A New of , with a Discussion of the and an Appended Glossary'

MARY NEEDLER ARAI 2 ABSTRACT: A new species of Pachycerianthus from southern California is de­ scribed and the status of the genus is discussed. A glossary of the terms used in the of the order is appended. THE ORDER Cerianth aria consists of long, soli­ plete specimen, and redescribed Botruanthus tary, anemone-like , without a pedal benedeni from four specimens, all taken from disc or external skeleton, with numerous single the Gulf of California. tentacles in two crowns, coupled mesenteries, Child ( 1908) reported on regeneration in and a single siphonoglyph. Earlier authors re­ Pachycerianthus aestuari. Oth er authors have lated these anemones to the Actiniaria, but they referred to forms collected in California, but are recognized by recent authors, such as Wells have not identified them. When specimens and Hill (1 956 ), as being most closely related were obtained by divers near Los Angeles it to the Antipatharia or "black corals." was found that they belonged to previously Previous knowledge of the taxonomy and mor­ undescribed species. One of these species is phology of the Californian Ceriantharia is mea­ described herewith a discussion of the genus gre. McMurrich (1893: 202-203 ) described a Pachyceriam hus in which it is placed. species, Cerianth us vas, from one poorly pre ­ As indicated by Torelli 0960:373) , the served specimen that had been collected at Isla terminology used for the various anatomical de Cedros, Baja California, Mexico. The where­ stru ctures in the Ceriantharia is greatly con­ abouts of this specimen is unknown and the fused in the literature. A glossary of terms information given is insufficient to allow iden­ used in the taxonomy of this group is there­ tification of the species or to place it in any fore appended, and the terminology used recent genus or family. Torrey and Kleeberger throughout this work has been standardized (1909) described three species: . for uni formity. aestuari, C. benedeni, and C. [obnsoni, basing their descriptions on collected from MATERIAL AND METHODS Mission Bay, San Diego Bay, and San Pedro The sea anemones on which the anatomical Harbor, California, respectively. Th e descrip­ descriptions are based were collected by Dr. tion of the latter species was based on two Rimond Fay and Mr. Philip Bernard of the specimens. Although these descriptions were Pacific Bio-marine Supply Company between incomplete, enough morph ological data were July, 1958 and August, 1960 and were main­ given so that McMurrich (1910: 11) placed tained in the sea water tanks of the Depart­ Cerianthus benedeni in the genus Botruanthus, ment of Zoology, University of California at which he erected for it. He placed the other two species in the genus Pachycerianthus Los Angeles. On removal from the tanks, the animals were anaesthetized for 2-8 hours in Roule. Carlgren (1 951 :433-436 ) described Pachycerianthus insignis from a single incom- equal parts of sea water and a solution of 0.36M MgCh.6H20. They were fixed for ap­ proximately 15 minutes in 10% neutral forma­ 1 Much of the data presented in thispaper was in- _ . cIuded in a dissertati on subm itted co the Uni versity of lin -(with an excess of MgC03 ), overn ight in California, Los Angeles, in parcial fulfillm ent of the picro-formol solution (one part concentrated requirements for the Ph.D . degree. Manu script re­ formalin: 3 parts saturated picric acid), and ceived Febru ary 3, 1964. stored in 70% ethanol. In each case the solu­ 2 Form er address: Department of Zoology, Univer­ sity of Californi a, Los Angeles, California. Present ad­ tions were pipened into the coelenteron. dress: D epartment of Biology, Uni versity of Alberta, Preliminary identification of nematocyst types Calgary, Albert a. was done on fresh material. The nematocyst

205 206 PAQFIC 50ENCE, Vol. XIX, April 1965

measurements given m this section were ob­ longer and rounder appendages near the pos­ tained from portions of five of the paratypes terior end of the mesentery which he believed of the species sectioned and stained with Hei­ to be true "acontie.' McMurrich ( 1910: 19) denhain's iron hematoxylin and eosin, Mallory's used the term . "craspedoneme" to distinguish triple stain, and acid haemalum and light Herrwig and H ertwig's "Mesenterialfaden" green. Ten nernarocysts of each type were meas­ from these "acontia." ured in each portion of each examined. Carlgren (1912a:68-72; 1912b:383-385), re-invesrigating the same species examined by RESULTS AND DISCUSSION van Beneden, showed that true acontia were not present even there but rather stru ctures Family homologous with the craspedonemes were pres­ DIAGNOSIS : Cerianrharia without acontioids ent in which the endoderm was reduced so as or cnidorages. to be surrounded by the two limbs of the fila­ ment. Bourne (1919:60-61 ) also noted differ­ Genus Pachycerianthus Roule, 1904. ences between the "acontia" of Ceriantharia DIAGN OSIS: Cerianthidae with the second and those of the Acriniaria, In some portions couple of protomesenteries short and sterile. of the collected works of van Beneden pub­ Arrangement of metarnesenteries in each quar­ lished posthumously in 1923, the term "aeon­ tette M,B,m,b ( 1,3,2,4) more or less distinct. tie" is used.However, in another portion he, in agreement with the above authors, also states DISCUSSION that the Ceriantharian structures are not ho- Torelli (1961: 25-27) claims that the genus mologous with those of the Adirii ~ria . Carl­ Pachycerianthus proposed by Roule ( 1904a: gren suggested the term "mucocraspedoneme" . 792- 793) is -invalid since 'it is not clearly dis- for the structures found in the Ceriantharia, tinguished from the genus Cerianthus. Her but in later papers adopted the term "aconrioid" opinion is par tly based on an incorrect inter- introduced by Pax (1914 :394), pretation of the terms in the literature. It is With this background, no purpose was therefore first necessary to trace briefly the served by Torelli's statement (1960:376-379) history of some of the terms used to describe that acontia are not present in Cerianrharia, the mesenterial filaments and their appendages. especially since her opinion was based on an Haime ( 1854:374) noted the pr esence of investigation of tWO species for which previous "cordons pelotonnes" along the edge of some authors had not even claimed the presence of of the mesenteries of Cerianthus m embrana- acontioids, i.e., Cerianthus m embranaceus and ceus. Von Heider (1879 :216-217) and Hert- Pachycerianthus dohrni ( the Ceriantbes viridis wig and Hertwig (1879:578-580) described of her paper). the "Mesenrerial-filamente" of the same species We may now consider the description of in more detail. The mesenteries of this species Pachycerianthus. R a u I e (190 4a: 792- 793; have numerous branching appendages, the 1904b:708- 710 ) described Pachycerianthus "Mesenrerialfaden," which are not homologous benedeni and designated it the type of the genus. to the acontia of the Actiniaria (as von Heider The description is poor but includes the follow­ thought ), since they possess ascending and ing characters of possible generic significance : descending limbs of the mesenrerial filament 1. Mesenteries are short; only the ventral 'separated by a portion of endoderm. Van Bene- - mesenteries (53 in the terminology proposed den (1897:27-37) further investigated the by van Beneden) reach the aboral extremity structure of the "filaments mesenreriques," or and border a narrow band which continues "craspedes" of Gosse (1860:xxiii) in speci- the sulcus and contai ns the vestiges of the D mens of several adult and larval species. He and 51 mesenteries. found "fils mesenterique" wit h a structure simi- 2. The mesenteries on either side of the 53 lar to Hertwig and Hertwig's "Mesenterial- diminish in lengt h towar d the dorsal region faden" and also, in several larval species, much but do not show a distinct quatro-septal dispo- Pachycerianthus torreyi n. sp.-ARAI 207

sition; the irregular diminution gives them, wrong reason, that these appendages are ordi­ rather, a biseptal arrangement. nary craspedonemes. Pachycerianthus is there­ 3. All the mesenteries, except those which fore retained in the Cerianthidae. occupy exactly the dorsal region, possess some Without re-examination of the original mate­ "craspedes" in the zone placed immediately rial, Carlgren (1912a:40-48 ) defined the genus below the enterostome. Some mesenteries have Pachycerianthus as "Cerianrhidae whose second "fils mesenteriques" on their "craspedes," and couple of protocnemes are short, steril e, and alternating ones do not carry them. The mesen­ provided with an extremely well-developed re­ teries deprived of "fils mesenteriques" are fer­ gion of the cnido-glandular tract. Arrangement til and possess "aconties'' on a rather large part of the metacnernes (=deuterocnernes McMur­ of their inferior extremity; their series begins rich ) in each quartette M,B,m,b (1,3,2,4) with the S3 and continues with the uneven more or less distinct." He originally included numbered mesenteries S5, S7, S9, etc. eight species in the genus and later (1924a: As Torelli (1961 :25-27) pointed out, this 182-186; 1940 :15; and 1951:435-436) added description is extremely incomplete, and hence three more. All of these species (even P. bene­ it is very difficult to place the genus in the deni, if we are considering the first quartette) modern classification of the order. Some of her have M,B,m,b metamesenterial arrangement. comments (1960:374-375; 1961:25) about All except P. benedeni are known to have the lack of clarity can be discounted, however, short, sterile second protomesenreries, and it since she has translated incorrectly the term is probable that in that species they are also "fils rnesenteriques" in the third item above as sterile since very small mesenteries are rarely mesenterial filaments. As noted earlier the term fertile. However, Carlgren has placed in the was used by van Beneden (1897:33) and other genus several species (including P. benedeni) French authors to refer rather to the "Mesen­ in which the extent of the cnido-glandular tract terialfaden" or craspedonemes. Correctly trans­ lated, the third passage states that some mesen­ is unknown, and also others, such as P. maua, teries have craspedonemes on their "craspedes" in which it is very short. In addition, as Torelli or mesenterial -filam enrs and alternate mesen­ (1 960:375-376) has indicated, the distin ctness teries have "aconties" on them . of the histological demarcation by Carlgren It is the interpretation of the term "aeon­ ( 1912a:59-67) of the mononerne into the ties" in the third passage above that is most cnido-glandular tract and the craspedion is problematical. McMurrich (1910: 10-14, 22­ questionable. (The mononeme is the aboral 24 ), in his classification of the order, placed monofilar portion of the mesenterial filament.) the genus in a family characterized by the It seems desirable therefore that this character presence in the of "acontia" or, as they be eliminated as a differential one not only in were later called, acontioids, but includ ed adult this but in all genera of cerianrhids. species both with and without aconrioids, Carl­ The 10 species (other than P. benedeni) gren (1912a:37-48) believed that the struc­ placed in the genus by Carlgren comprise a tures must be ordinary craspedonemes since distinct taxonomic entity with the remaining acontioids in all the better known species are characters with which Carlgren defined Pachy­ single, rather than scattered over a large part cerianthus and with no acontioids. C. bicycles of the lower end of the mesentery. He placed Torelli, 1961 and P. torreyi n.sp. of the present the genus in the family Cerianrhidae character­ work may also be added: This group' is sep-" ized by the lack of both acontioids and enid­ arared from the genera Cerianthus, Ceriantb e­ orages. Although Catlgren is probably correct, opsis, and Ceriantheomorphe by the possession the question can only be resolved definitely by of short, sterile second protomesenteries, and reference to the type specimen; unfortunately from Cerientbeopsis by. the MBmb arrange­ the location of this material is at present un­ ment of the merarnesenteries in Pachycerian­ known. In the meantime we may assume, as thus. Torelli (1961: 27) does not believe that Torelli (1%1: 26 ) also does, although for the the properties of the second proromesenteries 208 PACIFIC SCIENCE, Vol. XIX, April 1965 are imp ortant enough to serve as generic char­ Pacb ycerianthus torreyi n. sp. acters or to distinguish Pachycerianthus from DIAGNOSIS : Siphonoglyph running length of Cerianthus. These mesenteries, however, are actinopharynx.Directive mesenteries extending the first couple of mesenteries to develop and well beyond siphonoglyph. Second protomesen­ could therefore be expected to be the most teries between once and twice length of direc­ stable in the grown animal. McMurrich (1910: tives. Metarnesenteries having definite MBmb 23-24) considered the form of these mesente­ arrangement. M definitely longer than any ries to be characteristic, even at the familial I level. other mesentery. Ma longer than M2. m2 and In summary, it should be stated that an in­ ma shorter than mr , BI shorter than Pa. M I tensive search must be made for the type speci­ with trinerne almost to end of mesentery, eras­ men of P. benedeni. It is not in the collections pedonemes scattered over most of its length, of the Musee Oceanographique de Monaco, the very short or no mononeme. Musee de Bruxelles, the British Museum (N at­ LOCALITIES: Latigo Cove; and just inside ural History), or the Museum National d'His­ breakwater, Los Angeles Harbor (type local­ toire Naturelle, Paris, the most likely reposi­ ity) , California. tories. If it cannot be found, it will be neces­ sary to declare the species a nomen dubium. HABITAT: 12-18 meter depth; in mud, or Since .the other twelve species presently in­ mixed gravel and mud, bottom. cluded . in the genus appear to constitute a HOLOlYPE and PARAlYPES : To be deposited taxonomic entity, the declaration of the nomen in USNM. dubium would necessitate selection of a new type species and generic name for the taxon. DESCRIPTION (based on 10 adult specimens ): In the meantime, Pachycerianthus is retained Coloration of live animals. Column red­ in the Cerianthidae and includes the following brown , darkest JUSt below oral disc and toward species: P. aestuarii (Torrey and Kleeb erger, aboral pore, with narrow buff ring immedi­ 1909); P. benedeni Roule, 1904; P. bicycles ately around aboral pore. Marginal tentacles (Torelli, 1961) n. comb.; P. dohrni (van Bene­ buff (shading to bright pink towards tips in den, 1923) ; P. fimbriatus McMurrich, 1910; one specimen) . .Oral tentacles darker , with P. insignis Carlgren, 1951; P. johnsoni (Tor­ purplish tinge. rey and Kleeb erger, 1909 ); P. maua (Carl­ Coloration of a1zimals fixed in picro-jormol. gren, 1900 ); P. monostichus McMurrich, 1910; Red pigments of live animals faded, leaving P. multipli catus Carlgren, 1912 ; P. plicatus column light yellow-brown, tentacles yellowish Carlgren, 1924; P. solitarius (Rapp, 1829); buff. and P. torreyi n. sp. Length in anaesthetic. 13-31 em ( type 18.5 Pachycerianthus bicyclus is distinguished from em) . all other members of the genus by having two T entacles. Marginal arrangement 2 ( dr) 431. cycles of marginal tentacles but thre e or four 4231.4231... Labial arrangement 2 (de) 313.4232 . cycles of labial tentacles, and lacking craspe­ 4312 ...or 3 (dt)413.4232.4312... (type). In each donemes but possessing aboral appendices on crown division into third and fourth cycles in­ the second protomesenteries and first order distinct in some specimens. Directive labial metamesenteries.Torelli ( 1961:24-25 ) states tentacle _presenr..Marginal .tentacle numb er. ap­ that the specimens from which P. bicyclus was proxim ately 90-125 ( type approximately 100 ). described may not have reached full growth, Siphonoglyph . Running length of actino­ so that the species may pro ve to be an imma­ pharynx. Attached mesenteries 10 ( type) or ture stage of another. Since P. solitarius from 12. Hyposulcus indistinct, less than one-tenth the same area differs only in the number of length of siphonoglyph. Hemisulcus distinct, cycles of marginal tentacles, these species will flat orally, continued down directive mesentery probably prove to be synonymous. as a filament without ciliated tracts. Pachycerianthus torreyi n. sp.-ARAI 209

Protomesenteries. D ire c t i v e mesenteries c-.;"• ..--.-----.--...--:-..--.------....--=-.--.-.-n..-----,---.--,------­ , short er or longer (type) than siphonoglyph. <, : - .. , . , ,. . . . I , • :( : , I I , Second proromesenteries sterile, less than twice :1: I I I , I I , I I , I , I I :1:: T::I: , I I I I ----:.I.: : ~ , , , , I I :I : ::: ; I I I I , length of directive mesenteries, with aboral lobe :1:: , , I I ; ',~~: , I I I , I I , :c \ : l , , I , I , } varying greatly in length (long in type), tri ­ ,I ': ;1:;' I I : I ; : neme running to near tip of aboral lobe, usually --' ( type) with several craspedonemes at aboral I I end of trineme, convoluted mononeme running b2 b 3 b I around tip of lobe. Third protomesenteries ster­ 6 3 PI 6 2 ile, approximately same length ( type) or 6 , shorter than P2, longer than PI, rrinerne much P2 P3 shorten than in P2 with a num ber of craspedo ­ I--v~ nemes at aboral end, mononeme relatively long m m2 3 and highly convoluted along border of broad section of mesentery. - M etamesenteries. Definite MBmb arrange­ M2 m, ment ( Fig. 1). Reproductive organs in macro­ M 3 septa only. M I mesenteries at least five-sixths length of column , never reaching aboral pore; remaining mesenteries less - than two-thirds length of MI in most specimens (longest mes­ entery approximately seven-eights of MI in one specimen). Macrosep ta generally decreas­ ing in length toward multiplication chamber; Ma longer than M2, m2 and rna shorter than ml , rns longer or shorter (type) than m s, M, with rrineme almost to aboral end of mesentery, FIG. I. Diagram of the internal arrangement of craspedonemes scattered over most of its length, Pachycerianthus torreyi. View of half the animal cut sometimes with an aboral lobe of mesentery or longitudinally and pinned flat. Approximately Y2 X vertically, 5 V2 X horizontally. a, Oral disc; b, acrino­ bunch of craspedonemes at aboral end of tri­ pharynx; c, marginal tentacle bases; d, labial tentacle neme, short mononeme (type) or none, repro­ bases; e, mouth; i. siphonoglyph; g, hemisulcus; b, ductive region to above level of craspedon emes, protomesenteries; i, first quartette of metamesenteries; hermaphroditic. Macroseptum rni with long PI, directive mesentery; P2, second protomesentery; Ps, trinerne terminating much farth er from aboral third protomesentery; MI, M2, Ms, first order meta­ mesenteries; mi, rns, rna, second order metamesenteries; end than in M , trineme with scattered craspe­ I BI , B2 , Bs, third order merarnesenteries; b., bs, bs, donemes and sometimes with an aboral bunch, fourt h order merarnesenteries . mononeme well developed, convoluted or straight (slightly convoluted in type). Extent and form of rrinemes and mononernes of other X 10.0- 16.2 p. rare. Of marg inal tentacles, atrichs p. scarce, micro­ macrosepta variable. BI shorter than Ps, with 24.0-36.8 X 4.0-7.2 short rrineme with bunch of craspedonemes at basic b-rnastigophores 20.0-49.6 X 3.0-9.0 p. aboral end, highly convoluted mononeme along common, spiro cysts 7.6-43.2 X 1.2-5.6 p. nu­ broad section of mesentery. Oth er Band b with merous. Of labial tentacles, atrichs 24.0- 33.6 X structure similar to Bl although bunch of eras­ 3.6-6.4 p. scarce, microbasic b-mastigophores pedonemes of trineme may be reduced or ab­ 19.2-45 .2 X 2.0-8 .4 p. common, spirocysts sent. 13.2-33.6 X 1.2-5.2 p. numerous. Of actino­ N ematocysts (based on five adult specimens) . pharynx, atrichs 24.8-40.8 X 3.6-6.4 p. scarce, Of column, atrichs 29.6-83.2 X 5.0-29.6 p. microbasic b-mastigophores 20.8-44.4 X 2.8­ very 'common, microbasic b-mastigophores 18.5­ 7.6 p. common, spirocysts 11.2-26.8 X 1.6-4.4 43.2 X 3.2-8.0 p. scarce, holotrichs 42.2-67.6 p. very rare. Of ciliated tract of macroseprum, 210 PAOFIC SOENCE, Vol. XIX, April 1965 arrichs 21.2-32.8 X 2.4-4.4 p. rare or absent, only in the relat ive lengths of the M 2 and M3 microbasic b-mastigophores 18.4-50.8 X 2.8­ mesenteries. Pachycerianthus fimbriatus, from 9.2 p. scarce, spiro cysts 14.4-26.8 X 1.8-5.2 p. the same area as P. plicatus, differs in the same common. Of peloton of microseptum, micro­ character and possibly in the arrangement of basic b-mastigophores 18.4-30.0 X 4.4-7.6 p. the labial tentacles. Pachycerianthus bicyclus very common. and P. solitarius differ from P. torreyi in the Pachycerianthus torreyi is clearly distin­ absence of craspedonemes of the trineme and guished from the two species described by Tor­ in that several M mesenteries nearly reach the rey and Kleeb erger ( 1909) from southern Cali­ aboral pore. The latter character may also be fornia. The present species and P. aestuari dif­ used to distinguish P. m onostichus, P. 'dohrni, fer in that the latter possesses a siphonoglyph and P. multiplicatus from the present species. which does not reach the lips of the acrino­ Pachycerianthus maua differs by having the M 3 pharynx and has four mesenteries commonly mesentery shorter than the M 2, and also by pos­ reaching the terminal pore. Other characters of sessing an unusual type of aboral craspedoneme. P. aestuari which may differentiate it from P. The name, torreyi, is in honor of Dr. H. B. torreyi are a tentacle number which never ex­ Torrey, one of the earliest workers on cerian­ ceeds 34, a single cycle of marginal tentacles, tharian taxonomy in California. and a very broad siphonoglyph. Pachycerianthus johnsoni differs from P. tor­ GLOSSARY reyi in that the directive mesenteries do not As Torelli (1%0:373) has pointed Out, the extend beyond the siphonoglyph, as is the case terminology applied to the Ceriantharia has in the present species. Other characters of P. become exceedingly complicated through the johnsoni with possible diagnostic value are that multiplication of terms. In the following glos­ go nads are frequently borne on the third order sary the full definition of a structure has been mesenteries, that the siphonoglyph is narrow, placed with the presently accepted term and and that the Bi may be longer than the P3• other terms have been referred to this defini­ Among approximately one hundred cerianthid tion. English nomenclature or anglicised terms specimens taken and examined from deeper are used. Terms have been added from other water in Los Angeles Harbor, the type locality languages where these are not direct transla­ for P. johnsoni, no specimens of this species tions. Th e application of certain terms to re­ were found. It is probable that it is a shallow lated groups of animals has been given when water species and that the mud-flat habitat they are likely to cause confusion. where the two specimens were collected has 1 (rnetamesentery )(Carlgren, 1912b:368 )­ been eliminated by harbor installations. longest metamesentery in each quartette, i.e. The clarity with which the present species merarnesentery of the first order. can be distinguished from the other species in­ (tentacle) (Carlgren, 1900:25-26; McMur­ cluded in the genus depends, in good part, on rich, 1910 :17-18)-tentacle of innermost the detail in which they have been described. cycle of a crown. We have seen that the type species, P. bene­ subscript (rnetamesenrery ) (Carlgren, 1924a: deni, was very poorly described, but it is pos­ 169-170 )-belonging to first meramesente­ sible to distinguish it since there is a regular rial quartette ( q. v.) . diminution in the length of the rnetamesente­ 1 subscript (protornesentery) (Carlgren, 1912a: ries, i.e. M3 is shorter than M2• Pacbycerian­ 12 ) -belonging to firstprotomesenterial thus insignis, based on a single incomplete couple. specimen from the Gulf of California, has sev­ 1 superscript (Beneden, 1923:24-25 )- clos­ eral metamesenteries almost reaching the aboral est structure to siphonoglyph in metasome pore, and a P2 that is more than twice the ( q. v.) . length of Pl. l d superscript ( Beneden, 1897: lO-11 )-first Pachycerianthus torreyi is most similar to structure to right with reference to directive P. plicatus, an Indonesian species, differing plane at the siphonoglyph. Pachycerianthus torreyi n. sp.- ARAI 211

Ig superscript (Beneden, 1897: 10-11 )-first nerne, i.e. microseptum, of a meramesenterial structure to left with reference to directive quartette. plane at siphonoglyph. Biseptum (Beneden, 1897 :2 3)-two"longest or a superscript ( Beneden, 1923 :23 )- belonging two shortest mesenteries of a metamesente­ to second somatomere of the prosorne rial quartette. (q. v.) . b-mastigopbore (Carlgren, 1940 :3-4)- mas­ Acontioid ( Pax, 1914:394 )- long threadlike, tigophore with shaft tapering gradually to simple or slightly branched craspedoneme, term inal tube. rounded in cross section , with many mucous Botrucnid ( Beneden, 1897 :32; Carlgren, 1912a: cells and reduced endoderm, at or near aboral 72- 74 )- stalked bunch of cnidorages ( q.v.) . end of rnesenterial filament. Brachyneme (McMurrich, 1910 :14 )-micro­ Acontium (Gosse, 1858:125; 1860: xxiii- xxv, septum (q.v.) . 5; Herrwig and Hertwig, 1879 :562-565; Buccal cone ( Beneden, 1897 :9)-raised por­ Torelli, 1960 :376- 379 ) - thin threadlike tion of oral disc immediately surrounding structure of Actiniaria containing numerous mouth of some larval forms . nernatocysts, attached at one end to a mesen­ Buccal disc (Vogt, 1888 :29 )--oral disc (q . v.) . tery, usually below the filament. Buccal tentacle (Vogt, 1888 :9~1O)-labial ten­ Acontium (Beneden, 1897 :30-33; Bourne, tacle ( q.v.) . 1919 :60-61; Carlgren, 1912a:42, 68-72 )­ Capsule ( W eill, 1934:21, 27- 30 )-hollow case acontioid ( q.v.) . of nernarocyst. Acontium ( Heider, 1879 :217)-craspedoneme Cerinula (Beneden, 1897 :147; 1923 :19)-lar­ (q.v.). val stage with at least three couples of mesen­ Acromere ( Beneden, 1923 :22 )- "segment" of teries . body, including the directive chamber, di­ Ciliated tract ( Hertwig and H ertwig, 1879: rective mesenteries and directive tentacles 557 )- continuation of ciliated ectoderm and ( by analogy to Amphioxus) . mesogloea from a furrow of the actinophar­ Actinocoel (Bourne, 1919:35)-intermesente­ ynx along one side of a rnesenterial filament. rial chamber (q.v.) . Ciliated tract region (Ca rlgren, 1912a:59-60) Actinopharynx (Beneden, 1897:13-14)-tube - trineme (q. v.) . leading from mouth into coelenteron, lined Cloison ( Fisher, 1887: 383; Faurot, 1895 :50 ) by longitudinally ridged, invaginated, ecto­ -mesentery ( q. v.) . " derm which is supported by mesogloea. Cloison prequatroseptale (Beneden, 1897:24) Actinostome ( Beneden, 1897 :9-10, 13-14) ­ - protomesentery (q. v.). mouth (q .v.) . Cloison primitive ( Beneden, 1897:23)-one Animal length-external dista nce from base of of second couple of protomesenteries. marginal tentacles to aboral pore. Cloison quatroseptale (Beneden, 1897: 24 )­ Armature (Weill, 1930 :147; 1934:21-22)­ rnetamesentery (q. v. ) . spines on tube of nematocyst. Cnida (Gosse, 1858: 125; 1860:xxii, xxix-xxxiv, Atrich (Weill, 1930 :146; 1934: 37, 50-52, 269 ; Weill, 1934 :330 )-nematocyst sensu 624-626)-nematocyst with isodiametric, Haime (q.v.) . unarmed, tube open at tip. Cnida cochleata (Gosse, 1858 : 126-127; 1860 : Axenteron ( Beneden, 1897: 20 )-central por­ xxxiii -xxxiv; Bedot, 1896 :534; Weill, 1930 : tion of coelenteron undivided by mesenteries. 149)- spirocyst (q; v.) . Axial body (Moebius, 1866:387; Weill, 1934: Cnida glomi/era ( Gosse, 1858 :126; 1860:xxxii­ 22-24)- straight basal section of nernato­ xxxiii; Weill, 1934 :89-90) - nematocysr cyst tube within the capsule. with nearly oval capsule containing long, iso­ b (McMurrich, 1910 :14-15 )- shorter brachy­ diametric, loosely coiled tube (not in present nerne, i.e. microseptum, of a metamesenterial use). " quartett e. Cnidoglandular tract ( Hertwig and Herrwig, B (McMurrich, 1910:14-15 )- longer brachy- 1879 :557; Carlgren, 1949: 8 )-continuation 212 PAOFIC SOENCE, Vol. XIX , April 1965

of ciliated ectoderm from a ridge of the ring of tentacles all equidistant from mouth actinopharynx along the length of a mesen­ or from margin of oral disc. terial filament. d superscript (Beneden, 1897: 10-11 )-droit, Cnidoglandular tract (Carlgren, 1912a:59-66) i.e. right with reference to the directive plane -plectocraspedon ( q. v.) . at the siphonoglyph. Cnidom (Weill, 1926:1245; 1934: 35l)-ne­ d superscript (Beneden, 1923:23 )- directive. matocyst complement of coelenterate at any D (Beneden, 1897:20 )-cloison directrice, i.e. given moment of its existence. directive mesentery ( q.v.). Cnidoneme (Beneden, 1923:154 ) -acontioid Deuterocneme (McMurrich, 1910:6-1O)-met­ ( q. v.). amesentery (q. v.) Cnidorage (Beneden, 1897: 32; Carlgren, 1912a: Directive chamb er (Carlgren, 1900:27; 1906: 72-73 )-rounded projection of mesentery 66 )-Intermesenterial chamber between two containing numerous nernarocysrs. directive mesenteries. Coelenteron (Beneden, 1897:14)-main cav­ Directive mesentery (Herrwig and Hertwig, ity of the animal. 1879: 538, 572)-one of couple of mesen­ Column (Gosse, 1860:2, 268; Beneden, 1897: teries closest to directive plane at siphono­ 9) -body cylinder below oral disc and tenta­ glyph. cles. Directive plane (Carlgren, 1906:66) - plane Column length-internal distance from aboral symetrically bisecting the animal, running end of actinopharynx to aboral pole. through siphonoglyph and point diametri­ Commissural plane (Faurot, 1895:50, 59)­ cally opposite as seen in cross-section. directive plane. Directive tentacle (Carlgren, 1900:25, 1906 : Continuous mesentery (Heider, 1879:216-217; 66)-tentacle opening into directive cham­ McMurrich, 1890: 137)-mesentery reaching ber. the aboral pore, especially applicable to a dm. (Carlgren, 1924a:169)-directive mesen­ second protomesentery. tery (q. v.). Cordon pelotonne (Haime, 1854: 374 )-mes­ Dorsal-opposite to ventral (q. v.). enterial filament ( q. v.) . dt. (Carlgren, 1924a:169)-directive tentacle Couple of mesenteries (Faurot, 1895:103; Ben­ ( q.v.) . eden, 1897:20-21; Carlgren, 1906:66 )-two Ecthoraeum (Gosse, 1858: 126; 1860:xxxi­ corresponding mesenteries on opposite sides xxxv, 269 )-tube of nematocysr ( q. v.) . of directive plane. Enteroid (Lacaze-Duthiers, 1873: 302; Faurot, Craspedum (Gosse, 1858: 125; 1860:xxiii-xxiv, 1895:50-51, 234- 235) -mesenterial fila­ 5; Bourne, 1919:54-55; Torelli, 1960:374) ment (q. v.). -mesenterial filament (q. v.) . Enterostome (Beneden, 1897:14) - opening Craspedion (Carlgren, 1912a:60-68; Torelli, from cavity of actinopharynx into coelenteron. 1960: 375-376)-telocraspedon (q. v.). Filamentchen region (Carlgren, 1912b:365) ­ Craspedoneme (McMurrich, 1910:19-22) ­ telocraspedon (q. v.). threadlike' or flattened, sometimes branched, Filamentseptum (Heider, 1879:216; Carlgren, process of the mesogloea and entoderm of 1912a:51)-sterile mesentery (not in pres­ mesentery over which pass ascending and ent use) (d. Geniralseprum ). descending limbs of mesenterial filament. Fils mesenseriqaes (Beneden, 1897:33 )-cras­ Crown of tentacles (Haime, 1854: 350, 363; pedonerne (q .v. ) ._ Faurot, 1895: 218, 236-240 )-all cycles of g superscript (Beneden, 1897: 10-11 )- gauche, tentacles grouped around mouth or all cycles i.e. left with reference to directive plane at near edge of oral disc. siphonoglyph. Cycle of mesenteries (Carlgren, 1912a:51)­ Genitalseptum (Heider, 1879:216; Hertwig order of mesenteries ( q. v.) . and Hertwig, 1879:580-581)-fenile mesen­ Cycle of tentacles (Haime, 1854: 365-367; tery not reaching to aboral pore (d . Fila­ Heider, 1879:213; Faurot, 1891:70-71)- mentseptum). Pachycerianthus torreyi n. sp.-ARAI 213

Grenzstreijen (Carlgren, 1924b :347)-mesen­ M (Beneden, 1897: 10)- loge directrice, i.e. teret (q. v.) . directive chamber (q.v.). Hampe (Faurot, 1895 :50)-main portion of Macrobasic mastigophore (Weill, 1934 :38, 64­ mesentery excluding mesenterial filament. 68; Currress, 1955 :133-134 )-mastigophore Hampe (Weill, 1934:26)-shaft of nernato­ in which shaft is more than three times cap­ cyst (q. v.) . sule length. Hemisulcus (Beneden, 1897 :37-38; Carlgren, Macrobimesenterium (Carlgren, 1912a:51) ­ 1912a:50, 63 )-continuation of half hypo­ rnacrobiseprum (q. v.). sulcus along edge of a directive mesentery. Macrobiseptum (Beneden, 1897: 23-24)-two Holotrich (Weill, 1930:145; 1934:37-38,49­ longest mesenteries of a meramesenterial 50,64,624-626; Cuttress, 1955 :134-137)­ quartette. basophilic nematocyst with an isodiarnetric, MaC'l'ocneme (Stephenson, 1920: 456)- mesen­ open-ended, tube armed along its entire tery of Actiniaria attached to actinopharynx length . with strong retractors, gonads, and filaments . Hoplotelic terminal tube (Weill, 1930:147; Macrocneme (McMurrich, 1910 :14) macrosep­ 1934: 38 )-terminal tube bearing spines. tum of Ceriantharia (q. v.). Hyposulcus ( Beneden, 1897:15, 37-38; Carl­ Macroseptum ( Faurot, 1891: 68-69; 1895:233­ gren, 1912a:50,63 )-grooved structure con­ 234; Torelli, 1960 :374)-one of two longest necting the directive mesenteries, continua­ merarnesenteries in a quartette. tion of the siphonoglyph below the border Marginal tentacle (Haime, 1854: 363-368)­ of the acrinopharynx, tentacle of one of cycles near edge of oral Intermesenterial chamber (Faurot, 1895: 228) disc (d . labial tentacle) . - portion of coelentero n between two adja­ Mastigophore (Weill, 1930:146; 1934:38, 58) cent mesenteries . - nematocyst with well-defined isodiamerric Isorhize (Weill, 1930 :145; 1934: 37, 49, 624) shaft and terminal tube with open tip. -nematocyst with isodiametric tube open Median plane (Beneden, 1897:12)-directive at tip. plane (q.v.) . (Beneden, 1923:23 )-loge de multipication, Median streak (Carlgren, 1912a:60-67)-spi­ i.e. multiplication chamber. rocyst-glandular tract (q. v.). L (Beneden, 1897 :10)- loge, i.e. intermesen­ Median streak (McMurrich, 1910:20-21)­ terial Chamber, other than directive chamber. central tract of trineme. L (Beneden, 1923: 23 )- loge directrice, i.e. di­ rective chamber. Median tentacle ( Beneden, 1897:10)- direc­ L with superscript (Beneden, 1923: 23 )- loge tive tentacle (q. v.) . other than directive chamber. Mesenterial length-distance from aboral end Labial tentacle-(Haime, 1854:368- 369)- tenta­ of actinopharynx to aboral end of mesentery. cle of one of cycles surrounding mouth (d. Mesemerialfaden (Herrwig and Herrwig, 1879: marginal tentacle). 579-580)- craspedoneme (q. v.) . Lame (Haime, 1854:371, 374) - mesentery Mesenterial filament (Frey and Leuckart, 1847: (q. v.) . 12-18; Hertwig and Hertwig, 1879:556)­ Loge (Haime, 1854: 371) - inrermesenterial thickened rim running along free border of chamber (q.v. ). mesentery from end of actinopharynx abor­ m (McMurrich, 1910: 14-15) -shorter mac­ ally. rocneme, i.e. macroseptum, of metamesenter­ Mesenteret (Beneden, 1897:27; Carlgren, 1931: ial quartette. 8)-projection of mesentery bearing the m (Be neden, 1897 :113 )-multipication cham­ peloton, ber (q. v.) . Mesentery (Frey and Leuckart, 1847: 11; La­ M (McMurrich, 191O:14-15) - longer mac­ caze-Duthiers, 1873:301-302; Bourne, 1900: rocneme, i.e. macroseptum, of metamesenter­ 8-9)- longitudinal extension into coelen­ ial quartette. teron from body wall, attached to acrino- 214 PACIFIC SCIENCE, Vol. XIX, April 1965

pharynx and bearing thickened filament upon Oral disc (Faurot, 1895:47-48, 218 )-disc at its free inner edge below acrinopharynx. one end of animal bearing mouth and tenta­ Mesogloea (Bourne, 1887:303, 311-320; Tor­ cles. elli, 1952: 154-156)-supporting layer be­ Order of mesenteries (Carlgren, 1912b: 375­ tween ectoderm and endoderm. 376, 386; Torelli, 1960:374)-all merames­ Metacneme (Duerden, 1900:47, 1902:388­ eateries of same relative lengths in their 397) -metamesentery (q. v.) . respective quartettes. Metamesentery (Carlgren, 1912a: 50-51, 59; Orthocraspedon (Bourne, 1919:55-57) -tri­ 1912b:361 )-any mesentery other than pro­ neme ( q.v.) . tomesenteries. p. superscript (Beneden , 1923:23 )- belonging Meta some (Beneden, 1923: 21-22 ) - central to third somatomere of prosome ( q.v.) . P (Carlgren, 1912:8 )-protomesentery ( q.v.). portion of body excluding prosome and telo­ Peloton ( F a u r o t, 1895:235-236; Beneden, mere (q. v .). (by analogy to Amphioxus) . 1897:27,46; 1923:51-52, 79 )-highly con­ Microbasic mastigopbore (Weill, 1934:38,58­ voluted section of mononeme. 64, 624-626)-mastigophore in which shaft Penicillus (Stephenson, 1929: 178-179; Weill, is at most three times capsule length. 1934:91-92 )-nematocyst containing short, Mic1"obimesenterium (Carlgren, 1912a: 51)­ stout, armed cube (not in present use). microbiseptum (q. v.) . Peristome (Vogr, 1888:29)-portion of oral Microbiseptum (Beneden, 1897: 23-24 )-two disc between tentacle crowns. shorter mesenteries of meramesenterial quar­ Plectocraspedoa (Bourne, 1919:55-57)-por­ tette. tion of mononeme indistinctly delimited by Micros eptum ( Faurot, 1891:68-69; 1895:233­ abundance of large, thick-walled nernato­ 234; Torelli, 1960:374 )-one of rwo short­ cysts and large coarsely-granular gland cells, est rnetamesenteries in quartette. usually anterior to telocraspedon. Mononem e (Beneden, 1923:79-80) --'-aboral p-mastigophore (Carlgren, 1940:3-4; Cuttress, monofilar region of mesenterial filament. 1955:129-131, 133-134; Hand, 1961:189­ Mouth (Rapp, 1829:654; Haime, 1854:350, 190, 193)-mastigophore in which abrupt 372) -opening from exterior to cavity of reduction of shaft to terminal tube allows funnel-shaped invagination of axial body. actinopharynx. Protocnem e Muco craspedoneme (Carlgren, 1912a :71 )­ (Duerden, 1900:47' McMurrich 191O:6-1O)-protomesentery ( q.v. ) . ' acontioid ( q. v.) . Protomesentery (Carlgren, 1912a:50-51, 59, Multipication chamber (Vogr, 1888:24 )-in­ 1912b:361)-one of mesenteries arising in rermesenterial chamber opposite directive first embryological stage (presently consid­ chamber, in which new rnesenterial couples ered to be one of the first three mesenteries are formed. on either side of directive plane at siphono­ Mundscheibe (Hertwig and Hertwig, 1879: glyph). 471; Heider, 1879:212)-oral disc (q. v.) . Prosome (Beneden, 1923:21)-anterior por­ Nematocyst (Haime, 1854:354-357, 365; Be­ tion of body including three cycles of proto­ dot, 1886:51-52; Cuttress, 1955:124-126) mesenteries and associated tentacles, inter­ -capsule containing an expellable coiled rnesenrerial chambers adjoining and portions cube. of. columnar wall, peristome and actinophar­ Nematocyst (Bedor, 1889:607-608; 1896:535­ ynx (by analogy to Amphioxus ). 536; Weill, 1934:330)-nematocyst sensu Quart ett e of m etamesenteries (Faurot, 1891: Haime -excluding spirocysts. 67-68; 1892:238; 1895:2 31-234) -any Odd tentacle (Agassiz, 1863:527; Fischer, group of four mesenteries in a series as 1889: 24)-directive tentacle (q.v.) . counted from prorornesenteries, each group Oesophagus (Haacke, 1879:276,293; Jourdan, showing similar pattern of relative rnesenter­ 1879: 103, 108-109)-actinopharynx (q. v.). ial lengths; Pachycerianthus torreyi n. sp.-ARAI 215

Quatromere (B eneden , 1924 :43 )-four suc­ Sulculus (Haddon, 1889: 300)-siphonoglyph cessive somarorneres (q. v.). other than the sulcus of Actiniaria (q. v.) . S. (Beneden, 1897 :20 )-cloison, i.e. mesentery, Sulculus ( Carlgren, 1912a:55-56) - siphono­ other than directive mesenteries. glyph of Ceriantharia if considered homolo­ S with superscript ( Beneden, 1923:25 )-sarco­ gous to sulculus of Actiniaria ( q.v.) . septum, i.e. mesentery. Sulcus ( Haddon, 1889 :300 )-ventral siphono­ Sarcoseptum ( Haacke, 1879 :277, 293)- mes­ glyph of Actinaria, i.e. that one homologous entery ( q.v.). to single siphonoglyph of Peachia. Sorcosepte antipathoide (Beneden, 1923:18)­ Sulcus (Kingsley, 1904:347-351; Torelli, 1960 : prorornesentery ( q. u. ) . 383) siphonoglyph of Ceriantharia if con­ Sarcosepte cerianthoide ( Beneden, 1923: 19)­ sidered homologous to sulcus of Actiniaria metamesentery (q.v.) . ( q.v.) . Schlundrinne ( Herrwig and Hertwig, 1879: .t (Beneden, 1897: 10-11 )- labial tentacle 513, 572 )-siphonoglyph (q.v.)'. other than directive tentacle. Septum (Gosse, 1860 :xiii-xiv, 5, 268)-mesen­ ( Beneden, 1923: 23)-labial directive tenta­ tery ( q. v.) . cle (q. v.) . Shaft (Carlgren, 1940 :3) - thickened basal t with superscript (Beneden, 1923:23)-labial part of tube, of some nematocysts. tentacle other than directive tentacle. Siphonoglyph ( Hickson, 1883:693- 694; Mc­ T ( Beneden, 1897:10-11 )-marginal tentacle Mur rich, 1890 :131, 136 )-smooth ciliated ocher than directive tentacl e. groove of acrinopharynx, T (Beneden, 1923:2 31-median marginal ten­ Somatomere (Beneden, 1923:21-22 ) - "seg­ tacle, i.e. marginal directive tentacle . ment " of body including a couple of mesen­ T . with superscript (Beneden, 1923:23)-mar­ teries, the inrermesenterial chamb ers 'ante­ ginal tentacle other than directive tentacle. rior' to them , and the associated tentacles T.M. ( Beneden, 1897:10-11 ) -median mar­ and adjoining portions of columnar wall, ginal tentacle, i.e. marginal directive tentacle . peristome and actinopharynx ( by analogy to Te locneme (McMurri ch, 1910 :11 ':"' 13 ) -con­ A mphioxus ) . tinu ous mesentery ( q.v. ). Spiral cnida (Gosse, 1858: 126-127; 1860: T elocraspedon (Bourne, 1919: 55-57) - por­ xxxiii-xxxiv; Weill, 1934: 326 )-spirocyst tion of mononeme indistinctly delimited by ( q. v.) . abundance of spirocysts and eosinophilous Spirocyst (Bedor, 1889: 607-608 ; 1896 :534­ mucus gland cells, usually posterior to plec­ 536; W eill, 1934:325-342, 624; Cutrress, rocraspedon. 1955:124-126, 136 )-capsule conta ining an T elomere ( Beneden, 1923:22 )-"segment" of acidophilic, very regularly spiralled, tube of body including multipication chamber (by unif orm diameter which irregularly everts analogy to Amphioxus) . ( d . Holotrich ). T erminal tube (Weill, 1934 :26, 37)-thin dis­ Spirocyst-glandular tract (C arlgren, 1912a:60­ tal part of tube, in some nemarocysts, 67)- telocraspedon and histologically simi­ T hread ( Gosse, 1858 :126; 1860: xxix-xxxiv) lar medi al tract of trineme. -tube of nernarocyst ( q. v.) . Spirula- (Stephenson, 1929:179; Weill, 1934 : T hread ( Carlgren, 1940 :3 )-terminal tube of 91- 92 )- nematOcyst containing long slender nernatocyst or entire tube of thickened basal tube with enlarged or armed basal portion portion is absent. (not in present use) . Trineme (Beneden, 1923 :79-80)-adoral tri­ Stomatodaeum (McMurrich, 1890 :136) -ac­ filar region of mesenterial filament. tinopharynx ( q. v.) . T ube (Weill, 1934:21-22, 36-38 )-eversible Stomodaeum ( Bourne, 1900:7, 51) - actino­ hollow stru cture coiled within capsule of pharynx ( q.v.). nernatocysr. 216 PAOFIC SOENCE, Vol. XIX, April 1965

V entral (Kolliker, 1870-1872 )-end of direc­ --- 1889. Observation sur les nernatocysres. tive plane on which single siphonoglyph of Arch. Sci. Phys. N at. Ser, 3, 22:606-608. pennarulids is placed ( by convention ). --' - 1896. Note sur les cellules urticantes. V entral (Beneden, 1897 :12) - aboral portion Rev. Suisse Zool. 3:533-539. of column of Ceriantharia (by analogy with BENEDEN, E. VAN. 1897. Les Anthozoaires de the orientation of Amphioxus ). la "-expedition: ' Ergeb. Plankton­ Ventral ( Haacke, 1879:294; Carlgren, 1893: Exped. Humbolt-Stiftung 2e:1-222. 242-246; 1912a:54-58)- end of directive --- 1923. Travaux posthumes d'Edouard plane opposite single siphonoglyph of Ceri­ Van Beneden sur les Cerianthaires collation­ antharia if siphonoglyph is not considered nes par Paul Cerfontaine, Arch. BioI. Vol. homologous to that of the pennatulids. hors ser.: 1-242. V entral (H errwig and Hertwig, 1879: 572; Fau­ BOURNE, G. C. 1887. The anatomy of the rot, 1895:62, 228, 250; McMurrich, 1910:9; madreporarian coral Fungia. Quart. ]. Micro­ Torelli, 1960 :383)-end of directive plane scop. Sci. New Ser., 27:293- 324. on which single siphonoglyph of the Ceri­ -- 1900. The Anthozoa, pp. 1-84. In: E. antharia is placed if it is considered homolo­ R. Lankaster, [ed.], A Trea tise on Zoology. gous to siphonoglyph of pennatulids. Part II. The Porifera and Coelenterata. Adam and Charles Black, London. ACKNOWLEDGMENTS --- 1919. Observations on Arachnactis al­ I would like to acknowledge the guidance of bida M. Sars. Quart. ]. Microscop. Sci. Ser. Dr. B. C. Abbott throughout this work. Collec­ 2,64: 27- 65. tion and preliminary identification were made CARLGREN, O. 1893. Zur Kenntnis der Septen­ at the Department of Zoology, University of muskulatur bei Ceriantheen und der Schlund­ California, Los Angeles. The taxonomy and rinnen bei Anthozoen. Ofvers. Kgl. Svenska histology were completed at the Departm ent Verenskapsakad, Forh. 50:239-247. of Zoology, University of British Columbia. --- 1900. Ostrafrikanisch e Actinien, ge­ I am most grateful for the facilities and aid sammelt von Herrn Dr. F. Stuhlmann 1888 given me at the Universiry of British Columbia, und 1889. Mitt. Narurh. Museum Hamburg especially by Dr. W. Hoar and Dr. N. Band. 17:21-144. The technical assistance of Mrs. J. Eales and - - - 1906. Die Actinien-Larven, p. 65-89. Mrs. ]. McDonald is appreciated. I am in­ In: Nordisches Plankton, Zoologischer Teil, debted to Dr. C. Hand for reading the manu­ 6, Coelenterata, Part XI. Lipsius and Tischer, script. Kiel and Leipzig. Most of the animals used were collected ---' 1912a. Ceriantharia. Danish Ingolf­ under a University of California, Los Angeles, Exped. 5 (3 ):1-76. Patent Fund Grant to the author. Necessary --- 1912b. -aber Cerianrharien des Mittel­ supplies and equipment were purchased under meers. Mitt. Zool. Sta. Neapel, Berlin 20: a United States Public Health Service Grant 356-394. No . H-4019 to Dr. B. C. Abbott and a Sigma --- 1924a. Papers from Dr. Th. Morten­ Xi Grant-in-aid to the author. sen's Pacific Expedi tion 1914-16, XVI. Ceri­ antharia, Vidensk. Medd. N aturf. Foren, Kjobenhavn 75 :169-195. REFERENcES --- 1924b. Die Larven der Ceriantharien, AGASSIZ, A. 1863. On Arachnactis brachiolata, Zoanrharien, und Actiniarien der Deurschen a species of floating Actinia found at Na­ Tiefsee-Expedirion mit einem Nachtrag zu hant, Massachusetts. Boston ]. Nat. Hist. 7: den Zoantharien. Wiss. Ergeb. Deut. Tiefsee­ 525-531. Exped. Valdivia 19:341-476. BEDOT, M. 1886. Recherches sur les cellules --- 1931. On some Ceriantharia. Arkiv. urticantes. Rec. Zool. Suisse 4:51-70. Zool. 23A (2 ):1-1O. Pachycerianthus torreyi n. sp.-ARAI 217

--- 1940. A contribution to the knowledge --- 1860. AH istory of the British Sea­ of the structure and distribution of the cnidae Anemones and Corals. Van Voorst, London. in the Anthozoa. Lunds Univ. Arsskr, Ser. 362 pp. 2, 36 (3) :1-62. HAACKE, W . 1879. Zur Blastologie der Koral­ --- 1949. A survey of the Prychodactiaria, len. Eine morphologische Studie. Jena. Z. Corallimorpharia, and Actiniaria. Kgl. Sven­ Naturw, 13:269-320. ska Vetenskapsakad. Handl. Ser. 4, 1( 1) : HADDON, A. C. 1889. A revision of the British 1-121. Actiniae, Part 1. Sci. Trans. Roy. Dublin Soc. --- 1951. The Actinian fauna of the Gulf Ser. 2,4:297-361. of California. Proc. U.S. Natl. Museum 101: HAIME, M. J. 1854. Mernoire sur le Cerianthe 415-449. (). Ann. Sci. Nat. CHILD, C. M. 1908. Form regulation in Ceri­ Zool. Ser. 4,1:341-389. anthus aestuarii. BioI. Bull. 15:27-53. HAND, C. 1961. Presenr state of nematocyst re­ search: types, structure and function, p. 187­ CUTRESS, C. E. 1955. An interpretation of the 197. In: H . M. Lenhoff and W . F. Loomis, structure and distribution of cnidae in An­ [ed.], The Biology of Hydra and of some thozoa. Systematic Zool. 4 :120-137. other Coelenterates. Univ, Miami Press, Coral D UERDEN, J. E. 1900. Order of appearance of Gables. the mesenteries and septa in the Madrepora­ HEIDER, A. R. VON. 1879. Cerianth us mem­ ria. Johns Hopkins Univ. CiK.19, No . 146: branaceus Haime. Ein Beitrag zUlP!'~natomie 47-53. der Actinien, Sirzber, Akad. Wiss. Wien --- 1902. Relationships of the Rugosa Math. Naturw. Kl. Abr, I, 79:204-254. (Tetracoralla ) to the living Zoanrheae, Ann. HERTWIG, 0., and R. HERTWIG. 1879. Die Ac­ Mag. Nat. Hist. 109:381-398. rinien anaromisch und histologisch mit be­ FAUROT, 1. 1891. Sur Ie Cerianthus membrana­ sonder Berilcksichrigung des Nervenrnuskel­ ceus (Gmelin ). Mem, Soc. Zool. France 4 : systems untersucht. Jena. Z. Naturw. 13: 66-74. 457- 640; 14:39-89. --- 1892. Sur le developpement du Ceri­ HICKSON, S. J. 1883. On the ciliated groove (si­ anthus membranaceus. Bull. Soc. Zool. France phonoglyphe) in the stomodaeum of the AI­ 17:238. cyonarians. Trans. Roy. Soc. London B, 174: 693-705. --- 1895. Etudes sur l'anatomie, l'hisrolo­ JOURDAN, E. 1879. Recherches zoologiques et gie et le developpernent des Actinies. Arch. histologiques sur les zoanrhaires du Golfe de Zool. Exptl. Gen . Ser. 3, 3:43-262. Marseille. Ann. Sci. Nat. Zool. Ser. 6, 10: FISCHER, P. 1887. Contribution a l'actinologie 1-154. francaise. Arch. Zool. Exptl, Gen. Sec. 2, 5: KINGSLEY, J. S. 1904. A description of Ceri­ 381-442. anthus borealis Verrill. Tufts Coll, Studies --- 1889. Sur la disposition des tenracules 8:345-361. chez Ies Cerianthes. Bull. Soc. Zool. France KOLLIKER, A. 1870-1872. Anatornisch-syste­ 14:24-27. matische Beschreibung der Alcyonarien. Ab­ FREY, H., and R. LEUCKART, 1847. Ueber den handl. Senckenberg. Naturforsch. Ges. 7: Bau der Actinien und Lucernarien, im Ver­ 111-255; 7:487-602; 8:85-275. gleich mit dem ubrigen Anrhozoen,pp. 1­ LACAZE-DuTHIERS, H. DE. 1873. Developpe­ 18. In: Beirrage zur Kenntniss Wirbelloser menr des Coralliaires. Deuxierne memo ire. Thiere mit besonderer Beriicksichtigung der Actiniaires a polypiers. Arch. Zool. Exptl. Fauna des Norddeutschen Meeres. Friedrich Gen. 2:269-348. Vieweg und Sohn, Braunschweig. MaMURRICH, J. P. 1890. Contributions on the GOSSE, P. H. 1858. Researches on the poison­ morphology of the Actinozoa, 1. The struc­ apparatus in the Actiniadae. Proc. Roy. Soc. ture of Cerianthus americanus. J. Morphol. London 9: 125-128. 4:131-152. 218 PACIFIC SCIENCE, Vol. XIX, April 1965

--- 1893. Report on the Actiniae collected TORELLI, B. 1952. Su alcuni particolari aspetti by the United States Fish Commission steamer della istologia di Cerianthus. Pubbl. Staz. "Albatross" during the winter of 1887-1888. Zool. Napoli 23:141-162. Proc. U.S. N at. Museum 16:119-216. --- 1960. Chiarimenti sur alcune strutture --- 1910. Actiniaria of the Siboga Expedi­ dei Ceriantharia, Pubbl. Staz, Zool. Napoli tion, Part 1. Ceriantharia. Siboga Exped. 31:373-385. 15a: 1-78. --- 1961. Un Cerianthus del golfo di Na­ MOBIUS, K. 1866. On the urticating capsules of poli : C. bicyclus n. sp. (Anthozoa) . Pubbl. some Polypes and Acalephs. Ann. Mag. Nat. Sraz. Zool. Napoli 32:17-28. Hist. Ser. 3, 17: 387. TORREY, H. B., and F. 1. KLEEBERGER. 1909. PAX, F. 1914. Die Actinien, Ergeb. Fortschr, Three species of Cerianthus from Southern Zool. 4:339-642. California. Univ, Calif. Pubis. Zool. 6:115­ RAPP, W . 1829. Untersuchungen tiber den Bau 125. einiger Polypen des Mirrellandischen Meeres. VOGT, C. 1888. Des genres A rachnactis er Ceri­ Verhandl. kaiserlichen Leopoldinisch-Caro­ anthus. Arch. Biol, 8: 1-41. Iinischen Akad. Narurforscher 14:643-658. WEILL, R. 1926. Une categoric speciale de ROULE, 1. 1904a. Note preliminaire sur quel­ nernarocysts commune aux seuls hydrides, ques formes nouvelles de Cerianthaires, gyrnnoblastides et siphonophores. Compt. Compt. Rend. Assoc. Franc. Avance. Sci. 32: Rend. 182:1244-1247. 791-793. --- 1930. Essai d'une classification des --- 1904b, Sur un Cerianthaire nouveau. nernatocystes des cnidaires. Bull. BioI. France Compt. Rend. 138:708-710. er Belg. 64:141-153. STEPHENSON, T. A. 1920. On the classifica­ --- 1934. Contribution a l'etude des enid­ tion of Acriniaria. Part 1. Forms with aeon­ aires et de leurs nernarocystes. Trav. Sta. ria and forms with a mesogloeal sphincter. Zool. Wimereux 10:1-347; 11:348-701. Quart . J. Microscop. Sci. New Ser., 64:425­ WELLS, J. W ., and D. HILL. Anthozoa. Gen­ 574. eral features, pp. 161-165. In: R. C. Moore, --- 1929. On the nernarocysts of sea anem­ [ed.}, Treatise on Invertebrate Paleontology, ones. J. Marine BioI. Assoc. United King­ F. Coelenterata. Geol. Soc. Amer. and Univ , dom 16:173-200. Kansas Press, Lawrence, Kansas.