23 August 1988 PROC. BIOL. SOC. WASH. 101(2), 1988, pp. 234-250 NOMENCLATURE AND BIOLOGY OF ASTRANGIA POCULATA i=A. DANAE, =A. ASTREIFORMIS) (CNIDARIA: ANTHOZOA) Esther C. Peters, Stephen D. Caims, Michael E. Q. Pilson, John W. Wells, Walter C. Jaap, Judith C. Lang, Carol E. (Cummings) Vasleski, and Lauren St. Pierre GoUahon Abstract. ?The encrusting to ramose northern star coral, Astrangia poculata, is a common inhabitant of rocky shorelines, jetties, and pilings along the north- west Atlantic and Gulf of Mexico coastlines of the United States. Despite its popularity as an experimental laboratory animal, there has been considerable confusion as to the number of species which may exist and the proper name(s) of the species. A reexamination of material from throughout this range confirms that there is only one species, the correct name of which is Astrangia poculata (Ellis & Solander, 1786). A neotype has been deposited in the National Museum of Natural History. The same neotype is also deposited as the type of A. michelinii Milne Edwards & Haime 1849, type species oi Astrangia. The genus Astrangia includes over 30 Re- cent nominal species within the family Rhi- zangiidae, and is represented by 20 fossil species, some of which lived as early as the Middle Cretaceous, approximately 100 mil- lion years ago (Wells 1956). Members of the genus are all relatively small and are usually insignificant components of the fauna where they occur. The various species of Astran- gia, however, collectively occupy a vast geo- graphic range, possibly broader than that of any other genus of coral. Unfortunately, the genus Astrangia has never been completely reviewed. It is prob- able that many species names are syn- onyms, and also that a number of yet un- discovered species may exist (Pilson 1975). The confusion that exists within this genus became evident when a group of scientists at the University of Rhode Island began a series of studies on the ecology, physiology, and life history of the coral found in Nar- ragansett Bay. Whereas several previous studies had referred to the northern star cor- al as Astrangia danae, a number of other papers had been published on the very sim- ilar Astrangia astreiformis from the south- eastern coasts of the United States. How- ever, other texts suggested that only one species o{ Astrangia existed along the entire North Atlantic and Gulf coasts of North America (Hyman 1940, Wells 1956, for re- views see Cummings 1983, and Jaap, in prep.). A. astreiformis was also reported to occur off the west coast of Africa (Thiel 1928, Chevalier 1966). Throughout its range, the northern star coral appears remarkably tolerant of a wide range of environmental conditions. Colo- nies oi Astrangia are generally considered to be ahermatypic (non-reef-building) and hence azooxanthellate (=asymbiotic, i.e., lacking zooxanthellae), but some colonies of the northern star coral contain these algal cells abundantly within their tissues (Bosch- ma 1925a). These two factors, adaptability and variability in algal symbiosis, have con- tributed to the increasing popularity of this coral as an experimental laboratory animal. Therefore, in order to aid future researchers, we were motivated to clarify the taxonomic status of this coral. VOLUME 101. NUMBER 2 235 Family Rhizangiidae d'Orbigny, 1851 Genus Astrangia Milne Edwards & Haime, 1848a Diagnosis.?Co\on\a\, usually azooxan- thellate. Colonies encrusting or subplocoid, formed by extratentacular budding from edge zone; corallitcs united basally by thin coenosteum, septa dentate; columella pap- illary. [Type species: .-1. michelinii Milne Edwards & Haime, 1848b, by monotypy (=A. poculala Ellis & Solander, 1786).] Astrangia poculala (Ellis & Solander, 1786) Figs. 1-6 "Stony coral found on the shore near New York.": Ellis, 1755:84, pi. 32, figs. A. 1, A.3. 'IMadrepora calycularis Pallas, var. li, 1766: 319 note 186B.-Espcr, 1790:283. Madrepora poculala Ellis & Solander, 1786: 165.-Gmelin, 1791:3766.-Esper, 1790: 283 (=.V/, calycularis 0). [Original type locality "near New York." Type speci- men lost (P. F. S. Cornelius, pers. comm.).Type locality (ncotype) "off At- lantic City, New Jersey." Neotype de- posited in the National Museum of Nat- ural History USNM 80350.] Madrepora porculaia: Turton, 1806:623 [Misspelling of/?on7)f?v? VOLUME 101, NUMBER 2 n '??? 237 vival, Texas).-St. Pierre, 1986:1-55 (re- production, Texas). (Type locality "United Stales." Type found by Wells in Paris ir Coll. Michelin in 1934, but not found by Chevalier in 1980 (Cairns 1981).] Astrangia danae Agassiz, 1850:68-77, pi. I, fig. 7 (feeding behavior, digestion, colony growth, new polyp formation, nemato- cyst function, Massachusetts). ?Verrill, 1863:40 (North Carolina); 1864:47 (=A. astreiformis in Leidy, 1855, not A. danae ME & H 1849); 1866a:324 (not A. danae ME&H 1849); 1866b:335; 1872:436,437 (=A. astraeifonnis ME & H 1849); 1873: 408; 1874:446 (Vineyard Sound and ad- jacent waters).?Agassiz& Agassiz, 1865: 16, figs. 16-20.-Pourtales, 1871:80.- Dana, 1872, 1890:67-68, figured p. 68, a, b,c.-Damon, 1882:22 l.-Fewkes, 1889: 7-10, pis. 1-5 (natural history illustra- tions); 1891:53.-Hargitt, 1914:250, pi. 43, fig. 9 (natural history, Casco Bay, Maine). ?Boschma, 1925a:65-67 (zoo- xanthellae symbiosis); 1925b:407-439 (feeding behavior, digestion, zooxanthel- lae symbiosis). ?Cowles, 1930:333 (Chesapeake Bay, Cape Henry, Cape Charles).-Pratt, 1935:159, fig. 239.- Pearse, 1936:178.-Richards & Harbi- son, 1942:178, pi. 22, figs. 2 & 3 (Mio- cene, New Jersey). ?Vaughan & Wells, 1943:177 (see footnote 52: =1A. michelini ME & H I848).-Field, 1949:6, 18, pi. I. fig. 7.-Waterman, 1950:127-131, fig. pp. 120-121 (structure of corallum and pol- yps, zooxanthellae, feeding, reproduc- tion).-Alloiteau, 1952:623, fig. 74 (=.1. michelini ME & H 1848).-Hand, 1964: 25, pi. 3, fig. 4 (description). ?Driscoll, 1967:633-641.-Robertson et al., 1970: 55_65.-Calder, 1972:101 (Chesapeake Bay, Hog Island and Eastern Shore).? Sassaman & Mangum, 1973:1313-1319 (anerobic metabolism).?Jacques et al., 1977:455-461 (physiology, calcification, Rhode Island). ?Hayes & Goreau, 1977: 26-40 (calcification, histology). ?Bach- and, 1978:283-284 (habitat, life histo- ry).-Calder & Hester, 1978:93 (Charles- ton, South Carolina). ?Weslon, 1980:1- 93 (zooxanthellae distribution, Chesa- peake Bay). [Type locality "Massachu- setts." Deposition unknown.] Astrangia aslraeformis: ME & H: Leidy, 1855:139-140, pi. 10, figs. 9-16 (synon- ymy, nematocysts, Rhode Island). [Mis- spelling of A. astreiformis ME & H 1849]. Astrangia astraeiformis: ME & H: ME & H, 1857:614.-Verrill, 1863:39; 1866a:324; 1866b:335; 1872:436 (=A. danae Agassiz I850).-Pourtales, 1871:80.-?of Thiel, 1928:283-285, pi. 2, fig. 22 {=A. mer- catoris Thiel 1941, West Coast of Afri- ca).-Thiel, 1941:5, 15, pi. 1, figs. 5-6 (Martinique). ?Pearse & Williams, 1951: 136 (incorrectly attributed to Leidy, 1855, Maine to Florida and the West Indies).? Chevalier, 1966:926-930, pi. 3, figs. 2-3, pi. 4, fig. 6 (synonymy, description. Sierra Leone, Africa). [Misspelling (or corrected spelling) of .4. astreiformis ME & H 1849.) Astrangia danai: ME & H, 1857:614. [Mis- spelling (or corrected gender) of .4. danae ME& H 1849.] Astrangia edwardsii Verrill, 1866a:324 (=nom. nov. for .4. danae ME & H 1849, see Remarks); 1866b:335. not Astrangia mercatoris Thiel, 1941:16- 17, pi. 2, fig. 1 (=.4. astraeiformis Thiel, 1928:283-285, pi. 2. fig. 22; Angola. Bay of Elephants, West Coast of Africa).? Figs. 1-6. Aslraniiia pocidatu: I, Ncotypc colony (oirAtlarmc City, New Jersey. USNM 80350). > 0.88; 2- 3, Colony from off Jacksonville, Florida, 15 m. USNM 78509, xl.20, x3.40, respectively; 4, Deep-water branched colony (off Western Dry Rocks, Sand Key. Honda, 263 m. USNM 78508). >'0.86; 5, SEM of coe- nosteum adjacent lo calice showing lines of perforations constituting intercostal striae and cocnosteal granules, x49; 6, SEM of calice with 24 septa. < 23 (5-6. R/V Gerda station 1002, USNM 78507). 238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Chevalier, 1966:929-930 (discusses syn- onymy, see Remarks). Astrangia astraeiformis: no authority cited: Hedgpeth, 1953:164 (Texas). [Misspell- ing oi A. astreiformis.] not Astrangia astraeformis: no authority cit- ed: Buchanan, 1954:85-86 (=A. astraei- formis Thiel 1928 =A. mercatoris Thiel 1941, Ivory Coast, French Congo, An- gola, Africa). [Misspelling of,'!, astreifor- mis.] - - . ? - 1- :? Astrangea danae: no authority cited: Go- reau & Bowen. 1955:1 188 (calcification). [Misspelling of Astrangia danae.] .Astrangia asireformis: Milne Edwards & Haines(sic): Pampe, 1971:399. pi. 2. fig. 7 (Pleistocene, Texas). [Misspelling, spelled .t. astreiformis later on same page.] Astrangia asteriformis: no authority cited: Rudloc, 1971:154-157, 167, 172-174, figured on p. 174 (natural history, har- diness in aquarium. Florida). [Misspell- ing.] Astrangia a.streiformis: no authority cited: Ladd. 1951:137, 139, 149 (Texas'coastal waters). ?Slephenson & Stephcnson, 1952:35 (distribution, taxonomy).? Hedgpeth, 1953:164 (Texas).-Gunter & Geyer, 1955:40,44,45, 56. 57, 65 (Texas, Louisiana).-Parker, 1956:310, 315, 329, 354, 370 (dead specimens collected in east Mississippi delta region). ?Courtney & Edmond, 1974:63 (Marco Island, Flori- da).?Folheringham & Bruncnmeister, 1975:36. 161. fig. 2.24 (natural history). .istrungia danae: no authority cited: Wil- son, 1900:350 (North Carolina).-Allee, 1923:176 (Woods Hole. Buzzards Bay, and Vineyard Sound, Massachusetts).- H\man, 1940:610-611 (morphology, natural history). ?Storer. 1951, figured on p. 321. ?Stephenson & Stephcnson, 1952: 35 (distribution, taxonomy).-Blake. 1953:23 (Pleistocene, Maryland). ?Zinn. 1964:15 (Rhode Island).-Goreau et al.. 1971:254.-Williams& Murdoch. 1973: 551-563 (elTccts of gamma radiation on). ?Cummings, 1976:1-115 (physiol- ogy, Rhode Island). ?Szmant-Froelich & Pilson, 1977:417^24 (nitrogen excre- tion, symbiosis, Rhode Island). ?Hoff- man et al., 1978:165-170.-Jacques, 1978:1-169 (calcification). -Gosner, 1979:93-94,fig. 20.-Storer et al., 1979, figured on p. 320, 372, 390.-Szmant- Froelich, 1980b: 1-3 (popular account).? Szmant-Froelich el al., 1980:257-269 (reproduction, Rhode Island). ?Mein- kolh, 1981:391.-Cummings & Mc- Carty. 1982:1125-1129 (calcification, stable carbon isotopes, Rhode Island).? Jacques et al., 1983:135-148 (ecology, metabolism, Rhode Island). ?Swart, 1983:72 (stable carbon isotopes).? Szmant-Froelich & Pilson, 1984:153-162 (feeding frequency, nitrogen metabolism, respiration, Rhode Island). Astrangia sp.: Ferguson, 1948:17 (salinity tolerance. Louisiana). ?Keith & Weber, 1965:500 (isotopic composition, calcifi- cation, in part: Woods Hole, Massachu- setts).-?of Rabalais, 1978:62 (= Oculina diffusa, G\x\f of Mt\ico). Diagnosis. ?EncTU%\\n%, massive, or ra- mose irregularly shaped colonies. Corallum cerioid to plocoid. new polyps produced by extratentacular budding, having coralliles united basally by thin peritheca (vesicular coenosteum). Corallites round, irregular, or polygonal; tightly packed or spaced several millimeters apart, highly variable even within a single specimen. Caliccs 1-7 mm in diameter, with distinct equal costae on many corallites. Septa hcxamerally ar- ranged in 4 cycles, however, complete fourth cycle (48 septa) only attained in very large calices. Most calices have 24 septa, the S, merging basally with the S^ about 'A distance to columella. With increase in calicular di- ameter, pairs of Sj form flanking some S,, often with one pair per system (the space between two S,) resulting in 36 septa. Com- plete fourth cycle (48 septa) attained only in caliccs over 6 mm in diameter. Septa VOLUME 101, NUMBER 2 Strongly dentate, particularly S, at region of S, fusion, and slope gradually toward col- umella. Fossa shallow. Columella trabecu- lar or papillose. Polyps up to I cm tall and translucent white (azooxanthellate = asymbiotic) to brown (zooxanthellate = symbiotic, see Schuhmacher&Zibrowius 1985. for review of this terminology). Numerous nematocyst batteries on surface of tentacles; tip of ten- tacle rounded. Directive mesenteries pres- ent. Corallum often invaded by endolithic red or green boring algae, burrowing poly- chaetes, or sponges (e.g., Cliona cclata Grant, see Cummings 1983). Cnidocyst batteries principally composed of spirocysts. Terminal knob packed with large microbasic-p-mastigophore ncmato- cysts (about 29 tim long x 8 Mm in diameter in fixed, paraffin embedded sections) and spirocysts. Occasional small microbasic-p- mastigophores scattered in epidermis (about 20 Mm long x 7 ^m in diameter). Cnido- glandular band of mesenterial filaments possesses large microbasic-p-mastigo- phores (about 40 iim long x 7 ^m in diam- eter), holotrichous isorhizas (about 45 ^m long X 15 Mm in diameter), and small mi- crobasic mastigophores (?) (about 11 Mm long X 5 ^m in diameter). The distribution of these nematocysts vanes with location on the filament (ECP. unpublished). Histolog- ically distinguished by well-developed lon- gitudinal retractor muscles along the me- soglcal pleats. Gastrodermis lining calicoblastic epidermis features large cells with basophilic globules (osmiophilic, non- PAS positive, non-mucin) up to 4 Mm in diameter, which increase in number and density towards base of the polyps. Cali- coblastic epidermis thickened with fine eo- sinophilic granules (Peters 1984). Colonies gonochoric (=dioecious). gametes spawned annually (Szmani-Froelich el al. 1980. St. Pierre 1986). Remarks.-Madrepom poculata was de- scribed by Ellis & Solander (1786), based on the illustration of a specimen from "'near 239 New York" figured by Ellis (1755). Another reference to this "American" coral ap- peared in Pallas (1766) under the name Madrepora calycularis. var. 0. Typical M. calvcularis Pallas (1766) is now known as Astroides calvcularis (Pallas, 1766), found only in the Mediterranean (Zibrowius 1980). Therefore, the name calycularis. even if ap- plied to a specimen of western Atlantic As- irangia as form /3, is preoccupied by the Mediterranean species. Esper (1790:283) equated M. poculata with the M. calycularis li of Pallas, describing a corallum encrusting an American gastropod, which may have been the same as the astrangid-encrusted specimen of Conus spurius from the West Indies, figured by Seba (1758:152, pi. 55, figs. 11. 12). The last known reference to M poculata was Turton (1806). ME & H (1848b) described Astrangia michelinii as an illustration found in Mem- oire 4, Monographic des Astreides; this is a valid "indication" (International Code of Zoological Nomenclature (ICZN) 1985: Ar- ticle 16, vii), and therefore it is a valid species and the type species oi Astrangia by mono- typy. It was described without a type local- itv. In 1849, ME & H described (in order) 3 types in the Museum National d'Histoire Naturelle, Paris, (MNHNP), .istrangia dan- ae. .-istrangia michelini. and Astrangia as- ireiformis. The descriptions of the type specimens were very similar. They noted that .1. michelini was closely related to A. danae. Unfortunately, only the type locality of one of the 3 species, .1. astreiformis. was given, as from the "'United Stales." Agassiz (1850) described the same species from the United States and used one of the same names used by ME & H. .Astrangia danae. in a paper presented before the American Association for the Advance- ment of Science meeting in 1849. but it was not published until 1850. He made no com- parisons and apparently did not know that ihe name had alreadv been used for a species in the previous vear (ME & H 1849). and 240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON therefore was a junior primary homonym. It is curious that Dana (1872,1890) referred to A. danac Agassiz, suggesting that he had not been in correspondence with ME & H m the late 1800's. Dana noted that he him- self had proposed the name Pleiadia for the genus, alluding to the crowd of stars (the appearance of the corallum), in his report on Zoophytes (Dana 1846:722, Wells 1986: 50, nom. nud.). However, Agassiz (1850: 69) made no reference to this particular re- port. That he chose to call his species As- irangla presumably means that he had seen the figure (if not the specimen on one of his European trips) of A. michelinii published in 1848. Later authors, who had compared the type specimens and descriptions, commented on the relations of these species. Vaughan (1901:300) noted that the specimen of As- trangia astreiformis (corallites free above their bases), which he had collected in Puer- to Rico, was not typical of the specimens of A. astreiformis then in the USNM, and ob- served that the figure of A. michclini ME & H "suggests great similarity to the specimen from Puerto Rico." Alloiteau (1952) be- lieved that A. michelini ME & H was the same species as A. danae Agassiz. Chevalier (1966) observed that A. astraeiformis ME & H was similar to A. michclini of ME & H. Thus, A. michelinii Milne Edwards & Haimc is "similar" to A. danae Agassiz (Vaughan & Wells 1943, Alloiteau 1952), .1. astreiformis ME & H (Vaughan 1901. Chevalier 1966). and A. danae ME & H (Milne Edwards & Haime 1849). Leidy (1855) considered Astrangia as- treiformis ME & H to be the same as .1. ?/(3/2ac Agassiz. In 1863, Verrill recorded that .-1. danae Agassiz was equal to the A. as- treiformis descnbed by Leidy in 1855, and noted that A. danae Agassiz was not the same as A. danae ME & H. Verrill (1866a) reported that .1. bella E. & H. [ME & H], known from the Miocene formations in Maryland, was vcrv similar to both A. danae Agassiz and A. astreiformis ME & H, and noted that some authors would unite all three species. Pourtales (1871:80) remarked "The differences between these two species (A. as- treiformis M. E. & H. and A. danae Ag.) are of the slightest character, being chiefly de- rived from the mode of grouping, which is denser in the first than in the second. I am strongly inclined to believe that they will ultimately be united." In 1872, Verrill united .(. astreiformis ME & H with A. danae Agas- siz. Cummings (1983) and Jaap (in prep.) united A. astreiformis ME & H with A. dan- ae Agassiz, noting the great variability in specimens collected from throughout its range along the coastline of the eastern United States. Only Vaughan (1901) felt that there were sufficient distinctions based on the skeletal structures to maintain A. as- treiformis ME & H and A. danae Agassiz as separate species. Thus, these observations link Astrangia astreiformis ME & H with .i. danae Agassiz. .Although A. danae ME & H appeared to be very similar to .1. astreiformis ME & H (Lei- dy 1855. Chevalier 1966) and A. michelini ME & H (Milne Edwards & Haimc 1849), subsequent authors (Verrill 1863, 1866a) did not connect .1. danae ME & H with .4. danae Agassiz. Verrill 1 866a:324) stated again that ,1. danae ME & H ("the corallites being scat- tered and connected only at the base") was different from .4. danae Agassiz, and there- fore proposed the name .4. edwardsii as a replacement name for .4. danae ME & H. He erroneously thought that the date for .Agassiz's name was 1849 and the dale of ME & H's was 1850, because this latter date was cited in error by ME & H (1857:614). A number of later references attributed the publication date of .Agassiz's name to 1847 (e.g., Calder 1972) or 1848 (e.g.. Field 1949), further adding to the confusion. Fewkes (1889:8) noted that "The polypdom of .4. Danae (sic) M. E. & H., is held by Prof Joseph Leidy to be different from that of our New England species of .4strangia and more like .4. astraeformis of the same au- thors .... It is not possible to determine VOLUME 101, NUMBER 2 241 from the description of A. Danac given by Milne Edwards & Haimc whether our species differs from that which they describe under that name or not." J. W. Wells examined the type specimens in the MNHNP in 1934. Of Aslrangia mi- chelim ME & H 1848 (No. 412, Michelin Coll.), he noted "Does not look like the fig- ure but has M.E. & H.'s label on the wooden base. Very like .-1. danac [no authority cited] except for smaller [corallites] (2.5-3.0 mm). It is attached to a bit of igneous rock. This is the type species of Asirangia by mono- typy. No loc." (J WW notes, February, 1934). He also observed that in the Paris collection "Specimens labeled .1. JanaclME & H 1849] are from Si. Thomas and are A. soliiaria. coll. Duchassaing ca. 1846, except for one that appears to be the type. It encrusts a Tellina and fits the description well and is evidently conspecific with A. michelini as latter was figured in 1848 by M.E.&H. Has M.E.&H.'s label. No loc." Wells noted that the type of .-t. astreiformis ME & H 1849 was in the Paris collection in 1934, with the locality designated "Etats-Unis," but did not make any other observations on this spec- imen (JWW notes, February, 1934). From all indications (Cairns 1981), all of the types are now missing from the collections of the MNHNP. Wells considered Astrangia danae ME & H 1849 to be the proper name for the Amer- ican northern star coral (pcrs. comm. to A. Szmant-Froelich, 13 December 1977). not- ing that "Over the years 1 have examined many specimens from throughout this range [Massachusetts to Texas] and although there is much variation in skeletal morphology, the variation seems not be geographical or provincial, for specimens from one locality frequently show the full range of vanation in compactness or isolation of corallites. size of corallites. equality of septa, etc." Al- though the names A. danae Agassiz and .1. astreiformis ME & H had been used nom- inally to separate specimens from north of Cape Hatteras to New England from those specimens collected from south of Cape Hatteras into the eastern and .vestem Gulf of Mexico, respectively. Wells observed that A. danae ME & H had priority as the senior synonym. Cairns (1981) argued that Astrangia as- treiformis was the appropriate name for the American species. He noted that ME & H described both A. danae aind A. astreiformis in the same paper. Although danae\\did page priority, the type was lost, the description poor, and the type locality not given. He could not confirm that A. danae ME & H was the same as A. danae Agassiz and A. astreiformis ME & H. hence, he considered A. danae ME & H to be a nomcn dubium. The locality for A. astreiformis was given as "United States." Although the type of .^. astreiformis was also lost, it at least had a type locality, which is why Cairns consid- ered it as the senior synonym. Because Agassiz's description of the same species, which he named A. danae, was not pub- lished until 1850, Cairns also considered this name a junior synonym of .1. astreifor- mis and junior homonym of .-1. danae ME & H 1849. Based on our present examinations of specimens and recent biological data (see below), we concur that there is only one species of star coral of the genus Astrangia along the Atlantic and Gulf coasts of North America. Although the type specimens are now missing from the Michelin collection, earlier observations by Leidy, Verrill. Pour- tales. Fewkes, Wells. Alloitcau. and Che- valier suggest that the Astrangia types es- tablished by ME & H were very similar. The latter four authors thought that these types were probably only morphological variants of the colonial northern star coral. There is no doubt that this species is the same as the Madrepora poculata described by Ellis & Solander over 200 years ago. Although the names .(. danae and A. astreiformis have been used widely in the recent 'iterature (73 and 41 references, respectively), these names must be considered junior subjective syn- 242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON onyms. We conclude that the correct name, based on the Principle of Priority (ICZN 1985), is Astrangia poculata (Ellis & Solan- der, 1786). The type specimen of Madrepora pocu- lata is not present at the Glasgow or the British Museum (Natural History) and is believed to be lost (P. F. S. Cornelius, pers. comm.). In order to help clarify the complex nomenclatural issues concerning the names Astrangia poculata. A. michelinii ME & H 1848, A. danae ME & H 1849, A. astreifor- mis ME & H 1849, and A. danae Agassiz 1850, a neotype is chosen for A/, poculata. It isdeposited at the USNM (USNM 80350) and was collected from New Jersey, rela- tively close to the original type locality of "ofTNew York." As mentioned before, there is no type specimen of the type species Astrangia mi- chelinii (see Chevalier 1966). Furthermore, it was described without a type locality and its original description and figures do not reliably distinguish it from the other ap- proximately 30 Recent species in the genus. Subsequent references to A. luichelinii have always been in its capacity as the type species of the genus, not as a new record. Therefore, in order to redefine A. michelinii. and the genus Astrangia by virtue of its status as the type species of the genus, a neotype is cho- sen for .1. michelinii. Because several au- thors (e.g., Vaughan 1901, Alloiteau 1952, Chevalier 1966) have suggested that .1. michelinii is synonymous with the United Slates east coast species, the neotype chosen for.(. michelinii is a specimen of .1. pocu- lata. in fact the same specimen chosen as the neotype of .\/. poculata. a specimen from New Jersey (USNM 80350). A. michelinii thus becomes a junior objective synonym of .1. poculata. Thiel (1928) described a colonial .Astran- gia from the Atlantic coast of Africa, and identified it as .4. astraeiformis ME & H. In 1941, Thiel doubted the existence of the American species on the west coast of Af- rica, and established the species .\. merca- loris as the African species. Buchanan (1954) reported A. astreiformis from the Ivory Coast and Gulf of Guinea off western Africa. Bu- chanan (1954:86) remarked that his speci- mens agreed well with Thiel's (1928) de- scription of A. astreiformis from Africa. Chevalier (1966), however, believed that the American and African specimens that he examined were sufficiently similar to merit recognition as the same species. Unfortu- nately, as we know nothing of the African Astrangia. Thiel's (1928) specimens may in- deed represent a different species. Material examined. ??rom the USNM, 85 lots, the specimens representing the fol- lowing localities: Martha's Vineyard Sound, Buzzards Bay, Woods Hole, Massachusetts; Newport. Rhode Island; 19 lots from R/V (iosnold collected off northeastern Florida and Georgia (13-54 m); Chesapeake Bay, Virginia Beach, Virginia; Cape Henry, North Carolina; Myrtle Beach, South Carolina; Nassau County, Florida; northeastern coast of Florida; Dry Tortugas, Western Dry- Rocks off Sand Key, Florida; Marco. Gulf coast of Florida; Rockport, Galveston, Port Aransas. Texas. Specimens from the Texas coast depos- ited in the Texas Memorial Museum, Aus- tin, Texas (Numbers 1568-1573. 1753). Specimens from the Hourglass Cruises, deposited in the collection of the Depart- ment of Natural Resources, Bureau of Ma- rine Research. St. Petersburg, Florida (Numbers FSBC I 10027. 11360, 17984. 19858-19862) and 49 other cataloged spec- imens (various numbers from FSBC I 10028-31954, eastern Gulf of Mexico and Florida east coast), as well as specimens on loan from Texas A&M University (College Station. Texas), and Museum of Compar- ative Zoology (Harvard University, Cam- bridge, Massachusetts), representing loca- tions from New England to Florida. Louisiana, and Texas. (jcologic and geographic disinhution.? .Astrangia poculata is recorded from the Miocene (Richards & Harbison 1942). VOLUME 101, NUMBER 2 243 Pleistocene (Blake 1953, Pampe 1971) and Recent (Alloiteau 1952, Wells 1956). It oc- curs at depths from 0 to 263 m. It has been reported off Maine (Hargitt 1914) and from Cape Cod south along the eastern United States, around the Gulf coast of Florida to the southern Gulf coast of Texas and Mex- ico (E. Jordan pers. comm. to WCJ). It is very rare in the coastal region from south of Palm Beach to the Ten Thousand Islands, Florida, and is disjunct in the Mississippi delta region. It is also noted from Puerto Rico (Vaughan 1901) and Martinique (Thiel 1941). Thiel (1928) and Buchanan (1954) reported it from the Ivory Coast, French Congo, Angola, Africa, (although its exis- tence off Africa based on these reports has been questioned, see Remarks), and Che- valier (1966) reported it from the Bight of Biafra, Freetown, Sierra Leone, Konakry, Guinea, Dakar and Senegal (Gulf of Guinea), Africa. Discussion.?Corci\ species traditionally have been determined on the basis of their skeletal morphologies (since these are pre- served most easily, and appear in the fossil record). Veron (1982) stated that biological species in corals are "the sum of a series of forms of variation which differ in relative importance from species to species." Recent attempts to determine the range of variation which may exist within a single species have revealed the relative influence of phenotyp- ic and gcnotypic variation in skeletal de- velopment, and the contributions of each may vary between species or genera (see re- views by Wijsman-Best 1974; Foster 1979. 1980). Lang (1984) reviewed the utility of non-skeletal characters to aid in the differ- entiation of species when skeletal characters are ambiguous or environmentally variable. Recent studies have revealed a wide range of variation in the types of skeletal struc- tures produced by colonies of Astrangia pocii/aia. Field (1949) reported that branch- ing forms might be found when the colonies were located in a protected spot. Cummings (1983) investigated patterns of morpholog- ical variability in colonies of A. poculata from Rhode Island and the Gulf coast of Florida and found a high degree of mor- phologic plasticity. She noted that the pa- rameters measured (height/length, branch- ing, and polyp density) were correlated with exposure (light and wave action), and did not differ significantly between zooxanthel- late and azooxanthellale colonies. Colonies from Panacea, Florida were, however, more branched than those from Fort Wethenll, Rhode Island, and contained more zooxan- thellae per unit area of tissue. The Fort Wclherill colonies possessed more corallites per unit area and were smaller than else- where. As depth increased, colonies became taller and more branched, and polyp density decreased. Brunet (1982) described colonies from various sites off the Texas coast. En- crusting, mound and digitate colonies were found at the Port Aransas jetty on the chan- nel at the end of the barrier island. Mostly mounding and a few encrusting colonies, only rarely with early digitate morpholo- gies, were present on the pilings of the near- shore oil rigs he examined. He suggested that the digitate growth form was a response to sedimentation, to allow the polyps to ex- tend above the substratum. He also noted that the appearance of filamentous algae growing on dead portions of the corallum might force adjacent polyps to continue growing and overtop the algae to form char- acteristic knobby branches. The large ex- pansive polyps of the star coral are well- suited to removing sediment particles that may fall on its surface (Peters 1984). How- ever, we (ECP, CEV. LSG) have observed that during periods of cold weather (in both Rhode Island and Texas corals) the coe- nosarc tissues of .-t. poculata often contract and expose the coenosteum (as the polyps contract and forego feeding), which may be covered with sediment or coloni/ed by al- gae. These lesions may contribute to the formation of digitate colonies when the cor- al resumes skeletal deposition in the spring, although this has not been confirmed ex- 244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON perimentally. St. Pierre (1986) also ob- served high variability in colony morphol- ogy from encrusting to ramose or both types on the same colony, at the Port Aransas jetty. Jaap (in prep.) noted that substratum may also control colony morphology, with encrusting forms found on mollusk frag- ments or shells and thicker or branching forms on igneous rock bases. As for other skeletal characters. Jaap (in prep.) reported that Gulf of Mexico specimens generally had smaller corallite diameters and there were reduced numbers of septa in specimens tak- en from Texas or the western Gulf sites. He observed a minimum annual growth rate (basal diameter) of 38.4 to 46.8 mm for two clusters of Aslrangia which attached to a stone crab trap between April and Septem- ber 1977 (Jaap. in prep.). The largest branching specimen in the USNM (USNM 78508, Fig. 4) was collected from 263 m depth off Western Dry Rocks, Sand Key, Florida. The cellular composition and structure of Astrani^ia poculata arc distinctive, particu- larly in the development of the mesogleal pleats and the granules in the cells of the aboral gaslrodermis and calicoblast epider- mis, and differ from other members of this and closely-related families, such as the Oculinidae (see Peters 1984). .1. poculata is one of the few subtropical to temperate cor- als that may possess symbiotic dinoflagel- late algae known as zooxanthellac within its gaslrodermal tissues, and is the only mem- ber of the genus reported to do so. Cum- mings (1976). Jacques et al. (1977), and S/mant-Froelich (1980a) showed that zoo- xanthcllac densities in some colonies from Narragansett Bay may be equal to or greater than those reported to occur in tropical reef corals. In Narragansett Bay. colonies with /ooxanthellae are found directly adjacent to those without, and there are colonies that are only partially pigmentcd (spotted) brown by the algae. All varieties are found through- out the year. Colonies which appear to lack zooxantheiiae usuaih possess small popu- lations of the algae (less than 1 X 10'cells per polyp). Hence, Swart's (1983:72) argu- ments that colonies of/I. danae (=.-1. poc- ulata) that lack zooxanthellac might be a different species, or might be found under stress conditions, appear to be invalid. Algal populations do vary seasonally with changes in light intensity and temperature (Cummings 1983), and under sedimenta- tion stress (Peters & Pilson 1985). The algal populations also increase under conditions of high light and temperature in the labo- ratory (Cummings & McCarty 1982, Cum- mings 1983, Peters & Pilson 1985). Cum- mings (1976, 1983) suggested that this change was caused by differences in phys- iological tolerances between zooxanthellate and azooxanthellate colonies, with the for- mer belter adapted to summer (high tem- perature and light) and the latter better adapted to winter (low temperature and light) conditions. Possibly, because of the seasonal fluctuations of these conditions, fully zooxanthellate colonies are not nu- merically dominant in Narragansett Bay. Jacques (1978) observed that at high light and temperature conditions, the zooxan- thellate colonies calcified significantly faster than a/ooxanthellate ones. Peters & Pilson (1985) found that the tissues of zooxan- thellate colonies were more easily damaged by combined sedimentation and starvation stress than azooxanthellate colonies, and suggested that genetic as well as environ- mental factors influence the symbiosis. Weston (1980) found only zooxanthellate colonies in the relatively well-illuminated surface waters of Chesapeake Bay and azooxanthellate colonies deeper. Both kinds were found inhabiting a transition zone, the depth of which correlated with 89 to 95% attenuation of surface incident light. He also sugested that both genetic and en\ironmen- tal factors might influence the symbiosis. The colonies which Brunei (1982) found on shallow substrata (less than 2 m water depth) at thr Port .Aransas, Texas, jetty site con- tained zooxanthellac. At depths greater than VOLUME 101, NUMBER 2 245 2 m, and at the oil rig and the Liberty ship, specimens of Astrangia always lacked zoo- xanthellae. Colonies oi Astrangia poculata tolerate a wide variety of environmental conditions. In nature, populations have been found in salinities ranging from oceanic (36%o) sea- water (Cummings 1976) to salinities of 16- 19%c in Texas bays (Ladd 1951). Experi- mentally, this coral can adapt to salinities ranging from 10 to 40%c (Ferguson 1948, Cummings 1976) and can survive for more than six days under low oxygen conditions (Sassaman & Mangum 1973). In Naragan- sett Bay, Rhode Island,.). poculata survives an annual temperature range of - 1.5?C to 22?C, and up to 35?C under experimental conditions in the laboratory (Cummings 1976, Jacques 1978, Peters & Pilson 1985), St. Pierre (1986) observed annual temper- ature fluctuations from 17?C to 30?C during a study of Astrangia from Port Aransas, Texas, The unusual hardiness of .4. poculata is illustrated by the fact that MEQP (un- published) often kept colonies in small con- tainers in his office. In one case a small col- ony was maintained for more than three years in a one-liter container, often without feeding it or changing the water for months. The sexual reproduction of .4. poculata in Narragansett Bay, Rhode Island, was ex- amined by S/mani-Froelich et al. (1980). They demonstrated that each colony is dioecious, with a 1:1 male : female sex ratio for the population. Fertilization is external, with development of planktonic planula lar- vae, and zooxanthellac are not passed on in the eggs or sperm but must be acquired after settlement. .An annual reproductive cycle appeared to be tied to the seasonal maxi- mum temperature. Well-fed colonies, ex- perimentally maintained year-round under high light and temperature (20-22?C or 26?C) conditions continuously produced gametes (Szmant-Froelich 1981. Peters & Pilson 1985). St. Pierre (1986) studied the repro- ductive cycle of a population of Astrangia from a jetty ofi' Port .Aransas, Texas. She also found a 1:1 sex ratio. Gametogenesis began in February, with spawning occurring in late August to September, corresponding to the maximum water temperatures at this Gulf of Mexico site. Another cycle of ga- metogenesis appeared to begin in Novem- ber. Similarly, a second cycle was reported for the Rhode Island colonies beginning in October. Because the reproductive cycle of this coral appears to be essentially the same throughout its range, St. Pierre supported the synonymy of A. danae with A. astrei- formis (Si. Pierre 1986). Acknowledgments We thank all of the "Friends nf Astran- gia" whose interest in this coral over the years prompted this work. We are also grateful to C. J. Durden and H. B. McCarty for helpful discussions and manuscript re- view, and to F. M. Bayer and C, W. Sa- brosky for advice on nomenclature leading to the choice of neotype for both poculata and michelinii. Publication n.s.-3 of the Texas Memorial Museum. University of Texas. Literature Cited Agassi/, A.. & E. C. Agassi/.. 1865. 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National Museum of Natural His- tory, Smithsonian Instutition, Washington, D.C, 20560; (MEQP) Graduate School of Oceanography, University of Rhode Island, Narragansett, Rhode Island 02882-1197; (JWW) Department of Geological Sciences, Cornell University, Ithaca, New York 14853-1504; (WCJ) Florida Department of Natural Resources, Bureau of Marine Re- search, 100 Eighth Avenue S.E., St. Peters- burg, Florida 33701;(JCL) Texas Memorial Museum, University of Texas, Austin, Tex- as 78705, and Department of Zoology, Uni- versity of Texas, Austin, Texas 78712; (CEV) Amoco, 200 East Randolph Drive, Chicago, Illinois 60601; (LSG) Biology De- partment, Texas A&M University, College Station, Texas 77843. ; .'i; i.. i L