are, almost without exception, limited to Ircinia are quite common as are Verongia, Sphecio­ the deeper waters of the Gulf beyond the 100­ spongia, and several Axinellid and Ancorinid fathom curve. These sponges possess siliceous sponges; Cliona is very abundant, boring into spicules in which (typically) six rays radiate from molluscan shells, coral, and the rock itself. The II. central point; frequently, the spicules are fused sponge population is rich both in variety and in ~gether forming a basket-like skeleton. Spongin number of individuals; for this reason no attempt 18 never present in this group. is made to discuss it in taxonomic detail in this In contrast to the Hyalospongea, representa­ r~sum~. ti\Tes of the clasa Calcispongea are seldom, if Some of the sponges of the Gulf are of world­ ~\Ter, found in deep water. These sponges, unique wide distribution, i. e., Dysidea jragilis, while 1ll having spicules of calcareous material, are usu­ others are typically West Indian, and a few are a.l1y restricted to shallow water. They are not probably restricted to the Gulf. The West Indian conspicuous; typically, they are encrusting forms sponge fauna may be a single regional population Or tubiform in shape but only a few centimeters in with only minor locational differences. Water height. currents of the Caribbean, the Gulf, and the The major sponge group in the Gulf of Mexico, southern portion of the western Atlantic off Florida both from number of genera and from the range and the Bahama Islands have been shown to of distribution, is the class Demospongea. Pos­ to carry a sponge disease from one point to many sessing more or less spongin, and when spicules others in this region; it is therefore permissible to are present having unfused spicules of siliceous suggest that the same currents would be equally lllaterial, these sponges occur throughout the Gulf effective in distributing sponge larvae. eJctending from the shallow coastal waters down The commercial sponges of Florida, the Ba.­ to the deepest off-shore sections of the basin. hamas,,Cuba. and British Honduras are all quite The sponge bars or reefs of tho eastern Gulf are similar: the sheepswool sponge, Hippwspongia quite typical of the habitat and ecology of the lachne, is the most valuable sponge now available SPonges in shallow to moderate depths. These for the market. Reef, glove, yellow, and grass sO-called reefs are sections of rocky outcroppings sponges, all members of the genus Spongia, are of that are elevated a few inches to a few feet above less value and are therefore less eagerly sought by the general bottom profile. They are more or less the sponge fishermen. densely covered with commercial and non-com­ Two methods, in general, are used in the W ~st lllercial sponges, coral (usually Oculina) , and oth~r Indian region for the collection of commerCIal .t\.lcyonaria. The bottom-dwelling marine sponges. Hooking is practiced in waters of less groups (such as mollusks, annelids, Crustacea, than 7 or 8 fathoms throughout the entire zone, ltscidians) are associated with the dominant ""'------and light-weight, full diving rigs are used in the I COntribution No. 105, Marine Laboratory, UniversIty or ~flaml. Gulf of Mexico in depths of less than 20 fathoms. 21195340-114-18 259 260 FISHERY BULLETIN OF THE FrBH AND WILDLIFE SERVICE In addition to these two major methods a few DE LAUBENFELS, M. W.-Continued native islander fishermen skiI1:'dive for sponges in 1934. New sponges from the Puerto Rican Deep. Smithsonian Misc. ColI. 91 (17): 1-28. shallow water, and a few Florida sponge fishermen 1936a. A discussion of the sponge fauna of the Dry have begun in the past few years to drag tines Tortugas in particular and the West Indies in gen­ and other types of dredges in order to obtain more eral with material for a revision of the families and sponges with less work. orders of the Porifera. Carnegie Inst. Washington As a result of the constantly decreasing supply Pub. 467, Papers Tortugas Lab. 3: 1-225, pIs. 1-22, text figs. of natural sponges there have been repeated 1936b. A comparison of the shallow water sponges ne,", attempts in the past half century to establish the Pacific end of the Panama Canal with those at sponge farms in the shallow coastal waters of the Caribbean end. Proc. U. S. Nat. Mus. 83 (2993): Florida and the other Caribbean sponge producing 441-466, 6 figs. areas. Practical as they now a.re from a biological 1939. Sponges collected on the presidential cruise of. 1938. Smithsonian Misc. CoIl. 98 (15): 1-7, 1 fig. point of view, these farms have always failed 1948. The order KeratOBa of the phylum Porifera, a because of economic difficulties. It is now entirely monographic study. Allan Hancock Found. Oco.· possible, however, to produce artificially propa­ Paper 3, pp. 1-217, pIs. 1-30, text figs. 1-31. gated sponges as a sound commercial venture. 1949. Sponges of the western Bahamas. Am. Mus. In addition to the biological studies on the dis­ Novitates 1431, pp. 1-25, illus. tribution and growth of the Gulf of Mexico DUCHASSAJNG, DE FONBRESSIN P., and MICHELOTrJ, G. 1864. Spongiaires de la Mer Caraibe. Memoire publi6 porifera, there have been biochemical studies car­ par Ill. SocieM hollandaise des Sciences A Haarlem. ried out on the lipids, the carbohydrates, and the Natuurk. Verh. Mij. Haarlem, 21: 1-124, pis. 1-25. general chemical constitution of several of the more GALTSOFF, P. S. common marine sponges of this region. 1940. Wasting disease causing mortality of sponges in the West Indies and Gulf of Mexico. Proc. Eighth BIBLIOGRAPHY Scientific Congress 3: 411-421, 2 pis., 2 text figs. 1946. Sponges. U. S. Fish and Wildlife Service, Fishery AGASSIZ, ALEXANDER . Leaflet 4, pp. 1-7. 1888. Characteristic deep sea types. Sponges. (Prepared HIGGIN, T. from the memoirs of O. Schmidt on Atlantic and 1875. On a new sponge of the genus LujJaria, from' Caribbean sponges.) Bull. Mus. Compo Zool., 15: Yucatl1n, in the Liverpool Free Museum. With re­ 17G-179,25 figs. marks by H. J. Carter. Ann. and Mag. Nat. Rist. BARTSCH, P. Ser. 4, 16: 223-227, pI. 6. 1936. Sponge cultivation. Science, 83 (2164): 507. HYATr, A. BERGMANN, WERNER 1875. Revision of the North AmericanPoriferae; with 1949. Comparative biochemical studies on the lipids of remarks upon foreign species. Part I. Mem. Boston marine invertebrates, with special reference to the Soc. Nat. Hist. 2: 399-408, pl. 13. sterols. Sears Foundation Jour. Mar. Res., 8 (2): 1877. Revision of the North American Poriferae; witb 137-176, figs. 1-2. remarks upon foreign species.. Part n. Mem. Boston BROWN, H. H.; GALTSOFF, P. S.; SMITH, L. C.; and SMITH, Soc. Nat. Rist., 2: 481-554, pis. 15-17. F.G. W. KENT, W. S. 1939. Sponge mortality in the Bahamas. Nature, 143 • 1883. Report on the sponges of the Bahama Islands. (3628): 807-808. Aubert's Steam Printing Works. Pp. 1-18. CARTER, H. J. 1884. Catalogue of marine sponges, collected by Mr. MOORE, R. F. JOB. Willcox, on the west coast of Florida. Proc. 190811.. Commercial sponges and the sponges fisheries. Aca.d. Na.t. Sci. of Philadelphia. 36: 202-209. Bull. Bur. Fish. 28: 399-512, pIs. 28-66. COBB, J. N. MOORE, H. F. 1903. The sponge fishery of Florida in 1900. Rept. 1908b. kpractical method of sponge culture. Bull. Bur· U. S. Fish Camm. 1902. Fish. 28: 545-586, pIs. 67-76. CRAWS HAY, L. R. OLD, M. C. 1939. Studies in the market sponges. 1. Growth from 1941. The taxonomy and distribution of the boring" the planted cutting. J. M. B. A. United Kingdom sponges (Clionidae) along the Atlantic coast of North 23 (2): 553-574, 1 fig. America. Chesapeake BioI. Lab. ContT. 44, pp. 1-30, DE LAUBENFELS, M. W. 1 fig., pIs. 1-10. 1932. Physiology and morphology of Porifera exempli­ OSORlo-TAFALL, B. F., and CARDENAS, MAURO fied by Iotrochota birotulata Higgin. Carnegie Inst. 1945. Sobre las esponjlL.'1 comerciales de Quintana RoO '! Washington Pub. 435, Pap. Tortugas Lab. 28: W-66, una enfermedad que las destruye. Cienoia 6 (1): 2 pIs., 6. figs. 25-31, 2 figs. GULF OF MEXICO • 261

PtARSE, A. S. SMITH, HUGH M.-Continued 1932. Inhabitants of certain sponges at Dry Tortugas. 1899. The Florida commercial sponges. Bull. Fish. Tortugas Lab. Papers 28, Carnegie Inst. Washington Comm. 17: 225-240, pIs. 12-31. Pub. 435, 28: 'U7-124, 2 pis., 1 text fig. . STUART, A. H. aolG, MARIO SANCHEZ 1948. World trade in sponges. U. S. Dept. Comm. 1945. Sponge riches of Cuba. Rev. Agric. (Havana) Indust. Ser. 82, pp. -1-95, figs. 1-6, pIs. 1-26. 28 (1): 25-32. TIERNEY, J. Q. 8CII~nDT, OSCAR 1949. The sponge industry of Florida. Florida State 1879. Reports on the dredging, under'the supervision Bd. of Conserv., Ed. Ser. 2, pp. 1-19, figs. 1-4. of Alexander Agassiz, in the Gulf of Mexico. Die 1951. A report on the sponges collected during the spongien des meerbusen von Mexico. Erstes Heft.• University of Miami Marine Laboratory Gulf of 32 pp., pis. 1-4, Gustav Fischer, Jena. Mexico 1947-48 sponge investigation. Univ. Miami 1879. Die Fortsetzung meiner "Spongien des Moorbusens" Marine Lab. Pub. In press. Von Mexico. ZooI. Anz. 2, 379-380. VON LENDENFELD, R; 1880. Reports on the dredging, under the supervision of 1889. A monograph of the horny sponges. Truebner Alexander Agassiz, in the Gulf of Mexico. Die spon­ and Company, London. 936 pp., pIs. 1-50. gien des meerbusen von Mexico (und des Caraibisohen VOSMAER, G. C. J. moores). Zweites Heft., pp. 33-110, pis. 5-10. Gustav 1887. Porifera. In: Bronn, H. G., Die Klassen und 8 Fischer, Jena. ordnungen des Thier-reichs-Zweiter Band. Pp. lilITH, F. G. WALTON 369-496, pis. 26-34. 1941. Sponge disease. in British Honduras anll its 1933. The sponges of the Bay of Naples. Martinus transmiBBion by water currents. Ecology 22: 415-421, Nijhoff, The Hague. Vol. 1 and vol. 2, pp. 1-828. figs. 1-3. . Vol. 3, pIs. 1-71. 1949. Report on a survey of the sponge grounds north of WILSON, H. V. Anclote Light. Mimeographed. Florida State Board 8 of Conservation, pp. 1-29, figs. 1-4. 1902. The sponges collected in Porto Rico in 1899 by "ITH, HUGH M. the U. S. Fish. Corom. Bull. for 1900, 2: 375-411. 1899. Notes on the Florida sponge fishery in 1899. 1925. Studies on dissociated sponge cells. Carnegie Bull. Bur. Fish. 19: 149-151. Inst. Yearbook No. 24, 1925, pp. 242-246. Blank page retained for pagination BIOLOGY OF THE COMMERCIAL SPONGESI

By F. G. WALTON SMITH, Marine Laboratory, University of Miami

All commercial sponges of the Gulf of Mexico contains iodine and the amino'acids lysine, argi­ belong to the family Spongiidae. They are char­ nine, cystine, phenylalanine, and glycine. Very acterized by a lack of spicules and by the presence small amounts of histidine and tyrosine are also of a skeleton consisting of a network of anastomo­ present. sing spongin fibers. The canal system is of the Sexual reproduction takes place at all times of leucon type. Subdermal cavities which act as the year but most intensively during April, May, \Testibules to either the incurrent or excurrent and June, and in November and December, ac­ Canal systems, or both, are present to a varying cording to observations made by the author in degree and are particularly noticeable in the the Bahamas Islands. Eggs are found in sponges \Telvet sponge, Hippiospongia gossypina. as small as 2 inches in diameter. They are im­ The external form is rougWy subspherical in bedded in the tissues between the flagellated the case of sheepswool sponge, Hippiospongia chambers and are about .0.25 em. in diameter. lachne, the yellow sponge, Spongia zimocca ss. In their early stages of development they are barbara, hard head, Spongia officinalis ss. dura, white but become a dark olive green as the and the velvet sponge. The grass sponge; embryo develops. Whereas eggs are easily visible Spongia graminea, may be in the form of an up­ to the naked eye when a sponge is sliced open, the right cylinder, somewhat wider and slightly con­ spermatozoa are not recognizable except under caVe in the upper surface. Large sponges of this the microscope. No information is available as type in the Gulf of Mexico waters, however, are to whether spermatozoa and eggs are produced by USUally cup-shaped. Other sponges sometimes the same sponge at different times. Since they fOund are the reef sponge, Spongia officinalis ss. have not been observed in the same sponge at the obliqua. These are small sponges somewhat same time, it is possible that commercial sponges CYlindrical in general shape but with lobes ter­ are dioecious. minated by oscula on the upper surface. The Toward the end of embryonic development the glOve sponge, assigned to Spongia graminea, is embryo develops a circle of cilia at the anterior, stoutly columnar with fluted sides. The shape less strongly pigmented end. The larva then and appearance varies considerably according to escapes into the excurrent canal system and the environment. Detailed descriptions of the thence to the exterior. Further development has \Tarious forms are given by Moore (1910). not been observed in detail in commercial sponges. The outer surface is covered with a thin but There is little definite evidence as to the food tOugh skin which is usually dense black in color. of sponges. Ingestion of carmine particles by the Portions of the sponge buried in mud and the choanocytes has been observed by Bidder (1896) basal portions are more or less deficient in pig­ and subsequent authors, in the non-commercial ment. The flagellated chambers of all commer­ sponges. According to Bidder, the flagellated cial sponges are small in size and are pyriform or cells of Sycon raphanus, which are 5 microns in 8ubspherical in shape with a diameter of no more width, contain rod-shaped bodies similar to bacilli than 0.03 mm. The choanocytes are approxi­ between 1 and 2 microns long. Pourbaix (1931) mately 4 microns in cross section. describes the transference of carmine grains from . Spongin fiber has been analyzed by Block and choanocytes to adjacent amoebocytes. Pourbaix o 13 11ing (1939). It is composed of keratin, an in­ (1931) also states that after feeding with carmine, ert-----product related to the collogens. Keratin granules of this appeared in the amoebocytes of 1 Contribution No. 109 from the Marine Laboratory. University of MiamI. the Tunisian commercial sponge. 263 264 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

It seems fairly certain that any solid food used Commercial sponges appear to flourish where by the sponge must be very small in size and that there is a good flow of water but not under verY it is carried into the flagellated chambers by the exposed conditions. They are rarely found upoll internal water circulation. The large quantities bottoms consisting entirely of rock or, of course, of. bacteria fo;nd in the calcareous muds of the sediments. In the Gulf of Mexico they are found Bahamas in the vicinity of commercial sponges down to a depth of at least 150 feet and in water suggests that these organisms may be an impor­ close to low tide mark. They are not tolerant tant constituent of the food of these sponges. to reduced salinity except for short periods. They It has been suggested, however, that sponges may appear to be more resistant to temperature also be able to absorb dissolved organic nutrients. changes, however. Very little is known about the physiology of A considerable number of other organisms are commercial sponges. Bidder (1923), Parker associated with sponges. The surface is frequently (1914), and others have studied the water cur­ covered with encrusting Bryozoa and colonial rents. These are set up by movements of the tunicates. All species of the piling fauna normallY flagella of the choanocytes. These appear to found in the vicinity may be epizoic upon coJJl' operate independently of each other. The enor­ merical sponges. The green alga, Batophora mous number of flagellated chambers provides the oerstedi, and species of Acetabularia also becoJJle energy whereby a considerable volume of water attached to the sponge surface. The starfish, is pumped through the sponge. In noncommer­ Echinaster, is sometimes observed upon sponges cial species, such as Spinosella, it has been shown which have lesions of the outer skin, but it is not that although the volume of water is great, the certain to what extent the damage is caused bY pressure does not amount to more than 5 mm. this animal. Nudibranchs are sometimes found'ill in height of water. The jet stream leaving the small pits or irregularities of the surface, and it is osculum may be detected at several inches dis­ possible that they may feed upon the sponge tance and is sufficient so that on a calm day a tissue. Larger holes may be inhabited by varioUS distinct disturbance of the water surface may be species of small crabs. Dromia is often a perIllll' seen above commercial sponges which are near nent inhabitant of such places. The green algll, to the surface. The oscula may be closed by Dictyosphaeriajavulosa, grows in convenient nicheS muscular acfffbn when stimulated locally. and may become almost enclosed by the subse­ Bergmann (1949) and his associates have de­ quent growth of the sponge. A number of the scribed in a series of papers the extraction of smaller and commoner gastropods are also found sterols from sponges. There is a considerable inhabiting cavities of the sponge surface which degree of specificity in the type of sterol found in may have been caused by local necrosis, the various sponges, and this bears a close relation activities of carnivores or overgrowth of the to taxonomic classification. Although the com­ sponge around sedentary organisms. The barna' _ mercial sponge~so far investigated have a high cle, Balanus declivis; which is more commonl! fat content, the sterol content of the unsaponifi­ found in noncommercial species of sponges, ~ able matter is lower than the average. The sterols occasionally seen imbedded in the surface 0 of commercial sponges have not yet been ana­ commercial types. lyzed in detail. A considerable fauna inhabits the larger passages Commercial sponges are typically found asso­ of the canal system. Pearse (1934) lists a large ciated with muddy bottom sediments, particu­ number of organisms inhabiting a reef sponge i~ larly where rocky outcrops or reefs provide a Dry Tortugas. The most commonly found ill a suitable substrate upon which the larva can settle commercial sponges are the snapping shriJJlP' without being destroyed by silt. They are some­ Synalpheus brooksi, and, the polychaete worJll' times found growing attached to dead corals or Leodice spongicola., Other species of polychsetes gorgonians and may occasionally grow upon living are also found, particularly syllids and occssionn,llY green algae, such as Penicillus. Under these con­ Amphitrite. ditions the alga may eventually become com­ Among other Crustacea occurring here are the pletely imbedded in the sponge except for the pontonid shrimp, Coralliocaris pearsei, and nuJll lower portion of its stalk. erous amphipods of the genera, Leucothoe stld GUIJF OF MEXICO 265 Colomaestix. The stomatopod, Gonodactylm oer­ sponge cultivation are described by Smith (1949), . ~tedi, and small crabs may inhabit the larger Moore (1910b),Crawshay (1939), and Cahn Passages. (1948). Ophiuroids, particularly Ophiactis savigny, are "found both in surface depressions and in the larger BIBLIOGRAPHY :Portions of the canal system. The anemone, BERGMANN, WERNER. Aiptasia, appears to form depressions in the 1949. Comparative biochemical studies on the lipids of Surface of sponges into which it is able to retrea.t. marine invertebrates, with special reference to the The most destructive organism is ~pparently a sterols. Jour. Mar. Res. 8 (2): 137-176. BIDDER, G. P. ,/ ?pecies of fungus which caused widespread and 1896. The. collar-cells of Heterocoela. Quart. Jour. lntenflive mortality among the Gulf of Mexico aod Micro. Sci., N. S., 38: 9-42. Caribbean sponges in 1939. Galtsoff (1940) ten­ 1923. The relation of the form of a sponge to its cur­ tatively identifies the organism as Spongiophaga rents. Quart. Jour. Micro. Sci., N. S., 67 (2); communis which was first observed to be parasitic 293-323. BLOCK, R. J., and BOLLING, D. uPon sponges by Carter (1878). Since it was not 1939. The amino acid composition of keratins. Jour. Possible to culture the organism, it cannot defi­ BioI. Chem., Vol. 127. llitely be assigned to any particular group of CAHN, A. R. fungi. Furt.her accounts of this sponge disease 1948. Japanese sponge culture experiments ill the are also given by Walton Smit.h (1941) and by South Pacific Islands. U. S. Fish and Wildlife Serv­ ice, Fish. Leaflet 309, 9 pp. Osorio-Tafall (1945). CARTER, H. J. A number of observations have been made by 1878. Parasites of the Spongida. Ann. Mag. Nat. Galtsoff, Wilson, and others, upon the ability of Hist., Ser.5, 2: 165-169. disassociated sponge cells to re-aggregate. This CRAWSHAY, L. R. earlier work is also referred to by de Laubenfels 1939. Studies in the market sponges. I. Growth from . the planted cutting. Jour. Mar. BioI. Assn., U. K., (1934) in experiments upon the regeneration of 23 (2): 533-574. Iotrochota. GALTSOFF, P. S. The growth rate of sponges has been measured 1929. Heteroagglutination of dissociated sponge cells. by Crawshay (1939) who measured the increase in BioI. Bull. 57 (5): 25(}-260. size of small cuttings sliced from commercial 1940. Wasting discase causing mortality of sponges in the West Indies and Gulf of Mexico. Proc. Eighth SPOnges. He found that the velvet sponge grows Am. Sci. Cong. 3: 411-421. ~t Such a rate as to approximately'double or treble HYMAN, L. H. ,Its volume in the period of a year. Extensive 1940. The invertebrates: Protozoa through Ctcnophora. unpUblished series of measurements by the present Vol. 1, pp. 284-364, McGraw-Hill Book Co., New author have shown that the growth expressed as York. LAUBENFELS, M. W., DE .a. percentage of size diminishes with increasing 1934. Physiology and morphology of Porifera exem­ SIZe in the case of both sheepswool and velvet plified by Iotrochota birotulata Higgin. Carnegie SPonges. Whea these sponges reach approximately Inst. Washington Pub. 435, Papers Tortugas Lab. 12 inches in diameter the central portion of the 28 (2): 37-66, 2 pIs., 6 text-figs. upper surface begins to undergo local necrosis so 1936. A discussion of the sponge fauna of the Dry Tortugas ... Carnogie Inst. Washington Pub. 467, that the larger sizes become somewhat doughnut­ Papers Tortugas Lab. 30: 1-225. s~aped. This is apparently due to the loss of effi­ 1948. The order Kcratosa of the phylum Porifera. A CIency in respiratory, excretory, and nutritive monographic study. Allan Hancock Found. Occas. ~)(changes related to a diminution in the sur- Papers, vol. 3: 1-217, pIs. 1-30, figs. 1-31. ace/volume ratio. LENDENFELD, n., VON 1889. A monograph of the horny sponges. 936 pp., The increase in percentage growth rate with 50 pIs. TrUbner Co., London. decreasing size makes it possible to cultivate LINDNER, M. J. SPonges by cutting them into small pieces and by 1948. Mexican sponge fishery. Sponge Inst. Trade ~anting each piece upon a stone or cement base. Rept.64. egeneration of the cut surfaces takes place MOORE, H. F. 1910a. The commercial sponges and the sponge fish­ rapidly, and the sponge quickly attaches itself eries. Bull. U. S. Bur. Fish., 1908, 28: 399-511. to the stony surface. Other methods used in pIs. 28-66. 266 FISHERY BULLETIN OF THE FISH· AND WILDLIFE SERVICE

MOORE, H. F. POURBAlX, N. 1910b. A practical method of sponge culture. Bull. 1931. Notes sur Hippiospongia equina au voyage U. S. Bur. Fish.. 1908, 28: 545-586. d'~tude ~ Adjim-Djerba. Stat. Oceanog. SalamrobO, OSORIO-TAFALL, B. F. Bull. 22. 1945. Sobre lase sponjas commerciaies de Quintana SMITH, F. G. WALTON. Roo y una enfermedad que las destruye. Ciencla 1941. Sponge disease in British Honduras and its 6 (1): 25-31, 2 figs. transroiBSion by water currents. Ecology 22 (4): PARKER, G. H. 415-421. 1914. On the strength and volume of the water currentA 1949. Sponge cultivation. Mar. Lab., Univ. of Miami, produced by sponges. Jour. Exp. Zool. 16 (3): Spec. Servo Bull. No.3. (Mimeo.) 443-446. TIERNEY, J. Q. PEARSE, A. S. 1949. The sponge industry of Florida. State of 1934. Inhabitants of certain sponges at Dry Tortugas. Florida, Bd. of Conserv., Mar. Lab., Univ. MianU Carnegie Inst. Washington Pub. 435, Papers Tor­ Pub., Educ. Ser. No.2, pp. 1-19, figs. 1-4. tugas Lab. 28: 117-124. HYDROIDS OF THE GULF OF MEXICO

By EDWARD S. DEEVEY, Jr., Osborn Zoological Laboratory, Yale University

The life cycle of hydroids "typically" alternates of Mexico is a peculiarly appropriate setting for ~etween a sessile, asexually reproducing polyp or such a catalog of ignorance, for what we do not Ydroid stage and a free-swimming sexual medusa know about Gulf hydroids should be especially stage. One generation or the other may be re­ obvious against the massive backdrop of what we duced, however, or even suppressed entirely. know about the biology of the Gulf. Because of this fact and because hydroids are COLLECTING obtained by shore collectors or by dredging,· "'hereas medusae form part of the plankton and Hydroid collecting in the Gulf of Mexico has are studied by different zoologists, the taxonomy been chiefly undertaken in the Tortugas and·the of the group is in an unhappy state. Florida Keys, but if we except the work of Mayer Closely related types may differ in a respect (1910) as dealing almost entirely with medusae, that at first sight would appear fundamental: the the only paper specifically discussing the Tortugas Polyp may produce free-living medusae, or the fauna is that of Wallace (1909). Gulf of Mexico lnedusa may remain permanently attached to the. records are scattered through many papers (Fraser polyp, even degenerating completely except for 1943, 1945; Jiiderholm 1896, 1903, 1920; Leloup Its sex products, which then appear to be the sex 1935, 1937; Perkins 1908; Pourtales 1869; Stechow br?ducts of the polyp. This distinction, far from 1912, 1919, 1923, 1926) and through the mono­ .elllg of subordinal rank, may split a genus down graphs of Nutting (1900, 1904, 1915), but system­ the middle or, at most, may divide closely related atic dredging in the region has not been under­ genera from each other, according to one's view taken since the days of the Corwin and the Blake. of What constitutes a generic character. That is, The collections of L. de Pourtales on the former Unless the structural characters of the hydroid are vessel were reported by Allman (1877), and those to be given no weight whatever, the mode of of A. Agassiz on the latter were published by reproduction cannot provide the basis for erecting Clarke (1879) and Fewkes (1881). taxonomic categories higher than the genus. The monograph of Fraser (1944) recapitulates d' Thus, the hydrozoan systematist faces a real all the earlier records and adds many more, for Uemma, and he is not helped by the fact that Fraser examined the large collections in the ~any typically reproducing species have received U. S. National Museum and the Museum of ifferent names as medusae and as polyps and Comparati.ve Zoology whither most of the Amer­ ~Ust continue to bear them until proof of their ican hydroids collected in the nineteenth century Identity is obtained. The writer is not a student eventually made their way. It is possible to ~f Inedusae, and in fact his knowledge of hydroids extract from Fraser's book an essentially com­ hlargely confined to specimens preserved in alco­ plete picture of the hydroid fauna of the Gulf as tot He is in no position to do anything about far as it is known. In fact, thi.s procedure is the he fact that even the families are differently basis of the present account. Fraser's list has Collstituted in hydroid and medusa systematics, been expanded in only one noteworthy respect. and the task of fusing the two systems awaits an The survey of fouling on navigation buoys, a~ler zoologist. This chapter deals exclusively conducted by the Woods Hole Oceanographic "'Ith hydroids. . Institution during World War II, yielded several Ii The hydroid fauna of the Gulf of Mexico is hundred records· of hydroids from the Bahamas; . ttle known, and the chief purpose of this account both coasts of Florida, and Texas. Most of these 18 to document this fact. A treatise on the Gulf are still unpublished, but those that are new 267 268 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

, records from the Gulf proper are starred in the shallow water boreal species occurring at consid-' check list. The Texas records are of exceptional erable depths in the Gulf (and Caribbean): interest, however, as only three hydroids had Ewlendrium tenellum, LaJoea dumosa, L. gracillim(J,· . previously been recorded from those waters; Another way of looking at the facts is this: of they were included in the writer's account of the 156 species known from the Caribbean, 61 are not hydroids of Louisiana and Texas (Deevey 1950), "known from the Gulf; 29 of them, at least, are a zoogeographic discussion that was founded common enough in the Caribbean to have been . primarily on a small collection made by J. W. taken at more than one station. Hydroid sta­ Hedgpeth. tistics are scarcely necessary to prove the point, ZOOGEOGRAPHY but it is obvious that the Gulf is not a strictlY tropical body of water. Low winter temperatureS The list of 183 species looks impressive, but it and low and variable salinity, particularly'along would be idle to pretend that the hydroids of the northwestern shore, are only some ,of the faC~ the Gulf are well-known. Tropical regions gen­ tors that must be responSible for maintaining ~ erally have a weiLlth of species, but hydroid different "fauna" in the Gulf of Mexico. habitats are probably no more extensive in the The 95 species that are common to the Gulf tropics than elsewhere. The rarity of the rarest and the Caribbean present different probleIDS. species is correspondingly greater, and it is un­ Most of them are definitely warm water types, likely that more than half the hydroids living in although a high proportion belong to the tropical the Gulf have yet been found there. Partly flotsam (especially sargassum) fauna and so maY because of inadequate collecting and partly be­ not be true residents of the Gulf. Seventy of the cause shallow water hydroids are always under Gulf hydroids and 59 of the 156 Caribbean species suspicion as fouling organisms, little can be said are represented in the much larger list of 312 about the meaning of their geographic distribution. species recorded from the eastern tropical Pacific. Of the total of 183 hydroid species known from The richness of the Pacific fauna, which is knoWJl the Gulf of Mexico, 95 are also found in the almost exclusively from the Allan Hancock colleC' Caribbean and another 18, though not yet re­ tion (Fraser 1948), is another indication that the corded from the Caribbean, are' known from the Caribbean hydroids have been inadequately col~ eastern tropical Pacific. The remaining 70 species lected; it is surely correct to suppose that the ~8 include some of the ~ost interesting. species common to the Gulf and the eastern trop' Some of the 70, of course, are known only from ical Pacific will appear in the Caribbean, along the Gulf, but while a few true endemics are to be with many others. What is surprising is that expected it is too early to say which ones they are; nearly half of the tropical Atlantic hydroids croSS at any rate, no common species is known to be the Isthmus of Panama without undergoing endemic. The interesting members of the group specific differentiation. If, one cares for statis· of 70 are those whose main range is otherwise tical statements, the "strength of the relatio~' boreal. The boreal species among the hydroids ship" between the Gulf and Pacific faunas IS of Texas and Louisiana have already been dis­ nearly as great (38 percent) as is that, between cussed (Deevey 1950) . They include several, the Gulf and Caribbean faunas (52 percent). d such as Podocoryne carnea, that are unknown in the The Isthmus of Panama is not very old, an. warmer parts of the Gulf, and at least one, many biogeographers have supposed that it IS Tubularia crocea, whose ecology is well enough younger than most of the species of marine organ' understood to indicate that it could not flourish isms (Schuchert 1935). The problem is related to in southern Florida. To the list of supposed that of Tethyan distributions; pan-tropicalspeci~ relicts of a glacial age of the Pleistocene given in (more usually genera) are supposed to have hB f the earlier paper may be added the name of their ranges established in the Tethys Sea, 0 Oladocarpus jlexilis, a moderately deep water Cretaceous and early Tertiary age, only to hBV'e species taken at three stations off Mobile but not them sundered by the rise of central Asia (see otherwise known south of Cape May. In several Ekman 1934, 1935, for review). If disjunct other cases we have the familiar phenomenon of a distributions in the Mediterranean and the GULF OF MEXICO - 269 Indian western Pacific regions have had this CHECK LIST OF GULF OF MEXICO origin, there is no 'reason to doubt that the HYDROIDS Isthmus of Panama was crossed about the same Geographic distribution is indicated by the tUne by the same species, or some of them. The following symbols: . sPecies that are perhaps most likely to have spread K, Florida Keys, including Cay Sal Bank and 80 widely and to have .crossed modern land bar­ southern Florida as far east as MiarrM, but not riers so freely are now truly pan-tropical species, the Bahamas. but the evidence they provide, according to· the T, Tortugas. COnventional canons of biogeography, is ruled C, Cuba. invalid by the possibility that they are spreading Y, Yucatan. today. Unhappily, if one chooses to follow the NW, northwestern Gulf (Texas, Louisiana). rules and exclude the pan-tropical species, it can NE, northeastern Gulf (including 'Fampa Bay). only be said that the remaining species prove Co., Caribbean Sea. nothing, at least as far as hydroids are conc~rned. EP, eastern tropical Pacific Ocean, south Of 1'he reason is not so much biogeographic as tllJc:onomic. United States-Mexico boundary, and including the oceanic islands. The number of hydroids common to the two starred names are new records for the Gulf of sides of Central America is large, but an even * Me~ico, found in the Woods Hole Oceanographic larger number is common' to th~ two coasts of Institution fouling collection. ~orth America taken as a whole. According to Fraser's tabulation (1944), 123 species are known Suborder GYMNOBLASTEA (Anthomedusae), from east and west coasts of the Americas, and by athecate hydroids no means all of these are circumpolar. Neither· are the tropical species all pan-tropical. The Cordylophora lacustru Allman, 1844. NW; Ca. Turritopsu jascicularis Fraser, 1943. K. "American" distribution pattern is far too com­ *Turritopsis nutricula McCrady, 1856. K; Ca, EP. !ll.on to be accidental, but its commonness raises Syncoryne eximia 1 (Allman, 1859). NW. dOUbts about the taxonomy. Fraser was a sound, Syncoryne mirabilis L. (Agassiz), 1862. K; EP. careful worker with a "good eye" for specific Zanclea costata Gegenbaur, 1856. T, NW; Ca, EP. differences. However, his experience, though Zanclea gemmosa McCrady, 1858. T; EP. enormous, was largely confined to the Americas. Bimeriafranciscana Torrey, 1902. NW; B. Bimeria humilis Allman, 1877. T, NW; Ca. When one remembers that the hydroids of the Bougainvillia carolinensis (McCrady, 1858). T, NW. ~ll.8t Indies are poorly known (only three families Bougainvillia inaequalis Fraser, 1944. NW. of the Siboga hydroids having been reported by Bougainvillia rugosa Clarke, 1882. NW; Ca. nillard before his death), one cannot escape the Eudendrium album Nutting, 1898. K; EP. SUspicion that many species apparently endemic Eudendrium attenuatum Allman 1877. T; EP. Eudendrium carneum Clarke, 1882. T; Ca, EP. to the American tropics are still to be collected, Eudendrium distichum Clarke, 1879. K.

01' are already· known under other namesj from Eudendrium exiguum Allman, 1877. K; Ca, EP. other parts of the world. Apart from this tax­ Eudendrium eximium Allman, 1877. K, NE; EP. onOmic difficulty, inadequate knowledge of the Eudendrium jmticosum Allman, 1877. K. h:Ydroids of the western Pacific and Indian Oceans Eudendrium gracile Allman, 1877. K. Eudendrium hargitti Congdon, 1906. T. itnposes another limitation on the case for Tethyan Eudendrium laxum Allman, 1877. K; Ca. l>aleogeography, for western. Atlantic~western Eudendrium speciosum Fraser, 1945. N~. l>acific disjunctions have often been used (however Eudendrium tenellum Allman, 1877. K; Ca, EP. llnWisely) in building such a case, and we know Eudendrium tenue A. Agassiz, 1865. NW?; Ca, EP. of no certain examples among hydroids. Hydractinia echinata Fleming, 1828. K, NW. Podocoryne carnea Bars, 1846. NW. Dntil the hydroids of the world have been given Pennaria tiarella (Ayres, 1854). K, T, C; Ca, EP. !ll.uch more study and some monographic revision, then, it is unsafe to use them for many zooge­ I N. J. BerrllJ. In a letter to the author, has given good reasons lor suspeot· Ing that tho species reported under this name (rom Texas and (rom western • Ographic purposes. Florida (Deevey 1950). Is an undescribed spt'cles. 270 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Ectopleura grandis Fraser, 1944. K, NW. Family LAFOEIDAE Tubularia crocea (L. Agassiz, 1862). NWi EP. Acryptolaria abies (Allman, 1877). K; Ca. Cladonema mayeri Perkins, 1906. T. Acryptolaria conferta (Allman, 1877). T, C; Ca, EP. Acryptolaria elegans (Allman, 1877). K, Tj Ca. Suborder CALYPTOBLASTEA (Leptomedusae), Acryptolaria longitheca (Allman, 1877). K, Tj Ca. thecate hydroids Acryptolaria pulchella (Allman, 1888). K; EP. Eucryptolaria pinnata Fraser, 1938. C; Ca, EP. Family CAJI,1PANULARIDAE Filellum serpens (Hassall, 1852). T, NW; Ca, EP. Campanularia amphora (L. Agassiz, 1862). T? Filellum serratum (Clarke, 1879). C; Ca. Campanularia (?) brevicaulis Nutting, 1915. Y. Lafoea coalescens Allman, 1877. K. Campanularia jlexllosa (Hincks, 1861). T? Lafoea dumosa (Fleming, 1828). T; Ca, EP. *Campanularia (?) hummelincki (Leloup, 1935). Kj Ca. Lafoea gracillima (Alder, 1857). Tj Ca, EP. Campanularia (?) macroscypha Allman, 1877. K, T, Y. Lafoea tenellula Allman, 1877. K; Ca. Campanularia marginata (Allman, 1877). K, T, C; Lafoea venusta Allman, 1877. K, T, C; Ca. Ca. Lictorella convallaria (Allman), 1877. K, T, C; Ca, EP. Clytia coronata (Clarke, 1879). K, NW; Ca, EP. Zygophylax rigida (Fraser, 1940). Y. Clytia cylindrica L. Agassiz, 1862. K, NW; Ca, EP. Clytia fragilis Congdon, 1907. T, NW; Ca. Family SYNTHECIDAE Clytia johnstoni (Alder, 1856). K, T; EP. Synthecium ? gracile Fraser, 1937. T; Ca, EP. Clytia longicyatha (Allman, 1877). K, T, NW; Ca, Synthecium ? marginatum (Allman, 1877). K; Ca. EP. Synthecium ? nanum Fraser, 1943. Ti Ca. Clytia macrotheca (Perkins, 1908). K, T; Ca. Synthecium ? rectum Nutting, 1904. C. , Clytia minuta (Nutting, 1901). T? Synthecium tubithecum (Allman), 1877. K, T, C; Ca. Clytia noliformis (McCrady, 1858). T, NW; Ca, EP. *Clytia raridentata (Alder, 1862). K; Ca, EP. Family SERTULARIDAE Gonothyraea gracilis' (SarB, 1851). K, NW; Ca, EP. Diphasia digitalis (Busk, 1852). K, T, C; Ca. Obelia bicuspidata Clark 1876. K, NW, NE; Ca. l Pasya quadridentata (Ellis and Solander, 1786). '1', Obelia commissuralis McCrady, 1858. T?; EP. Obelia dichotoma (Linnaeus, 1758). K, T, C, NW, NWj Ca, EP. "/. Sertularella amphorifera Allman, 1877. K, T, C, , NE; EP. Ca, EP. Obelia equilateralis Fraser, 1938. NW; Ca, EP. Sertularella areyi Nutting, 1904. C. E' Obelia geniculata (Linnaeus, 1758). NW; Ca, EP. Sertularella conica Allman, 1877. KT NW N ' Obelia hyalina Clarke, 1879. K, Cj Ca, EP. Ca, EP. Obelia obtusidens (Jaderholm, 1904). NW; EP. Sertularella distans (Allman, 1877). T, C, Y; Ca. Family CAMPANULINIDAE Sertularella formosa Fewkes, 1881. C; Ca, EP. W. Sertularella gayi (Lamouroux, 1821). K, C, Y, N I *Cuspidella costata HinckB, 1868. K. Ca. Cuspidella humilis (Alder, 1862). NW; EP. Sertularella humilis Fraser, 1943. K. Eucuspidella pedunculata (Allman, 1877). T. Sertularella megastoma Nutting, 1904. Y; Ca. *Lafoeina tenuis SarB, 1873. K, NE. Sertularella pinnigera Hartlaub, 1900. K; Ca. Oplorhiza parvula Allman, 1877. K. Sertularella quadrata Nutting, 1904. C; Ca. Stegopoma fastigiata (Alder, 1860). T. Sertularella sieboldi Kirchenpauer, 1884. C. Thyroscyphus ramosus Allman, 1877. K, T; Ca. Sertularella speciosa Congdon, 1907. K; Ca. Family HALECIDAE Sertularella tenella (Alder, 1856). Cj Ca, EP. , Sertularia cornicina (McCrady, 1858). K?, Y; Ca, Halecium bermudense Congdon, 1907. K, T, NEj Ca, EP. ~. ~ Sertularia dalmasi (VerBluyB, 1899). T, NW, Halecium dyssymetrum 2 Billard, 1929. K, T. Ca,EP. Halecium filicula Allman, 1877. K. Sertularia exigua Allman, 1877. K; Ca, EP. Halecium gracile Verrill, 1874. K; EP. Sertularia jlowersi Nutting, 1904. C. NE; Halecium macrocephalum Allman, 1877. K, Ti EP. Sertularia injlata (Versluys, 1899). K T NW, Halecium nanum Alder, 1859. K, T, NW; Ca, EP. Ca, EP. Halecium tenellum HinckB, 1861. T, NE; Ca, EP. Sert'ularia mayeri Nutting, 1904. K, T; Ca, EP. ElP. Family HEBELLIDAE 8edularia pourtalesi Nutting, 1904. K, T; Clio, Sertularia stookeyi Nutting, 1904. Kj EP. Hebella calcarata (A. Agassiz, 1865). NE; Ca, EP. Sertularia tumida Allman, 1877. T. . Ca. Scandia mutabilis (Ritchie, 1907). T; Ca, EP. Sertularia turbinata (Lamouroux, 1816). K, T, Ell'. 7'huiaria crisioides (Lamouroux, 1824). Kj Clio, • Preliminary study of this species In the Woods Hole Oceanographic In­ stitution fouling collection Indlcetes that It Is probably the II. dvaavmdrum Thuiaria tropica (Stechow, 1926). T. of LeJoup 119361 but not of Billard. and that It needs 1\ Dew name. Idiella pristis (Lamouroux, 1816). K, T; Ca. GULF OF MEXICO 271

Family PLUMULARIDAE Plumularia geminata Allman, 1877. K; Ca. Aglaophenia allmani Nutting, 1900. K, T; Ca. Plumularia inermis Nutting, 1900. T; Ca, EP. Aglaophenia aperta Nutting, 1900. C, NE. Plumularia macrotheca Allman, 1877. C. Aglaophenia bicornuta Nutting, 1900. C. Plumularia margaretta (Nutting, 1900). K, T; Ca, Aglaophenia? constricta Allman, 1877. K. EP. Aglaophenia cristifrona Nutting, 1900. C, NW. Plumularia megalocephala Allman, 1877. K, C; Ca, ? Aglaophenia dichotoma Kirchenpauer, 1872. NE. EP. Aolaophenia dubia Nutting, 1900. K, T, C; Ca, EP. Plumularia paucinoda Nutting, 1900. C. Aglaophenia elonoata Meneghini, 1845. NE. Plumularia setacea (Ellis, 1755). R, T, NW, NE; Aglaophenia ftowersi Nutting, 1900. K; Ca. EP. Aglaophenia late-carinata Allman, 1877. K, T, NW; Plumularia setaceoides Bale, 1882. K, Ca. Ca. Plumularia strictocarpa Pictet, 1893. Kj Ca. Aglaophenia longiramosa Fraser, 1945. NE. Schizotricha dichotoma Nutting, 1900. K. Aglaophenia lophocarpa Allman, 1877. K, T, C, NEj Schizotricha lenella (Verrill, 1874). NW?; Ca, EP. Ca, EP. ThecocarpU8 bispinosus (Allman, 1877). K. Aglaophenia ? mercatoris Leloup, 1937. NE. Thecocarpus distans (Allman, 1877). K, T. Aglaophenia perpusilla Allman, 1877. K, T, NW; Ca. SUMMARY Aglaophenia raridentata Fraser, 1944. K. A total of 183 species of hydroids, 31 athecate Aglaophenia rhynchocarpa Allman, 1877. K, T, C; Ca. and 152 thecate, are known from the Gulf of Aglaophenia rigida Allman, 1877. K, NW; Ca, EP. Mexico, mostly from the Tortugas and the Florida Aglaophenia tridentata Versluys, 1899. K, T; Ca. Keys. Medusae are not considered. The Gulf Aglaophenoides mammiUata (Nutting, 1900). T. and the Caribbean have 95 species in common, Aglaophenopsis hirsuta Fewkes, 1881. K. but 61 Caribbean species are unknown in the Gulf. Antennella gracilis Allman, 1877. K, Cj Ca, EP. Antennella quadriaurita Ritchie, 1909. K, C. Seventy Gulf species also occur in the eastern Antennella secundaria (Gmelin, 1788). T; Ca. tropical Pacific, including 18 not yet known from Antennopsis distans Nutting, 1900. C. the Caribbean. Taxonomic difficulties, as well as Antennopsis hippuris Allman, 1877. K. inadequate collecting, make hydroid geography an Antennopsis longicorna Nutting, 1900. C. unsatisfactory subject, and it is uncertain how far Antennopsis nigra Nutting, 1900. C. ,Antennularia simplex Allman, 1877. K, C. the apparently common "American" distribution Cladocarpu8 carinatus Nutting, 1900. T. pattern should be taken seriously. What is espe­ Cladocarpus dolichotheca Allman, 1877. K, T. cially interesting is the occurrence in the Gulf of Cladocarpus ftexilis Verrill, 1885. NE. a significant number of boreal species, some of Cladocarpus ftexuosu8 Nutting, 1900. NE. them seemingly disjunct in the northwestern Gulf. Cladocarpus longipinna Fraser, 1945. NE. Cladocarpus obliquu8 Nutting, 1900. C. LITERATURE CITED Cladocarpu8 paradisea Allman, 1877. K. Cladocarpus 8igma (Allman, 1877). K, C. ALLMAN, G. J. Cladocarpus tenuis Clarke, 1879. T; Ca. 1877. Report on the Hydroida collected during the Cladocarpus ventricosus Allman, 1877. K. exploration of the Gulf Stream by L. F. de Pourtal~s, Diplopteron longipinna Nutting, 1900. K. Assistant United States Coast Survey. Mem. Mus. Diplopleron quadricorne Nutting, 1900. C; Ca. Compo Zoo1-., Harvard College 5 (2): 1-66, 34 pIs. Halicornaria sinuosa Fraser, 1925. K, NE. Cr,ARKE, S. F. Halicornaria speciosa Allman, 1877. K; Ca. 1879. Report on the Hydroida collected during the Halopteri8 carinata Allman, 1877. K, T; Ca. exploration of the Gulf Stream and Gulf of Mexico Lylocarpus clarkei Nutting, 1900. C, Yj Ca. by Alexander Agassiz, 1877-78. Bull. Mus. Compo Lytocarpus grandis (Clarke, 1879). Kj Ca. Zool., Harvard College 5 (10): 239-252, 5 pIs. Lytocarpus philippinu8 (Kirchenpauer, 1872). K, T; DEEVEY, E. S. Ca, EP. 1950. Hydroids from Louisiana and Texa.'l, with remarks Monostaechas quadridens (McCrady, 1858). K, T, Y; on the Pleistocene biogeography of the western Gulf NW, NEj Ca, EP. of Mexico. Ecology 31: 334-367. Plumularia attenuata Allman, 1877. Kj Ca, EP. EKMAN, SVEN. Plumularia clarkei Nutting, 1900. C. 1934. Indo-Westpazifik und Atlanto-Ostpazifik, eine Plumularia diaphana (Heller, 1868). K, T, NW; tiergeographische Studie. Zoogeographica 2: 320­ Ca, EP. 374. Plumularia filicula Allman, 1877. K; EP. 1935. Tiergeographie des Meeres. Leipzig: Akade­ Plumularia ftoridana Nutting, 1900. K, T, NWj EP. mische Verlagsges. xii +542 pp. 272 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

FEWKES, J. W. NUTTING, C. C. 1881. Reports on the results of) drt'Jdging under the 1900. American hydroids, Part 1. The Plumularidall' supervision of Alexander Agassiz ... by the U. S. Smithsonian lnst., U. S. Nat. Mus., Spec. Bull., HZ Coast Survey steamer Blake. XI. Report on the pp., 34 pis. Acalephae. Bull. Mus. Compo Zool., Harvard Col­ 1904. American hydroids, Part II. The Sertularidall' lege, 8: 127-140. Smithsonian lnst., U. S. Nat. Mus., Spec. Bull., 151 FRASER, C. McL. pp., 41 pis. 1943. Distribution records of some hydroids in the col­ 1915. American hydroids, Part III. The Campanularl­ lection of the Museum of Comparative Zoology at dae 8Qd the Bonneviellidae. Smithsonian lnst., U. S. Harvard College, with description of new genera and Nat. Mus., Spec. BulL, 118 pp., 27 pis. new species. Proc. New England Zool. Club 22: 75­ PERKINS, H. F. 98, pis. 15-20. 1908. Notes on medusae of the western Atlantic. Carne­ 1944. Hydroids of the Atlantic coast of North Americ~. gie lnst. Washington, Pap. Tortugas Lab. 1 (8): 451 pp., 94 pis. Toronto: Univ. Toronto Preas. 133-156, pis. 1-4. 1945. Notes on some recently collected hydroids in the DE POURTALils, L. F. United States National Museum, with descriptions of 1869. Contributions to the fauna of the Gulf Stream at three new species. Jour. Washington Acad. Sci. 35: great depths. Bull. Mus. Compo Zool., Harvard 21-23. College, 1 (6): 103-120. 1948. Hydroids of the Allan Hancock Pacific Expedi­ tions since March, 1938. Allan Hancock Pacitic SCHUCHERT, CHARLES. Exped. 4 (5): 179-334. 1935. Historical geology of the Antillean-Caribbean

JXDERHOLM, ELOF. region or the lands bordering the Gulf of Mexico and 1896. Ueber aussereuropli.ische Hydroiden des Zoologi­ the Caribbean Sea. John Wiley, New York: 811 pp. schen Museums der Universitll.t Upsala. K. Svensk. STECHOW, E. Vet.-Akad., Handl., Bihang, 21, Aft. IV, No.6: 1-20. 1912. Hydroiden der MQnchener Zoologischen Staats­ Hl03. Aussereuropil.ische Hydroiden im schwedischen llammlung. Zool. Jahrb., Abt. Syst. 32: 333-318, Reichsmuseum. Ark. Zool. 1: 259-312. pis. 12, 13. 1920. On some exotic hydroids in the Swedish Zoological 1919. Zur Kenntnis der Hydroidenfauna des Mittel­ State Museum. Ark. Zool. 13 (3): 1-11. moores, Amerikas, und anderer Gebiete. 1. ZoOI. LELOUP,' EUGENE. Jahrb., Abt. Byst. 42: 1-172. 1935. Hydraires calyptoblastiques des lndes Occiden­ 1923. Zur Kenntnis der Hydroidenfauna des Mittel­ tales. Mem. Mus. Roy. d'Hist. Nat. Belg., ser. 2, meeres, Amerikas, und anderer Gebiete. II. Zool. fase. 2, 73 pp. Jahrb., Abt. Syst. 47: 1-270. 1937. Resultats scientitiques des croisieres du Navire­ 1926. Einige neue Hydroiden aus verschiedenen MooreS­ 11:cole Beige Mercator, Vol. 1, No. VI. Hydroidea, gebieten. Zool. Anz. 68: 96-108. Siphollophora, Ceriantharia. Mem. Mus. Roy. d'Hist. WALLACE, W. B. Nat. Belg., ser. 2, fasc. 9: 91-127. 1909. A collection of hydroids made at the Tortug&S' MAYER, A. G. during May, June, and July, 1908. Carnegie lost. 1910. Medusae of the world. Carnegie lnst. Washing­ Washington Yearbook 7 (1908): 136-138. ton Pub. 109, 3 vols. HYDROMEDUSAE OF THE GULF OF MEXICO

By MARY SEARS, Wood.f Hok Oceono,rophic lmtitution

,So few plankton collections have been made in Crady, among the Anthomedusae; Laodicea cruiciatG the Gulf of Mexico and so few of these have been ForskAI; Phalidium di8coida Mayer; Phialucium carolinGe studied by hydromedusae specialists that it is Mayerj Eucheilota ventricularia MoCrady; E. duodecimali8 A. Ag8BBiz; Emima mira McCrady; E. elephtu Haackel; n.ecessary to rely almost entirely on Mayer's Emimalphe8 coerulea L. Ag8BBiz; PhortiB pyramidGli8 (1900, 1904, 1910) reports from the Tortugas. L. Ag8BBizj P. lactea Mayer; Aequorea ftoridana Mayer, It is to be noted that most names listed in Mayer's among the Leptomedusae; Gerl/onia probo8cidaliB ForskAI; first (1900) paper were reduced to the synonomy Liriope tetraphylla Chamisso and Eysenhardt; Olindiaa of, other species in his monograph of 1910. Sub­ pho8phorica tenuiB Fewkes j Rhopalonema velatum Gegen­ baur, among the Trachymedusae; and AeginurG grimaldii sequent writers, expecially Bigelow (1913, 1918, Maas; Cunoctantha octonaria McCrady; Solmundella 1919, 1938, 1940), Kramp (1919, 1920, 1926, bitentaculata Quoy and Gaimard (using the names which ~930, 1932, 1933, 1939, 1942, 1947, 1948), and appear to conform with present usage). anson (1936) have also made certain revisions A goodly number of species, however, do not ap­ resulting in a slight reduction in the number of pear to have been subject to critical review in sPecies originally enumerated by Mayer in his. recent years, so that it is uncertain whether they b:l.onograph (1910). are good species or merely synonyms of others. In comparing the resulting list with that in Hence, they are not listed at this time. .the section on the hydroids it is striking to note that few species (i. e., Turritopsis nutricula LITERATURE CITED ~cCrady, Zanclea costata Gegenbaur, Bougain­ BIGELOW, H. B. ':ia carolinensis McCrady, Pennaria tiarella 1913. Medusae and Siphonophorae collected by the U. cCrady, and Oladonema mayeri Perkins among S. Fisheries steamer Albatr088 in the northwestern the Anthomedusae) appear on both lists. In Pacific, 1906. Proc. U: S. Nat. Mus. 44 (1946): ~e part this is due to the fact that many 1-119, 6 pIs., 2 text figs. hYdroids do not liberate free-swimming medusae, 1918. Some Medusae and Siphonophorae from the west­ ern Atlantic. Bull. Mus. Compo Zoo!., Harvard Col­ hOod that many medusae have.very much reduced lege, 62 (8): 365-442, 8 pIs. Ydroid stages. Indeed, in many instances the 1919. Hydromedusae, siphonophores, and otenophores hYdroid stage is completely unknown, if it exists. of the Albatro88 Philippine Expedition. Bull. 100, Iosofar as it has been possible to determine U. 8. Nat. Mus. 1 (5): 279-362, pis. 39-43. ~ll1y 1938. Plankton of the Bermuda Oceanographic Expedi­ about 70 species have been recorded (30 tions. VIII. Medusae taken during the years 1929 nthomedusae, 34 Leptomedusae, 5 Trachy­ and 1930. Zoologica, New York, 23 (2): 99-189, 23 medusae, and 3 Narcomedusae) from the Gulf text figs. of Mexico region. Some of these, not already 1940. Eastern Paoific Expeditions of the New York ~ntioned above, are good species also known Zoological Society XX. Medusae of the Templeton e eWhere: Crocker and Eastern Pacific Zaca.Expeditions, 1936­ 1938. Zoologica, New York, 25 (3): 281-321, text S Corllmorpha nutan8 Hartlaub j H ybocodon forbuii Mayer; figs. 1-20. ~tBia mirabiliB L. Ag8BBiz; Ectopleura minerva Mayer; KRAMP, P. L. ( ~purena ophiogaater Haeckel; ZancleopsiB dichotoma 19HI. Medusae. Pt. 1. Leptomedusae. Danish In­ II'yetj Amphinema dinema P~ron et LeSueur; A. octaedra golf Expedition 5 (8): 111 pp., 5 pis., 14 charts, 17 lIio~ckel; A. rug08a Mayer; A. turrida Mayerj M erga text figs. t IQcea Ag8BBiz and Mayer; Podocoryne minuta Trinci; 1920. Anthomedusae and Leptomedusae. Rept. 80i. 1(lIt":,,.orea alexandri Mayer; Bougainvillia niobe Mayer; Res. Michael BarB Exped., 1910, Zoo!. 3' (2): 13 pp., 4Uikerina elegan8 Mayer; Prob08cidactl/IG ornatG Mc- 1 pI., 6 text figs. 273 274 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

KRAMP, P. L.-Continued KRAMP, P. L.-Continued 1926. Medusae. Pt. 2. Anthomedusae. Danish In­ 1947. Medusae. Pt. III. Trachylina and ScyphoZOIi golf Exped. 5 (10): 102 pp., 2 pis. with geographical remarks on all the medusae of tbe 1930. Hydromedusae co,llected in the southwestern part northern Atlantic. Danish Ingol! Exped. 5 (14): 66 of the North Sea and in the eastern part of the channel pp., 6 pis., 20 text figs. in 1903-1914. Mem. Mus. Roy. d'Hist. Nat. Belgi­ 1948. Trachymedusae and Narcomedusae from the que No. 45: 55 pp. Michael Sars North Atlantic Deep-Sea Expeditioll 1932. A revision of the medusae belonging to the family with additions on Anthomedusae, Leptomedusae, and Mitrocomidae. Vidensk. Medd. Dansk. Naturh. Scyphomedusae. Rept. Sci. Res. Michael SaTs Nortb Forening 92: 305-383, pI. 10, 52 text figs. Atlantic Deep-Sea Exped., 1910, 5 (1): 23 pp., 1 pl., 1933. Craspedote Medusen. Teil 3. Leptomedusen. 7 maps. Nordisches Plankton Zoo!. 6, Coelenterata: 541-602, MAYER, A. G. text figs. 1-68. 1900. Some medusae from the ToTtugas, Florida. Bull MUS· 1939. Occasional notes on Coelenterata. III. A. On the Compo Zoo!., Harvard College, 37 (2): 82 pp., 44 piS. systematical position of the Williidae. B. Genus 1904. Medusae of the Bahamas. Mus. Brooklyn Ins!' Koellikerina nom. nov. C. Tentacles on the manu­ Arts and Sci., Mem. Nat. Sci. 1 (1): 1-33, 7 pis. brium of SaTsia. D. On two new species of hydroids 1910. Medusae of the world. Carnegie Inst Washing­ from the Kara Sea. Vidensk. Medd. Dansk. Naturh. ton Pub. 109, 3 vols., 735 pp., 76 pis., 428 text figs· Forening 103: 503-516, 9 text figs. RANSON, G. 1942. The Goothaab Expedition 1920. Medusae. 1936. Meduses provenant des campagnes du Prince Medd. om Gr~nland 81 (1): 1-168, 27 maps, 11 text Albert 1 or de Monaco. Res. Camp. Sci., MonaCO figs. 92: 248 pp., 2 pis. SIPHONOPHORES IN THE GULF OF MEXICO

By MARY SEARS, Woods Hole Oceanographic Institution

The siphonophores most often recorded from In the Gulf of Mexico, one might expect to find the Gulf of Mexico are the two large conspicuous possibly 50 other species of Calycophorae, which species with floats above the surface of the water, have been taken in the tropical Atlantic and per­ Physalia physalis L. and Velella· velella L. Pos­ haps as many more among the Physophorae, sibly Porpita umbella O. F. Muller 1 should also Rhizophysaliae, and Chondrophorae combined. be included with these (Whitten, Rosene, and Most of these species have been taken at one time Hedgpeth, 1950). As early as 1886, Fewkes wrote, or another in the tropical Atlantic and might be III have many new localities for this medusa expected to be carried by the currents into the [i. e., Velella] in the Gulf of Mexico." The Gulf of Mexico. The depth of the sill at the A.tlantis in the winter of 1951 sailed through entrances to the Caribbean and Gulf of Mexico SWarms of Physalia together with large quantities is sufficiently great to permit entry of even the of Velella some miles in extent (Stetson, personal species that live at considerable depths, a factor communication) out in the Gulf off the northwest which, for example, apparently prevents some Coast of Florida, and newspapers give frequent siphonophore species from entering the Mediter­ account of the contamination of west Florida ranean (Bigelow and Sears 1937). bathing beaches. In short, it would not be surprising to find any The smaller, more common species, however, one of the 140 or more siphonophore species, now have scarcely been noted in the Gulf except at a known, in the Gulf of Mexico. few localities around its periphery, chiefly at the l'ortugas (Mayer 1900) and in adjacent bodies of LITERATURE CITED Water such as the Straits of Florida (Bigelow 1918), the Caribbean (Fewkes 1889), and the Gulf BIGELOW. H. B. 1911a. Biscayan plankton collected during a cruise of Stream proper (Bigelow 1918; Fewkes, 1882, 1886, H. M. S. Research, 1900. XIII. The Siphonophora. 1889). These records are indicative that about Trans. Linn. Soc., London, Ser. 2, Zool., 10 (10): 25 of the better known species in all probability 337-358, pI. 28. OCcur in the Gulf of Mexico proper: 1911b. Reports on the scientific results of the Expedi­ tion to the Eastern Tropical Pacific, 1904-1905. Abyla carina Haeckel; Abylopsis letragona Otto; A. XXIII. The Siphonophorae. Mem. Mus. Compo eschscholtzii IIuxley; A galma okeni Eschscholtz; Zool., Harvard College, 38: 171-402,32 pIs. Amphicaryon acaule Chun; Bassia bassensis Quoy 1913. Medusae and Siphonophorac collected by the and Gaimard; Ceratocymba sagittata Quoy and U. S. Fisheries Steamer Albatross in the Northwestern Gaimard; Chelophyes appendiculata Eschscholtz; Pacific, 1906. Proc. U. S. Nat. Mus, 44: 1-119, pIs. Diphyes bojani Chun; Diphyes dispar Chamisso and Eysenhardt; Enneagonum hyalinum Quoy and 1-6. Gaimard; Eudoxoides spiralis Bigelow; Galetla 1918. Some Medusae and Siphonophorae from the australis Quoy and Gaimard; Hippopodius hip~ Wcstern Atlantic. Bull. Mus. Compo Zool., Harvard papua ForskAI; Lenaia fowleri Bigelow; Rhizophysa College, 62 (8): 365-442, 8 pIs. eysenhardti Gegenbaur; Rhizophysa filiformi8 1919. Hydromedusae, siphonophores, and ctenophores Forskll.1; Sphaeronectes truncata Will; Stephanomia of the Albatross Philippine Expedition. Bull. U. S. rubra Vogt; Sulculeolaria monoica Chun; Sulcule~ Nat. Mus., No. 100, 1 (5): 279-362, pis. 39-43. olaria quadridentata Quoy and Gaimard; Vogtia 1931. Siphonophorae from the Arcturus Oceanographic glabra Bigelow; Vogtia pentacantha Kollikcr (as Expedition. Zoologica, N. Y., 8 (11): 525-592, text they are now named).' figs. 185-220. ----- BIGELOW, H. R, and M. SE,6.RS. I 8t>e Digelow and Scars (1937) for U80 01 this name. 1937. H. 2. Siphonophorae. Rept. Danish Ocean. Int Relerences which arc especially helplul and readily accessible In estahlish. Exped., 1908-10, to the Medit-erranean and adjacent ~ g the acceptcd names are; DJgelow, 19Ua, 191Jb, 1913, 1918, 1919, 1931; 1ge1owand Scars. 1937; Bcars, In press; Totton, 1932, 1941, in press. seas, BioI. 2: 1-144. 83 text figs.

251Hi84 0-54-·-10 275 276 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

FEWKES, J. W. SEARS, MARY 1882. No.7. Explorations of the surface fauna of the 1953. Notes on siphonophores. 2. A reVISIon of the Gull Stream, under the auspices of the U. S. Coast Abylinae. Bull. Mus. Comp. Zool., Harvard College, Survey ... 1. Notes on acalephs from the Tortugas, 109 (1): 1-119, 29 text figs. with a description of new genera and species. Bull. Mus. Compo Zool., Harvard College, 9: 251-289,7pls. TOTTON, A. K. 1886. Report on the medusae collected by U. S. Fish 1932. Siphonophora. Brit. Mus. (N. H.) Great Bar­ Commi88ion Steamer Albatro88 in the region of the rier Reef Expedition 1928-29, Sci. Repts.4 (10): 317­ Gulf Stream in 1883-84. Ann. Rept. Comm. Fish 374, 36 text figs. and Fish. for 1884: 927-980, 10 pIs. 1941. New species of the siphonophoran genus Lenaia 1889. Report on the medusae collected by the U. S. Totton, 1932. Ann. Mag. Nat. Hist., Ser. 11. 8: Fish Commission Steamer Albatro88 in the region of 145-168, 29 text figs. the Gulf Stream in 1885 and 1886. Ann. Rept. WHITTEN, H. L., ROSENE, H. F., and HEDGPETH, J. W. Comm. Fish and Fish. for 1886: 513-536, 1 pl. 1950. The invertebrate fauna of Texas coast jetties: ,. MAYER, A. G. preliminary survey. Pub. lnst. Mar. Sci. 1 (2): 1900. Some medusae from the Tortugas, Florida. Bull. 53-86, 1 pl., 4 text figs. Mus. Comp. Zool., Harvard College, 37 (2): 82 pp., 44 pis. SCYPHOZOA

By JOEL W. HEDGPETH, Scripps In.rtitution ojOceano,raphy

With the exception of Mayer's studies at Tor,.. lornetra quinquecirrha, a semaeostome medusa' ~ugas nearly ~Q y,ears ago there has beenno serious This medusa may cause a mild rash or unpleasant attempt to study· the medusae of the Gulf of sting/ but severe cases of jellyfish poisoning by Mexico. While there are.a few.scatter.ed records this species are rarelyreported. (Hedgpeth 1945). of medusae along the coast,. little is· available on An occasional denizen of bay waters is the moon the occurrence of jellyfish in the pelagic 'regions of jelly, 4ureUia aurita. While it is usually more a the Gulf or of tne deep-wate'rforins.: frequenter of the lower bays and gulf waters, it There are a few large medusae which charac­ at times outnumbers Dactylometra in the bays. teristically 'occur in the neriticwa~ers of the Gulf Another large Medusa in Gulf waters is the of Mexico from. Florida to;the Rio Grande. 'Fore­ lion's mane, Cyanaea capillaUJ, var. ·versicolor. most of these is StomolophttS meleagris,.the cab­ Mayer gives ~he southern limit of this variety as bagehead. This rhizostome. oftj3noccurs in vast Cape Canaveral, FloridaI and did not find it at numbers in lower bays and around passes at the Tortugas. R. O. Christenson, of Alabama Poly­ end of the summer. Such swarms were observed technic Institute, identified this medusa from at Port Aransas, Texas, on August 4-5, 1947, and Mobile Bay, on February 20, 1938. A large, September 20, 1948. The latter swarm was esti­ reddish, striped medusa was observed on an out­ mated to be drifting through the channel on an going tide at Port Aransas, Texas, on March 16, incoming tide at the rate of 2 milli~n an hour. 1948, which appeared to be this species. It is of 1'he bobbing white domes of these jellyfish seemed interest to note that both these recor.ds are to­ to be packed almost solidly across the 800-yard ward the p.nd of the Gulf coast winter. Burken­ width of the channel. Occasionally, StomolophttS road (personal communication) considers it com­ is caught by the ton in shrimp trawls, and there mon in Louisiana waters. has been some speculation about a possible eco­ According to Mayer, some 75 species of medu­ nomic use for these animals. Mayer mentions sae occur at Tortugas. Nine of these are scypho­ Several oriental species which are pickled, but such zoans, and the great majority are hydromedusae, a specialty delicatessen use would hardly cut into more pr~perly considered under hydroids (p. 267). the unwanted surplus. With the possible excep­ There are probably more species to be found in tion of a few hypersensitive individuals, Stomolo­ the Gulf. Following is a list of the Scyphomedu­ phus is not a dangerous species to bathers and may sae known to occur in the Gulf of Mexico, prin­ be handled with impunity. It is a fine animal for cipally at Tortugas. This list includes the 9 PhYsiological experimentation. species discussed by Mayer anc:I the 2 species Another rhizostome, Rhopilema verrilli, may be whose occurrence in the Gulf was not known to more common than suspected. Specimens have him; 2 been taken from Mobile Bay, and one was col­ lected at Port Aransas. This is a much larger Carybdaea aurifera Mayer, pp. 510, fig. 328. llledusa than StomolophttS, and Mayer suggested A rare form taken only twice at Tortugas. that it is a southern form which occasionally I The 1'0rtugue!!8 Man-or·War, PIIw,azla, Is more dangerous and probably e'ttends as far north as Long Island. Burkenroad causes some dlstr688 to unwary bathers every 96890n. 'I'blB slpbonophore (personal communication) considers it common in Is olten washed up on the outler boachea In groat numbers, and Ita atranded poeumatophores are heard poppIng under the wheels by motorists drIving Chandeleur Sound. on the beach. Two amallor slphonophoros, PorpUa linneana and Veld/a One of the most common jellyfish in the bays, Del/ela. are often stranded on the beach. I All page and ftgU1'O numbers given alter the name 01 the species .eler to especially during the summer months, is Dacty- Mayer's pUblication, 1910. 277 278 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Nausithoe punctata Kalliker, pp. 554-556, pI. 60, figs. 4-5, Cassiopea xamachana R. P. Bigelow, pp. 641-646, pis. text fig. 352.1 69-72. C. frondosa Lamarck. Pp. 647-648, pI. 69 This medusae is noteworthy for the peculiar, branched and 72. scyphistoma which lives commensally in sponges. These interesting medusae are the subject of several Found in all tropical and warm seas. papers in the Tortugas Laboratory series. The first is known from Tortugas and Jamaica; the second is Linuche unguiculata Eschscholtz, pp. 558-559, pI. 59, figs. more widely distributed throughout the West Indies. 1-10. Rhopilema verrillii (Fewkes), PP. 707-709, pI. 7, fig. 1, A West Indian species. Forms vast swarms in spring text tfg. 424. in the Florida-Bahama region according to Mayer. From New Haven to Port Aransas, but not (7) Dactylometra quinquecirrha L. Agassiz, pp. 585-588, pls. Tortugas. 62-64a, .text figs. 370-372. Stomolophus meleagris L. Agassiz, pp. 710-711, pis. 75-76. Widely distributed from New England to the tropics; Abundant along southern Atlantic and Gulf of MexicO possibly also Pacific. Abundant in Tampa Bay in (not north of Chesapeake Bay), and West Indies. August (p. 586). AlBO on Pacific side of Isthmus, and north at least to San Diego. Cyanaea capillata var. versicolor L. Agassiz, pp. 6OG-601, pI. 65, figs. 1-2. LITERATURE CITED Westeru Atlantic and Gulf, south of Cape Hatteras. HEDGPETH, JOEL W. 1945. Reexamination of the adventure of the Lion's Aurellia aurita Lamarck, PP. 623-626, pI. 67, figs. 1-4, fig. Mane. Sci. Mont~y 60: 227-232. 4; 68. MAYER, A. G. In east American waters, common from Greenland to 1910. Medusae of the world. Carnegie Inst. Washing­ West Indies. World wide. ton Pub. 109, 3: 499-735.

I See footnote 2, page 277. ANTHOZOA:ALCYONARIAI

By FREDERICK M. BAYER, U. S. National MUleum

The Alcyonaria of the Gulf of Mexico 2 are little the two Siboga supplements (1935, 1937), adds to known. No systematic work treats them in the list of species known from the Tortugas area. detail, and the preparation of such an account Explorations in the Gulf of Mexico have not Inust await more extensive collections from' the been extensive, and collections are correspondingly entire region. Even papers mentioning occasional inadequate. The exploratory vessels, Albatross, Gulf species are few and, with perhaps two or Fish Hawk, Pelican, Blake, Bibb, and Bache have three exceptions, deal only with those found in the all made dredge hauls in the Gulf, but the records extreme sout.heastern part ",

TABLE I.-Geographical di8tribution of alcyonarian8 known from the Gulf of Mexico ~. Arctlc to Cape Cod. F. Sombrero Key to Tortugas Bank; Straits of J. Galveston to Veraoruz. · C. Cod to C. Hatteras. Florlda;~. W. coast of Cuba. J. Vora Crut to C. Catoche. g. C. Hatteras to C. Canaveral. a. C. Sable to Apalachee Bay. K. Ccntrsl Gulf Bastn. · Bermuda. H. Apalachee Bay to Galveston. L. West Indies. E. C. Camavcral to Sombrero Key. c. 101>-49\) Iathoms. M. Caribbean Sea. g. Low water to 10 fathoms. d. 500-9llll fathoms. N. Brazil. ( to-99 fathoms. 10. Kukenthalll1111. o. 1000 fathoms and deeper. 2' Bayer 111411. 11. Kllkenthal1ll24. x. No depth given. · Bayer 1U52. 12. Kunr.c 11116. 20. Stlasny 111Uc. ~. Blclschowsky 11l21l. 13. Moser 11121. 21. Stlasny 11141d. 6' Cary 1006. 14. de pourtl1les 1868. 22. Thomson 11127. · Cary 11118. 15. Wcss 1920. 23. Toeplltz 1920. ~. Detchmann 1930. 16. Stlasny 11135. 24. Verrill 1864. · Gordon 11125. 17. Stlasny 1937. 25. Verrill 1869. 8. lielJprJn 1800. 18.StJasny 11141a. 26. Verrill 1883. 9. Kllkcnthal1111l\. 111. Stlasny 11141b. 27. Verrill 1007. ------,....------,.------...,------West Indies to Arclle to Sombrero Key Gulf of Mexico Brad\ Species A B CDE FGH J K LMN ------1------ORDER TELESTACEA 'r \ 'I'ltLII8TIDA:& e b ...... " __ " ...•.•..••.• "'" . "'-" 'rel~:~ ~;~~in;ti,6i:iJ~::::::~::::::::::~:::::::::::::::::::~:::: ~::::: :::::: :::::: :::::: --\i-- g b ORDKR ALCYONACEA ~Idali ALCYOSIIDAJI: a occldcntalls, 2, 6•. _ _"'" b...... b b •..••• _•.... -._ .••.••..••••••• b ••.•••.•••.• ~~~iHif~~afo~~~l;~~~,~:~~~~~~:::::::~::::::::~::::::::::::::: :::::: :::~:: :::::: ::~:: ~ :::::: :::::: :::::: :::::: :::::: If :::::: :::::: t I Published with the permlaslon of the Secretary of the B~lthsonlan Instl· II Hne drawn (rom Cape Sable, Fla., due south to the coast 01 Cuba, and Utfon. another from Cape Snn Antonio, Cuba., to Cape Catoche, YucatAn. This QI For the purposes of thIs summary, the geographical boundaries of the delimitation Is purely arbitrary and docs not coincide with faunIstio bound. lUI 01 Mexico wlll btl taken to Include, In addition to tb~ usua1land masses, aries. 279 280 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

TABLE I.-Geographical distribution of alcyonarians known from the Gulf of Mexico-Continued

West Indle.. to Arctic to Sombrero Key Gulf of Mexico Brazil Species ABC DEFGHI r K L MN ------1---".------.-f------ORDn GORGONAOEA

SUBORDER SCLERAXONIA BRIARICIDAJ: ~ro~~\':=tt~~'l~'_~::::::::::::::::::::::::::::::::::::::::::::: :::::: :::::: :::::: __ ~ __ a a a b b b b ~~ig:g~~~:~~,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ ~~~.~~~ :~~~~~ ::~:: ~~~~~~ ::~:: a a b

8VBOBDJiR HOLAXONlA

ACANTBOOOBOllDAJ: Acanthogorgla 88per1l, 2, 6, 26•••. ••_._ ••_. __ .: ._. ••__• •• d 0 b,o XVBJClClDAJ: •• • • .__ b, c o b • • ._ b, c 0 c ____ . , __ •• •• • b,c - b S, b ._._. -. .____ 0 b ; • .__ a· _. __ •__ •' a b b • __ . . ._' a ._. __ •• _. ._••__ .__ a a o - • - • • ---._. _._ .._ b ~ . :::::: --boo :::::: :::::: :::::: ~: ~ :::::: :::::: C -.------li------.• ---.-. -..--- d ~----- :::::: C b :::::: :::::: :::::: ---b-- ---Ii-- -'-d-' :::::: -. . -_. __ . .--___ C --.--- •••-.- t· ----.- ---.-- -"'-- .----. --- ..---'b-- -..•.- ----. b ----.- --b-- ---b-.· :::::: :::::: , __ ,_, -.-•. - -•. -.- ~ :::::: :::::: :::::: :::::: :::::: --ii-- :::::: ::~::; =rr~v~r1n~~,66!~::::::::::::::::::::::::::::::::::::::: :::::: :::::: :::::: :::::: :::::: b,o ._._. •- • - ._ b,o 0 PLEXAUIlIDAE a a a a a a a a a a a a a a. a. a a a a a a a

OOROONUDAJ: a a b a a ...... a x :::::: :::::: :::::: :::::: :::::: --b-- a a a a __ ,_,_ ..__ ._ ._. •.. a ...... ______• a, b a

--b"-- .------.--._-.__ .•------.• ------.- a ._ -. • __ ------•• --.- a .--- ...-__.-.____ a - a a __ . .___ a a Pterogorgla cltrlna, 6, 11, 'ZT ••• __ •• __ •__ ._. -._. • ••• __ a a, ba ____ . . ._. . __ ._. __ . a Pterogorgh~ guadalupllWlls, 2 ••_. __ ••_. •__ ••• • • •• • a •• __ •• •• ._ ._._._ a

OOROONICLLlDAIC c ______...... __ b,o 0 ...... '.... c b 0 _.. --" ~lei£~~~6~~~~~~~~~~~~~~~~~~~~:~~~~~~~~~~~~~~~~~~ ~~~~~~ ~~~~~~ ~~~~~~ ::~:: ~~~~~~ b :::::: --b" :::::: :::::: :::::: b;o -T- ...:::. b

CBIlYBOOOIlOUDAIC c c d l§~~i:::::::::::::::m:::::::::::::: x ::::: :m::::::: mm ::m b GULF OF MEXICO 281

T A.BLE I.-Geographical distribution of alcyonariana known from the Gulf of Mezico-Continued

Arotlc to Sombrero Key . Gulf of Mexloo . West Indies to BI'lll:U .. Species A BCD E F GH 1 K L M N ------'--1------.-----, . SUBORDJ:R HOLAxoNiA-Contlnued

0 'T' :::::: :::::: "0" :::::: be, . :::::: ::::::

""_' _••••• "_'_' __ •••••_._.. J: 0 • __ ...: ~~Ji·f~·0'~:~~~:-~i~~~~i~~~-:~~~~:~,i-~~[, ~i~~, ::~:: ~~-[,~ ::~:: .•••.. """ .....••.••.• _..... d .••. _. """ ...... -. ------.------.-- .... _-- ...._------_ .. I__ ••, ••"._. _••__ •••••-'••••••_ 0 ••••_•••••.• A . I8IDIDAJ: . .' , , . canella eburnea, 2, 6, 26 ._ ...._•..._..•._••....•.•.__ .:_.__ .._. -'..: ...... •__ • __ •.._ ...__ . ' d ...:.. d o,d o,d o Order PENNATULAOEA a RJ:NlLLlDAJ: -.. ------_ .. -.. ---_ .... -_ ... -- -.- .. ------.. - ae~::u~~~,~':··:::::::::::::::::::::::::::::::::::::::: :::::: :::::: ...~ .. :::::: :::::: :::::: :::::: : a •..•••.•_... a a a rumCULlNIDAJ: F . IlDlcuJlna quadranguJarls, 2, 6, 26. __._._•.•.•.•.•..•.•._•..•_. 0 '0 d ._ •.•. c••••• """ """ 0 ""'_ •••••• e 0, d .•••_••_•••_ PROTOP'lILlDAJ: l'rotoptUum sp. ct. thomsonl, 2•••. __ ._ •.•.._•.••-'_ .•..-'.•.•.._. , •••..••..'••.•..•• _••••.•,._.••" ••_ ." ••_ 0 o _"'" ...•_•..,•.. ""_' .•.... UI4IlJ:LLUUDAJ: ' U .' d ••.••••••••, t1:::l:U~: f~=r:: :: ~::::;:::::::::::::::::::::::::::::::::::: '--i" d~ e :::::: :::::: :::::: :::::: :::::: :::::: :::::: :::::: : e . VmOULARIlDAJ:

\'lrguJarla m1rabllls, 2, 6...._•._•••••._.•••.••_•••.••••••••••: __ • J: ._••_, """ ••__ •••'_'" """ ••_._. •••••.• a ~------_--:.._-'-_..:.--:.---'---'-_..:.-_'----:..--'---'--'---..:.-

1'his table has been compiled from the literature on either hard or soft bottom. The few gorgonian and from collections in the U. S. National MUseum, species which have been investigated in regard to inCluding unpublished records from the Albatross, temperature tolerance (L. R. Cary, Papers from li'ishHawk, and Pelican expeditions. Published the Dept. of Marine Biology, Carnegie Institu­ locality records within the Gulf of Mexico ,as tion of Washington, v. 12, No.9, 1918) can with" defined above have been located' for 72 Speciesi stand from 5° to 9° C. (approximately) above the records of only 9 species from Gulf localities exclu­ average maximum surface temperature of the area sive of the lower Florida Keys and Tortugas have (at the Tortugas, about 29° C.), but it is unlikely been found. Another 19 species haV'e been added that colonies would establish or thrive outside of by records in the collections olthe U. S. National a rather limited temperature range. In the Museum, bringing the total number of'species to absence of experimental evidence, it is impossible 91. These species repreSent 18 familiesin40f the to state the limits of the salinity and oxygen 6'known 'orders.' , . . . variation which the alcyonarians can tolerate. ,, Although little is known of the physiology of A few species can live in situations where the the alcyonarians, it is clear that bottom condi­ salinity is occasionally somewhat reduced, but Fons, t,emperature, salinity, available oxygen, most, including the West Indian reef-dwelling ~nd sedimentation play important· parts in limit­ forms, are never found where appreciable dilution ltlg their distribution. Limits of' tolerance are regularly occurs. Certain species are limited, to aPparently quite' na~ow but not equally so for all outer reef situations, and oxygen may be the factors. A solid substrate providing satisfactory critical factor in such cases. As a rule, alcy­ Conditions for the attachment of larvae 'is almost onarians are not found in continuously muddy ~niversally required among all alcyonarian groups waters, but some can tolerate very muddy con" eJtcepting the pennatulids. A very few gorgona­ ditions for short periods. Cean species are able to live unattached, and The reef areas'of the Tortugas and lower Florida ~ number, especially of the families Chrysogorgi­ Keys support a typically West Indian gorgonian Idae and Isididae, can adapt themselves to live assemblage. The predominant families are the 282 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Plexauridae and the Gorgoniidae; while not all of The shallow-water gorgonian fauna of the the known West Indian members of these families northern Gulf of Mexico is clearly identical with have been recorded from this area, most are to be but discontinuous from that of the Carolina coast. expected. This community does not extend This interrupted distribution pattern has been northward undiminished for any appreciable dis­ pointed out by Deevey (Ecology, vol. 31, No.3, tance, although a few of the hardier species range pp. 334-367, 1950) for some hydroids and other about halfway up the Florida west coast. The invertebrates and is described for fishes in this scarcity of suitable reef-like situations along this volume (Rivas, p. 503). Deevey suggests thatre­ coast seems to account in part for their reduction duced temperature during periods of glaciation in numbers, ,and temperature may be of equal permitted continuity of the cool-:water fauna importance in limiting the northward distribution around south Florida, but it would seem fully as of shallow-water gorgonians. Antillogorgia acer­ plausible that this continuity existed when Flor­ osa, A. americana, and Pterogorgia anceps are ida was submerged and that subsequent dispersal characteristic reef forms which extend some dis­ around the peninsula has been prevented by a tance up the west coast of Florida, and they thermal barrier. Since apparently favorable situ­ probably occur wherever there is solid bottom ations exist all along the east coast of Florida, suitable for attachment and permanent support. the southward dispersal of these discontinuously The predominant West Indian genera of reef­ distributed gorgonians is probably not limited by dwelling gorgonians, PlexaureUa, Eunicea, Antil­ bottom conditions but by some other environ­ logorgia, Gorgonia, Pterogorgia, and Phyllogorgia, mental factor of which temperature seems to be are restricted to. the warm western Atlantic, the most likely. In any event, it can hardly be while a few, such as Pacifigorgia and Muricea, doubted that the present-day distribution re­ are most numerous on the Pacific coast of the flects a former continuity of the Gulf and Caro­ middle Americas, and at least one, Leptogorgia, lina faunas, but a satisfactory explanation must is found also in the eastern Atlantic, the Mediter­ await the study of some group with an extensiv'e ranean, east Africa, and the East Indies. fossil record. The alcyonarian fauna of the lower west coast Although its southern limit is not known, the of Florida is thus a decimated West Indian as­ shallow water temperate assemblage is probably semblage. To the northward it merges with and present along most of the Texas coast, somewhere soon, perhaps near Tampa, is replaced by a dis­ along the coast of Mexico mingling with and giving tinctly temperate fauna the predominant gorgo­ way to the hardier elements of the West Indian nians of which are Leptogorgia virg'lLlata and L. fauna which encroach upon" it from the south. setaeea (both of which are referable (Bayer 1952) At least one gorgonacean, Leptogorgia setacea, eS­ to Verrill's genus Eugorgia) , and Muricea pendula. tends as far south as the Guianas and Trinidad. These species are especially abundant along the The presence of actively growing coral reefs coast of the Carolinas and south perhaps to north­ at Veracruz and along the coast of Yucatan haS ern Florida; L. virgula!a extends north to New long been recognized, but the composition of their York in moderately deep water, but all three seem fauna is little known. Heilprin (1890) reports to be lacking from the lower east coast of Florida. only one species of gorgonian from the Veracruti The short-stemmed sea pansy, Renilla muUeri, is reefs and remarks ,that the vast gorgonian 8e& common in the northel'n Gulf and extends south­ gardens so typical of the Bermudas are lacking­ wa.rd to Bra.zil; it likewise occurs along the Pacific The Bingle species that he records,. Ple~U1'(J, Coa.st from Central America to Chile. It has not flexuosa, belongs to the West Indian fauna, and been recorded from the Atlantic coast of Nortb it seems likely that other West Indian specie;s America where the only species appears to be occur there. Heilprin notes further that XipM­ Renilla renijormis, the common long-stemmed sea gorgia (now Pterogorgia) anceps was found Bot pansy. The latter species occurs also in the Gulf Progreso, Yucatan, another record indicative of of Mexico with a variety e:x;tending south to the the West Indian fauna. The occurrence of the Straits of Magellan and another in California. West Indian reef species Gorgonia :flabeUum oll GULF OF MEXICO 283 the Texas coast (one unpublished reco~d in the BIBLIOGRAPHY 0. S. National Museum) needs to be verified. BAYER, FREDERICK M. In the deeper waters (lO-500 fathoms) of the 1949. Chalcogorgiinae, a new subfamily of Chrysogor~ Southeastern Gulf, practically all of the alcyo­ giidae (Coelenterata: Alcyonaria), and a description of ChalcO(lorgia pellucida, new genus and new species, narians are West Indian species belonging to from the Straits of Florida. Jour. Washington Acad. genera of wide distribution. The Gorgonellidae, Sci. 39 (7): 237-240, 1 fig. Chrysogorgiidae, Primnoidae, and M uriceidae re­ 1952. New western Atlantic records of octocorals pI,ace in predominance the plexaurids and gor­ (Coelenterata: Anthozoa), with descriptions of three goniids of very shallow water. Most of the species new species. Jour. Washington Acad. Sci. 42 (6): BIELSCHOWSKY, EVA. are widespread throughout the Antilles and prob­ 1929. Die Gorgonarien Westindiens. Kap. 6: Die. ably also in the Caribbean. From the occurrence Familie Gorgoniidae. ZooI. Jahrb. Supp. 16 (1): of such characteristic forms as Beb'ryce grandis 63-234, figs. 1-40, pIs. 2-5. '-- and Scleracis guadalupensis in the extreme north- CARY, LEWIS R. .ern Gulf, it is probably safe to assume that a 1906. A contribution to the fauna of the coast of Loui­ siana. Gulf BioI. Sta., Cameron, La. (Louisiana good proportion of the West Indian species are State Bd. of Agric. and Immig.), Bull. 6: 50-59. present throughout the Gulf of Mexico wherever 1918. The Gorgonaceae as a factor in the formation of bottom conditions permit. There is no evidence coral reefs. Carnegie lost. Washington Pub. 213: as to the composition of the slcyonarian fauna of 341-362, pis. 100-105. this bathymetric range in the western part of the DEICHMANN, ELISABETH. 1936. The Alcyonaria of the western' part of the At­ Gulf, and intensive collecting should be done in lantic Ocean. Mem. Mus. Compo ZooI. Harvard 53: that region to clarify the distribution patterns of 1-317, pis. 1-37. the West Indian species as they enter the Gulf of GORDON, ISABELLA. Mu~. \ 1925. Gorgonids from Curacao Island. Het Koninklijk Zoo!. Genootschab Leyden. Bijdragen tot de Dier­ The limited deep-sea dredging which has been· kunde 24: 15-24, pIs. 3-4. done in the Gulf of Mexico has resulted in very HEILPRIN, ANGELO. few alcyonarian records. The isidid gorgonian, 1890. The corals and coral reefs of the western waters ..1.canella eburnea, which was taken in depths of the Gulf of Mexico. Proc. Acad. Nat. Sci. Phila­ ~anging from less than 200 to above 950 fathoms delphia. 1890, 42: 303-316. KttKENTHAL, WILLY. 10 the Gulf of Mexico, is also known from the 1916. Die Gorgonarien Westindiens. Kap. 1, Die ~orthwestern Atlantic, the West Indies and Car­ Scleraxonier; Kap. 2, tiber den Venusfacher; Kap. Ibbean, the coast of Brazil, and the eastern Atlan­ 3, Die Gattung XiphifJoTfJia H. M. Edw. Zool. Ja.hrb. tic, always at considerable depths: Beyond the Supp. 11 (4): 444-503, 26 figs., pI. 23. 1,OOO-fathom contour, three pennatulid species 1919. Gorgonaria. Wissenscahftliche Ergebnisse der deutschen Tiefsee-Exped. auf dem Dampfer Valdivia ha"e been dredged: Umbellula guntheri, U. lin­ 1898-99, 13 (2): 1-946, 318 figs., pis. 30-89. dahlii, and Funiculina quadrangularis, all of which 1924. Gorgonaria. Das Tierreich 47: xxviii+478, 209 also occur at extreme depths in the northern and figs. Berlin u. Leipzig. eastern Atlantic. • KUNZE, G. There seems to be no truly endemic element in 1916. Die Gorgonarien Westindiens. Kap. 4.-Die Gllttung Eunicea Lamouroux; Kllp. 5.-Die Gattung the alcyonarian fauna of the Gulf of Mexico. Plexaurella. Zoo!. Jahrb. Supp. 11 (4): 505-586, 55 The strictly shallow·water forms of the northern figs., pIs. 24-28. half are also the predominant species along the MOSER, JOHANNES. Carolina-Georgia coast, while those of the south­ 1921. Ergebnisse einer Revision der Gattung Plexaura Ctn part are typically West Indian. The species Lamouroux. Zoo!. Anzeiger 53 (5/6): Uo-UB. ~ POURTALts, L. F., DE moderate depths throughout the Gulf a're 1863-1869. Contributions to the fauna of the Gulf est Indian, and a northern element does not Stream at great depths (2d series). Bull. Mus. Camp. ~})l>ear to be present. Finally, the character­ ZooJ. Harvard College 1 (6): 103-120, 1867; 1 (7): Istically deep-sea forms thus far known from the 121-142, 1868. ~f RIESS, MARGOT. are of wide distribution at similar depths 1929. Die Gorgonarien Westindiens. Kap. S.-Die t oughout the Atlantic and are possibly even Familie Muriceidae. Zoo!. Jahrb. Supp. 16 (2): cOSlnopolitan. 377-420, 4 figs., pi. 8. 284 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

8TIASNY, GUSTAV. TOEPLITZ, -CHARLOTTE M. 1935. Die Gorgonacea der Siboga-Expedition. Supple­ 1929. Die Gorgonarien Westindiens. Kap. 7.-Die ment 1, Revision der Plexauridae. Siboga-Exped. Familie Gorgonellidae, zugleich eine Revision. Zool. Monog. 13b7 : vi+l06, 27 figs., pis. 1-7. Jahrb. Supp. 16 (2): 235-376, 26 figs., pIs. 6-7. 1937. Die Gorgonacea der Siboga-Expedition. Supple­ VERRILL, ADDISON EMERY. ment 2, Revision der Scleraxonia mit Ausschluss der 1864. List of the polyps and corals sent by the MuseuJ'll Melitodidae und Coralliidae. Siboga-Exped: Monog. of .comparative Zoology to other institutions in eX­ 13bB: vi+ 138, 38 figs., pIs. 1-8. change, with annotations. Bull. Mus. Compo Zool., 194111.. Studien fiber Alcyonaria und Gorgonaria, II. Harvard College 1 (3): 29-60. _, _. Zool. Anzeiger 134(3/4): 53-71, 8 figs. 1869. Critical remarks on the halcyonoid polyps with 1941b. Studien fiber Alcyonaria und Gorgonaria, III. descriptions of new species in the museum of Yale Zool. Anzeiger 134(11/12): 254-268, 11 figs. College, No.4. Am. Jour. ScI. (2) 48: 419-429. 1941c. Studien fiber Alcyonaria und Gorgonaria, IV. 1883. Report on the Anthozoa, and on some additional Zool. Anzeiger 135(1/2): 13-25, 10 figs. species dredged by the Blake in 1877-79, and by the 1941d. Gorgonaria von Venezuela (Inseln Blanquilla U. S. Fish Commission steamer Fish Hawk in 1880-82. und Los Frailes). Arch. N&!rl. de Zool. 6 (1): 101­ Bull. Mus. Compo Zool., Harvard College 11 (l): 116, figs. 1-4, pis. 1-2. -1-72, pis. 1-8. THOMSON, J. ARTHUR. 1904-1907. The Bermuda Isiands. Part 5.-'-Charac­ 1927., Alcyonaires provenant des campagnes scientifiques teristic life of the Bermuda coral reefs. Trans. Con­ du Prince Albert Ier de Monaco. R~s. Camp. Scient. necticut AlJad. Sci. 12: 160-304, 28 pis. i index and Albert de MonA.Co 73: 1-77, pis. 1-6. errata,pp. 311-316. ANTHOZOA: THE ANEMONES

By JOEL W. HEDGPETH, Scripps Institution of Oceano,raphy, University of California There is as yet no systematic study of the collections reported in this work indicate a mixture anemones of the entire Gulf of Mexico including' of tropical, West Indian forms and species of the the Tortugas region. The papers of McMurrich, Middle Atlantic coast. Of particular interest especially 1889, and Watzl (1922) on Bahamas is the finding of Aiptasiomorphia luciae at Port actinians, together with Duerden's (1902) report Aransas, adding yet another locality for that on Porto Rican species, are useful aids to the ubjquitous species. The accompanying table, study of the Tortugas anemone fauna which (table 1), compiled principally from the literature, apparently is about the same as that of the indicates the affinities of the cornmon species Bahamas. For the Gulf of Mexico proper there found at Bahamas and Tortugas. This is sup­ is only the recent paper by Carlgren and Hedgpeth plemented by brief notes on some of 'the more (1952) on species from Texas and Louisiana. The interesting forms.

TABLE L-Synop8i8 and known di8tribution of anemone8 in Bahamas, Tortuga8, and the Gulf of Mexico [CompUed from the literature; .ynonymy (In parentheses). after Oarlll'lln, 11149)

Gulf of Mexloo

Species WoodB Beau· Ber· Baha· West Depth Remarks Hole fort muda Ibas Indies Torlu· Loul. gas (or .Iana Tems Key!') ~------II------I------I------Order CORALLIMORPHARIA

OOBALLUlOBPWDAB ~nodf' babmemll WatzL______X rdea jlMlda Duch. et Mlch. •• •• ._____ X ---coj--- :::::::: :::::::: :::::::: .~~~~ci:::::::::::: ·St. Thomuand.JamaIoa. (HderanUUI florldu.)

ACl'INODI8CIDAB J>lIradl.COIoma nel/lma (Duoh. ot Mich.). • ... X CO) •• .do••• °St. Thomu, Balti,' .Ja· malca. J>(radllCOIoma carlgrml CWatEI)-.-----.-- •• . __ ._... __ X ______._. ..do.__ ••__ •_ ~c=~~~:tr'coe:::Je (Duoh. et Mloh.). • __ • __ .____ XX C·) • • .do•• ·.Jamalca. A.ctlnothrlz .andl 16omllt) Order AOTINIARIA; Suborder ENDOCOELANTBBAB HALCUBUDAB lillleurllll ¢lotUi McMurrlch __ ._ .. •• -.------_ X 100 fatholllB •__ Suborder NYNANTBBAB

AuCIlDAB tebrCLunl1l danae (Ouoh. et Mlch.)••• ------_ X X (0) X Shore °O~80, 8t. Thomu, . lUl/llcta: a-adactl, l>arlabllll) .Jamaica. AcmnIDAI: "'~~Ia bermudemll cMcMurr
ch)-.------XX (0) •• __ •• • _ ~~~*:~:~c:::::::::::::: !lemonlll.arl/auemll Hargltl .____ ::::::::XX:::::::: -':x--- .______i :::::::: :::::::: :::::::: ::::::::X ::::::::::::::::::::Pelagia, on BIU" " ' PlllIUID. f;jtlodOBoma clWernllta (Bosc)------. X ::______X. .__ XX __ ,tCrJc:::m=)(McMurrlch)•••••-.-. ---•••-- .-•••--•••--•••- .--..... ------X .------+100fathomB••• __ :.~ra Icrebal (Duoh. et Mloh.)••---- • __ • _. "_" ,__ CO) , ••_ X Shore _ St. Thomas, .Jamaica. COiid pleura rarfllllrmllta Watd. ._. . ._.___ X -- -. ------_. __ .do._••_. _ lj(o.r=~J::)'" CWetland?------~, X, XX ------. ----..------~~~~1~~~~~OMurrIOh)---.-... .___ XX (0) __ C~,.Jamaica.

285 286 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

T ABLE I.-Synopsis and known distribution of anemones in Bahamas, Tortugas, and the Gulf of M exico---Contillued

Gulf of Mexico

Species Woods Beau· Ber· Baha· West Depth Remarks Hole fort muda mas Indies Tortu· Loui. l(as (or slana Texas Keys) ------1------1------1------ORDD CORALUMORPHARIA-Con. ACTINIIDAE-Continued Phl/U4df&jlOlcull!tra (LeSueur) • ._ XX (0) St. Thomas. (A,ltractf& !lOlculi!tra; Oulactf& falcicu- lata MeM.) PAI/U4c1i, conquilegla (Ouch. et Mleh.)_••. •• .••• • X (AI/tradf& upama Ouerden) MINYADIDAE Minllal olirocea (LeSueur)•• _•••• ._•• _ X x x Pelagic.. _ STOICBAlmIDAE Slolclladf&hdia7ltAua (EllIs). • __ •._ x XX (Df&eoIoma anemone) PBYKANTHIDAB Phl/manthua crucifer (Le 8ueur) •• _••_ X X X ACTlNOSTOLIDAB Para7ltAua rapiformf& (LeSueur) New x X X Shallow water (AmmopAilactf& raplformf&) Haven +10 faths.? Subtr. AconUaria ISOPHELUDAE Telmatactf&roaeni (Watz1) -______X (PAeUia rOlen!) HORHATHDDAE Actinauge langieornf& Verrill •. X X x X 100-250 faths ._ CaUiactf&lrieolor (Le Sueur) •_ X X ? X Shore-l0/20 laths__ SlepAenauge ,pongleola (Verrill) .______X X 100 faths ._ (Sagarlia ,pangieola) AIPTASnDAE Aip/lUiapaUlda (Verrill)______X ? X X Aiplcuialagetea (Ouch. et Mleh.)•• __ ._ •• _. .. _. __ .___ X BorlholomDiadumene leucolena (Verrill)______X X Shore ._ (Sagari/a Uucolena) Order CERIANTHIARIA • Cerianlhlapril ammcanua VerriIL . __ •. x Shallow W(lu-r to about 10 laths. Cnianthromorp1le brlUUremf& (Carlgren).• •. __ .•• .• .• __ . 1(}-14 laths. Brazll._New Mexico ------_._------'------NOTE: The fOllowing spl'Cles, SO far known only from the northwestern Gull 01 Mexico, havo recently boon described by Carlgren and Hedgpl'th (1952): Andwaklldae, Andwakla f&abel/ae; Actinlidae, Bunodactl, /aae...i,; AiptllSiomorphldae. Alp/alimorpho /aaen,I,; 8agartlidae?, Bolrl/onlubercu[alua; zoan­ tharia, Zoantnidae. Pall/thaa luatmf&.

Of all the animals living in the sea the anemones of various parts of each species. Each marine are at the same time among the most beautiful and laboratory or station should compile a set of color most difficult to study. A sound basis for the photographs, camera lucida drawings of the study of anemones is a detailed series of notes on nematocysts (under oil immersion) making ~p the living animals including color sketches or the cnidom, and serial sections for each species If photographs, measurements, and descriptions of its fauna. A collection of huddled lumps 0 the nematocysts, and a set of well-prepared slides coelenterate flesh is almost useless to all but the GULF OF .MEXICO 287 lllost thorough-going specialist in this group, and Bunodactis texaensis Carillren and Hedllpeth. in the absence of satisfactory material our knowl­ A conspicuous gray anemone, superficially edge of Gulf coast actinians will remain in its resembling Bunodosoma cavernata, but with ver­ present fragmentary state. rucae instead 'of vesicles on the column and a pattern of darker gray or greenish to light brown NOTES ON COMMON SPECIES splotches on the disc (see color plate, Calgren and LUbrunia danae (Duchassalnll and MIchelotti). Hedgpeth). It occurs on the jetties at Galveston This anemone is conspicuous for the large, and offshore near Port Aransas. branched outgrowths ("pseudotentacles" or Minyas olit/acea (LeSueur.) "fronds") at the top of the column just below the A pelagic antillean species which occasionally tentacles. The animal is brownish and lives in drifts ashore on the coasts of Texas and Louisiana, hollows in coral rock. It is common at Tortugas sometimes in considerable numbers. The animal and was described from there by Hargitt (1911) is an olive brown color, with the tentacles ap­ as Oradactis variabilis. McClendon (1911), in a parently reduced to knoblike processes. The Paper on habits.of several invertebrates, provides animal remains at the surface by means of a float a color plate. in the pedal disc. According to observations of M. D. Burkenroad, Minyas will shed its float in ~nemonia sargassensis Hariitt. an aquarium, but does not produce.a new one A well characterized anemone both in habit and under these conditions. It may be that the appearance. Originally described from sargassum mature Minyas (as yet unknown) is a sessile form. drifting into Woods Hole, it is found on that Condylactis gigantea (Welnland). plant as it drifts ashore along the Texas coast and The "passion flower" anemone is common at is recorded from Beaufort by Field (1949). It is a Tortugas, the Bahamas, Miami, and various pla~es BlUaH, rather squat, velvet-brown species. 1 The. in the West Indies. The color of the column varlCS tentacles may be tinted green and are occasionally from bright scarlet to brownish, ele tentacles are branched (fig. 60). brownish or paler than the column and usually Bunodosoma cavernata (Bose). tipped with scarlet. The common jetty form of the Texas coast, Stoichactis helianthus (Ellis). especially at Port Aransas and Port Isabel. Cary The "sun flower" anemone is a characteristic (1906) found it common on the Cameron jetties. West Indian species common in the Bahamas and It was originally described from the Carolina coast at Tortugas. It is easily identified by the broad, and is a characteristic member of the Beaufort incompletely retractile disc with its large number fauna. It is a muddy to dull brown colored anem­ of short, stubby tentacles. The disc is greenish or one with pearl gray vesicles on the column, with with green patches, and the peristome is usually reddish to brownishor bluish tentacles, but usually bright yellow. The tentacles are greenish to With a red stripe on the back of the larger tentacles yellow. (fig. 60). Some specimens are entirely cherry red. Paranthus rapi/ormis (LeSueur). 'rhe West Indian B. granulifera is considered to be A characteristic member of the shallow water ll. synonym of this species (Carlgren 1952). bottom assemblage along the Texas coast. Cary ~nthopleura krebsi (Duchassainll and Michelotti). found it washed ashore along the Cameron beach. Previously known from St. Thomas and Ja­ It is found off Beaufort and along the coast north­ JUaica, a colony of small individuals occurs on the ward to New Haven. Although this species has Port Isabel jetties. The column and tentacles are a moderately well developed basal disc, it is a White, with rows of bright red velTucae which are bUlTowing form. 'fhe column is whitish, the disc larger and more regular toward the top of the green with faint salmon markings (fig. 60). Speci­ column (fig. 60). mens brought on deck in a trawl or dredge con­ tract to a spherical shape resembling peeled onions. I M. D. Burkenroad contends (In IItterls) that there is another common sPecies on Gulf Sarq/U,um, smaller than Aflemonfa, and reproduolng com· Calliactis tricolor (LeSueur). tnonly hy longitudinalllssion. Carlgren (in litterla) thinks it posslhle that Common on the shells of the gastropod Rehderia this may be a Bundeop8l,. This problem cannot be oIarlfied until the anemones 01 the Atlantic Sargassum are critically studied. off the southern Atlantic coast and sometimes 288 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Bunodosoma cavernata

..' Anthopleura krebsi

.... .i :;

.J

FlOUR. 6O.-Anemonee from the northern coaet of the Gulf of Mexico. GULF OF MEXICO 289

pa 21 ida.

FIOUIlIl 61.-Anemonea from the northern ooast of the Gulf of Mexico. abundant on hermit crab shells and oxystome crab are bright orange; the tentacles are pinkish to red carapaces (fig. 61) in comparatively shallow water with darker bandings. The directives are often along the Texas coast. Occasionally, a specimen more deeply colored than the other tentacles. This is found living in shallow water attached to some is a common West Indian species. immobile substrate. The column is whitish, AipttuiG pollidG (Verrill). cream colored or brownish with darker vertical According to Cary, this species (fig. 61) was stripes and darker brown spots marking the common on the jetties at Cameron in 1906; I cinclides at the base of the column; the acontia have not seen it on Texas jetties but found it 290 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE commonly on oysters near Port Isabel. It is BIBLIOGRAPHY usually rich brown with darker stripes. The CARLGREN, OSKAR tentacles may be solid brown, or whitish. The 1949. A survey of the Ptychodactiaria, Corallimorpharia brown color is due mainly to zooxanthellae. Off­ and Actiniaria. K. sven ka vetensk. Handl. (4), 1 shore, in about 10 fathoms, there is a pale powder (1), 121 pp., 4 pIs. 1952. Actinaria from North America. Arkiv for Zool., blue phase found in old Pinna shells. The species 3 (30): 373-390, 10 figs. is known from the Carolina coast, especially at ---, and HEDGPETH, JOEL W. Beaufort, where it occurs in large colonies (Carl­ 1952.' Actinaria, Zoantharia and Ceriantharia frorn gren 1952). shallow water in the northwestern Gulf of Mexico. Bortholomeo onnulato (LeSueur). Pub. Inst. Mar. Sci. Texas, 2 (2): 143-172, 9 figs., 4 The tentacles appear to be ringed because of the color plates. annular swellings or incomplete bands of nemato­ CARY, L. R. cyst batteries. The column is whitish at the base, 1906. A contribution to the fauna of the coast of Louisi­ ana. Gulf Biologic Sta. Bull. 6: 50--59. (p. 51). darkening to brown near the top. The tentacles are brown. The species is found in Bermuda, CONKLIN, EDWIN G. 1908. Two peculiar actinian larvae from Tortugas, Bahamas, and the West Indies, and a specimen Florida. Papers Tortugas Lab. 2: 171-186, 4 pIs., 5 has been collected at Port Isabel, Texas. figs.

Aiptasiomorpha lucioe (Verrill). DUERDEN, J. E. Easily identifiable by its olive green column 1902. Report on the actinians of Porto Rico. U. S. with orange vertical stripes (when not in a color­ Fish. Comrn. Bull. 20 (2): 321-374, 12 pis. less phase), this little anemone (fig. 60) is almost FIELD, LOUISE RANDALL cosmopolitan. It was first observed by Verrill at 1949. Sea anemones and corals of Beaufort, North Caro­ Woods Hole but may have spread originally from lina. Duke Univ. Marine Sta. Bull. 5: 1-39., 94 figs. [Cuutlon: The corrections made by Carlgrcn (1952) and CnrlKrcn nnd Japan. It is found on the Pacific coast of North Hedgpeth (1952) concern only the nnemone section.] America, at various places in Europe, and now HARGITT, CHARLES W. from the Texas coast (Port Aransas). 1911. Cradactis variabilis: an apparently new TortugaB Aiptosimorpho texaensu Carlgren and Hedgpeth. actinian. Papers Tortugas Lab. 3: 51-53, 1 pI. \ A small, salmon pink to whitish species locall.v MCCLEND"ON, .J. F. common in bays of Texas and Louisiana. It has 1911. On adaptations in structure and habits of some been recorded from salinities as low as 9 parts per marine animals of Tortugas, Florida. Paperll TorW­ thousand, and seems to be an estuarine species. gas Lab. 3: 55-62, 2 pis. It is usually found on oysters and piling. McMuRRICH, J. PLAYFAIR Cerwntheomorphe brasiliensis Carlgren. 1889. The actiniaria of the Bahama Islands, W. 1. Jour. Morph. 3 (1), 80 pp., 4 piB. This cerianthid has been collected near the 1896. Notes on some actinians from the Bahama IslandS, coast of Texas and northeastern Mexico, and was collected by the late Dr. J. 1. Northrup. AnnBls originally described from San Sebastian, Brazil. New York Acad. Sci. 9: 181-194. Cary reported a colony of cerianthids from the 1898. Report on the actiniaria collected by the Baharna 225~ Chandeleur Islands, which may be this species, or Expedition of the State Univcrllity of Iowa, 4: 249,3 pis. possibly Cerianthiopsis americanus, which occurs at Beaufort. These large burrowing anemones are WATZL, OTTO 1922. Die Actiniarien der Bahamainseln. Auf Grund difficult to collect, and no specimen from the der Sarnmlung deB Hern Dr. N. Ros6n (1908-09)· Chandeleurs has come to notice. Arkiv fiir Zoo!. 14 (24),89 pp., I pI., 10 text figs. GULF OF MEXICO MADREPORARIAI

By F. G. WALTON SMITH, Marine Laboratory, University of Miami

By reason of the great difference in their normal found where vigorous water circulation exists habitat corals fall into two very distinct groups. The branching corals grow more readily in com­ The hermatypic or reef corals are usually, but not paratively still water than do the massive types. always, large and massive or branching in form. A few species such as Porites jurcata and Manicina They are usually associated with other corals in areolata may be found on muddy bottoms. bUilding considerable masses of living coral reef. Growth rates of corals at Dry Tortugas have The deep-water or ahermatypic corals, on the been measured by Vaughan. Non-porous species other hand, are usually small and solitary though grew at an average rate of 9.0 mm. in diameter sometimes branching in form. and 5.00 mm. in height per year. Porous species Hermatypic corals grow in water up to 50 fath­ increased at an annual rate of 40 mm. in diameter OIns in depth but are only active in reef building and 25.0 mm. in height. Montastrea annularis, in depths to 25 fathoms.· Most reef growth occurs a massive type, showed an annual increment in in less than 15 fathoms. Ahermatypic corals are weight of 54.8 percent, whereas, a branching coral, found mostly in deeper water from the edge of Acropora palmata, increased 194.9 percent in the continental slope to the neighborhood of 3,000 weight per annum. Both specimens were approxi­ fathoms. The majority live between 90 and 300 mately 100 grams at the beginning of the experi­ fathoms. The temperature range for reef corals ment. Growth of corals is greater at higher is approximately 19° C. to 36 0 C. (63° F. to 97° temperatures. Since temperatures in the Gulf of F.) with an average minimum, however, of 22° C. Mexico are generally close to the lower limit of (7.2° F.). Ahermatypic corals live best within a the range, reef growth is accordingly slower than range from about 8° C. to 21 0 C. in the West. in the warmer seas. Larvae of species found in Indian region. the Gulf of Mexieo have a planktonic life of be­ The distribution of reef corals in the Gulf of tween 1 and 3 weeks. Mexico and the relation of the coral fauna of this Winter temperatures in HIe Gulf of Mexico are area to those of neighboring areas is dependent close to the lower limit for vigorous reef growth. upon the physiological requirements of corals. There are therefore no strongly developed reefs 'fhese have been studied in detail and are dis­ except for those of the Dry Tortugas, the Florida Cussed by Vaughan in a series of papers describing Keys, and the Alacran and oth~r reefs of the experiments carried out at Dry Tortugas. Campeche Bank. Less vigorous r~ef develop­ The average optimum salinity for reef corals is ment is found at Veracruz and at a few other 36 parts per thousand, although a range of 27 to plac(~s within the warmer, more southerly waters 40 parts per thousand may be tolerated. Expo­ of the Gulf (fig. 62). SUre to air is also tolerated to a variable extent. Scattered coral heads which fail to form true SPecies with more porous skeletons are eonsidcr­ reef structures are found elsewhere in the Gulf of ably more resistant to exposure. Strong light is Mexico, particularly off the west coast of ~loridll. essential to vigorous growth. This is apparently Because of the amount of sediment present, they the result of the zooxanthellae which are normally are rarely found close inshore but usually at some Present in the tissues of reef corals. Corals are distance from the coas't in more than 5 fathoms of carnivorous in habit. water. Scattered heads are also found in deeper Reef corals do not, as a rule, withstand any water in a line running from south of New Orleans great amount of sediment and are accordingly toward the Texas coast and in another line run­ -I Contribution No. 100 from the Marlnr Labomtory. University of Miami. ning southward parallel to the Texas coast. Sur- 2591134 O-M-20 291 292 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

_. Co...l ....1'.

Alacran I ••r ~0;"·;· .$ •• · ~. Merida GUL' 0' MIXICO

JAIlUARY 180TIIJ:ltllll AIlD COUL DISTIlISUTIOJ POW!

FIG. 62-January isotherms and distribution of corals in the Gulf of Mexico prisingly few published records exist regarding Indies have been recorded from Florida. Only 11 the presence of reef corals in the Gulf of Mexico. of these are reported from Veracruz (Heilprin The Alacran reef, described by Agassiz (1878, 1890) . 1888), the Veracruz reef, described by Heilprin The Caribbean reef fauna consists of about 26 (1890), and the Dry Tortugas and Florida Keys genera and 51 species (table 1). The species reefs, described by Agassiz (1888), Vaughan, and listed here include a number which are undoubt­ others complete the list. From these, and various edly varieties or growth forms, such as Acropora unpublished sources, Joubin (1912) prepared a prolijera and certain species of Porite8. They are map of coral reefs of this area. Data concerning included here, however, since they are recognizably patches of scattered coral heads are also shown on different, and such a list is convenient to field the U. S. Hydrographic Office Bottom Sediment workers who are more concerned with accurate Charts 1125 BS and 1126 BS published in 1943. and speedy identification rather than the some­ All available records to date are indicated in the times debatable questions of taxonomy. For accompanying chart which shows approximate purposes of identification, the handbook of At­ locations only. lantic reef corals by Smith (1948) is most useful Except for reefs of the Florida Keys and Vera­ since it contains complete keys and descriptions cruz, published accounts of Gulf reefs are insuffi­ and is weIl illustrated. ~"or an authoritative cient to give an adequate list of species. The monograph on coral taxonomy, reference should fauna' is nevertheless typically West Indian. be made to Vaughan and Wells (1943) who pro­ Forty-two of the 51 species known to the West vide an extensive bibliography. GULF OF MEXICO 293

TABLE I.-Western Atlantic hermatypic specUls of corals Trochosmiliidae, Mussidae, and Caryophylliidae~ rp'lndlcates recorded from Florida; V indicates recorded from Veracruz] None of the Fungiidae are represented and only Madracis among the Seriatoporidae. Montipora, SUBOOlDER ABTROCOENIIDA Astreopora, Goniopora, and Trachyphyllia, all of FaroU)' ABTROCOENnDAE 1. A'lrocoenla pect/nata PourtalM •.• __ ••••_•••_•••.•••••_... F which exist as fossils of the Caribbean Neogene, 2. Stephanocomla mlc1lellnl Edwards and Halme•.•_•••••: ••••• are now absent. Genera known only to the West BERIATOPORIDAE • A;RO:::f;=~ decactll (Lymanl ..••••-•••••.• ~.~ •••••••-••••••--. F Indian fauna are ColpophyUia, Dendrogyra, Mean­ 4. Acropora emncornll (Lamarckl _ •• __ ._. __ F drina, Mussa, Mycetophyllia, Manicina, 1so­ 6. A. palmala (Lamarckl __ _ __ FV 6. A. pro/I/era (Lamarckl ._ __ •__ __ FV phyllia, 1sophyllastrea, Eusmilia, Dichocoenia, SUBORDER FUNOIIDA and Agaricia. All of the West Indian genera AOARICIIDAE occur on the Florida reefs. Their absence from 7. Agar/cia Cl9aricUu (Llnnaeusl •• _....__ •__ ....__ .... • F F other parts of the Gulf of Mexico must be ascribed ~: ~ll~ Qe~k======::::::::=: F 1: to the existence of unfavora.bletemperature BIDERAllTRBIDAE 10. ~Ullrea radlam (Pallasl ...... __ •__ ... F conditions which permit only the more hardy 11. S.,i1derea (Ellis and Bolanderl • , __ .. FV 12. S. •Iellota Verrill.•• __ __ _ to live there since their presence on the Florida PORITIDAE reefs, their known length of larval life, and the 13. PorUu Ullreoldu Lamarck ••• •••••~ .•__ ._.______FV 14. P. branlUrl Rathbun.__ ...... __ ..., __ . existence of fa.vorable currents are sufficient for -16. P. dlDar/eala LeBueur .._.. .. : F 16. P./ureala Lamarck. __ __ .. FV dispersal throughout the area. 17. P. porile, Wal168I ...--..------.--••------F 18..P. Derrllll ebbefll..__ • __ ...__ ...__ .. · The ahermatypic corals are much less restricted SYBORDER FAVIIDA by geographical boundaries than the hermatypic FAVDDAE forms since the required conditions for their 19. FaDla conferta VerrilL...... __ • .. •_ F existence are widespread throughout the deeper ~: ~~£rili::::::::::::::::::::::::::::::::::::::: waters of the ocean. The extent of distribution 23. f:Dlp~~iDoIaE:fJ: (Ellis and Bolanderl. .. F 24. D. laby,r'nlhlformll (Llnnoousl _.. __ __ FV of any species depends partly upon the depth' 26. D. ,Ir go,a (Danal. __ • __ ..______FV 26. Col~PhVUla amaranlhUl (Mullerl. .. F range inasmuch as the deeper the normal habitat, 71. C. alam (Mullerl __ .... _.....__ __ .._••_.... F 28. ManleillCl areolata (Llnnaeus} __ __ • .. F the greater the continuity of suitable environ­ 29. M. mCIVor' Wells ~ __ .. ~_ .._ •• F 30. Cladocora arbUlcu.la LeSueur .. ..__ • __ •.• FV ments. On account of the generalized distribution 31. SoltllClllrea baurnonl Edwards and Halme__ _: __ F 32. S. /lJIadu (Dana}...__ ...... __ .. ; ______•F of ahermatypic corals and the lack of adequate 33. MonlClllrea annularll (Ellis and Solanderl. •• FV 34. M. brazU/allCl (VerrUlI • __ .. __ surveys of the greater part of the deeper waters 36. M. caDernCla (Llnnaeusl __ __ ... FV of the Gulf of Mexico, it has seemed more useful ABTRANOnDAE 36. Altrang'a .0lUarla (LeSueurl ..__ ...__ ...... •• F to provide a list of ahermatypic corals known

OCULINIDAE from the West Indies together with their tempera­ 37. Ocu.llna dl!!Ula Lamarck__ ...... __ .... .______FV 38. O. Dalenclennul Edwards and Halme ." __ ture and depth ranges. It is reasonable to expect 39. O. Darleola LeSueur __ ....__ ...__ __ ••_..__ ... F that where the proper temperature, depth, and TRO CHOSIlILIIDAB 40. MeandrlllCl meandrUu (Llnnae\llll ...__ .. ••• ....__ F bottom conditions exist in the Gulf, the ap­ 41. M. brllllliemil (Edwards and mlme}_... ..__ ... .._ F 42. Dlc1locoenlClllolcull Edwards and Halme •• __ • F propriate West Indian species may be found 43. Dendrowra cv"ndrllol Ehrenberg ..__ . ...__ ...... F eventually. Existing accounts of Gulf of Mexico MU88IDAE «. MUlmmllia brUlll1tmil (Verrll1} ... ..__ __ ...__ .. corals are mainly confined to those of Pourtales 46. M. harllf (Verrllll • ...... 46. MUI,a angulola.\Pallas). _. _•• . __ ••: •••.• F dealing with the deep-water fauna of the Florida 47. I,opAl/lUulrea rig da (Danal ... ..__ •. __ ...__ • __ • __ F 48. MvcnOChVlllalamarcJcana (Edwards and Halme} .. F Keys and between Dry Tortugas and the 49. I,opAV la ,lnuCla &EllIs and Solanderl __ .._•_ F 60. I. muUI!lora Verrll . __ ...... __ .... . F Campeche Bank. SUBORDER CARYOPHYLLIIDA Descriptions of West Indian ahermatypic species

CARYOPHYLLUDAB are to be found in Agassiz (1888), Pourtales (1868, 61. EUlmUla/Ulllglala (Pallasl ...__ ...__ .....__ ..__ ..__ ..... F 1871, 1874, 1879, 1880), and Verrill (1883, 1908). - A key to Western Atlantic genera is given in Smith . The West Indian fauna includes species belong­ (1948). A list of West Indian genera with ltlg to the .A.strocoeniidae, Acroporidae, Agariciidae temperature and depth ranges is given in table 2, ~i~er~treidae, PoritidaejFaviidae, Oculinidae, compiled from data in Vaughan and Wells (1943). 294 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

TABLE 2.-Ahermatypic genera of corals found in West SUBORDER DENDROPHYLHDA Indies and Gulf oj Mexico with approximate temperature DENDROPIlYLLllDAE and depth ranges BalanopAvllia Wood D-I,100 meters; 7-280 C. SURORDER ASTROCOENIIDA DendropAvllia ID-~ meters; 11-210 C. TUba'/rea Lesson SERIATOPORlDAlt '. D-400 meters; 15-280 C. . Modroel' Milne Edwards and Halme 0 TrocholMammla Pourtalils 0-800 meters; 10-21 C. liOO-l,.lQO metNs; 5-7° C. EnaUop,ammill Michelotti St'RORDER FUNGIIDA 27D-2,000 meters; 3-150 C. TAecop.llmmia Pourlalils FUNGIIDAE 150-1,000 meters; D-)40 C. Fung/oevo/hu, Sars BalAvp.ammia vonMarenteller IJ(Hl,OOO meters; 1-210 C. 226-330 meters; Insufficient data for temperature range. SUBORDER FAVIIDA In shallow water the only ahermatypic corals ASTRANOIlDAE A,/rang/o MUne Edwards and Halme are 3 or 4 species of Astrangia, Madracis, and D-lJO meters; 8-280 C. PhVUonglo Milne Edwards and Halme Phyllangia. Most species are found in deeper 0-90 meters; 23-280 C. down~ Colanglo Pourtales water from the edge of the continental slope D-580 meters; &-27" C. ward. A total of 84 species is known, According OCULlNII>AE to Vaughan and Wells (1943), 13 of these are Modreporo Llnnaeus 6O-1,~ meters; 4-16° C. identical with living species of the northern and TROCIlOSHILnDAl: eastern Atlantic; 59 are endemic. Cosmopolitan D

AJOo'TJlEHIPIIYLUlI>AE italicus, Desmophyllum cristagalli, Fungiacyath'1.lS An/hem/phvllla Pourtaills )7D-730 meters; 8-170 C. symmetricus, Lophelia prolijera, Madrepora oculata, and Stephanocyathus nobilis. SUBORDER CARVOPIIYLLlIDA

CARYOPHYLLIIDAE TABLE 3.-West Indian fossil genera found living elsewhere, Carl/a phI/ilia Lamarck D-2,BOO meters; 2-270 C. SUBORDER ASTROCOENIIOA CoenocvolhUIJ Milne Edwards and Halme (HI,BOO meters; 11--26° C. ASTROCOENIIOAE CvoJhoura, Moseley Sll/locoenlelia Yobe and Suglysma 70-1,100 meters; 3-210 C. Eocene-Oligocene; Recent-Japan, Mauritius Ozv,mllia Duchassaing SEJUATOPORIDAE 75-300 meters; lo-lllo C. SllilopAora SchwelKKer Balhl/cl/olhu, Milne Edwards and Halme Eocene-Miocene; Recent-Red Sea, Indo-Pacific ~5-145 meters; 10-13° C. Trochoel/albu, MUne Edwards and Halme SUBORDER FUNGIlOA 15-1,~ melers; 4-27" C. Peponocl/alhu, Gravler AOAJUCIIDAE 160-1,100 meters; 7-21 0 C. Parona Lamarck Telhocl/athu. Kuhn Ollgocene-Mlocene; Recent-lndo-PacUlc 6s-1.100 meters; 3-120 C. Leplo,.,i, Milne Edwards and Baine DeltocvalAu. Milne Edwards and Halmc Oligocene-Miocene, Recent-Indo-Pacific 16-4,300 meters; 4-280 C. THA)I)/A5TER1DAIt Cera/o/rochu. MUne Edwards and Balme P.ammocora Dana M-7oo meters; b-17° C. Miocene; Recent-indo-Paclllc Stephanocvalhu, Sequenza AOATHlPnYLLlII>AIC 360-2,200 meters; 3-12" C. Diploa,/rea Matthal Turblnolla Lamarck Cretaceous-Olfgoccne; Recent-lndo-PacWc ss Bato/rochn. Wells POR1TlI>AE 180-~70 meters; 11-21 0 C. Gonlopora de Blalnvllle SpA.nolrocA", Mllne Edwards and Halme Cretsccous-Mloccne; Recent-Indo-Pacific 22-27S meters; 11--260 C. D18mophl/Uum Ehrenberg SUBORDER FAVnDA ab-2,OOO meters; 3-23° C. Paroamilia Milne Edwards and Halme FAVlllJAE 310-370 melers; 8-12° C. PaDi/.. Link Co.no,mllia Pourtales Eoccne-Mloccn~Rect'nt-Indo-Pael1le, Red Sea 145-400 meters; 13-23° C. OO1lia,lrea Milne 1O;dwards and Halne Anomocora Studer Eocene-Ollgocene, Recent-Indo-Pacific, Red Sea 90-820 meters; 4-170 C. A,/erolmilia Duncan 80-260 meters; 13-16° C. BIBLIOGRAPHY OUYNIIDAE GUI/nia Duncan 17()-6,jQ meters; 7-12" C. AOASRIZ, Al.EXA:o.·PER. Stmocvalhu, l'ourtal~s 1888. Three cruises of the Blake. Houghton, MilJl irl 8O-8111 melers; 7-)40 C. Sehlzocl/alhu, Pourlallls and Co., Bostoll and New York. Also: Bull. MuS. lOO-l,~ meters; !D-!8° C. Compo ZooI., Harvard College, vols, 14 and 15. FlA9ELLIDAE --- and DE POURTALES, L, F. Flabellum Lesson 1874. 8. The zoological results of the Hassler 0-3.260 meters; 2-270 C. 1'0. MOl/oml/eel Ellrenberg Expedition. I. Echini, crinoids, and corals. Me l1l 0-1,000 meters; 6-280 C. Mus. Compo Zool.. Harvard College, 4: 1-50, 15 Oardlnala VBughnn 70-000 meters; 6-180 C. cuts, 10 pis. GULF OF MEXICO 295

bUERDEN, J. E. VAUGHAN, T. W. 1902. West. Indian madreporarian polyps. Nat. Acad. 1901. The stony corals of Puerto Rican waters. Bull. Sci. Mem. (Washington), Vo!. 7. U. S. Fish. Comm, 1900, 2' 289-320. FIELDEN, H. W. 1911. Recent Madreporaria of southern Florida. Car­ 1893. Transportation of coral by the Gulf Stream. The negie Inst. Washington Yearbook 9: 135-144. Zoologist (3), 17: 352-353. 1912. Madreporaria and marine bottom deposits of GREGORY, J. W. Florida. Carnegie Inst. Washingt.on Yearbook 10: 1900. On the West Indian species of Madrepora. Ann. 147-156. Mag. Nat. Hist. (7) 6: 20-31. 1913. Studies of the geology and of the Madreporaria of fIEILPRIN, A. the Bahamas and of southern Florida. Carnegie 1891. The corals and coral reef of the western waters Inst. Washington Yearbook 11: 153-162. of the Gulf of Mexico. Acad. Sci. Philadelphia Proc. 1914. The reef corals of southern Florida. Carnegie 42: 303-316. InsL Washington Yearbook 12: 181-183. JO UBIN , M. L. 1915a. Reef corals of the Bahamas and southern Florida. 1912. Carte des bancs et des r~cifs de Coranx (Madre­ Carnegie Inst. Washington Yearbook 13: 222-226. pores). Ann. de !'Inst. Oceanog. (Old Ser.), vo!. 4, 1915b. The geological significance of the growth-rate of No.2. the Floridian and Bahamian shoalwater corals. bE POURTAI.ES, L. F. Washington Acad. Sci. Jour. 5: 591-600. 1863-69. Contributions to the fauna of the Gulf Stream 1916a. Results of investigations of the ecology of the at great depths. Bull. Mus. Compo Zoo!., Harvard Floridian and Bahamian shoal-water corals. Nat. College, 1 (6): 103-120; 1 (7): 121-142: Acad. Sci. Proc. 2: 95-100. 1871. Deep-sea corals. IlIus. cat. Bull. Mus. Compo 1916b. On recent Madreporaria of Florida, the Bahamas Zoo!., Harvard College, 4: 1-93. and the West. Indies, and on collections from Murray 1878. Reports on the results of dredging, ... in the Island, Australia. Carnegie Inst. Washington Year­ Gulf of Mexico, by the U. S. Coast Survey Steamer book 14: 220-231. Blake. II. Crinoids and corals. Bull. Mus. Compo and WELLS, JOHN WEST. Zoo!., Harvard College, 5 (9): 197-216, 2 pIs. 1943. Revision of the suborders, families and genera of 1880. Reports on the result.s of dredging, ... in the the Scleractinia. Geo!. Soc. Am., Spec. Pap. 44. Gulf of Mexico, by' the U. S. Coast Survey Steamer' Blake. VI. Report on the corals and Antipatharia. VERRILL, A. E. Bull. Mus. Compo Zoo!., Harvard College, 6: 95-120. 1902a. Variations and nomenclature of Bermudian, 1883. Report on the Anthozoa and some additional West Indian and Brazilian reef corals, with notes on species dredged by the Blake, 1873-79; and Fish various Indo-Pacific corals. Trans. Connecticut Hawk, 1880-82. Bull. Mus. Compo Zoo!., Harvard Acad. Arts & Sci. 11: 63-168. College, 11: 1-72. 1902b. Comparisons of the Bermudian, West Indian and SMITH, F. G. W. Brazilian coral fauna. Trans. Conn. Acad. Arts & 1948. Atlantic reef corals. University of Miami Press. Sci. t 1: 170-206. D. S. NAVY HYDROGRAPHIC OFFICE and DIVISION of WAR WELLS, J. W. RESEARCH of the UNIVERSITY of CALIFORNIA. 1932. Study of the reef corals of the Tortugas. 'fortugas 1943. Bottom conditions. Bottom Sediment Chart Lab. Ann. Rept., Carnegie lust. Washington Year­ 1943. H. O. Chart Nos. 1125 BS, 1126 BS. book 31: 290-2!l I. Blank page retained for pagination CTENOPHORES IN THE GULF OF MEXICO

By MARY SEARS, Woods Hole Oceanolraphic Institution

Ctenophores are so fragile that they are not been reported from neighboring parts of the readily preserved or, if they are, certain diagnostic Atlantic. Whether other species will be found characters may become obscured. Thus, many in this area seems problema.tical. At any rate, records of their occurrence are somewhat un­ although ctenophore species are more numerous certain. Equally uncertain are their names be­ in the Gulf, they apparently do not occur in dense cause about 40 years ago four important papers swarms as is so characteristic of them in more appeared almost simultaneously (Bigelow 1912; northern waters. Mayer 1912; Mortensen 1912; Moser 1912), and LITERATU~E CITED insofar as I can ascertain, nobody has reviewed the group critically since that time. Itis probable, BIGELOW, H. B. 1912. Reports on the scientific results of the expedition however, that the ctenophore fauna of the Gulf to the eastern Tropical Pacific ... XXVI. The of Mexico is as well known as that of any neigh­ ctenophores. Bull. Mus. Compo Zoo!., Harvard boring area due to Mayer's (1900, 1912) and College, 54 (12): 369-404, 2 plB. Fewkes' (1882) observations at the Tortugas. FEWKES, J. W. :Nevertheless, only about a dozen species have been 1882. No.7. Explorations of the surface fauna of the Gulf Stream, under the auspices of the U. S. Coast recorded with any certainty in the Gulf: Beroe Survey by Alexander Agassiz. 1. Notes on acalephs ovata Bosc; BolinopSis vitrea L. Agassiz j Gest'Um· from the Tortugas with a description of new ge~era veneris LeSueur j E'Urhamphaea vexiUigera Gegen­ and species. Bull. Mus. Compo Zoo!., Harvard baur; Folio parellela Folj Hormiphora hormiphora College, 9: 251"':'289, 7 pIs. MAYER, A. G. Gegenbaur; u'UCothoe ochracea 1 Mayerj Mne­ 1900. Some medusae from the Tortugas, Florida. tniopsis mccradyi Mayer; Ocyropsis crystaUina Bull. Mus. Compo Zoo!., Harvard College, 37 (2): Rang; Ocyropsis maculata Rang; Tinerje beeMen 13-82, 44 pIs. Mayer; Tinerfe lactea Mayer (using the names 1912. Ctenophores of the Atlantic coast of North Amer­ that appear to be acceptable today). This is a ica. Carnegie Inst. Washington Pub. No. 162: 58 8light reduction in the number originally described pp., 17 pIs. MORTENSEN, TH. because a number proved to be identical with 1912. Ctenophora. Danish Ingol! Exped. 5 (2): 95 8pecies which had been described earlier. This pp., 10 pIs. list also includes most of the species that have MOSER, F. 1912. Die Ctenophoren der Deutschen SQdpolar Ex­ - pedition 1901-1903. Deutsche SQdpolar Exped. 11 1 Fewkes' (1882) record or EueIIarf, muilkornu QUOY and Oafmard Is con· dered by Mayer (1912, p.ll6) to have been his new specIes, LeucotAoe ochraua. (Zoo!. 3): 117-192, 1 text fig., pIs. 20-22.

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