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- ilar Astrangia astreiformis from the south- cent nominal species within the family Rhi- eastern coasts of the United States. How- zangiidae, and is represented by 20 fossil ever, other texts suggested that only one species, some of which lived as early as the species o{ Astrangia existed along the entire Middle Cretaceous, approximately 100 mil- North Atlantic and Gulf coasts of North lion years ago (Wells 1956). Members of the America (Hyman 1940, Wells 1956, for re- genus are all relatively small and are usually views see Cummings 1983, and Jaap, in insignificant components of the fauna where prep.). A. astreiformis was also reported to they occur. The various species of Astran- occur off the west coast of Africa (Thiel 1928, gia, however, collectively occupy a vast geo- Chevalier 1966). graphic range, possibly broader than that of Throughout its range, the northern star any other genus of coral. coral appears remarkably tolerant of a wide Unfortunately, the genus Astrangia has range of environmental conditions. Colo- never been completely reviewed. It is prob- nies oi Astrangia are generally considered able that many species names are syn- to be ahermatypic (non-reef-building) and onyms, and also that a number of yet un- hence azooxanthellate (=asymbiotic, i.e., discovered species may exist (Pilson 1975). lacking zooxanthellae), but some colonies The confusion that exists within this genus of the northern star coral contain these algal became evident when a group of scientists cells abundantly within their tissues (Bosch- at the University of Rhode Island began a ma 1925a). These two factors, adaptability series of studies on the ecology, physiology, and variability in algal symbiosis, have con- and life history of the coral found in Nar- tributed to the increasing popularity of this ragansett Bay. Whereas several previous coral as an experimental laboratory animal. studies had referred to the northern star cor- Therefore, in order to aid future researchers, al as Astrangia danae, a number of other we were motivated to clarify the taxonomic papers had been published on the very sim- status of this coral.

VOLUME 101. NUMBER 2 235

Family Rhizangiidae d'Orbigny, 1851 1986:6 (cited as the type species). [Emended spelling of .4. michelin;'.] Genus Astrangia Milne Edwards .istrangia danae ME & H, 1849:180.-Ver- & Haime, 1848a rill, 1863:40; \i6AAl (not A. danae .\g'Si%- Diagnosis.—Co\on\a\, usually azooxan- siz 1850); 1866a:324 (=A. edwardsii); thellate. Colonies encrusting or subplocoid, I866b:335; 1872:432-*38; 1874:446 (ge- formed by extratentacular budding from ography ).—Pearse & Wharton, 1938: edge zone; corallitcs united basally by thin 636.-Cairns, 1979:207.-Jacques & coenosteum, septa dentate; columella pap- Pilson, 1980:167-178 (physiology, illary. [Type species: .-1. michelinii Milne symbionts, Rhode Island). —Szmant- Edwards & Haime, 1848b, by monotypy Froelich, 1980a: 1-191 (natural histoo', (=A. poculala Ellis & Solander, 1786).] biochemistry, Rhode Island). —Szmant- Froelich & Pilson, 1980:85-97 (feeding Astrangia poculala frequency, biochemistry, Rhode Is- (Ellis & Solander, 1786) land). -Szmant-Froelich, 198 i: 1 33-144 Figs. 1-6 (nutrition, Rhode Island). —Cummings, 1983:1-147 (physiology, morphology, "Stony coral found on the shore near New Rhode Island, Honda).-Peters, 1984:61- York.": Ellis, 1755:84, pi. 32, figs. A. 1, A.3. 63. figs. 6b, c. d, I Od, e, 13g, 1 7a, b, 22h, 'IMadrepora calycularis Pallas, var. li, 1766: 26f g (histology, Rhode Island). —Wise- 319 note 186B.-Espcr, 1790:283. man, 1984:1-75 (biochemistry).-Peters Madrepora poculala Ellis & Solander, 1786: & Pilson, 1985:215-230 (sedimentation 165.-Gmelin, 1791:3766.-Esper, 1790: stress, histopathology, Rhode Island). 283 (=.V/, calycularis 0). [Original type [Type locality unknown. Type specimen locality "near New York." Type speci- last seen by Wells in 1934, now missing men lost (P. F. S. Cornelius, pers. from Coll. Michelin (Cairns 1981).] comm.).Type locality (ncotype) "off At- .-Istrangia astreiformis ME & H, 1849:181.— lantic City, New Jersey." Neotype de- Verrill, 1864:47; 1866a:324.-Vaughan, posited in the National Museum of Nat- 1901:300, pi. II, figs. 2, 2a, 2b (Puerto ural History USNM 80350.] Rico). —Whittcn et al.. 1950:75 (Texas Madrepora porculaia: Turton, 1806:623 channels). - Parker, 1959:2130, 2131, fig. [Misspelling of/?on7)f»v»

VOLUME 101, NUMBER 2 n '••• 237

vival, Texas).-St. Pierre, 1986:1-55 (re- 55_65.-Calder, 1972:101 (Chesapeake production, Texas). (Type locality Bay, Hog Island and Eastern Shore).— "United Stales." Type found by Wells in Sassaman & Mangum, 1973:1313-1319 Paris ir Coll. Michelin in 1934, but not (anerobic metabolism).—Jacques et al., found by Chevalier in 1980 (Cairns 1977:455-461 (physiology, calcification, 1981).] Rhode Island). —Hayes & Goreau, 1977: Astrangia danae Agassiz, 1850:68-77, pi. I, 26-40 (calcification, histology). —Bach- fig. 7 (feeding behavior, digestion, colony and, 1978:283-284 (habitat, life histo- growth, new polyp formation, nemato- ry).-Calder & Hester, 1978:93 (Charles- cyst function, Massachusetts). —Verrill, ton, South Carolina). —Weslon, 1980:1- 1863:40 (North Carolina); 1864:47 (=A. 93 (zooxanthellae distribution, Chesa- astreiformis in Leidy, 1855, not A. danae peake Bay). [Type locality "Massachu- ME & H 1849); 1866a:324 (not A. danae setts." Deposition unknown.] ME&H 1849); 1866b:335; 1872:436,437 Astrangia aslraeformis: ME & H: Leidy, (=A. astraeifonnis ME & H 1849); 1873: 1855:139-140, pi. 10, figs. 9-16 (synon- 408; 1874:446 (Vineyard Sound and ad- ymy, nematocysts, Rhode Island). [Mis- jacent waters).—Agassiz& Agassiz, 1865: spelling of A. astreiformis ME & H 1849]. 16, figs. 16-20.-Pourtales, 1871:80.- Astrangia astraeiformis: ME & H: ME & H, Dana, 1872, 1890:67-68, figured p. 68, a, 1857:614.-Verrill, 1863:39; 1866a:324; b,c.-Damon, 1882:22 l.-Fewkes, 1889: 1866b:335; 1872:436 (=A. danae Agassiz 7-10, pis. 1-5 (natural history illustra- I850).-Pourtales, 1871:80.-«of Thiel, tions); 1891:53.-Hargitt, 1914:250, pi. 1928:283-285, pi. 2, fig. 22 {=A. mer- 43, fig. 9 (natural history, Casco Bay, catoris Thiel 1941, West Coast of Afri- Maine). —Boschma, 1925a:65-67 (zoo- ca).-Thiel, 1941:5, 15, pi. 1, figs. 5-6 xanthellae symbiosis); 1925b:407-439 (Martinique). —Pearse & Williams, 1951: (feeding behavior, digestion, zooxanthel- 136 (incorrectly attributed to Leidy, 1855, lae symbiosis). —Cowles, 1930:333 Maine to Florida and the West Indies).— (Chesapeake Bay, Cape Henry, Cape Chevalier, 1966:926-930, pi. 3, figs. 2-3, Charles).-Pratt, 1935:159, fig. 239.- pi. 4, fig. 6 (synonymy, description. Sierra Pearse, 1936:178.-Richards & Harbi- Leone, Africa). [Misspelling (or corrected son, 1942:178, pi. 22, figs. 2 & 3 (Mio- spelling) of .4. astreiformis ME & H 1849.) cene, New Jersey). —Vaughan & Wells, Astrangia danai: ME & H, 1857:614. [Mis- 1943:177 (see footnote 52: =1A. michelini spelling (or corrected gender) of .4. danae ME & H I848).-Field, 1949:6, 18, pi. I. ME& H 1849.] fig. 7.-Waterman, 1950:127-131, fig. pp. Astrangia edwardsii Verrill, 1866a:324 120-121 (structure of corallum and pol- (=nom. nov. for .4. danae ME & H 1849, yps, zooxanthellae, feeding, reproduc- see Remarks); 1866b:335. tion).-Alloiteau, 1952:623, fig. 74 (=.1. not Astrangia mercatoris Thiel, 1941:16- michelini ME & H 1848).-Hand, 1964: 17, pi. 2, fig. 1 (=.4. astraeiformis Thiel, 25, pi. 3, fig. 4 (description). —Driscoll, 1928:283-285, pi. 2. fig. 22; Angola. Bay 1967:633-641.-Robertson et al., 1970: 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- on). —Cummings, 1976:1-115 (physiol- onymy, see Remarks). ogy, Rhode Island). —Szmant-Froelich & Astrangia astraeiformis: no authority cited: Pilson, 1977:417^24 (nitrogen excre- Hedgpeth, 1953:164 (Texas). [Misspell- tion, symbiosis, Rhode Island). —Hoff- ing oi A. astreiformis.] man et al., 1978:165-170.-Jacques, not Astrangia astraeformis: no authority cit- 1978:1-169 (calcification). -Gosner, ed: Buchanan, 1954:85-86 (=A. astraei- 1979:93-94,fig. 20.-Storer et al., 1979, formis Thiel 1928 =A. mercatoris Thiel figured on p. 320, 372, 390.-Szmant- 1941, Ivory Coast, French Congo, An- Froelich, 1980b: 1-3 (popular account).— gola, Africa). [Misspelling of,'!, astreifor- Szmant-Froelich el al., 1980:257-269 mis.] - - . • - 1- :• (reproduction, Rhode Island). —Mein- Astrangea danae: no authority cited: Go- kolh, 1981:391.-Cummings & Mc- reau & Bowen. 1955:1 188 (calcification). Carty. 1982:1125-1129 (calcification, [Misspelling of Astrangia danae.] stable carbon isotopes, Rhode Island).— Jacques et al., 1983:135-148 (ecology, .Astrangia asireformis: Milne Edwards & metabolism, Rhode Island). —Swart, Haines(sic): Pampe, 1971:399. pi. 2. fig. 7 (Pleistocene, Texas). [Misspelling, 1983:72 (stable carbon isotopes).— spelled .t. astreiformis later on same page.] Szmant-Froelich & Pilson, 1984:153-162 (feeding frequency, nitrogen metabolism, Astrangia asteriformis: no authority cited: respiration, Rhode Island). Rudloc, 1971:154-157, 167, 172-174, figured on p. 174 (natural history, har- Astrangia sp.: Ferguson, 1948:17 (salinity diness in aquarium. Florida). [Misspell- tolerance. Louisiana). —Keith & Weber, ing.] 1965:500 (isotopic composition, calcifi- Astrangia a.streiformis: no authority cited: cation, in part: Woods Hole, Massachu- Ladd. 1951:137, 139, 149 (Texas'coastal setts).-«of Rabalais, 1978:62 (= Oculina waters). —Slephenson & Stephcnson, diffusa, G\x\f of Mt\ico). 1952:35 (distribution, taxonomy).— Diagnosis. —EncTU%\\n%, massive, or ra- Hedgpeth, 1953:164 (Texas).-Gunter & mose irregularly shaped colonies. Corallum Geyer, 1955:40,44,45, 56. 57, 65 (Texas, cerioid to plocoid. new polyps produced by Louisiana).-Parker, 1956:310, 315, 329, extratentacular budding, having coralliles 354, 370 (dead specimens collected in east united basally by thin peritheca (vesicular Mississippi delta region). —Courtney & coenosteum). Corallites round, irregular, or Edmond, 1974:63 (Marco Island, Flori- polygonal; tightly packed or spaced several da).—Folheringham & Bruncnmeister, millimeters apart, highly variable even 1975:36. 161. fig. 2.24 (natural history). within a single specimen. Caliccs 1-7 mm .istrungia danae: no authority cited: Wil- in diameter, with distinct equal costae on son, 1900:350 (North Carolina).-Allee, many corallites. Septa hcxamerally ar- 1923:176 (Woods Hole. Buzzards Bay, ranged in 4 cycles, however, complete fourth and Vineyard Sound, Massachusetts).- cycle (48 septa) only attained in very large H\man, 1940:610-611 (morphology, calices. Most calices have 24 septa, the S, natural history). —Storer. 1951, figured on merging basally with the S^ about 'A distance p. 321. —Stephenson & Stephcnson, 1952: to columella. With increase in calicular di- 35 (distribution, taxonomy).-Blake. ameter, pairs of Sj form flanking some S,, 1953:23 (Pleistocene, Maryland). —Zinn. often with one pair per system (the space 1964:15 (Rhode Island).-Goreau et al.. between two S,) resulting in 36 septa. Com- 1971:254.-Williams& Murdoch. 1973: plete fourth cycle (48 septa) attained only 551-563 (elTccts of gamma radiation in caliccs over 6 mm in diameter. Septa 239 VOLUME 101, NUMBER 2 New York" figured by Ellis (1755). Another Strongly dentate, particularly S, at region of reference to this "American" coral ap- S, fusion, and slope gradually toward col- peared in Pallas (1766) under the name umella. Fossa shallow. Columella trabecu- Madrepora calycularis. var. 0. Typical M. lar or papillose. calvcularis Pallas (1766) is now known as Polyps up to I cm tall and translucent Astroides calvcularis (Pallas, 1766), found white (azooxanthellate = asymbiotic) to only in the Mediterranean (Zibrowius 1980). brown (zooxanthellate = symbiotic, see Schuhmacher&Zibrowius 1985. for review Therefore, the name calycularis. even if ap- plied to a specimen of western Atlantic As- of this terminology). Numerous nematocyst irangia as form /3, is preoccupied by the batteries on surface of tentacles; tip of ten- Mediterranean species. Esper (1790:283) tacle rounded. Directive mesenteries pres- equated M. poculata with the M. calycularis ent. Corallum often invaded by endolithic li of Pallas, describing a corallum encrusting red or green boring algae, burrowing poly- an American gastropod, which may have chaetes, or sponges (e.g., Cliona cclata Grant, been the same as the astrangid-encrusted see Cummings 1983). specimen of Conus spurius from the West Cnidocyst batteries principally composed Indies, figured by Seba (1758:152, pi. 55, of spirocysts. Terminal knob packed with large microbasic-p-mastigophore ncmato- figs. 11. 12). The last known reference to M cysts (about 29 tim long x 8 Mm in diameter poculata was Turton (1806). ME & H (1848b) described Astrangia in fixed, paraffin embedded sections) and michelinii as an illustration found in Mem- spirocysts. Occasional small microbasic-p- oire 4, Monographic des Astreides; this is a mastigophores scattered in epidermis (about valid "indication" (International Code of 20 Mm long x 7 ^m in diameter). Cnido- Zoological Nomenclature (ICZN) 1985: Ar- glandular band of mesenterial filaments possesses large microbasic-p-mastigo- ticle 16, vii), and therefore it is a valid species and the type species oi Astrangia by mono- phores (about 40 iim long x 7 ^m in diam- typy. It was described without a type local- eter), holotrichous isorhizas (about 45 ^m long X 15 Mm in diameter), and small mi- itv. In 1849, ME & H described (in order) 3 crobasic mastigophores (?) (about 11 Mm long X 5 ^m in diameter). The distribution of types in the Museum National d'Histoire Naturelle, Paris, (MNHNP), .istrangia dan- these nematocysts vanes with location on ae. .-istrangia michelini. and Astrangia as- the filament (ECP. unpublished). Histolog- ireiformis. The descriptions of the type ically distinguished by well-developed lon- specimens were very similar. They noted gitudinal retractor muscles along the me- soglcal pleats. Gastrodermis lining that .1. michelini was closely related to A. danae. Unfortunately, only the type locality calicoblastic epidermis features large cells of one of the 3 species, .1. astreiformis. was with basophilic globules (osmiophilic, non- PAS positive, non-mucin) up to 4 Mm in given, as from the "'United Stales." Agassiz (1850) described the same species diameter, which increase in number and from the United States and used one of the density towards base of the polyps. Cali- same names used by ME & H. .Astrangia coblastic epidermis thickened with fine eo- danae. in a paper presented before the sinophilic granules (Peters 1984). Colonies American Association for the Advance- gonochoric (=dioecious). gametes spawned annually (Szmani-Froelich el al. 1980. St. ment of Science meeting in 1849. but it was not published until 1850. He made no com- Pierre 1986). Remarks.-Madrepom poculata was de- parisons and apparently did not know that ihe name had alreadv been used for a species scribed by Ellis & Solander (1786), based on the illustration of a specimen from "'near in the previous vear (ME & H 1849). and 240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

therefore was a junior primary homonym. noted that some authors would unite all three It is curious that Dana (1872,1890) referred species. Pourtales (1871:80) remarked "The to A. danac Agassiz, suggesting that he had differences between these two species (A. as- not been in correspondence with ME & H treiformis M. E. & H. and A. danae Ag.) are m the late 1800's. Dana noted that he him- of the slightest character, being chiefly de- self had proposed the name Pleiadia for the rived from the mode of grouping, which is genus, alluding to the crowd of stars (the denser in the first than in the second. I am appearance of the corallum), in his report strongly inclined to believe that they will on Zoophytes (Dana 1846:722, Wells 1986: ultimately be united." In 1872, Verrill united 50, nom. nud.). However, Agassiz (1850: .(. astreiformis ME & H with A. danae Agas- 69) made no reference to this particular re- siz. Cummings (1983) and Jaap (in prep.) port. That he chose to call his species As- united A. astreiformis ME & H with A. dan- irangla presumably means that he had seen ae Agassiz, noting the great variability in the figure (if not the specimen on one of his specimens collected from throughout its European trips) of A. michelinii published range along the coastline of the eastern in 1848. United States. Only Vaughan (1901) felt that Later authors, who had compared the type there were sufficient distinctions based on specimens and descriptions, commented on the skeletal structures to maintain A. as- the relations of these species. Vaughan treiformis ME & H and A. danae Agassiz as (1901:300) noted that the specimen of As- separate species. trangia astreiformis (corallites free above Thus, these observations link Astrangia their bases), which he had collected in Puer- astreiformis ME & H with .i. danae Agassiz. to Rico, was not typical of the specimens of .Although A. danae ME & H appeared to be A. astreiformis then in the USNM, and ob- very similar to .1. astreiformis ME & H (Lei- served that the figure of A. michclini ME & dy 1855. Chevalier 1966) and A. michelini H "suggests great similarity to the specimen ME & H (Milne Edwards & Haimc 1849), from Puerto Rico." Alloiteau (1952) be- subsequent authors (Verrill 1863, 1866a) did lieved that A. michelini ME & H was the not connect .1. danae ME & H with .4. danae same species as A. danae Agassiz. Chevalier Agassiz. Verrill 11866a:324) stated again that (1966) observed that A. astraeiformis ME ,1. danae ME & H ("the corallites being scat- & H was similar to A. michclini of ME & tered and connected only at the base") was H. Thus, A. michelinii Milne Edwards & different from .4. danae Agassiz, and there- Haimc is "similar" to A. danae Agassiz fore proposed the name .4. edwardsii as a (Vaughan & Wells 1943, Alloiteau 1952), replacement name for .4. danae ME & H. .1. astreiformis ME & H (Vaughan 1901. He erroneously thought that the date for Chevalier 1966). and A. danae ME & H .Agassiz's name was 1849 and the dale of (Milne Edwards & Haime 1849). ME & H's was 1850, because this latter date Leidy (1855) considered Astrangia as- was cited in error by ME & H (1857:614). treiformis ME & H to be the same as .1. A number of later references attributed the £/(3/2ac Agassiz. In 1863, Verrill recorded that publication date of .Agassiz's name to 1847 .-1. danae Agassiz was equal to the A. as- (e.g., Calder 1972) or 1848 (e.g.. Field 1949), treiformis descnbed by Leidy in 1855, and further adding to the confusion. Fewkes noted that A. danae Agassiz was not the (1889:8) noted that "The polypdom of .4. same as A. danae ME & H. Verrill (1866a) Danae (sic) M. E. & H., is held by Prof reported that .1. bella E. & H. [ME & H], Joseph Leidy to be different from that of known from the Miocene formations in our New England species of .4strangia and Maryland, was vcrv similar to both A. danae more like .4. astraeformis of the same au- Agassiz and A. astreiformis ME & H, and thors .... It is not possible to determine VOLUME 101, NUMBER 2 241

from the description of A. Danac given by specimens collected from south of Cape Milne Edwards & Haimc whether our Hatteras into the eastern and .vestem Gulf species differs from that which they describe of Mexico, respectively. Wells observed that under that name or not." A. danae ME & H had priority as the senior J. W. Wells examined the type specimens synonym. in the MNHNP in 1934. Of Aslrangia mi- Cairns (1981) argued that Astrangia as- chelim ME & H 1848 (No. 412, Michelin treiformis was the appropriate name for the Coll.), he noted "Does not look like the fig- American species. He noted that ME & H ure but has M.E. & H.'s label on the wooden described both A. danae aind A. astreiformis base. Very like .-1. danac [no authority cited] in the same paper. Although danae\\did page except for smaller [corallites] (2.5-3.0 mm). priority, the type was lost, the description It is attached to a bit of igneous rock. This poor, and the type locality not given. He is the type species of Asirangia by mono- could not confirm that A. danae ME & H typy. No loc." (J WW notes, February, 1934). was the same as A. danae Agassiz and A. He also observed that in the Paris collection astreiformis ME & H. hence, he considered "Specimens labeled .1. JanaclME & H 1849] A. danae ME & H to be a nomcn dubium. are from Si. Thomas and are A. soliiaria. The locality for A. astreiformis was given as coll. Duchassaing ca. 1846, except for one "United States." Although the type of .^. that appears to be the type. It encrusts a astreiformis was also lost, it at least had a Tellina and fits the description well and is type locality, which is why Cairns consid- evidently conspecific with A. michelini as ered it as the senior synonym. Because latter was figured in 1848 by M.E.&H. Has Agassiz's description of the same species, M.E.&H.'s label. No loc." Wells noted that which he named A. danae, was not pub- the type of .-t. astreiformis ME & H 1849 lished until 1850, Cairns also considered was in the Paris collection in 1934, with the this name a junior synonym of .1. astreifor- locality designated "Etats-Unis," but did not mis and junior homonym of .-1. danae ME make any other observations on this spec- & H 1849. imen (JWW notes, February, 1934). From Based on our present examinations of all indications (Cairns 1981), all of the types specimens and recent biological data (see are now missing from the collections of the below), we concur that there is only one MNHNP. species of star coral of the genus Astrangia Wells considered Astrangia danae ME & along the Atlantic and Gulf coasts of North H 1849 to be the proper name for the Amer- America. Although the type specimens are ican northern star coral (pcrs. comm. to A. now missing from the Michelin collection, Szmant-Froelich, 13 December 1977). not- earlier observations by Leidy, Verrill. Pour- ing that "Over the years 1 have examined tales. Fewkes, Wells. Alloitcau. and Che- many specimens from throughout this range valier suggest that the Astrangia types es- [Massachusetts to Texas] and although there tablished by ME & H were very similar. The is much variation in skeletal morphology, latter four authors thought that these types the variation seems not be geographical or were probably only morphological variants provincial, for specimens from one locality of the colonial northern star coral. There is frequently show the full range of vanation no doubt that this species is the same as the in compactness or isolation of corallites. size Madrepora poculata described by Ellis & of corallites. equality of septa, etc." Al- Solander over 200 years ago. Although the though the names A. danae Agassiz and .1. names .(. danae and A. astreiformis have astreiformis ME & H had been used nom- been used widely in the recent 'iterature (73 inally to separate specimens from north of and 41 references, respectively), these names Cape Hatteras to New England from those must be considered junior subjective syn- 242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

onyms. We conclude that the correct name, loris as the African species. Buchanan (1954) based on the Principle of Priority (ICZN reported A. astreiformis from the Ivory Coast 1985), is Astrangia poculata (Ellis & Solan- and Gulf of Guinea off western Africa. Bu- der, 1786). chanan (1954:86) remarked that his speci- The type specimen of Madrepora pocu- mens agreed well with Thiel's (1928) de- lata is not present at the Glasgow or the scription of A. astreiformis from Africa. British Museum (Natural History) and is Chevalier (1966), however, believed that the believed to be lost (P. F. S. Cornelius, pers. American and African specimens that he comm.). In order to help clarify the complex examined were sufficiently similar to merit nomenclatural issues concerning the names recognition as the same species. Unfortu- Astrangia poculata. A. michelinii ME & H nately, as we know nothing of the African 1848, A. danae ME & H 1849, A. astreifor- Astrangia. Thiel's (1928) specimens may in- mis ME & H 1849, and A. danae Agassiz deed represent a different species. 1850, a neotype is chosen for A/, poculata. Material examined. —¥rom the USNM, It isdeposited at the USNM (USNM 80350) 85 lots, the specimens representing the fol- and was collected from New Jersey, rela- lowing localities: Martha's Vineyard Sound, tively close to the original type locality of Buzzards Bay, Woods Hole, Massachusetts; "ofTNew York." Newport. Rhode Island; 19 lots from R/V As mentioned before, there is no type (iosnold collected off northeastern Florida specimen of the type species Astrangia mi- and Georgia (13-54 m); Chesapeake Bay, chelinii (see Chevalier 1966). Furthermore, Virginia Beach, Virginia; Cape Henry, North it was described without a type locality and Carolina; Myrtle Beach, South Carolina; its original description and figures do not Nassau County, Florida; northeastern coast reliably distinguish it from the other ap- of Florida; Dry Tortugas, Western Dry- proximately 30 Recent species in the genus. Rocks off Sand Key, Florida; Marco. Gulf Subsequent references to A. luichelinii have coast of Florida; Rockport, Galveston, Port always been in its capacity as the type species Aransas. Texas. of the genus, not as a new record. Therefore, Specimens from the Texas coast depos- in order to redefine A. michelinii. and the ited in the Texas Memorial Museum, Aus- genus Astrangia by virtue of its status as the tin, Texas (Numbers 1568-1573. 1753). type species of the genus, a neotype is cho- Specimens from the Hourglass Cruises, sen for .1. michelinii. Because several au- deposited in the collection of the Depart- thors (e.g., Vaughan 1901, Alloiteau 1952, ment of Natural Resources, Bureau of Ma- Chevalier 1966) have suggested that .1. rine Research. St. Petersburg, Florida michelinii is synonymous with the United (Numbers FSBC I 10027. 11360, 17984. Slates east coast species, the neotype chosen 19858-19862) and 49 other cataloged spec- for.(. michelinii is a specimen of .1. pocu- imens (various numbers from FSBC I lata. in fact the same specimen chosen as 10028-31954, eastern Gulf of Mexico and the neotype of .\/. poculata. a specimen from Florida east coast), as well as specimens on New Jersey (USNM 80350). A. michelinii loan from Texas A&M University (College thus becomes a junior objective synonym Station. Texas), and Museum of Compar- of .1. poculata. ative Zoology (Harvard University, Cam- Thiel (1928) described a colonial .Astran- bridge, Massachusetts), representing loca- gia from the Atlantic coast of Africa, and tions from New England to Florida. identified it as .4. astraeiformis ME & H. In Louisiana, and Texas. 1941, Thiel doubted the existence of the (jcologic and geographic disinhution.— American species on the west coast of Af- .Astrangia poculata is recorded from the rica, and established the species .\. merca- Miocene (Richards & Harbison 1942). VOLUME 101, NUMBER 2 243

Pleistocene (Blake 1953, Pampe 1971) and ical variability in colonies of A. poculata Recent (Alloiteau 1952, Wells 1956). It oc- from Rhode Island and the Gulf coast of curs at depths from 0 to 263 m. It has been Florida and found a high degree of mor- reported off Maine (Hargitt 1914) and from phologic plasticity. She noted that the pa- Cape Cod south along the eastern United rameters measured (height/length, branch- States, around the Gulf coast of Florida to ing, and polyp density) were correlated with the southern Gulf coast of Texas and Mex- exposure (light and wave action), and did ico (E. Jordan pers. comm. to WCJ). It is not differ significantly between zooxanthel- very rare in the coastal region from south late and azooxanthellale colonies. Colonies of Palm Beach to the Ten Thousand Islands, from Panacea, Florida were, however, more Florida, and is disjunct in the Mississippi branched than those from Fort Wethenll, delta region. It is also noted from Puerto Rhode Island, and contained more zooxan- Rico (Vaughan 1901) and Martinique (Thiel thellae per unit area of tissue. The Fort 1941). Thiel (1928) and Buchanan (1954) Wclherill colonies possessed more corallites reported it from the Ivory Coast, French per unit area and were smaller than else- Congo, Angola, Africa, (although its exis- where. As depth increased, colonies became tence off Africa based on these reports has taller and more branched, and polyp density been questioned, see Remarks), and Che- decreased. Brunet (1982) described colonies valier (1966) reported it from the Bight of from various sites off the Texas coast. En- Biafra, Freetown, Sierra Leone, Konakry, crusting, mound and digitate colonies were Guinea, Dakar and Senegal (Gulf of Guinea), found at the Port Aransas jetty on the chan- Africa. nel at the end of the barrier island. Mostly Discussion.—Corci\ species traditionally mounding and a few encrusting colonies, have been determined on the basis of their only rarely with early digitate morpholo- skeletal morphologies (since these are pre- gies, were present on the pilings of the near- served most easily, and appear in the fossil shore oil rigs he examined. He suggested record). Veron (1982) stated that biological that the digitate growth form was a response species in corals are "the sum of a series of to sedimentation, to allow the polyps to ex- forms of variation which differ in relative tend above the substratum. He also noted importance from species to species." Recent that the appearance of filamentous algae attempts to determine the range of variation growing on dead portions of the corallum which may exist within a single species have might force adjacent polyps to continue revealed the relative influence of phenotyp- growing and overtop the algae to form char- ic and gcnotypic variation in skeletal de- acteristic knobby branches. The large ex- velopment, and the contributions of each pansive polyps of the star coral are well- may vary between species or genera (see re- suited to removing sediment particles that views by Wijsman-Best 1974; Foster 1979. may fall on its surface (Peters 1984). How- 1980). Lang (1984) reviewed the utility of ever, we (ECP, CEV. LSG) have observed non-skeletal characters to aid in the differ- that during periods of cold weather (in both entiation of species when skeletal characters Rhode Island and Texas corals) the coe- are ambiguous or environmentally variable. nosarc tissues of .-t. poculata often contract Recent studies have revealed a wide range and expose the coenosteum (as the polyps of variation in the types of skeletal struc- contract and forego feeding), which may be tures produced by colonies of Astrangia covered with sediment or coloni/ed by al- pocii/aia. Field (1949) reported that branch- gae. These lesions may contribute to the ing forms might be found when the colonies formation of digitate colonies when the cor- were located in a protected spot. Cummings al resumes skeletal deposition in the spring, (1983) investigated patterns of morpholog- although this has not been confirmed ex- 244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

perimentally. St. Pierre (1986) also ob- lations of the algae (less than 1 X 10'cells served high variability in colony morphol- per polyp). Hence, Swart's (1983:72) argu- ogy from encrusting to ramose or both types ments that colonies of/I. danae (=.-1. poc- on the same colony, at the Port Aransas ulata) that lack zooxanthellac might be a jetty. Jaap (in prep.) noted that substratum different species, or might be found under may also control colony morphology, with stress conditions, appear to be invalid. encrusting forms found on mollusk frag- Algal populations do vary seasonally with ments or shells and thicker or branching changes in light intensity and temperature forms on igneous rock bases. As for other (Cummings 1983), and under sedimenta- skeletal characters. Jaap (in prep.) reported tion stress (Peters & Pilson 1985). The algal that Gulf of Mexico specimens generally had populations also increase under conditions smaller corallite diameters and there were of high light and temperature in the labo- reduced numbers of septa in specimens tak- ratory (Cummings & McCarty 1982, Cum- en from Texas or the western Gulf sites. He mings 1983, Peters & Pilson 1985). Cum- observed a minimum annual growth rate mings (1976, 1983) suggested that this (basal diameter) of 38.4 to 46.8 mm for two change was caused by differences in phys- clusters of Aslrangia which attached to a iological tolerances between zooxanthellate stone crab trap between April and Septem- and azooxanthellate colonies, with the for- ber 1977 (Jaap. in prep.). The largest mer belter adapted to summer (high tem- branching specimen in the USNM (USNM perature and light) and the latter better 78508, Fig. 4) was collected from 263 m adapted to winter (low temperature and depth off Western Dry Rocks, Sand Key, light) conditions. Possibly, because of the Florida. seasonal fluctuations of these conditions, The cellular composition and structure of fully zooxanthellate colonies are not nu- Astrani^ia poculata arc distinctive, particu- merically dominant in Narragansett Bay. larly in the development of the mesogleal Jacques (1978) observed that at high light pleats and the granules in the cells of the and temperature conditions, the zooxan- aboral gaslrodermis and calicoblast epider- thellate colonies calcified significantly faster mis, and differ from other members of this than a/ooxanthellate ones. Peters & Pilson and closely-related families, such as the (1985) found that the tissues of zooxan- Oculinidae (see Peters 1984). .1. poculata is thellate colonies were more easily damaged one of the few subtropical to temperate cor- by combined sedimentation and starvation als that may possess symbiotic dinoflagel- stress than azooxanthellate colonies, and late algae known as zooxanthellac within its suggested that genetic as well as environ- gaslrodermal tissues, and is the only mem- mental factors influence the symbiosis. ber of the genus reported to do so. Cum- Weston (1980) found only zooxanthellate mings (1976). Jacques et al. (1977), and colonies in the relatively well-illuminated S/mant-Froelich (1980a) showed that zoo- surface waters of Chesapeake Bay and xanthcllac densities in some colonies from azooxanthellate colonies deeper. Both kinds Narragansett Bay may be equal to or greater were found inhabiting a transition zone, the than those reported to occur in tropical reef depth of which correlated with 89 to 95% corals. In Narragansett Bay. colonies with attenuation of surface incident light. He also /ooxanthellae are found directly adjacent to sugested that both genetic and en\ironmen- those without, and there are colonies that tal factors might influence the symbiosis. are only partially pigmentcd (spotted) brown The colonies which Brunei (1982) found on by the algae. All varieties are found through- shallow substrata (less than 2 m water depth) out the year. Colonies which appear to lack at thr Port .Aransas, Texas, jetty site con- zooxantheiiae usuaih possess small popu- tained zooxanthellac. At depths greater than VOLUME 101, NUMBER 2 245

2 m, and at the oil rig and the Liberty ship, also found a 1:1 sex ratio. Gametogenesis specimens of Astrangia always lacked zoo- began in February, with spawning occurring xanthellae. in late August to September, corresponding Colonies oi Astrangia poculata tolerate a to the maximum water temperatures at this wide variety of environmental conditions. Gulf of Mexico site. Another cycle of ga- In nature, populations have been found in metogenesis appeared to begin in Novem- salinities ranging from oceanic (36%o) sea- ber. Similarly, a second cycle was reported water (Cummings 1976) to salinities of 16- for the Rhode Island colonies beginning in 19%c in Texas bays (Ladd 1951). Experi- October. Because the reproductive cycle of mentally, this coral can adapt to salinities this coral appears to be essentially the same ranging from 10 to 40%c (Ferguson 1948, throughout its range, St. Pierre supported Cummings 1976) and can survive for more the synonymy of A. danae with A. astrei- than six days under low oxygen conditions formis (Si. Pierre 1986). (Sassaman & Mangum 1973). In Naragan- sett Bay, Rhode Island,.). poculata survives Acknowledgments an annual temperature range of - 1.5°C to 22°C, and up to 35°C under experimental We thank all of the "Friends nf Astran- conditions in the laboratory (Cummings gia" whose interest in this coral over the 1976, Jacques 1978, Peters & Pilson 1985), years prompted this work. We are also St. Pierre (1986) observed annual temper- grateful to C. J. Durden and H. B. McCarty ature fluctuations from 17°C to 30°C during for helpful discussions and manuscript re- a study of Astrangia from Port Aransas, view, and to F. M. Bayer and C, W. Sa- Texas, The unusual hardiness of .4. poculata brosky for advice on nomenclature leading is illustrated by the fact that MEQP (un- to the choice of neotype for both poculata published) often kept colonies in small con- and michelinii. Publication n.s.-3 of the tainers in his office. In one case a small col- Texas Memorial Museum. University of ony was maintained for more than three Texas. years in a one-liter container, often without feeding it or changing the water for months. Literature Cited The sexual reproduction of .4. poculata in Agassi/, A.. & E. C. Agassi/.. 1865. Sea-side studies Narragansett Bay, Rhode Island, was ex- in natural history. Boston, 155 pp. amined by S/mani-Froelich et al. (1980). Agassi/. L. 1850. On the structure of coral animals.— They demonstrated that each colony is Proceedings of the American Association for the dioecious, with a 1:1 male : female sex ratio Advancement of Science 2:68-77. for the population. Fertilization is external, Alice. W. C. 1923. Studies in marine ecology 1. The distribution of common littoral Invertebrates of with development of planktonic planula lar- the Woods Hole region. —Biological Bulletin vae, and zooxanthellac are not passed on in 44(4):167-191. the eggs or sperm but must be acquired after Alloiteau, J. 1952. Generalites sur les Coclcnlercs; settlement. .An annual reproductive cycle Hydro/oaires; Scypho/oaires; Alcyonaires: ge- appeared to be tied to the seasonal maxi- neralites sur les Madreporaires; Madrcporaircs post-paleo/oiques. Pp. .^6^10. 5.19-684 in). mum temperature. Well-fed colonies, ex- Pivetcau. ed.. Traile do paleontologie. Volume perimentally maintained year-round under I. Masson. Pans. high light and temperature (20-22°C or 26°C) Bachand. R. G. 1978. Cold water coral —Sea Fron- conditions continuously produced gametes tiers 24:283-284. (Szmant-Froelich 1981. Peters & Pilson Blake. S. F. 1953. The Pleistocene fauna of Wailes blutfand Langleys bluff. N'arvland.—Smithson- 1985). St. Pierre (1986) studied the repro- ian Miscellaneous Collections l2l(l2):l-32. ductive cycle of a population of Astrangia Boschma. H. 1925a. The natire of the association from a jetty ofi' Port .Aransas, Texas. She between antho/oa and /oo.xanthcliac. —Pro- 246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ceedings of the National Academy of Sciences United States Bureau ofFisheries 46 (for 1931): 11:65-67. 277-381. . 1925b. On the feeding reactions and diges- Cummings, C. E. 1976. The effects of temperature tion in the coral polyp Aslrangia danae. with and salinity on the survival and respiration of notes on its symbiosis with zooxanthellae. — Bi- Asinngia danae with and without zooxanthel- ological Bulletin 49:407^39. lae. M. S. Thesis, University of Rhode Island. Brunet, C. 1982. Coral feeding rates and food re- 115 pp. source selectivity in response to various densi- . 1983. The biology of/l.s/ran^/a«/a«a<'(Milne ties of naturally occumng zooplankton. M. S. Edwards & Haime 1849): Studies of biotic and Thesis. University of Texas at Austin, 150 pp. abiotic factors affecting the physiology and mor- Buchanan. J. B. 1954. The zoographical significance phology of the northern star coral. Ph.D. Dis- of the Madreporaria collected in the Gold Coast, sertation. University of Rhode Island, 147 pp. West Africa. —Revue dc Zoologie et dc Bota- . & H. B. McCarly. 1982. Stable carbon iso- niquc Africaincs 49( l-2):84-88. tope ratios in Aslrangia danae: evidence for al- Cairns. S. D. 1977. Stony corals 1. Caryophylliina gal modification of carbon pools used in calci- and Dendrophylliina. Memoirs of the Hourglass fication.—Geochemica et Cosmochimica Acta Cruises (Marine Research Laboratory, Depart- 46:1125-1129. ment of Natural Resources. St. Petersburg. Ror- Damon. W. E. 1882. Coral in an aquarium —Amer- ida| .^(4). 27 pp. ican Microscopy Journal 3:221. . 1978. A checklist of the ahcrmatypic scle- Dana. J. D. 1846. Zoophytes. United States Explor- ractinia of the Gulf of Mexico, with the descrip- ing Expedition (1838-1842). 740 pp. tion of a new species.— Gulf Research Reports . 1872. Coral and coral islands. Dodd and 6;9-15. Mead. Publishers. New York. 398 pp. . 1979. The deep-water scleractinia of the Ca- . 1890. Corals and coral islands, third edition. ribbean Sea and adjacent waters. Studies on the Dodd. Mead, and Company. New York. 440 Fauna of Curasao and Other Caribbean Islands. pp. No. 180. 341 pp. Defenbaugh. R. B. 1976. A study of the benthic mac- . 1981. Marine flora and fauna of the north- roinvcrtebrates of the continental shelf of the eastern United States: Scleractinia. —NOAA nonhem Gulf of Mexico. Ph.D. Dissertation. Technical Report. NMFS Circular 438. U.S. Texas A&M University. 476 pp. Department of Commerce, 15 pp. Driscoll. E. Ci. 1967. Attached epifauna-substrate re- Caldcr, D. R. 1972. Phylum Cnidaria. Pp. 97-102 in lations—Limnology and Oceanography 12:633- M. L. Wass ct al.. compilers. ,\ checklist of the 641. biota of lower Chesapeake Bay. Sp>ccial Scien- Ellis. J. 1755. .An essay towards a natural history of tific Report No. 65. Virginia Institute of Marine the corallines, and other manne productions of Science. the like kind, commonly found on the coasts of .& B. S. Hester. 1978. Phylum Cnidaria. Pp. Great Britain and Ireland. London, for the au- 87-93 in R. G. Zingmark. ed.. An annotated thor. 103 pp.. 37 [381 pis. checklist of the biota of the coastal /one of South . & D. Solander. 1786. The natural history of Carolina. Belle W. Baruch Institute of Marine many curious and common zoophytes collected Biology and Coastal Research. University of from various parts of the globe Benjamin White South Carolina Press, Columbia. South Caro- and Son. London. 206 pp. lina. [isper. J. C. I 790. Die Planzenthierc in Abbildungen Chevalier. J.-P. 1966. Contribution a Tcludc des nach der Natur mit Farben erlcuchlcl nebst Be- madreporaircs des cotes occidentales de TAfn- schreibungen. volume 1. parts 5-6:193-320. que Iropicalc (I partiej. —Bulletin dc I'Inslitule Ferguson. A. H. 1948. Experiments on the tolerance Frani^ais .Afrique Noire 28 (scries .A):912-975. of several marine invertebrates to reduced sa- Cooley. N. R. 1978. An inventory of the estuarine linit\—Proceedings of the Louisiana Academy fauna in the vicinity of Pcnsacola. Flonda.— of Science 11:16-17. Florida Marine Research Publication No. 31. Feukcs. J.W. 1889. The anaiomy ofAstrangia dan- 1 1 3 pp. ae.—S\\ lithographs from drawings by A. Son- Courtney. C. M.. & T. D. Edmond. 1974. The mac- rel. Natural history illustrations prepared under roinvertebrates of Marco Island. Flonda. Pp. the direction of L. Agassiz. 1849, Explanation 51-159 //( Marco .Applied Marine Ecolog\ Sta- of plates by Fcwkes. Published bs the Smith- tion Progress Rcpon. July 1 1974-Dec3l 1974. sonian Institution. 20 pp. Marco Island. Florida. . 1891. An aid to a collector of the Coelenierala Cowles. R. P. 1930. .A biological study of the odshorc and Echinodcrmata of New England—Bulletin waters of Chesapeake Bay. —Bulletin of the of the Essex Institute 23:1-92. VOLUME 101, NUMBER 2 247

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continuous low-level gamma radiation on ses- tory, Smithsonian Instutition, Washington, sile marine invertebrates. Pp. 1-786 in Anon- D.C, 20560; (MEQP) Graduate School of ymous, ed.. Radioactive contamination of the Oceanography, University of Rhode Island, manne environment. International Atomic En- ergy Agency, Vienna. Narragansett, Rhode Island 02882-1197; Wilson, H. V. 1900. Marine biology at Beaufort.— (JWW) Department of Geological Sciences, American Naturalist 34(401):339-360. Cornell University, Ithaca, New York Wiseman, R. W. 1984, Light intensity effects on su- 14853-1504; (WCJ) Florida Department of peroxide dismutasc activity in symbiotic colo- nies of,-li/ra^i^/ai/awap. M.S. Thesis, University Natural Resources, Bureau of Marine Re- of Rhode Island, 75 pp. search, 100 Eighth Avenue S.E., St. Peters- Zibrowius. H. 1980. Les scleractiniares de la Medi- burg, Florida 33701;(JCL) Texas Memorial tcrranee et de I'Atlantique nord-oriental. Me- Museum, University of Texas, Austin, Tex- mories de ITnslitute Oceanographique Monaco, as 78705, and Department of Zoology, Uni- no. 11, 284 pp. versity of Texas, Austin, Texas 78712; Zinn, D. J. 1964. Yes, there are corals in Rhode Island.-Maritimes 8(4);15-I6. (CEV) Amoco, 200 East Randolph Drive, Chicago, Illinois 60601; (LSG) Biology De- (ECP)(SDC) Depanment of Invertebrate partment, Texas A&M University, College Zoology. National Museum of Natural His- Station, Texas 77843.

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