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Oe ieee 1h) 3 i UNIVERSITY OF CALIFORNIA PUBLICATIONS ZOOLOGY WILLIAM EMERSON RITTER EDITOR VOLUME II WITH 19 PLATES BERKELEY THE UNIVERSITY PRESS 1905-1906 ee Ao coma lngtis ~ ey R\\o5 R Lae ~ ef GN & a . \ } } ; | ; f / iA NCL CONTENTS. Introduction. A General Statement of the Ideas and the present No. 1. No. 3. No. 4. Aims and Status of the Marine Biological Association of San Diego, by Wm. E. Ritter. With 2 maps ............ Mhe Jidex of a Marine Biological Survey -...---—---.._.._- iibhempAtreais TOM Wee SUTVC ye Cn = ee mee ee eae eee a we were store em SIM Tre) etdurarits\ eI MRSS G0 eee eee ene een me Order of Advance on the numerous Lines of Investigation Knowledge of the Physical Conditions of the Area —....... Instrumentalities for Prosecuting such a Survey -..............- z IMG CS SiiyarOls van ala TICs bath 25 cteeee cs acer ee Present Status, asi to Ways and Means —...______.--...---- ns GORI C ale NO GCS See ee ete ke eso, cee ee ee eee ee ee Remarks on the present Condition of marine Biology in (Ger eral) eee oe oat mee eee ec gee Ses eee ee ce The Hydroids of the San Diego Region, by Harry Beal BIRO TEE Yeats ofl PLL CS pee eee ee eee eee Hern O CUT CAO Ting see eee Sees eek Dee ee a ee Pees eat ee (Maal ke: Gre ADM perel ove eno al SU ace ene et soe Key to Families, Genera, and Species -............ eo eee tA IDESERIp LOM ot NSP CClCSm eee = aene eee ee een See Salty ey er eas nn rn se esa es ohn eS ee The Ctenophores of the San Diego Region, by Harry Beal TR rs Cy emi ea tl nent ese ne eee en cre ee A estereilOme Ol POCLOS eee aes Mee eee reat cee cece TEST GUI oeat 290) 0h 6 ce Ae ee ns ee oe Rae eee See RR eee ee The Pelagic Tunicata of the San Diego Region, excepting the Larvacea, by Wm. E. Ritter. With figures 9-31. DEAE iS SS ee ee Pee ee JUTGETROXO KUO TKO A eee io ee ec tn RRs See TOP Cab SS] OLSON OTS) «eas cea ese se ne mera eae IDWSOWISISMOT OIE (S]OC OME | aos acess occ Sra ace co eee SEE PAST riVa een NeON VBS OVINGyegt2 4 a) eh) A eee cee eee neec cee eae ne fee ee Hee Ree eer ID eXsrobiaty over a) ea Coes, 6 2b, 5C6 = Nee eee Pe eee nee ee PP Per The Pelagic Copepoda of the San Diego Region, by C. O. Bisbee eg Walhbl GOO UIE OS exes nes coe: oe tener enceat odes nace VETa HATED LU VOLET OI ee ees Soe oe oe ce sore ee Beebe eee Eee Ee erm Ol Giyg sees Ze et Dee Aas es See ae eee TICES HCC BOTS | 0270) a = a a a As HA ad JANUARY. Nantucket. 2 .se% se. 34 22 5.98 16 11,849 60 Key West: 4).2% << one. 73 82 1.42 7 7,834 31 Parallone.. A Ss a a As Pereira 39 29 2. 1d 10 Bre istets 79 70 1.94 4 Be eas 60 49 6.30 24 aie siete Sif AT 6.01 2a Sieaetanchs 63 50 PA ANTS 10 Sd Siti 48 38 4.08 iif Saat anton 80 (Al iL yl of sta epets 57 50 2.29 13 Digroneraee 63 50 1.29 8 Stee 66 52 15 S EON anio 61 50 2.39 1 See ae 82 ike 13.01 12 a cecke 55 51 ss il Be 3 eres 66 52 30 1 eet 65 56 12 3 Naowres 65 54 2.38 12 Nor stewen 86 ath 1 740) 112 satan aue 55 51 OL il Soe 66 52 Trace 0 Siteeeanrc 69 60 0 0 Sie ects 74 62 2.09 9 aiayaeateats 87 77 1.40 i131 omer th tia 56 52 0 0 Bronce 62 52 .02 1 eoctere F/il 62 0 0 Meesaneys 71 61 2). 29 12 Cea aie ae 88 (tat 4.24 13 ae ath 56 53 Trace 0 sane ate 62 52 .06 2 erste ts 76 66 Trace 0 eee 67 56 .78 5 Se. eitSea 87 77 3} 515) 16 atthe state No records. BP coneats al S17 5 OT 5 76 64 Trace 0 WIND. ment in miles. Total move- 11,294 7,242 11,940 7,126 5,041 10,274 7,378 10,890 6,544 4,665 9,033 6,018 14,993 8,921 4,153 9,019 6,856 13,757 9,448 4,531 8,011 6,750 11,974 10,574 4,335 8.377 6,417 11,066 9,674 4,165 8,869 6,092 7,141 4,132 Max. velocity, miles per hour ow w Wo - Oo V vi University of California Publications. | ZooLocy 4 7 3 TEMPERATURE. PRECIPITATION. E je ee 4 = = 2's Ss o = Shs oF sey) = LOCALITY. = a as E : Ea e A = = = a AS oa aq OCTOBER. Nantucket) eS a-ecrne. D7 47 1.01 8 11,700 48 ey. SWiesb. eves ote 82 74 fi 14 8,675 71 Hamalloner ence see ese 61 56 2.01 7 10,791 50 San Francisco ...... 68 56 Deo 7 5,506 32 San Dieco: .f see ee: 74 59 Say 3 4,171 20 NOVEMBER. Nantucket. “tf. es. 46 35 3.29 8 11,394 60 Fieve WeSt) 2.2r¥. cancer 76 68 6.22 12 7,573 43 Harallones .e. . ssen 2 59 51 1.58 6 9,168 43 San Aran CIsGO sae. 63 es LOT 3) 3,851 22 Sans Diegourn a... . es 74 54 0 0 3,930 23 DECEMBER. Nantucket) Js%.4.. 35 25 4.67 17 13,184 64 Ney SWiOSUi:.. ern me's 74 65 . 34 2 6,841 26 Barsalloner eee. tetercis Sil 51 2.22 10 11,431 58 San Francisco ...... 55 46 1.59 10 4,876 38 san) DIe€Z0) «25 ca). 2 66 51 2.46 tC 3,884 19 Perhaps the most important fact, from the present point of view, exhibited by this table is that pertaining to winds. It will be noted, for San Diego, that the maximum velocity for the year was 36 miles an hour, in February. On the basis of the “*‘ Beaufort 9? Seale’’ of wind velocities, this is a ‘‘Strong Breeze.’’ February and March are the climax of the stormy season. For the months May to December, inclusive, the maximum velocities run from ‘*Centle Breeze’’ to ‘‘ Fresh Breeze.’’ La Jolla, the suburb of San Diego at which the laboratory is located, is on a rocky point jutting into the open sea with water of 200 fathoms attainable inside of five miles; so the ecological problems of oceanic plankton, and of bottom-forms can be here attacked under peculiarly favorably conditions. The western boundary of the area corresponds roughly to the edge of the continental shelf in this region, and immediately beyond this 2,000 to 2,300 fathoms are reached. While this extreme depth is distant about 200 miles from San Diego, by making San Nicholas Island a temporary base the 2,000 fathom oe RGUELLO Pt.CONCEPTION Santa Barbara O Los Angeles PrVINC ENTE de ee Hill PrFERMIN Santa Barbara Is.” Begqs Rock © Santa Catalina Is. San Nicolas as a9s San Clemente Is. PTLAJOLLA cheeT 4 >? los. 3g, 569 807 Tanner Bank Bae Gi hoa 4/1 660 324 AIG 69 Cortes Bank 2 743 344 7 : 15@. jo a o2 Bishop Rk;.. 5! Los Cornonados 1053 Map 1.—Showing the Area to be surveyed. Modified from United States Coast and Geodetie Survey Chart. ’ « i ® e i — ' ' . ' ’ i = | ow «aa tenant sles Sane —aii-aihatiiioate amo te ” i r , Crit 4 Garr it ah OP Pyadodnlnu Ase i ale Vou. 2] Ritter—Introduction. vu curve is only 65 miles away. Within the area is a wide range of depth and great variety of bottom. A basin 40 miles off Point Loma has a depth of over 1,000 fathoms. On the other hand, the Cortes Banks, just beyond the southern boundary, carry but 15 feet of water at low tide. There can be no doubt that deep sea and ’longshore investiga- tions have not yet been brought together to the extent they ought to be. 3.—-The Initial Step. The first step in such a survey would obviously be to find out what plants and animals inhabit the area; to establish a speaking acquaintance, as one may say, with the organisms that are later to be more intimately known. So far this has absorbed most of the effort, and it will of necessity demand the continu- ance of much effort for a long time in the future. The ideal being kept always in view, the mere description of the new spe- cies for the exclusive use of expert taxonomists in the several groups, would not be sufficient. The entire fauna and flora must be recorded in such a way as to make the records a good founda- tion for the broader and deeper studies to follow. These consid- erations have determined the character of the faunistic papers now published, and that will come hereafter. The present vo!- ume contains the following contributions to a knowledge of the fauna: No.1.—The Hydroids of the San Diego Region, by Professor iE Bs lorrey. No. 2.—The Ctenophores of the San Diego Region, by Pro- fessor H. B. Torrey. No.3.—The Pelagic Tunicata of the San Diego Region, except- ing the Larvacea, by Professor Wm. E. Ritter. No. 4.—The Pelagic Copepoda of the San Diego Region, by C. O. Esterly. No. 5.—The Nonencrusting cheilostomatous Bryozoa of the West Coast of North America, by Dr. Alice Robertson. No. 6.—The Dinoflagellata of the San Diego Region, by Pro- fessor C. A. Kofoid. vill University of California Publications. | ZooLoGy Not only are new species described, but all that have thus far been found in the area are characterized, and in most eases illus- trated by figures, so that these papers will constitute a series of hand-books, as far as they go, for the identification of the species treated. It is also intended that the bibhographical lists accom- panying the papers shall serve as useful guides to the literature of the several groups for those who may take them up for the study of special problems connected with them. 4.—Order of Advance on the Numerous Lines of Investigations. While there is no reason for attempting a rigorously laid out order of attack on the numerous problems, at natural sequence, within certain limits, will establish an order; and where practical administrative conditions conveniently adapt themselves to such sequence this order will be followed. For example. the species representing a given pelagic group having been got well in hand, a natural second step would be the determination of the seasonal distribution of the group, since the study of the collections for the taxonomy would surely bring together, incidentally, consid- erable data on this problem. Following close upon the treatment of seasonal distribution would come that of horizontal and ver- tical distribution, the chorology ; and inseparably linked with these would be the problems of food and reproduction ; and these again would lead to problems of migration, with their intimate depend- ence upon temperature and other environmental factors. And here, completeness of knowledge being ever the watchword, the demand would arise for applying experimental and statistical methods in the effort to get at the deeper significance of the facts observed, and generalizations reached from the observational investigations. The chain of questions hanging one to another is endless and, of course, completeness of knowledge in a literal sense, 1S an unattainable ideal. 5.—Knowledge of the Physical Conditions of the Area. It does not need to be said, in the light of general biolog- ical conceptions reigning in this day, that an aim at comprehen- siveness of knowledge cannot for a moment neglect the physical] conditions under which organisms live. What has to be consid- j : vp: A sta id hie - Be hamlet) abd eh 1 x aoe i= Sas Oe Lad are Dk R' ; : 7 iene | ve Boundary vSAN DIEGO Uv) 0) a =a O -2 a @ Fu Sin ll an oss Vis — (i Nit owing bottom contours in fathoms. We Hie F Whistle NAL A Wii -y = NEY AAyt= G yh WW 4 ies ALUN tpeogyy AAW or 72s » sk )» by Professor George Davidson. C iS pe os = po a al N ce al a S) > oy ~ a 9 Map —— Vou. 2] Ritter—Introduction. 1X ered in connection with a marine undertaking like the present, is the specific things that must be done, and the means for doing them. Oceanography is in position to hand over to the marine biologist, ready prepared, a large amount of the information he must have; and, likewise, physics and chemistry have important resources that can be drawn upon. But these general sources in nowise obviate the necessity for constant and searching studies on the sea water in connection with such a survey as that contem- plated. Conditions of the water as to temperature, and currents; mineral, gaseous, and albuminoid content, ete., must be known at the particular time and place to which the biological studies per- tain, and no general knowledge of this character can suffice. Physies, chemistry, and hydrography must, therefore, be integral parts of such a survey. 6.—Instrumentalities for Prosecuting such a Survey. It is obvious that no small outlay of money would be essen- tial for even a good beginning; and that considerable progress in it could be made only with large expenditures for both equipment and operation. The ideal laboratory building would not be large, but would be constructed with great care. Aquaria would con- stitute an important element in the plant for the work on shore. From $50,000 to $75,000 should build and equip an ample labora- tory and aquaria. Equipment for the work at sea would demand the greater portion of the capital. For the deep-water work a ship of the class of the U. S. Fisheries steamer Albatross would be essential. For less depths, say 1,000 fathoms and under, a much smaller vessel would be as efficient or even more so, since it can be han- dled so much more quickly. As noted above, our area is extremely favorable for this purpose. ) | as follows: second muscle not bifurcating ; third bifureating near dorsal line as on opposite side, the anterior branch again bifur- eating to send a branch forward which anastomoses with second muscle, the single band thus produced passing into the ventral pedunele, Pl. III, fig. 8. In the asymmetry of the muscles this species resembles SN. rostrata, S. punctata, S. magalhanica of Ap- stein. The climax of the phenomenon is reached in SN. rostrata. First and second muscles united on each side by two longi- tudinal bands, Pl. III, fig. 7. In addition to the dorsal hp muscle given off as a branch from first body muscle, an angular lip muscle on each side its dorsal limb reaching over and becoming continuous with its mate of the other side; another small upper lip muscle near edge of lip. The fourth body muscle of dorsal side, relatively very small, bifurcating on each side short distance from dorsal median line, the anterior branch anastomosing with posterior branch of third muscle, and pos- terior branch passing under atrial orifice. The fourth ventral muscle is really the posterior branch of third dorsal muscle. As already stated, the ganglio-hypophyseal complex and the intestinal tract are clearly immature in the largest zooids seen. I, consequently, refrain from including a characterization of them in the diagnosis of the species. Certain facts about the intestinal tract, however, should be mentioned. In the first place, the late period in the life of the zooid at which it becomes complete, at least as compared with C. affinis, is noteworthy. In the latter species, the intestine has assumed its final form and_ position while the bud is still in the chain, and even before the whorls are formed. In C. bakeri, on the contrary, the way in which the anal end of the intestine projects freely from the posterior end of the body as a whole, shows clearly that the organ is not vet complete, even in the oldest zooids found. With little doubt the final form is a cirele here as in affinis. An apparently wholly unique feature in bakeri, however, seems to be the two appendages of the intestine shown in the figures. The intestinal tract as a whole has the form of a horseshoe, the plane of the shoe being approximately at right angles with the sagittal plane of the zooid. The entire bow projects backward beyond the posterior end of the endostyle and gill. The mouth of the esophagus is 58 University of California Publications. [| ZooLocy situated a little dorsal and to the right of the end of the endo- style; and the anus enters the atrium to the left of the endostyle. The esophagus is marked off from the stomach by being distinctly less in diameter than the stomach. At the anterior end of the intestine a cecum nearly as large as the intestine itself is given off, which curves backward and upward and forms a very con- spicuous object in all the stages of development observed, co., figs. 7 and 8. From the posterior extremity of the intestinal bow a great finger-like outgrowth of the mantle extends backward and up- ward. This appendage is even longer and more conspicuous than the stomachal cecum described above. Into it extend prolonga- tions of the posterior branches of both dorsal and ventral body muscles ; and in addition it contains a well defined axial strand, the connections and nature of which are doubtful, p. d., figs. This appendage would seem to be comparable with the portion of the post-abdomen that extends beyond the intestinal loop in various compound ascidians. The axial strand is probably the testis, or a portion of it. The ovary is situated on the right side of the body at the extreme posterior end, midway between the atrial orifice and the esophagus; and the oviduct, which is unusually long, extends forward to a level with the third ventral muscle band. C. bakeri appears to have more in common with C. floridana, Apstein, than with any other known species. It is, however, very distinct from this latter, as is obvious from the following, among several other differences: The largest specimen of C. floridana seen by Apstein was 12 mm long. In view of the con- siderable number of specimens taken by the Plankton Expedi- tion, the great disparity in size thus indicated shows pretty con- clusively that C. bakeri is a much larger species than floridana. The lateral glandular organs of the solitary floridana are dis- tinctly less extensive than in bakeri, and are, according to Ap- stein’s statement, continuous on each side as in C. pinnata. The intestine of floridana has a single appendage, while that of bakeri has two. The closest resemblance between the two is in the mus- culatures of the solitary forms, but even here there are well marked differences, which, however, need not be dwelt upon, VoL. 2] Ritter—The Pelagic Tunicata. 59 since a comparison of my figures with that of Apstein will make them clear. It is worth noting that this adds another to the list of species in which the intestine of the aggregate genera- tion is in the form of a circle, these species being C. affinis, C. floridana and C. baker. The specimens of C. bakeri thus far obtained are few, and the variations in the muscle bands in these few suggest that fur- ther study of more ample material may modify somewhat the scheme given in the diagnosis; I cannot, however, believe that such modification can materially alter the results so far as concerns the definition of the species. In no species of salpa with which I have had experience have I found so much difficulty in tracing the muscles. This difficulty is due to the softness of the animal, and the extreme delicacy and transparency of the muscles them- selves. The separation of the muscles into body muscles on the one hand, and orifice muscles on the other, I recognize as being a particularly arbitrary matter in this species. For example, there would be almost as much reason for considering what I have enumerated as the first body muscle, a lip muscle; or, on the other hand, for calling what in my scheme is the third upper hp muscle, a body muscle. In fact, I have little doubt that what Apstein has designated number one in floridana corresponds to my third dorsal lip muscle. But the homologizing of the muscle bands in the different species of salpa is an exceedingly difficult, if indeed possible, thing; though comparison of the develop- mental stages would probably help in the matter. In all, about fourteen specimens of the species have been taken during the last three years, all on the coast of southern California, and all excepting one, which was taken in March, during the months of June and July. Cyclosalpa affinis (Chamisso). Salpa affinis Chamisso, 1819, p. 11, pl. figs. 2A-C, solitary genera- tion; 2D-H, aggregate generation. Cyclosalpa affinis Blainville, 1827. Salpa affinis Meyen, 1832, p. 407. Salpa affinis Traustedt, 1885, p. 357, Pl. I, figs. 6, 7, and 8. Cyclosalpa affinis Herdman, 1888, p. 86. Cyclosalpa affinis Lahille, 1890, p. 11. Salpa affinis Apstein, 1894, p. 4; Cyclosalpa, p. 24. 60 University of California Publications. | ZooLocy (a) Solitary (budding) generation—F ig. 9. Body distinetly larger at anterior end, and tapering nearly uniformly to the posterior end; the anterior end with a pronounced ventral bend. the posterior with a nearly equal dorsal bend. Test rather thin and soft, and highly transparent, without special thickenings or Fig. 9.—S. affinis, solitary generation. asperities; a pair of lateral and somewhat dorsal appendages at ‘ the posterior end. No lineaform ‘‘glandular’’ organs. Leneth of full grown zooids, 80 to 100 mm. Branchial orifice directed somewhat ventrad, lips prominent, upper overarching. Atrial orifice without lips, directed somewhat dorsad, armed, particu- larly in older specimens, with a pair of latero-dorsal appendages. Body muscles eight, all excepting last interrupted on ventral side. First and second interrupted on dorsal side (in older specimens only). First trending backward to some extent on ventral side. Lip muscles of branchial orifice complicated; two sphincters in upper and one in lower; two longitudinal bands on each side running forward from the first body band, one to the dorsal lip, the other to the ventral; a band on each side extend- ing from the angle of the orifice postero-dorsad; a pair of short, strong dorsal longitudinal bands in dorsal lip. Endostyle dis- tinetly curved in its anterior third to correspond with the ventral bend of the body as a whole; extending from far forward in the ventral lip back to the esophagus; both extremities turned abruptly up, the anterior more conspicuously so than the poste- rior. Ganglion slightly in front of anus. Hypophyseal mouth a large, highly convoluted, nearly closed ring, with opening to the left (fig. 11). Gill long and narrow, extending from slightly in front of the anus to the esophagus, terminating, consequently, —-. VoL. 2] Ritter —The Pelagic Tunicata. 61 ti Fig. 11.—Hypophysis of C. affinis. nearly coincidently with the endostyle. Intestine nearly straight and of uniform character, though turned a little dorsad and to the left at anal end; the anus somewhat trumpet shaped. Stomach in the sharp curve of the tract, flattened; a large cecum extending backward appearing as a direct posterior prolongation of the intestine. Entire intestine uniform orange, generally, but occasionally devoid of color. Pericardium-heart large, situated ventrad of the posterior end of the endostyle. Salpa chain reach- ing forward in mid-ventral line under the endostyle and emerg- ing to the outside far forward, between the ventral ends of the first body muscle band. Fig. 10.—C. affinis, aggregate generation. (b) Aggregate (sexual) generation.—Fig. 10. Agegregations containing from nine to twenty zooids, united radially about a common centre, by the large ventral peduncle, remaining intact until zooids are fully grown; in nature, six or eight or more of 62 University of California Publications. | ZOOLOGY the whorls united tangentially.- Body in general cylindrical, though somewhat arched dorsally, and tapering at the atrial end. Rather soft and of uniform consisteney, there being no thick- ened or specially stiffened areas in the test; the connecting peduncle and protruding intestinal tract forming very promi- nent projections from the ventral side. Full grown zooids from 60 to 70 mm. long. Body usually quite transparent and without pigment, but occasionally traces of pink on surface of test about anterior end. Lips of branchial orifice prominent, of approxi- mately equal size; atrial orifice without lips. Body muscles five on the dorsal side and six on the ventral, all continuous across the dorsal side, and all except sixth interrupted by a narrow interval ventrally; the fifth and sixth ventral muscles joining laterally to form the fifth dorsal. The second band giving off a longitudinal muscle laterally, which extends forward to the angle of the branchial orifice; the fifth giving off a small branch on each side, which extends back close under the intestine; the sixth, with a branch on each side connecting with the first atrial. A well developed band in each lip of the branchial orifice meet- ing in a wide angle on each side. First atrial band nearly as broad as body bands; the other atrials, about eight in number, very delicate. Endostyle extending from slightly in front of the first body muscle to behind the fifth muscle, and quite to the intestine. Gill extending from somewhat in front of the second muscle fully to the intestine, hence terminating near the poste- rior end of the endostyle. Ganglion and sense organ under the second body muscle, and a little posterior to the anterior end of the gill. Hypophyseal mouth close in front of the anterior end of the gill, large and much convoluted, becoming rosette shaped, the band narrowly open to the right. The intestine projecting from the ventral side like a hernia; forming almost a circle of relatively large size, in full grown zooids 1.5 em. in diameter, the curve extending downward and forward, so that the anus is very near to, and to the left of, the esophageal mouth. Stom- ach scarcely larger in diameter than intestine, and not distinctly set off from it. Esophagus short and much smaller in diameter. Deep yellow, nearly uniform throughout. Heart conspicuous, immediately in front of intestinal ring on ventral side. Placenta A A a eS Vou. 2] Ritter—The Pelagic Tunicata. 63 with embryos on right side between fourth and fifth muscle bands. Testis, an elongated white mass in the intestinal ring closely applied to the intestine. Vas deferens given off from the middle of the testis, passing across the intestinal circle, and opening near the esophageal mouth. Down to the present time this has been one of the rarer of the early known species of salpa. It was described by Chamisso, by whom it was taken in the region of the Sandwich Islands. It did not occur in the Challenger collections, and has not until now been reported again from the Pacific, so far as I am aware. Meyer observed it, though not closely, nor in great abundance, about the Canary Islands, and we have several other ref- erences to its occurrence in the tropical and_ subtropical Atlantic. The Plankton Expedition took, according to Apstein, a total of nine specimens at two stations; one in the Gulf Stream, Lat. 41.6, the other in the Sargasso Sea, Lat. 31.5. Voigt, 1854, includes it in his list of species of the Mediterranean in the vicinity of Nice, but gives no further information about it. Its constant abundance on the southwestern coast of North America, in a plankton area at least adjacent to, if not in reality part of, that from which it was originally described, and its apparent rarity in other parts of the world, would seem to indicate that the headquarters of the species is here, though such a suggestion relative to the distribution of strictly pelagic organisms can have little value until supported by much more data than we yet possess. Despite the considerable differences between the Salpa here treated as C. affinis and any of the published descrip- tions and figures of the species, I am convinced of the correct- ness of the identification. All the discrepancies of any moment may be accounted for from the fact that the descriptions and figures hitherto published have probably all been made from museum specimens. For example, the straight long axis of the solitary generation as shown in the figures of Chamisso and Traustedt, give a wholly erroneous impression of the general form of the species, but the true shape as shown in lateral view, fig. 9, ean be fully appreciated only by examining the living swimming animal. Preserved specimens have more the form of the figures of the authors above mentioned. Again, the two 64 University of California Publications. | ZooLocy processes at the atrial end are not adequately recognized in any of the published figures. This is due in part to the fact that they are very short in the younger zooids, and in part to their being, in adult preserved specimens, either broken or worn off, I have examined specimens of about 1.5 em. in length from the mid-Pacific, practically the region in which Chamisso obtained his, and find the processes short though distinctly indicated, essentially as they are shown in Traustedt’s figs. 7 and 8, Pl. I. Finally, Traustedt figures an arrangement and anatomosing of the seventh and eighth muscle bands of the solitary generation, somewhat different from anything I have seen, but the point is a trivial one, even if his representations are entirely correct. The species has been obtained at Santa Catalina Island and off San Diego during the months of March, June, July, and August. It was particularly abundant during March, 1904, in the last named locality, and was reproducing actively, both sex- ually and asexually. Salpa fusiformis-runcinata Cuvier.-Cham. Salpa fusiformis Cuvier, 1804, p. 23, fig. 10. Salpa runcinata Chamisso, 1819, p. 16, Pl. figs. 5A-51. Salpa runcinata-fusiformis Krohn, 1846, p. 112. Salpa runecinata-fusiformis, Leuckart, 1854, p. 3 ad seq. Pl. I, figs. 6, 8, 16, 17, 18; and Pl. IT, figs. 1, 3, 4; 5, 13, 15, and 18. Salpa runcinata-fusiformis Traustedt, 1885, p. 370, Pl. 2, figs. 29, 30, 31. Salpa runcinata-fusiformis Herdman, 1888, p. 76, Pl. 6, figs. 5-12. Salpa fusiformis Apstein, 1901, p. 1117, figs. 6a and 6b. Fig. 12.—S. fusiformis-runcinata, solitary generation. (a) Solitary (budding) generation —Fig. 12. Nearly eylin- drical, somewhat larger at the atrial end, both orifices terminal, truneate in general effect at both ends. Length of largest speci- os Si _ ee ee Vou. 2] Ritter—The Pelagic Twnicata. 65 mens, 70 mm., 76 mm., to 80 mm. Test variable, in some thin and soft, particularly anteriorly; in others, thicker and firmer, particularly posteriorly; a number, from eight to twelve, more or less regular, longitudinal, serrated ridges running from the atrial end forward a variable distance, but most prominent over the nucleus. Branchial orifice with two lips, of which the dorsal is distinetly the higher. Atrial orifice also with inconspicuous dorsal and ventral lips. Body muscles nine, confined to the dorsal side and reaching down laterally scarcely half way to the endostyle, where they terminate abruptly. First three anterior body muscles confluent dorsally; all the muscle bands distinctly broader dorsally. A constrictor muscle in the dorsal lip, and one in the ventral lip, both terminating behind the angle of branchial orifice, where the ends cross each other and extend a short distance beyond the point of crossing. Atrial orifice with six or eight muscles of a few fibres each, those of the dorsal and ventral lips terminating at the angles, where they cross one another. Endostyle slender, straight, extending from the level of the angle of the branchial orifice back to the intestine, on a level with the eighth muscle band. ‘‘Gill’’ long, narrow, and nearly straight, extending from a little in front of the first muscle band to the ninth band. Hypophysis horseshoe shaped, its plane nearly in the sagittal plane of the animal’s body, about midway between the muscle band of the upper lip and the first body band. Intestinal tract making a compact ‘‘nucleus,’’ cor- responding to the interval between the eighth and ninth muscle bands, and projecting somewhat on ventral side; the broad short end of the rectum projecting dorsad from the nucleus to open into the cloaca. Dark red generally, though not universally. Heart on ventral side, immediately in front of intestinal mass. Chain of buds extending forward along ventral median line from near the nucleus for a variable distance, then bending on itself and reaching back to emerge to the outside through an orifice behind the nucleus. (b) Aggregate (sexual) generation——Fig. 13. Body ellipt- ical in outline, with processes at each end, short and broad in the young, much longer in full grown zooids, where they become as long as the body. At the outset these processes are always 66 University of California Publications. [ ZooLoGY asymmetrical, the anterior being to the right, the posterior to the left, or vice versa (the one figured is anterior process right, and posterior left). Length of large specimen, 25 mm.; usually smaller. Branchial orifice inclined distinctly upward, though not wholly dorsal. Atrial opening nearly directly backward. Lips of branchial orifice not prominent, dorsal deeper, but ven- tral extending farther forward in correlation with the obliquity of the orifice. Atrial orifice scarcely lipped. Body muscles Fig. 13.—S. fusiformis-runcinata, aggregate generation. seven, never interrupted on dorsum, and never continuous across the ventral side. Anterior four, and posterior three, confluent on dorsum. Last of anterior group and first of posterior group confluent laterally, but the two disconnected at ends. Posterior two, the sixth and seventh, confluent well down the side, the two separating, the sixth to pass in front of the nucleus, the seventh — behind it. The seventh usually confluent with a smaller muscle belonging to the atrial orifice. A large muscle band in the dorsal lip of the branchial orifice a short distance back from the edge, a delicate one at the very edge, and a broad band in the ventral lip. An angular muscle band at the angle of the branchial ori- fice on each side, its angle directed toward the angle of the orifice, and its two limbs directed, the one ventrad, the other dorso- posteriad. Endostyle slender, extending far forward under the ventral lip to a level with the ventral ends of the sixth muscle, some distance in front of the nucleus. Gill rather shorter, rela- tively, than in the solitary generation, and making a wider angle with the endostyle. Anterior end about middle of the interval between the dorsal lip and the anterior group of muscle bands, posterior end at the nucleus, hence some distance behind the posterior end of the endostyle. ‘‘Nucleus’’ rather small, com- pact, egg shaped, situated far back, projecting somewhat from Vou. 2] Ritter—The Pelagic Twnicata. 67 the general surface of the animal. Color of nucleus orange, though not uniform in all parts. Heart between posterior end of endostyle and nucleus. This is by considerable the most abundant species of salpa of the western shores of North America, and probably of the whole Pacific Ocean, at any rate north of the equator. It has been taken at almost every point on the coast from Alaska to Lower California, and at many of these in large numbers. On the whole California coast it has been taken in nearly every month of the year, though the systematic collecting at San Diego thus far indicates it to be considerably more abundant during the summer than in the midwinter months. Salpa fusiformis-runcinata, form echinata. Fig. 14.—S. fusifornis-runcinata, echinate form. Postero- dorsal view, showing serrations of test, and muscle bands. pop. 1 ' 1 | i ! i} ; t , ' 1 pap. : i S772. s.m0.T lr Mig. 15.—S. fusiformis-runcinata, echinate form, ventral view of surface. 68 University of Califorma Publications. | ZOOLOGY Having worked over a large quantity of material of S. fusi- formis-runcinata with reference to the question of the status of echinata, I reach the conclusion that a well marked style, or form of the species, which may be ealled echinata, must be recognized, but that it is not a distinct, persistent variety, as Apstein has treated it: much less a species as Herdman concluded from his study of the Challenger material. There are two particulars by which, at their fullest expres- sion, the form is distinguished from the typical fusiformis- runcinata. One is the echination of the test, the other the arrangement of the body muscles. Figs. 14 and 15 are dia- erams, though made with special care, from the examination of three specimens which agreed almost perfectly in these partic- ulars. Fig. 14 is a postero-dorso-dextral ‘view. It shows two double rows of echinations on the dorsal surface that begin some distance behind the anterior end and extend to the posterior end, but do not terminate in spines. lLaterally from these is a row on each side on the edge of a prominent ridge—almost a fin— of test, this ridge extending the entire length of the body and terminating posteriorly in two prominent processes, 7. and 7. p. d. p. On the ventral side are two sets of submedian rows (Fig. 15, s. m. r. and s. m. r.’), an anterior and a posterior, each-set open anteriorly but joined posteriorly, the posterior junction of the posterior set being in a prominent posterior ventral process p. v. p., and laterally from these are again two more rows. As to the muscles, the three anterior ones, though converging some- what, scarcely touch one another as they do typically in fusi- formis-runcinata. The eighth and ninth do not even converge; they are entirely parallel. Herein is perhaps the most striking difference between echinata and the type of the species. This description applies exclusively to the solitary genera- tion. As to the aggregate generation, one finds an occasional lot of zooids that are unusually robust, this being especially apparent at the posterior end of the animal, where the posterior process of test becomes much thickened and solidified, with the serrated edges highly developed. In one lot of this sort observed off San Diego in March, 1904, the animals reached a total length of 50 mm. or more; and several zooids in one gathering made ar he een tases waaay a ig el Oa Vou. 2] Ritter—The Pelagic Tunicata. 69 by the Albatross (data as to time and locality lacking, but cer- tainly Pacific Ocean material) a total length of 60 mm. was reached, the body here, exclusive of the processes, being 40 mm. These last were somewhat larger than the largest Challenger specimens of echinata. That these robust aggregate zooids belong with the echinata form of the solitary generation may be held as probable, although nothing less than absolute proof of this will warrant associating them positively in classification. Now a few more words in support of my opinion that we have here a ease of extreme, for this group of animals, individual vari- ation, or fluctuation, rather than a true variety, or ‘‘elementary species.’’ In the first place, as to the echination of the test. It is doubtful if this is ever wholly absent in S. fusiformis- runcinata. Certainly if it is, it is so only exceptionally. So far as the evidence goes on this point, it is to the effect that the thickening of the test at the posterior end, the prominence of the ridges, and the serrations increase with the size, and pre- sumably with the age, of the zooids in both generations. But more extended and exact information is needed here. There are undoubtedly some observations opposed to this supposition. For example, I have one specimen of the solitary generation taken at Bolinas Bay, California, November 18, 1895, which, although seareely more than half the size of the largest echinata, yet possesses the longest, heaviest three posterior processes I have seen in any zooids whatever. But here the serrations are almost entirely wanting. In this specimen, too, the muscle plan is strictly that of fusiformis-runcinata—that is, the anterior three and posterior two are fully fused. And here I would say that the examination of a large number of specimens with reference to the point has failed to discover a single instance of the sepa- ration of these muscles in a small zooid. I consequently incline to the opinion that the separation of the muscles is an age char- acter. But here, too, more positive evidence is needed. My pro- visional conclusion is, then, than echinata is an old age form of fusiformis-runcinata. In view of the usually clear delimitation of species in Salpa, the question of the status of echinata is especially interesting. It well deserves more extensive and critical examination. Ap- 70 University of California Publications. | ZOOLOGY stein’s suggestion that we have here a variety that pertains to the solitary generation alone is interesting, but can hardly be regarded as of much value until established by direct evidence. This form has been taken several times at various places on the California coast during the last ten or twelve years. It would seem to be coincident with the typical fusiformis-runci- nata in distribution. Salpa tilesii-costata (Cuvier-Quoy et Gaim.). Salpa tilesii Cuvier, 1804, p. 375, figs. 3-6. Salpa costata Quoy et Gaim, 1834, Zool. t. 3, p. 587, Pl. 86, figs. 1-5. Salpa costata-tilesii Krohn, 1846, p. 114. Salpa costata-tilesii Traustedt, 1885, p. 379, Pl. 1, figs. 10 and 11; and Pl. Il. figs. 38-41, and 47. Salpa costata-tilesii Herdman, 1888, p. 60, Pl. 4, figs. 1. 4, 8. Salpa costata Brooks. 1893, p. 10 (particularly), Pl. IV, fig. 4; Pl. VIII, fig. 4, Salpa Tilesii Apstein, 1894, p. 16; 1901, p. 111 10, figs. lla, 11b. * \ ; Fi f | | eee hie AVP D> 2 = held Les ee ars : | Sper \ 9 ¢ “ f ooernad | Fig. 16.—S. tilesii-costata, solitary generation. (a) Solitary (budding) generation.—Fig. 16. Body much larger anteriorly, tapering gradually back to the region of the nucleus, then expanding again rather abruptly to the atrial ter- mination. A prominent hump on the ventral side corresponding to the nucleus. Anterior end rounded: posterior truncate but for slight lateral notches, and armed with a pair of firm com- pressed, green edged appendages arising laterally from a little in front of the edge of the atrial orifice. Both orifices distinctly terminal. Lips of branchial orifice prominent, the ventral some- what more so, upper finely serrated on its inner edge. Length of longest specimen, 19 em., exclusive of appendages, which were 47 mm. Test thick and firm, particularly on the ventral side, — ———'— —~a= a eee ep ———_————- = @& VoL. 2] Ritter.—The Pelagic Tuncata. 71 and most of all over the nucleus. Whole surface, except for a broad area at the anterior end, beset with low, spine like processes. Frequently a girdle of indefinite limitation of dark green in the mid body, across the dorsal side and extending well down toward the mid-ventral line, the color, which is on the surface of the test, being densest laterally. The edges of the posterior append- ages densely and uniformly colored with the same green. Body muscle bands 20, occasionally 19 or even 18, uniformly spaced and mostly all parallel, but the first and second inclined some- what backward on the dorsum. All (in old zooids only?) limited to the dorsal half of the body, and all interrupted in the mid- dorsal line, and typically, the 7th or 8th or 9th interrupted laterally. A single broad band in dorsal lip broadly interrupted on each side of median line; also a pair of short, longitudinal, nearly parallel, widely separated bands in this lip; two bands in ventral lip; a complicated and variable crossing of short bands at angle of orifice. Nine or ten or more delicate, wavy, more or less interrupted bands belonging to the atrial siphon. Endostyle slender, gently curved to conform to the outline of the body, reaching entirely back to the nucleus. Guill relatively rather short, extending from a little in front of the first muscle band to the nucleus. Hyphysis close to anterior end of gill, but short distance in front of ganglion, forming a distinctly pendant tuberele, on which the rather large, irregularly triangular mouth mouth is situated. Fig. 17.—Costata-tilesii, aggregate generaion. (b) Aggregate (sexual) generation.—F ig. 17. Body in gen- eral cylindrical, but irregular, especially posteriorly from the 72 University of California Publications. | ZOOLOGY projection of the intestinal mass and the atrial siphon. Remain- ing in the aggregated condition and firmly united at least until 7 em. long, and while thus united, body somewhat asymmetrical from the mode of aggregation, the branchial and atrial orifices being turned respectively to the right or left, depending on whether the zooid be right or left in the chain. Largest zooid seen, 14 em. Test rather thick and stiff, particularly on ventral side, and most of all over nucleus, where in old zooids it becomes opalescent. Surface in some, though not in all, beset with low, broad, scattered processes, these on the whole more pronounced on dorsum. An irregular area of yellowish green in the test over the nucleus, and occasional small patches of this on dorsum. Lips of branchial orifice prominent, of nearly equal height, the dorsal overarching; ventral projecting forward and below in a blunt prow. Atrial siphon narrow, thin walled, elongated, with- out lips. Body muscle bands five, limited to the dorsal side, and not extending more than half way down to the mid-ventral line. The first three drawing together, but not touching on the dorsum, and each interrupted by a narrow interval in the mid-dorsal line. These three muscles, and frequently the fourth, interrupted on the side toward the axis of the chain, but usually not on the other side. Fifth muscle forked on each side. A single strong band in the dorsal lip, widely interrupted on both sides of the middle, two bands in the ventral lip, the dorsal and ventral lp bands crossing and intermingling in a complex but somewhat variable way at the angles of the orifice. Two short longitudinal bands in dorsal lip. Numerous delicate bands in atrial siphon, all confluent with a longitudinal bantl on each side. Endostyle slender, nearly straight, reaching back entirely to the nucleus. Gill relatively short, scarcely reaching into the anterior third of the animal; hypophysis small, hardly recognizable without dis- section. Nucleus relatively large, compact, ovate, regular; rectum far back, projecting dorsad several millimeters above the gen- eral level of the nucleus, greenish brown at its anterior end, yellow posteriorly, with an irregular scarlet area on its dorsal side and extending somewhat on to the gill. Embryos normally four, situated dorsally to the right of the median line, between the fourth and fifth body muscles. a Se ee Vou. 2] Ritter—-The Pelagic Tunicata. 73 Although this, by far the largest, most magnificent of all our species of Salpa, has never been taken in great abundance on the California coast, it is by no means rare, since a few speci- mens at a time have been frequently collected at numerous points during the last fifteen years. The collections show it to have been taken in March, May, June, July, August, and November, with the largest numbers in March and July. Salpa democratica-mucronata Forsk. Salpa democratica Forskahl, 1775, p. 113, Pl. 36, fig. G (solitary gener. ) Salpa mucronata Forskahl, 1775, p. 114, Pl. 36, fig. D (aggregate gener. ) Salpa Cabotti Agassiz, 1886, p. 17, figs. 1-5. Salpa democratica-mucronata Krohn, 1846, p. 112-113. Salpa democratica-mucronata Leuckart, 1854, p. 3, et seq., Pl. I, figs. 1, 3, and numerous others. Salpa democratica-mucronata Traustedt, 1885, p. 365, Pl. 2, figs. 25 and 28. Salpa democratica-mucronata Herdman, 1888, Pl. VILI, figs. 1-10. Salpa democratica Brooks, 1893, pp. 6-16, particularly for anat- omy. Pl. 2, especially. Thalia democratica-mucronata Herdman, 1899, p. 748. Salpa mucronata Apstein, 1901, p. IIT 5, figs. 5a and 5b. Fig. 18.—S. democratica-mucronata, solitary generation. (a) Solitary (budding) generation—Fig. 18. Form ovate, the posterior end armed with a pair of long,slender, bilaterally placed processes, with a median ventral process, usually bifid, the ventral horn much shorter than its mate, which is sometimes nearly as long as the laterals; usually with a median dorsal process, sometimes of considerable length, but more frequently 74 Universily of California Publications. [ ZOOLOGY short or occasionally wanting. Length of body, exclusive of processes, about 8 mm. Test very thick and exceedingly trans- parent. A blunt pocket of the mantle reaching into the base of the long lateral processes. Branchial orifice inclined somewhat upward, with dorsal and ventral lips, the dorsal distinetly higher and broadly notched in middle. Atrial orifice nearly terminal, but inclined a little upward, without lps. 188 5. Calanus finmarchicus Gunner ..... 20-0. cccee eee sree een eeece 125 65 CALAIS OMGCTIS WANA ene ~io's et oeatel es ets ote ate take) eel acest ental) ciate 128 = GOlANUS MNANOT. GICSDEGGING) .'- cia) c ioe etacieuete leis Gia veetaiweatale tee wi tlerorete cs 126 8: OCalanus robustior Giesbrecht <<< <<... 223 His Savanna, WeMinuan Csr 565 o000cano sed condo o0b0G0de 222 GYMNOPLEA. Gymnoplea (sub-order) Giesbrecht, 1892, p. 41. Gymnoplea (tribe) Giesbrecht, 1898, p. 7. I Sub-order. The genital orifices lie in the first segment of the posterior division of the body; they are ventral and paired in the female, unpaired and lateral in the male. The fifth pair of feet in the female are like the preceding pairs, retrograded or lacking; in the male a pairing organ always present. The first segment cf the posterior division of the body (abdomen) never bears appendages. The abdomen of the male is 5 segmented (fig. 3a), and the female seldom carries the eggs in sacks hanging from the genital orifice. The anterior antennae in the male may be symmetrical, or one may form a grasping organ; in the female the antennae are symmetrical. 118 University of California Publications. [| ZooLoGy KEY TO THE GENERA OF THE GYMNOPLEA. (The genera marked with an asterisk have been found in the San Diego region.) 1. Inner ramus of third and fourth feet 3-jointed, figs. 12a, 19b..... 2 1. Inner ramus of third and fourth feet 2-jointed, fig. 44d.......... 19 2 inner ramus; OL Lrstsroot),.o-jOLULOd ot meme hte een iee eee eee 3 2.) Loner ramus ob first’ £006 -2-jOIntediaw. «yejac ners oe eae eee 32 2. inner ramus of first footie -joimbed) fia lod: eee lel ee ee 38 3. A black or brown knob on the first segment of the cephalothorax oO in the antero-lateral angle, on the right or left side; figs. 33a, ERLE eray dio niche a Sa ee yate Rea an ewe) er oe ee eA ts eee *Pleuromamma This! knob VAbSeMba ary. 5 ke cetietsesite eto eens ers erel o Siete cteke alertness t . First joint of inner ramus of second foot with proximally curved hooks ‘on thesinner margins fos SOD. oie eteres vesine «ieee *Metridia 2 Dhis' joint, Jike the rest, bearing a bristles: o/s... -\-11- se © brea 5 . Terminal joint of outer ramus of third and fourth feet with two spines or thorns on outer margin, and one terminal bristle; fig. ld 6 . Terminal joint as above, but with three spines on outer border; HPS). LTO LOD 2 coker ec tave sin morte Mayen Acie eee re averted eee 6a 6. Terminal bristle of outer ramus of third and fourth feet with nroad smooth borders ne: 1d oes cre aise ree eae eter *Calanus 6a. Terminal bristle bearing teeth or spines on outer border; figs. Oe DIP oxare ot a aha. 4, Bele Sick letgphe tc 1a ale seuel ore, s Lae atel s eitirstia tee ancl seee fea eae f 7. One bristle of left ramus of furca much longer and thicker than Ghee OLher wsitorale + erctero cls ane ereryeloe eects 8 feebuncals bristles: SymiMetrCala. ciccs sen 2 crete: wie eee Crete ste mele 9 8. Mandibular blade with three or four teeth, the ventral one hooked and separated from the others by a wide space; fig. ERS siesta seotels age cert oe Rieck eve euebcuhe. Sahu SECA ete, Sehiace *Heterorhabdus 8. Mandibular blade with at least 8 teeth..................... Disseta 9! AmtervoraAncenn ge SymMMenLICAl . hs spe sis o/es mm lois,ciets’s = fester stetaaiets 10 9.,Anterior antennae (as ymimenm cally i. chy- see cision wines an sae es 14 LO! Rami of ftth feet 3-jommteds mips S263 seo e ss wisis ees eiclege coeteereis 11 1O:) Rami of fitthy feet. a —jommbe Merwe ss ate eel vol ofc paras eo oot *Augaptilus? 10. Outer ramus 3-jointed, inner ramus 2-jointed............ Isochaeta? 10. Outer ramus 3-jointed, inner ramus 1l-jointed............... Isias? 10. Outer ramus 3-jointed, inner ramus lacking............. *Phyllopus? 10. Outer ramus 1-jointed, inner ramus rudimentary; fig. 42b *Arietellus? 11. Middle joint of outer ramus of fifth foot with a thorn-like pro- cess which is fused with the joint; fig 32c.......... *Centropages? 11. This joint with an awl-shaped or rudimentary bristle on the inner vhorder: figs sOU,eoRey ar ee cet peel scce ter acter thee leet tte 12 12. Terminal joint of inner ramus of fifth foot with five bristles; ik) Sasol Morea Sho arcane xi Gee oc Soo ae 0 *Lucicutia? 12. This joint, withvat, least) sixes bristles: orto ore artnet eterno eer 13 13: Abdomen with 4 seomentsi 7. )icte trey ieqie +o lislereiss)aiste)<' < 140. (c) Genital segment of female, lateral, X83. (d) Inner ramus of second foot of female 185. B.2, second basal of foot. Ri.1, first joint of inner ramus. Si., inner mar- ginal bristle. Allied to C. gracilis, but in the female the ventral surface of the genital segment is much more convex (ef. figs. 4a and 5c), and the first basal of the anterior maxilliped has a bulging protrusion on the outer border (ef. figs. 4b and 5d). 4 Bristles on anterior maxilliped longer than in C. gracilis, inner ramus of left fifth foot stylet-like, jointed and without bristles, outer ramus much elongated. Coloration: As in C. gracilis. Length: Female, 3.17 mm. Occurrence: San Diego, July 14, 1903, one female; December 21, 1904, December 29, 1904, one female each day. 130 University of California Publications. | ZOOLOGY Sub-fam. EUCALANINAE. Eucalanina Giesbrecht, 1892, p. 45. 2 Body elongate, head for the most part much lengthened (figs. 6a, b) and seldom distinct from the first thoracic segment. Rostral filaments slender, abdomen usually with three segments, seldom with four; furea often fused with the anal segment. First and second and eighth and ninth joints of anterior antennae fused. The swimming feet, and especially the rami, are short in comparison with the length of the body; inner ramus of first pair 1- or 2- jointed, 3-jointed in the following pairs. Terminal bristle of outer rami with smooth edge, that of the first pair as in the succeeding three pairs; fifth pair absent or uniramous; if present, with from three to five joints. 4 Body, especially the head, shortened (fig. 6c); anterior antennae without reduction in number of joints; furea as in the female. The mouth parts may be stunted. Fifth pair of feet not well developed, left foot uni- or biramous, right uniramous or lacking. 1. Genus Eucalanus Dana. Calanus Dana, 1848, p. 11; 1849, p. 278. Eucalanus Dana, 1852, p. 1047. Eucalanus (in part) Lubbock, 1856, p. 13; 1860, p. 160. Calanella Claus, 1863, p. 174; not Eucalanus Claus, 1881, p. 325. Eucalanus Brady, 1883, p. 37. Eucalanus Giesbrecht, 1888, p. 333; 1892, pp. 46, 131, 739; 1895, p. 246; 1898, p. 19. Eucalanus Wheeler, 1899, p. 166. Anal segment and furea fused, latter asymmetrical; head triangular, often elongate, fused with thorax; abdomen short, that of female with three or four segments, of the male with 5. Anterior antennae longer than body, 23-jointed in female, term- inal bristles plumose and colored. Outer ramus of mandible 7- or 8- jointed and shorter than inner. Mandible of female longer than maxilla; second basal of mandible makes with the outer ramus a eylindrieal body on which the inner ramus articulates proximally to the outer ramus (fig. 7¢). Inner ramus of posterior maxilliped with long bristles. Swimming feet short; outer rami 3-jointed, inner ramus of first pair 2-jointed, of second to fourth pairs 3-jointed. Fifth pair absent in female; in male (fig. 6d) Vou.2] Ksterly—Copepoda of the San Diego Region. Bul both are uniramous. The left 4-jointed, the right 1- to 4- jomted or lacking. Head appendages of male retrograded and modified, body shortened. KEY TO SPECIES. AN GON Will 8) OR Zh TSGUNTMIS. coco cobs asadococens0e000g500e g AN KOON Yanda (3) CeAMIEMISs bo a ooe conc cor oud ecooUnBU OD REDO ODOC 4 91. Two segments between anal and genital segments (fig. 6a)...... Jo sb ow eco dub noamomtinn Cmome bod Gd 6 ning. Go-cto molec E. elongatus 1. One segment between genital and anal segments (fig. 7b)....... 2 2. Inner border of second basal of mandible divided into two approximately equal portions by the insertion of the inner rep (Cie WO) soasasaccdescosouogeocecHodqon yon E. attenuatus Ze eroximalsporiion: much) lono er thangs taller pottetspeneltalcieleNelensl-i-r1 3 3. Two terminal bristles of left side of furca longer but hardly thicker than on the right side; genital segment (fig. 8d) much broader hans lone oniton-shia Peds eis itt ello sell lelelct en E. crassus 3. Two terminal bristles of left side of furca longer and much thicker than on the right side. Second basal of maxilla with four inner marginal bristles; forehead (fig. 9a) triangular, TROMUACNG! I IMO, Gosoopoaueade mss ugecgooSonoDoUDbE E. subtenuis $1. Both feet of fifth pair present ..........-.- 2. sees eee e ee eee eee 2 Ie Rireht foot of dikth pair aWSeMba ca. 3. Genus Mecynocera I. C. Thompson. Leptocalanus Giesbrecht, 1888, p. 334. Mecynocera Thompson, I. C., 1888a, p. 150. Mecynocera, Giesbrecht, 1892, p. 160; 1898, p. 23. Mecynocera Wheeler, 1899, p. 167. Furea symmetrical, articulating with anal segment ; mandible shorter than maxilla and less than half as long as the fourth pair of feet, similar in structure to that of Calanus, but inner ramus is nearly as long as second basal and twice as long as outer ramus; inner ramus of posterior maxillipeds at least as long as first or second basal. First pair of feet with outer ramus of three joints, inner of one joint; fifth pair present, with five joints on either side. & Unknown. 2 Head distinct from thorax; rostral threads delicate ; abdo- men short, with three segments; genital segment and furea sym- metrical. Anterior antennae of unequal length, more than twice as long as the body, with 23 joints, bristles few and very long. Inner ramus of posterior antennae nearly twice as long as outer ‘amus. The succeeding appendages, similar to those of Calanus: feet short, outer rami with three joints, inner ramus of first pair with one joint, of second to fourth with three; fifth pair with basals, outer ramus with three joints, inner ramus lacking. 1. Mecynocera clausi I. C. Thompson. Mecynocera clausii Thompson, I. C., 1888a, p. 150, pl. 11, figs. 1-4. Leptocalanus filiformis Giesbrecht, 1888, p. 334 Mecynocera clausii Giesbrecht, 1892, p. 160, pl. 5, fig. 1; pl. 11, figs. 43, 45; pl. 35, figs. 21, 22; M. clausi, 1898, p. 23. Mecynocera clausii Wheeler, 1899, p. 167, fig. 5. The only species of the genus. Coloration: Exceedingly transparent, without pigment in my specimens. 138 University of California Publications. [ ZooLocy Length: Female, 0.9-1 mm. Occurrence: The only specimens I have were collected Deeem- ber 30, 1903, on the ‘‘Banks’’ off Point Loma. NS <— a= f i jaa [) io \ ] / 1 a | || y, J \ A (ee \ (ease Pp | Srema Fig. 11.—Mecynocera clausi. Female, dorsal, x 45. Sub-fam. PARACALANINAE. Paracalanina Giesbrecht, 1892, p. 48. 2 Cephalothorax with four segments, abdomen with from two to four; rostrum ends in two soft filaments. Anterior antennae 25-jointed, with long terminal joints, but the division between the first and second and eighth and ninth joints may not be clear. Outer ramus of posterior antennae at most as long as the mner ramus; the mouth parts like those of Calanus. Terminal bristle of outer rami of feet with smooth border; basals and rami set with spines; fifth foot rudimentary (2- 4- jointed) or lacking. Vou.2] Esterly—Copepoda of the San Diego Region. 139 4 Characters as in the male of Calanus; the number of joints in the anterior antennae more reduced, the end-joint always shortened, and sometimes fused with the preceding one; fifth pair of feet weakly developed, the left 5-jointed, the right 4- or 2- jointed, or lacking. 1. Genus Paracalanus Boeck. Calanus Claus, 1868, p. 172. Paracalanus Boeck, 1864, p. 8. Paracalanus Claus, 1881, p. 326. Paracalanus Bourne, 1889, p. 145. Paracalanus Giesbrecht, 1892, pp. 48, 164, 757; 1898, p. 23. Paracalanus Dahl, 1893, p. 21. Paracalanus Wheeler, 1899, p. 168. Second basal of first pair of feet with an inner marginal bristle; proximal division of outer border of third joint of outer ramus of fourth pair (fig. 12a) over twice as long as the distal; outer border of the second joint not dentate; proximal division of the outer border of third joint of outer ramus in the third and fourth feet dentate; sealpelliform terminal bristle of the outer ramus in the third pair longer than the end joint; second joint of inner ramus of first pair with 5, third of same in second pair with 7 bristles. The abdomen of the female (fig. 120) with 4 seg- ments; the last joint of anterior antennae less than 114 times as long as the next to the last. Fifth foot of female short, 2-jointed (fig. 12c) ; right foot of male with 2 joints, left with 5 (fig. 12c). 2 Head fused with first thoracic segment, and fourth thoracic segment with fifth. Rostrum produced into two thin filaments. Genital segment and furea symmetrical, latter without bristle on outer margin. Anterior antennae with 25 joints. Outer ramus of posterior antennae shorter than inner; mandible with broad blade, the sack-like appendage on the first joimt of the inner ramus small. Maxilla with obscure segmentation of inner ramus, without bristle on the second lobe of outer border, and with but one on the first inner marginal lobe. Anterior maxilliped with outer marginal bristle. Inner ramus of the first swimming foot with 2 joints, of the second to fourth foot with 3 joints. 4 Abdomen with 5 segments. Number of joints of anterior antennae reduced through fusion of joints 1 to 6 and 7 to 8, end joint shortened but free. Aesthetasks enlarged and numerous. 140 University of California Publications. | ZOOLOGY Mandibular blade, appendages on inner border of maxilla and anterior maxilliped stunted, those of posterior maxilliped less so, its outer marginal bristles long and richly plumose. The swim- ming feet show slight peculiarities. 1. Paracalanus parvus Claus. Calanus parvus Claus, 1863, p. 173, pl. 26, figs. 10-14; pl. 27, figs. 1-4. Paracalanus parvus Claus, 1881, p. 327, pl. 3, figs. 1-16. Paracalanus parvus Bourne, 1889, p. 145, pl. 11, figs. 1-3. Paracalanus parvus Giesbrecht, 1892, pp. 164, 170; pl. 1, fig. 5; pl. 6, figs. 28-30; pl. 9, figs. 5-11, 25, 27, 31, 32; 1898, p. 2a. ps. Fig. 12.—Paracalanus parvus. (a) Fourth foot of female 195. fKe.3, third joint of outer ramus. (b) Female, dorsal, X83. (c) Fifth foot of female 410. (d) Second joint of inner ramus of second foot of female X195. (e) Fifth foot of male. Ps., left foot. Pd., right foot. 2 Inner bristle of furea barely longer than the furea. Ante- rior antennae reach, when brought to the sides of the body, per- haps to the posterior border of the third abdominal segment. , 4 , : : ‘ Vou.2] Esterly—Copepoda of the San Diego Region. 141 First joint of inner ramus of maxilla with two bristles on ante- rior face. Third lobe of second basal of posterior maxilliped with two bristles. Inner margin of first basal of the fourth pair of feet ends in one or two points (fig. 12a) ; anterior and posterior faces of first basal of second to fourth pairs set with hairs and spines; surfaces of first and second joints of outer ramus of the third pair and of second joint of the fourth, naked. Fifth foot rudimentary, symmetrical. 4 Fifth foot asymmetrical (fig. 12e); compare also generie¢ description. Coloration: Rather transparent, with red pigment in varying amounts and distribution, never very abundant. Length: Both sexes within 0.8-1.2 mm. Occurrence: Fairly common in hauls with smaller nets, both sexes being present summer and winter. Sub-fam. CLAUSOCALANINAE. Clausocalanina Giesbrecht, 1892, p. 49. @ Head usually fused with the first thoracic segment, fourth thoracic always fused with the fifth; rostrum ends in two short, soft filaments or is lacking; abdomen with four segments, furea symmetrical. Eighth and ninth joints of anterior antennae fused ; terminal joint short, seldom fused with the preceding one. Outer ramus of posterior antennae 6-jointed and always longer than the inner ramus. The other appendages of the head for the most part as in Calanus. Inner ramus of the first pair of feet 1- jointed, of the second pair 2-jointed, of the third and fourth 3- jointed; terminal bristle of the outer rami with dentate border ; third joint of outer ramus in second to fourth pairs with three bristles on outer border. Fifth pair rudimentary on each side, 3-jointed or lacking. & Unknown in Spinocalanus and Ctenocalanus. Abdomen with shortened anal segment; anterior antennae and head appendages in some eases like those of the female, in others as in the Paracalaninac. Fifth pair of feet: the right, 1- to 5-, the left 5- jointed. 142 University of California Publications. [ ZOOLOGY 1. Genus Clausocalanus Giesbrecht. Calanus Dana, 1849, p. 278; 1852, p. 1047. Calanus Claus, 18638, p. 172. Eucalanus Claus, 1881, p. 325. Drepanopus (in part) Brady, 1883, p. 76. Clausocalanus Giesbrecht, 1888, p. 334; 1892, pp. 50, 185, 733; 1898, p. 27. Rostrum with two points; second basal of second and third swimming feet with toothed distal margin and broad outer ramus. Mouth parts and number of segments of anterior antennae reduced. 2 Head fused with thorax and the fourth with the fifth thoracic segment. Abdomen with four segments, genital segment and furca symmetrical. Anterior antennae extend beyond the thorax, 23-jointed. Outer ramus of posterior antennae 114 times as long as the inner, the former 6-jointed, with short bristles on the proximal joints. First joint of inner ramus of mandible with a very small, sack-like appendage. Maxilla and maxillipeds as in Calanus, outer marginal bristle lacking on anterior maxilli- peds. Outer rami of swimming feet with 3 joints; inner ramus of first foot 1-jointed, of second 2-jointed, of third and fourth 3-jointed. End joint of outer ramus with finely dentate terminal bristle, and four bristles on inner border in second to fourth pairs; fifth pair uniramous, 3-jointed. 4 Head fused with first thoracic segment, and elongated at expense of free thorax rings. Rostrum suppressed ; abdomen with five segments, anal very short. Anterior antennae with joints 8-10, 13-16, 20-21, 24-25 fused. Outer ramus of posterior antennae twice as long as inner. Blade of mandible, appendage of inner border of maxilla, and anterior maxilliped suppressed ; less so the posterior maxilliped, the outer marginal bristle of which is not enlarged. Swimming feet elongated. Left fifth foot (fig. 13c) long, uniramous and with 5 joints, right short, 1- to 3-jointed. 1. Clausocalanus arcuicornis Dana. Calanus arcuicornis Dana, 1849, p. 278; 1852, p. 1056; 1855, pl. 72, fig. 9a-b. Calanus mastigophorus Claus, 1863, p. 173, pl. 27, figs. 5-8. Vou.2] Esterly.—Copepoda of the San Diego Region. 143 Clausocalanus arcuicornis Giesbrecht, 1888, p. 334; 1892, pp. 186, 1985 pl ios 1a pig; fig 7; pl.) 10; figs: 3-8, 14, 16, 17, 19; pl. 36, figs. 29-31, 34; 1898, p. 27. Clausocalanus arcuicornis Wheeler, 1899, p. 171, fig. 9. b Ee : 2) Fig. 13.—Clausocalanus areuicornis. (a) Male, lateral, X45. (b) Second basal of second foot to show toothed distal margin, 410. (c) Fifth foot of male X83. 2 Genital segment longer than the two following. Furea about as long as broad. No aesthetask on fourth, sixth, eighth, eighteenth or twenty-second joints of the anterior antennae. & Second segment of abdomen at least as long as the third and fourth together (fig. 13a) ; right foot of fifth pair with three joints (fig. 138c). Coloration: Not very transparent, with red pigment in vari- ous places on the posterior part of the body and on genital seg- ment. Occurrence: San Diego, June 25, 1904, one male. Sub-fam. AETIDIINAE. Aetidiina Giesbrecht, 1892, p. 52. Aetidiinae Wolfenden, 1903, p. 263. 2 Head sometimes distinct from first thoracic segment ; other- wise the cephalothorax always has four segments, as has the 144 University of California Publications. [ ZOOLOGY abdomen invariably. Rostrum strongly chitinized, usually with one point, seldom with two or lacking. Genital segment and furea usually symmetrical. In the anterior antennae the eighth and ninth and twenty-fourth and twenty-fifth joints are fused. Outer ramus of posterior antennae at least fully as long as the inner, and usually longer; the second and third joints of the. outer ramus are distinct. Mandible as in Calanus, with strong blade, and occasionally shortened inner ramus. Maxilla with well developed lobes on inner margin and usually with hooked bristles even on the second basal and the inner ramus; outer ramus relatively small. Bristles of anterior maxilliped short but strong, those of the inner ramus relatively slender and sparsely plumose; the articulation of the inner ramus is rather on the _ posterior surface of the second basal than at the end. Inner ramus of posterior maxillipeds at most 24 as long as the second basal. Inner ramus of first swimming feet always 1-jointed; that of the second almost always 1-jointed, while in the second and third the inner ramus is 3-jointed; the form of the swimming feet as in the Clausocalaninae; inner marginal bristle of first basal long and plumose. 4 Known in Aetideus, Euchirella and Undeuchaeta. Charac- ters like those of Clausocalanus; oceasionally the twentieth and twenty-first joints of one of the anterior antennae are fused. Left foot of fifth pair 5-jointed (if the right is lacking, or stylet- like, in which ease the right is claw-like). 1. Genus Aetideus Brady. Aetidius Brady, 1883, p. 75. Aetidius Thompson, 1888b, p. 142. Aetidius Giesbrecht, 1892, pp. 53, 213. Aetideus Wolfenden, 1903, p. 266; 1904, p. 116. Aetideus Giesbrecht, 1898, p. 31. 2 Cephalothorax and abdomen with four segments, symmet- rical; rostrum large, prolonged into two thick chitinous prongs; last thoracic segment produced into a spine on each side. Ante- rior antennae 23-jointed, reaching about to the end of body. Rami of posterior antennae about equal in length, outer ramus 7- jointed. Outer rami of all swimming feet 3-jointed, inner ramus of first and second pairs 1-jointed, of third and fourth 3-jointed. Fifth pair of feet absent. ———E————— Oe Vou.2] Esterly—Copepoda of the San Diego Region. 145 4 Anal segment very short, abdomen with five segments. Anterior antennae 20-jointed, joints 8-10, 12 and 13, 20 and 21, 24 and 25 fused. Blade of mandible, appendages of inner border of maxilla and anterior maxilliped stunted. Left fifth foot uni- ramous, 5-jointed; right lacking; swimming feet as in female. 1. Aetideus armatus Brady. Aetidius armatus Brady, 1883, p. 75, pl. 10, figs. 5-16. Aetidius armatus Giesbrecht, 1892, p. 213, pl. 2, fig. 6; pl. 14, figs. 1-13; pl. 36, figs. 6-9. Aetideus armatus Giesbrecht, 1898, p. 31. Aetideus armatus Wolfenden, 1903, p. 266. Fig. 14.—Aetideus armatus. Female. (a) Animal from side X20. (6) Second foot X83. St., terminal bristle of outer ramus. With the characters of the genus. Coloration: Rather transparent; there was no pigment in my specimens, but Giesbrecht says that red may occur in the body. Leneth: Female, 3 mm. Occurrence: San Diego, June 9, 1904, one female; June 14, two females. 2. Genus Gaidius Giesbrecht. Gaidius Giesbrecht, 1895, p. 249; 1898, p. 32. Gaidius Wolfenden, 1902, p. 365; 1903, p. 266; 1904, p. 114, pl. iO), eS 1G tsk Rostrum short, one point (fig. 15b), sides of last thoracic seg- ments produced into a sharp spine (fig. 15a). Inner ramus of posterior antennae three-fourths as long as outer. Outer ramus of first foot 2-jointed, of the second to fourth 3-jointed, inner ramus of first and second feet 1-jointed, of the third and fourth 3-jointed. | 146 University of California Publications. | ZOOLOGY May be distinguished from Aetideus by the form of the ros- trum, relatively shorter inner. ramus of the posterior antennae, and by the fusion of the proximal joints of the outer ramus of the first foot (fig. 15d). 1. Gaidius pungens Giesbrecht. Gaidius pungens Giesbrecht, 1895, p. 249, pl. 1, figs. 1-4; 1898, p- 32. Gaidius pungens Wolfenden, 1903, p. 266. ‘ Fig. 15.—Gaidius pungens. Female. (a) Dorsal, X18. (b) Head, lateral, x45. (ce) Tube-like processes on inner distal portion of second basal of fourth foot 195. (d) First foot «195. Ri., inner ramus. 2 Anterior antennae reaching at least to posterior border of thorax. The processes on the inner border of the first basals of the fourth feet are heavier and stiffer than in the preceding pairs, being almost tube-hke (fig. 15c). & Unknown. Coloration: Transparent, with little or no pigment. Length: Female, 3 to 3.5 mm. Vou.2] Esterly—Copepoda of the San Diego Region. 147 Occurrence: San Diego, May 31, 1904, eight females; two males which seem to be of this species were taken also’ at this time, but they are distinctly immature. 3. Genus Undeuchaeta Giesbrecht. Euchaeta (in part) Brady, 1883, p. 57. Undeuchaeta Giesbrecht, 1888, p. 335; 1892, pp. 54, 227, 766; 1898, p. 33. Undeuchaeta Sars, 1900, p. 58, pls. 15, 16. Undeuchaeta Wolfenden, 1903, p. 267. 2 Abdomen with four segments, the first with the genital open- ing on the convex ventral surface, at least as long as the second and longer than the last segment. Lateral angles of last thoracic segment rounded, or at least not produced into spines. Anterior antennae 23-jointed, outer ramus of first foot 2-joimted, inner ramus l-jointed. Outer ramus of posterior antennae at least 114 times as long as the inner; outer ramus of maxilla (fig. 16e) small, middle bristles shorter than the distal and proximal ones, outer marginal lobe with much elongated middle bristles. 4 Anterior antennae 21-jointed, cephalo-thorax with four segments, abdomen with five, anal segment very short. Head with rather high crest (fig. 16d), last thoracic segment prolonged into angles, but not pointed. Inner ramus of posterior antennae 34 as long as the outer. Mandible, maxilla and maxillipeds much reduced. Outer ramus of first foot indistinctly 3-jointed. Left foot of fifth pair uniramous (inner ramus reduced to a very small, rod-like projection), outer ramus (fig. 16f) end- ing in a short style (terminal joint of ramus). Right foot bira- mous. Terminal joint of outer ramus produced into a long stylet, inner ramus as in Euchaeta (cf. fig. 23a) ; outer ramus of each foot 3-jointed. The second joint of the outer ramus of the left foot (fig. 16f. Re. 2) bears a toothed process (fused with the joint) which flares distally ; at the base of this and on the second joint is articulated a process, which together with the terminal joint of the ramus and the toothed process forms a forceps. The abdominal segments are densely covered with fine spines or hairs, and the posterior margins of the segments are toothed. In the structure of the fifth pair of feet these male animals very closely resemble the males of the genus Huchaeta, but seem 148 University of California Publications. [ ZooLoGy to be distinct from the latter in bearing an articulating process on the second joint of the outer ramus of the left foot. There is a muscle attached to the process which serves to move it. The relative lengths of the rami of the posterior antennae dis- tinguish the animals from Fuchirella, as does the division (though indistinct) of the outer ramus of the first foot into three joints. In Euchaeta, the outer ramus of the first foot is distinctly 3-jointed in the male, and the rami of the posterior antennae are about equal in length. In several female speci- mens also the outer rami of the first feet are indistinctly divided into three joints, and the sexes correspond in this respect. Sars (1900, p. 59-63) has deseribed the male and female of Undeuchaeta spectabilis. So far as I know, his is the first record of the male of the genus. In his specimens the anterior antennae of the female are 24-jointed, while in Giesbrecht’s the number of joints is 23. In his description of the male, Sars gives the num- ber of joints of the anterior antennae as 22, but in his drawing (pl. 16, fig. 2) there are but 21. The fifth pair of feet in the male of U. spectabilis is very different from that in the San Diego specimens, a striking point being that both the right and left feet are biramous. The description of the male of the genus given above is based upon the San Diego specimens. 1. Undeuchaeta major Giesbrecht. Euchaeta australis Brady, 1883, p. 65, pl. 21, figs. 5-11. Undeuchaeta major Giesbrecht, 1888, p. 336; 1892, pp. 227, 232, pl. 37, figs. 56, 57, 59; 1898, p. 34. 2 Head with median crest, genital segment with protrusion on right side and a hooked pointed appendage at the right of the genital opening (fig. 16a). & (new) Compare generic description. Coloration: Female not especially transparent, with red pig- ment on basals of posterior maxillipeds, and in mouth region. Male: plumose bristles of furea steel-blue; those of posterior antennae and mandible red; on the feet the bristles are faintly red on the outer ends. ‘— }. 2 ee a ee ees Vot.2] EKsterly—Copepoda of the San Diego Region. 149 Length: Female, 4.5-5.5 mm.; male, 6-6.5 mm. Occurrence: Four females were taken from May 18 to June 23, 1904; five on December 23, 1904, on the ‘‘Banks.’’ One male was taken in July, 1904; another on November 1, 1904, off Point Loma. reg on Re eres d e c. Fig. 16.—Undeuchaeta major. (a) Female, lateral, X20. Ab.1, first abdominal segment. (b) Male, lateral, X9. (c) Head of male, dorsal, X83. (d) Head of male, lateral, X83. (e) Outer ramus of maxilla of female 195. (f) Distal portion of left fifth foot of male. Re.2, Re.3, respective joints of outer ramus. 5 2. Undeuchaeta minor Giesbrecht. Undeuchaeta minor Giesbrecht, 1888, p. 335; 1892, pp. 228, 232, pl. 14, figs. 31-34; pl. 37, figs. 55, 58; 1898, p. 34. OTe. S \\\ 2 . . < a Fig. 17.—Undeuchaeta minor. Female. (a) Lateral, x20. (b) Second foot X83. 150 University of California Publications. | ZooLogy 2 Head without crest (fig. 17a), genital segment with a spine on the dorsal surface. Coloration: Similar to that of U. major. The digestive tract of the single specimen was filled with orange red material. Length: Female, 3.18 mm. Occurrence: San Diego, June 14, 1904. 4. Genus Euchirella Giesbrecht. Undina (in part) Lubbock, 1856, p. 21. Calanus (in part) Lubbock, 1856, p. 15. Undina Claus 1863, p. 186. Euchaeta (in part) Brady, 1883, p. 59. Euchirella Giesbrecht, 1888, p. 336; 1892, pp. 54, 233, 743; 1898, p- 34. Euchirella Cleve, 1900, p. 4. Euchirella Wolfenden, 1903, p. 267. Rostrum present in most species, simple; lateral angles of last thoracic segment not pointed. Inner ramus of posterior antennae 14 to 14 as long as the outer ramus, the two proximal ‘ joints of which are fused. Inner and outer rami of the maxilla short, the former provided with heavy hooked bristles. Outer ramus of the first pair of feet 2-jointed, that of the second to fourth pairs 3-jointed. Inner ramus of first and second pairs 1-jointed, of the third and fourth 3-jointed. Right foot of fifth pair of male with shear-like formation of distal portion, the left foot stylet-like (ef. figs. 18d, 19d). 2 Head not always distinct from thorax, last two thoracic segments fused., Abdomen with four segments, genital segment and fureal bristles symmetrical or asymmetrical. Anterior an- tennae with 23 joints, reaching to the end of the thorax or some- what beyond. Second basal of posterior maxillipeds twice as jong as the 5-jointed inner ramus. Feathering on the inner margin of the proximal basal joint of the fourth pair of feet replaced by spines. Fifth pair of feet absent. & Head oceasionally with a median crest; abdomen with five segments, anal segment very short. Joints 20 and 21 of right anterior antennae fused, inner ramus of posterior antenna rela- tively longer than in the female. Blade of mandible, appendages on inner border of maxilla, and anterior maxillipeds reduced; Vou.2] Esterly—Copepoda of the San Diego Region. 151 posterior maxilliped slender. Spines on second basal of fourth foot unusual. Right foot of fifth pair biramous, with forceps; left stylet-like, with rudimentary inner ramus. KEY TO SPECIES. Brisa] ihe SE CUO) Pget2 C1S( 8) eet tase RE Red, Cates ae ee ye ee ea io) BE UEt Mab OGRE EESCIIE. 55 'crctae A rales cha dirceaeta cme rermeiies am Na cast wares aus 4 Q1. Head without crest, and rostrum one-pointed (fig. 19a) ........ 2 [Head wath erest, rostrum present (figs 200)06 $4.06 vs. 0secks ooo a 2. Outer ramus of posterior antenna about twice as long as inner.. POCAIDToOOoGoOCGoDonObDoOOOooCoO DTU o OOO OOo DUM COO OC Haga nS E. rostrata 2. Outer ramus nearly four times as long as inner (fig. 20c)........ 3 3. Genital segment with long sac-like appendage on left side (fig. HSE) Spire ny so RSRee PS cc sroy he cts enc ec OAR GNS SoS] gears Mtaton fl ohare es crer ance E. messinensis a Heaiewat he low erest (fe 20M) sa. ca catia ers sree) saa eee a, E. pulchra a Eleagewith iioh Cresta (Ho. 226)... vtec eo oe8.s Cccass ss he ene E. galeata @ JAG aA NONy Cressy (Ge PASO) Geis Gwomue bon GomoG oes onoood Gone 2 Meade wabhOut seresben(He: ZUG) jf. 2 sr. sis: oj eieiord-csceve: srnlelenet sue tere jacsichals 8 ats 3 2. Forceps-like terminal portion of right fifth foot longer than the JORISEUL joronron@in, (Galery ISA) s Gagaccncoucace Ruste cet ekareeMenene E. messinensis 2. Terminal portion (forceps) of right fifth foot shorter than the ORAL TNOMaON, Ciey HUM), aeiocs Secoonadaudoo bongo otas E. pulchra 3. Fifth foot short, the right about four times as long as the second basalts: broad Chios 20) oo ysncla erstcetsl arses cistla dita ehs vec. ee & E. amoena 3. Right fifth foot six times as long as the second basal joint. .E. rostrata 1. Euchirella messinensis Claus. Undina messinensis Claus, 1863, p. 187, pl. 31, figs. 8-18. Euchirella messinensis Giesbrecht, 1892, pp. 232, 244; pl. 15, figs. 12, 16, 21, 24; pl. 36, figs. 14, 15, 18, 24, 25; 1898, p. 35. 2 Forehead with rostrum, without crest (fig. 18a); genital segment asymmetrical, with sac-like appendage on left side of dorsal surface (fig. 18) ; third terminal bristle on right side of furea elongated. Inner ramus of posterior antenna 14 as long as outer, second joint of former with 5-4 bristles. First basal of fourth pair of feet with one or two spines on inner border, the longer of which reaches beyond the end of the joint. 4 Forehead with a low and rather long crest (fig. 18c) ; fifth foot slender, the right foot (fig. 18d) over seven times as long as the second basal is broad, the forceps longer than the basals. Coloration: Not very transparent; red pigment in body and on bristles of posterior antennae, and basals of swimming feet. Leneth: Female, 4.5 mm.; male, 4 mm. Le University of California Publications. [ ZOOLOGY Occurrence: July 9, 1903, one male; July 22, 1903, one female. Fig. 18.—Euchirella messinensis. (a) Head of female, lateral, X15. (b) Abdomen of female X15. (c) Head of male X30. (d) Fifth foot of male X20. 2, Euchirella rostrata Claus. Undina rostrata Claus, 1866, p. 11, pl. 1, fig. 2. Euchaeta hessei Brady, 1883, p. 63, pl. 20, figs. 1-13; pl. 23, figs. 11-14. Euchirella rostrata Giesbrecht, 1892, pp. 233, 245, pl. 15, figs. 3, 13, 25; pl. 36, figs. 19, 20; 1898, p. 36. Euchirella rostrata Cleve, 1900, p. 4, pl. 2, figs. 1-12. 2 Front without crest, with rostrum, abdomen symmetrical. Inner ramus of posterior antennae 14 as long as outer ramus; second joint of inner ramus with 8-6 bristles. First basal of fourth pair of feet (fig. 195; B. 1) with 6 or 7 triangular lamellae on the inner border. Bristle on outer margin of second joint of outer ramus of the second pair reaches at least to the point of the first bristle on the outer border of the third joint of the ramus. 4 Head without crest, with rostrum. Fifth foot six times as long as its second basal joint. Margin of second joint of outer ramus not denticulate, third joint smooth. Inner ramus of pos- terior antenna 14 as long as outer. First basal of fourth foot without triangular lamellae. (Cleve 1900). Vou.2] Hsterly—Copepoda of the San Diego Region. F533 The male was described by Cleve, 1900, and is identical with Euchaeta hesser Brady. Coloration: Red pigment as in EF. messinensis, but more abun- dant, especially on swimming feet. = Bil Fig. 19.—Euchirella rostrata. (a) Female, lateral, X18. (b) Fourth foot, female, X83. B.1, first basal, showing lamellar pro- cesses. Length: Female, 2.97-3.1 mm. Occurrence: San Diego, July 14, 16, 21, 1903, females; May 24, 1904, two females; June 2, 1904, one female. 3. Euchirella pulchra Lubbock. Undina pulchra § Lubbock, 1856, p. 26, pl. 4, figs. 5-8; pl. 7, fig. 6. Calanus latus 2 Lubbock, 1856, p. 15, pl. 2, fig. 12; pl. 11, figs. 8-11. 154 University of California Publications. [ ZOOLOGY Euchaeta pulchra Brady, 1883, p. 638, pl. 14, fig. 7; pl. 20, figs. ilbyealye ale Euchirella pulchra Giesbrecht, 1892, pp. 233, 244, pl. 15, figs. 22, 23, pl. 36; figs. 13, 27; 1898, p. 36. d Fig. 20.—Euchirella pulchra, (a) Female, lateral, X9. (b) Fifth foot of male X45. (c) Rami of posterior antennae, to show relative lengths; bristles omitted, 83. i., inner ramus. (d) First basal of fourth foot of female 140. 2 Front with low crest (fig. 20a) and small rostrum. Genital segment asymmetrical; left side strongly convex in front of the middle of the segment, right side indented. Inner ramus of pos- terior antennae about 24 as long as outer ramus; second joint of inner ramus with 6-5 bristles. First basal of fourth pair of feet (fig. 20d) with one or two thorns about equal in length on the inner border, which do not reach the distal margin of the joint. Outer bristle of the second joint of the outer ramus of the second | pair at most as long as the first outer bristle of the third joint. Vot.2] Hsterly.—Copepoda of the San Diego Region. fre On OV & Considerably like EF. messinensis, the chief difference being in the structure of the fifth pair of feet (fig. 206). The claw of the right foot is shorter than the basal (in messinensis longer). Coloration: About as in H. messinensis. Length: Female, 3.4-4 mm.; male, 3.5 mm. Occurrence: San Diego, May 31, 1904, two immature males, one female; June 23, 1904, one female adult ; December 23, 1904, ‘*Banks,’’ eleven females, all adult; one male adult, December 30, 1904, on the ‘‘ Banks.’’ 4. Euchirella amoena Giesbrecht. Euchirella amoena Giesbrecht, 1888, p. 336; 1892, pp. 233, 244; pl. 15, fig. 20; 1898, p. 36. Fig. 21.—Euchirella amoena. Male. (a) Lateral X20. (b) Fifth foot x 45. Q Unknown. & Front without crest. Fifth pair of feet shortened, the right about four times as long as the second basal is broad. Length: Male, 3.02 mm. Occurrence: San Diego, May 28, 1904, one male. 5. Euchirella galeata Giesbrecht. Euchirella galeata Giesbrecht, 1888, p. 336; 1892, p. 233, 244; pl. 15, fig. 18; pl. 36, figs. 22, 26; 1898, p. 36. 2 Head with high crest, and rostrum; genital segment asym- metrical, strongly protruding on the posterior portion of the dorsal surface. Inner ramus of posterior antennae about 25 as 156 University of California Publications. | ZOOLOGY long as the outer; basals of fourth foot about as in E. pulchra, the spines not reaching to the distal border of the joint. & Head as in the female. Coloration: Opaque, without pigment. Length: Female, 6.5 mm. Occurrence: San Diego, November 18, 1904, one adult female, two immature males. a Fig. 22.—Euchirella galeata. Female. (a) Lateral X9. (6b) First basal of fourth foot X83. Sub-fam. EUCHAETINAE. ~ Euchaetina Giesbrecht, 1892, p. 55. 2 Rostrum with one point; a pouch-like appendage in front of the upper labium. Inner marginal bristle of furca very long. Distal hooked bristles of anterior maxillipeds longer than the proximal. Outer ramus of first pair of feet 2-jointed, of the second to fourth 3-jointed; inner ramus of first and second pairs 1-jointed, of the third and fourth pairs 2-jointed. 3 Abdomen as in the Clausocalaninae. Outer ramus of first pair of feet 3-jointed; fifth foot on each side with 2-jointed basal, and biramous; inner ramus of left stylet-like, of right truncate; left outer ramus 3-jointed, right 2-jointed. 1. Genus Euchaeta Philippi. Euchaeta Philippi, 1843, p. 54, pl. 4, fig. 5. Euchirus Dana, 1846, p. 183. Euchaeta Dana, 1848, p. 20; 1849, p. 279; 1852, p. 1084. Euchaeta Claus, 1863, p. 163. Euchaeta Giesbrecht, 1892, pp. 55, 245, 740; 1895, p. 251; 1898, Dass 2 Cephalothorax with five segments, the last two thoracic segments fused; abdomen with four segments, genital segment more or less asymmetrical. Anterior antennae of varying rela- tive lengths, but of characteristic form, with 23 joints. Rami 4 (od) Vou.2] Hsterly—Copepoda of the San Diego Region. 1 of posterior antennae about equal in length, outer ramus with seven joints. Blade of mandible with few but strong teeth. Sec- ond basal joint of the posterior maxilliped at least three times as long as the inner ramus of five joints. Inner marginal bristle of first basal of the swimming feet long and richly plumose, terminal bristle of outer rami finely toothed; fifth pair absent. 4 Head fused with thorax; abdomen with five segments, ana! segment short; innermost bristle of furea shortened and bent at an angle. Blade of mandible, appendage of inner border of max- illa, and anterior maxilliped stunted ; less obvious differences also in the posterior antennae and maxillipeds and swimming feet ; outer ramus of first pair of feet 3-jointed. Feet of fifth pair long, strongly built, and of rather complicated structure (fig. 23a). KEY TO SPECIES. J Earlre(ol ne SOO RMEET OFS (CNN acto oer Cre MeC LG Ole o Abin ae Sie Gi OCIS AiO ean fe) JRA EGINE MHOYONE. OREN Ne ers-oetaln eae So ooo oid a ebitoe od colds ie OImeini eco 4 41. Terminal joint at each foot of fifth pair, with long straight or slightly curved stylet; elevation for frontal organ not pro- TIRUNGHB IED” 5% Gi oa ed cele haw BU CIO oo GOO OOmorn SLAG pion oro bane E. acuta 91. Hairs of frontal organ on a low elevation (fig. 25b)............. 2 1. Hairs of frontal organ on an elevation which extends toward the fer OMityp (UO LAD) cre tataye srevekete cick sie leks iene NsFo: eres oie 6] 4) let Cxaiyn siaeve ts o/s 4 2. Genital segment with asymmetrical outgrowths (figs. 25c, d); no bristle in middle of outer border of first joint of outer ramus of the first foot; terminal bristles of furca about equal in length, the dorsal (inner) bristle much longer and thicker Giles; BH) scpesgcanoodotonssaadgpqoornonebootegueseucnaune 3 3. Genital segment with a knob-like protuberance in front on the IGE BIC Gee cdcos at docls pbodsoD oud Gonos Con ocomopnednaT BE. acuta 3. Genital segment without such an outgrowth ............... E. media 4. Middle spine on outer border of terminal joint of second foot longer than the others, and the distal indentation in the border GReey asin (CUE MEU KO coe ec clan CIpIAeicO On Olceced Ok minInImicacieRcrO e Cria 5 SAnterionantennaesloneer tham boOdiye sc aces see E. spinosa Or 1. Euchaeta acuta Giesbrecht. Euchaeta acuta Giesbrecht, 1892, pp. 246, 262, pl. 16, figs. 6, 10, 14, 18, 21, 27, 39 ;pl. 37, figs. 47, 48, 52; 1898, p. 38. 2 Elevation on front of head flat; genital segment asymmet- rical, more strongly convex on the right side than on the left, and with more prominent process at the right of the opening; a knob-shaped outgrowth on anterior part on left side. Furea 158 University of California Publications. | ZooLoGy with four terminal bristles nearly equal in length, inner bristle of furea much thicker than end bristles. Anterior antennae reach a little beyond the posterior end of the thorax. Fig. 23.—Euchaeta acuta. (a) Fifth foot of male X37. Ri. dz., inner ramus of right foot. Re. 3, dx., third joint of outer ramus. Re. 1, 2, sn., first and second joints of outer ramus of left foot. Proc., process. Sph., spermatophore. (b) Second-and third joints of outer ramus of left fifth foot of male «140. Parts as in a. (c) Second foot of male X60. (d) Maxilla of male X60. B.1, first basal. B.2, second basal. Le.1, first lobe of outer margin. fi., inner ramus. FRe., outer ramus. First lobe of outer border of maxilla (cf. fig. 25f) with six bristles( one very small), second basal with three, fused second and third joints of inner ramus with four. Outer border of Vou.2] Esterly.—Copepoda of the San Diego Region. 159 first joint of outer ramus of first pair of feet concave; outer border of third joint of outer ramus of second pair and its outer bristles different than in the following pairs of feet; outer mar- ginal bristle of second joint of outer ramus reaches almost to end of the first outer marginal bristle of the third joint. Third joint of outer ramus of left fifth foot of male (fig. 23a) with a stil- etto-like process; second joint with a finely dentate, pyramidal and pointed process (proc.). Coloration: Rather opaque, a fleck of red pigment in the mouth; most of the pigment is found on the back and sides of the cephalothorax, and on the posterior maxillipeds. Length: Female, 4 mm.; male, 3.5-4 mm. Occurrence: July 31, 1903, one male; June 23, 1904, one male and one female. A good many (12-15) males were taken at one time on December 23, 1903, on the ‘‘ Banks.’’ 2. Euchaeta spinosa Giesbrecht. Euchaeta spinosa Giesbrecht, 1892, pp. 246, 263, pl. 16, figs. 12, 26, 34,47; pl. 37, figs. 31, 34, 35, 50; 1898, p. 39. 2 Elevation in front of head (fig. 24a) produced anteriorly ; genital segment almost symmetrical, with large, flap-like projec- tion at each side of the orifice (fig. 245). Second terminal bristle of the furea longer than the other terminal bristles, dorsal bristle much thicker than the terminal ones. Anterior antennae reach beyond the end of the furea by more than the end joints. First outer marginal lobe of maxilla with eight bristles, second basal with three, fused second and third joints of inner ramus with four. Outer border of first joint of outer ramus of first foot concave; outer border of third joint of outer ramus of second pair and its outer bristles different than in the following pairs; outer marginal bristle of second joint reaches to the end of the first marginal bristle of the third joint (fig. 24c). Basals and rami of posterior pairs of feet covered in places with short spines (fig. 24d). & Unknown. Coloration: Red in cephalothorax, sometimes on furea. plumose bristles of maxillipeds same color. Eggs blue. Leneth: Female, 6 mm. or over. 160 University of California Publications. [ ZooLocy Occurrence: July 21, 1903, one female; May 26, 1904, one female; July 5, 1904, four females; May 28, 1904, two females, one with egg cases, one without. i Senses * Fig. 24.—Huchaeta spinosa. Female. (a) Head, lateral, X15. (b) Abdo- men, lateral, X48. Gen. seg., genital segment. (c) Outer ramus of second foot X45. (d) Basals, inner ramus, proximal joints of outer ramus of fourth foot X45. 3. Euchaeta media Giesbrecht. Euchaeta media Giesbrecht, 1888, p. 337; 1892, pp. 246, 263, pl. 16, figs. 13, 36; pl. 37, figs. 39, 40; 1898, p. 39. 2 Elevation on front of head low (fig. 25d); genital segment asymmetrical with processes in the region of the orifice and a flap on the right side of the segment behind the orifice (figs. 25c, d) ; furea (fig. 25g) as in E. acuta. Anterior antennae extend a little beyond the posterior border of the genital segment. Vot.2] Esterly—Copepoda of the San Diego Region. 161 First outer marginal lobe of the maxilla (fig. 25f) with eight bristles, second basal with three, fused second and third joints of the inner ramus with four. Outer border of first joint of outer ramus of first foot concave; outer border of third joint of second pair and its outer bristles different than in the following pair. nor: S! let lth rsd Fig. 25.—Euchaeta media. Female. (a) Lateral, X18. (b) Head, lateral, X83. (c) Genital segment, dorsal, X45. (d) Genital seg- ment, from right side X45. (e) Second foot X83. (f) Max- illa X83. Le.1, first lobe of outer margin. Li.1, first lobe of inner margin. Ri.2, 3, fused second and third joints of inner ramus, bristles not shown. (g) Furea, dorsal, X45. Si., inner marginal bristle. 4 Unknown. Coloration: Rather transparent; there is no pigment in the preserved specimens I have seen. Length: Females average about 3.3 mm. Occurrence: Forty or fifty females, many with eggs, were taken December 23, 1903, on the ‘‘Banks’’ with males of EL. 162 University of California Publications. | ZooLoGy acuta. Three or four females were taken during June and July, 1904. The specimens which I have placed in this species correspond to Giesbrecht’s descriptions; but the females have hairs on the ventral sides of the abdominal segments, and Giesbrecht does not mention these nor figure them (1892, pl. 37, figs. 39, 40). The outer marginal lobe of the maxilla is always provided with eight bristles, but one of these is very inconspicuous and much shorter than the others. As these animals correspond very closely in other respects to Giesbrecht’s specimens, especially in the form of the genital segment, I have thought best to include them under his species, even though there are slight differences. Sub-fam. SCOLECITHRICIN AE. Scolecithricina Giesbrecht, 1892, p. 55. 2 Head commonly fused with first, and fourth with fifth thoracie segment; rostrum with two usually soft filaments; abdo- men with four segments, symmetrical. Eighth and ninth joints of anterior antennae always fused, and occasionally other joints. Outer ramus of posterior antennae 6-jointed. Blade of mandible with weak teeth; inner ramus of maxilla fused with second basal. The distal bristles of the anterior maxillipeds are modified into sac-like structures (fig. 306), which occasionally are pencillate at the end; lobes of appendages closely crowded together. Inner ramus of posterior maxillipeds at most only as long as the second basal. Inner rami of swimming-feet jointed as in the Clauso- calaninae and set with spines; fifth foot rudimentary or absent. & Abdomen with shortened anal segment, number of joints of anterior antennae reduced, the twentieth and twenty-first often fused only in one. Other head appendages like those of the female, or specifically modified. Left fifth foot 5-jointed, oceasionally with inner ramus, the right 4-jointed (rudiment of inner ramus sometimes present) or lacking. a Vou.2] Hsterly—Copepoda of the San Diego Region. 163 1. Genus Scolecithrix Brady. Undina (in part) Lubbock, 1856, p. 21. Scolecithrix Brady, 1883, p. 56. Scolecithriz Giesbrecht, p. 337; 1892, pp. 56, 265, 264; 1898, p. 42. Lophothriz Giesbrecht, 1895, p. 254. Amallophora (in part) Seott, T., 1893, p. 54. Neoscolecithrix Canu, 1896, p. 426. Scolecithrix Wolfenden, 1904, pp. 119, 120. Cephalothorax ellipsoidal, head fused with thorax, abdomen of female with four segments, of male with five; anal segment commonly short. Anterior antennae in female 19- to 24- jointed, in male 17- to 24- jointed, end joints (24 and 25) fused or dis- tinet, aesthetasks well developed, more numerous in male. Bitiny part of mandible and maxilla rather weak, inner ramus of max- illa mostly unsegmented and fused with the second basal. Distal bristles of anterior maxilliped thick, soft, in appearance some- thing like the aesthetasks of the antennae; these may be vermi- form, end in tufts (pencillate), or be pestle-shaped, and are usually present in both sexes. Head appendages of male like those of the female, but may in special cases be modified in par- ticular ways. Outer rami of first four feet 3-jointed, inner ramus of first foot 1-jointed, of second 2-jointed, of third and fourth 3-jointed; surfaces of both often set with spines and points. Fifth foot in female uniramous, 1- to 3-jomted, seldom absent; fifth foot of male uniramous on each side, or the left biramous and the right uniramous, or both biramous. KEY TO SPECIES. I JBIGEKCl TANOM CHES poeodcdoudeemoaenoadomacce 5 Ute OmbanO Ow oc 2 Il, JELeRGh WANN meh (Gi, Zs) Saakaoaccagcocbucb: usoboonoonuUdDe 4 2. Anterior antennae of female 19-jointed; right of the male 17-, acy Wein; TUSHSHOuUMHEh oc benossadcadodobs doa avo nGo cK UMaDUCma od 3 2. Anterior antennae of female 23-jointed............... S. subdentata 2. Number of joints unknown; for characters compare description anil 306%, BO) cos ona dab ogowoooonpde cas Coo ebe se Gboon nT cor S. pacifica 3. First joint of outer ramus of first foot with a thorn-like bristle Gm Omer mame (NE 26) os eccyo cle crere a ere wie oe ole «wis «els ate ee S. danae 3. This joint without the bristle ..........2.---.-..-2..-.0> S. bradyi 4. Anterior antennae of female 23-jointed; fifth foot (fig. 26e) ; right antenna of male 17-jointed, left 18-jointed; fifth foot (Cie PCED ae poise oo dot SenOie os Ole in Oreo a S. persecans 164 University of California Publications. [ ZooLoGy 1. Scolecithrix danae Lubbock. Undina danae Lubbock, 1856, p. 21, pl. 4, figs. 6-9. Scolecithriz danae Brady, 1883, p. 57, pl. 17, figs. 1-12. Scolecithrix danae Giesbrecht, 1888, p. 333; 1892, pp. 265, 283, pl. pl. 13, figs. 4, 9, 14, 17; pl. 37, fig. 6; 1898, p. 42. hin Fig. 26.—Scolecithriz danae. (a) Female, lateral, X20. (b) Genital seg- ment, female, lateral, X83. (c) Outer ramus of first foot of female X140. (d) Fifth foot of male X83. fRe.1dz., first joint of outer ramus of right foot. Re.sn., outer ramus of left foot. Ri.sn., inner ramus of left foot. 2 Fourth thoracic segment separate from fifth, latter with rather flat, rounded lateral angles. Third and fourth segments of the abdomen broader than long, genital segment with ventral, shovel-shaped process (fig. 265), anal segment short. Anterior antennae with nineteen segments, reaching beyond posterior border of the thorax but little. Outer ramus of posterior antennae 9/7 as long as the inner ramus, seventh joint of outer ramus without proximal bristle. Second basal of maxilla with five, inner ramus with six, outer with five bristles (ef. fig. 29c). First -—- —_e SS eae Vot.2] Ksterly.—Copepoda of the San Diego Region. 165 basal of fourth pair without inner marginal bristle, first joint of outer ramus of first pair (fig. 26c) with outer marginal bristle. Fifth pair of feet absent. 4 Mouth parts not retrograded; left fifth foot biramous, right uniramous, terminal joint very short (fig. 26d). Coloration: In formalin, both males and females have a light red or pink color. Length: Both sexes, 2-2.2 mm. Occurrence: June 28, 1904, one female; December 29, 1903, one female, surface tow at 2. a.m. One male, October 20, 1904. 2. Scolecithrix bradyi Giesbrecht. Scolecithrix bradyi Giesbrecht, 1888, p. 337; 1892, pp. 266, 283, plete. ple doy Hes. oe 7 bi 2d 28s pk Sitios. dl, 2, 9; 1898, p. 42. / Fig. 27.—Scolecithria bradyi. Female X31. 9° Line of separation between fourth and fifth thoracic seg- ments visible only on the back; lateral portions of last thoracic segment elongated into two flaps, on the right more than on the left. Third and fourth segments of the abdomen much broader than long, genital segment asymmetrical, anal segment as long as the preceding ones, furea twice as long as broad. Anterior antennae 19-jointed, not reaching the posterior end of the thorax. Outer ramus of posterior antennae longer than the inner, seventh joint of the outer ramus without a proximal bristle. Maxilla as in S. danae, except that outer ramus has four bristles. First basal of fourth pair without bristle on inner margin, first jomt of outer ramus of first pair without outer marginal bristle; fifth foot very small. 4 Right anterior antennae with 18 joints, left with 17. Left fifth foot longer than the right by the last joint. Third joint of the outer ramus of the right large and with a prong. Coloration: Yellowish pigment in body, mouth region, and on feet. 166 University of California Publications. [ ZooLoGy Length: Female, 1.4 mm. Occurrence: June 14, 1904, one female. 3. Scolecithrix persecans Giesbrecht. Scolecithrix persecans Giesbrecht, 1895, p. 253, pl. 3, figs. 6-12; 1898, p. 48, fig. 9. Fig. 28.—Scolecithrix persecans. (a) Male, lateral, X9. (b) Second foot, male, X45. (c) Left fifth foot of male X83. (d) Right fifth foot, male, X83. fe.3, third joint of outer ramus. (e) Fifth foot of female. 4 Tead with rather high crest (fig. 28a), last two thoracic segments fused; left anterior antenna 18-jointed, right 17- jointed, reaching beyond cephalothorax. Outer ramus of pos- terior antennae at least 114 times as long as inner; second basal of maxilla with five, outer ramus with eight, inner with seven bristles. appendages of anterior maxilliped in part pencillate. First basal of fourth foot with feathered inner border; middle of outer border of first basal of second and third feet with a small tooth, outer border of second basal of second to fourth feet with a tooth (fig. 285) ; spines on outer margin of the two prox- Vou.2] Esterly—Copepoda of the San Diego Region. 167 imal joints of outer ramus of first foot shorter and more slender than on the third joint; terminal saw of outer ramus of third foot indented at base, inner ramus of foot with three spines on posterior surface of second and joints; no spines on posterior surface of inner ramus of fourth foot. Anterior surface of outer ramus of second to fourth feet without spines, few on the anterior face of the inner ramus. Fifth foot fig. 28c, d. @ Anterior antennae 23-jointed, reaching to end of furea; abdomen symmetrical, ventral surface of genital segment convex. Posterior antennae mandible, maxilla and maxilliped and swim- ming feet as in the male. Fifth foot symmetrical, rather well developed (fig. 28e). Coloration: Opaque white in formation, eye spots red. Length: Male, 5.3 mm.; female, 4.6 mm. Giesbrecht gives the leneth of the male as 4.5 mm. Occurrence: Two males, one female collected at San Diego, May 31, 1904; obtained also May 18 and June 23, 1904. The female was not obtained by Giesbrecht, and has not since then been deseribed, as far as I am aware. There can be little doubt that the outer ramus of the right fifth foot in the male is 3-jointed, and that the terminal joint in Giesbrecht’s single specimen was broken off. I have seen a considerable number of males, and in all the outer ramus is 3-jointed as shown (fig. 28d). 4. Scolecithrix subdentata n. sp. © Last two thoracic segments fused, each side with a small in- dentation in the lateral margin. Anterior antennae 23-jointed, not much longer than the cephalothorax. Inner ramus of the pos- terior antenna 34 as long as the outer; second basal of maxilla with four bristles, rami each with five (fig. 29c). Appendages of anterior maxilliped vermiform. First basal of fourth foot with a small, non-plumose bristle on inner margin; inner mar- ginal bristle of second basal of third and fourth pairs long and plumose; outer margin of first basal of first, second and third pairs with a small tooth in the middle, inner margin with prom1- nent rounded process bearing the inner marginal bristle. First joint of outer ramus of first pair with outer marginal bristle. Fifth foot 2-jointed, leaf-like; terminal joint broad, oval, with 168 University of California Publications. [ ZOOLOGY a short distal spine on the outer border, and a longer proximal] spine in the middle of the outer border (fig. 290). & Unknown. ri ——\W Fig. 29.—Scolecithriz subdentata, n. sp. (a) Female, lateral, X31. (6) Fifth foot, female, X195. (c) Maxilla 140. B.2, second basal. i., inner ramus. fe., outer ramus. Approaches S. dentata Giesbrecht in form of last thoracie segment, but the indentation is not as deep as in that species. The fifth foot is much as in dentata, but more oval and rounded. Distinct from dentata in possessing an outer marginal bristle on the first joint of outer ramus of first foot, and in the number of joints of the anterior antennae. The bristles of the maxilla distinguish S. subdentata most sharply. S. subdentata has the same number of joints in the antenna as S. longicornis Scott and S. auropecten Giesbrecht. Length: Female, 1.48 mm. Occurrence: San Diego, May 31, June 14, June 23, 1904. 5. Seolecithrix pacifica n. sp. 2 Fourth and fifth thoracic segments fused, rounded later- ally. First segment of abdomen about as long as second and third together; the latter two are equal in length. Outer ramus of posterior antenna a little longer than the inner ramus. Sec- ond basal of maxilla with five bristles, inner ramus with eight, a Vou.2] Hsterly—Copepoda of the San Diego Region. 169 outer with five (fig. 30d). First basals of fourth feet without inner marginal bristle, inner border of second basal in second to fourth pairs ending in a sharp point. First joint of outer ramus of first foot with short, curved outer marginal bristle; first joint of outer ramus of fourth pair without outer marginal bristle. Fifth foot (fig. 830c) 2-jointed, with a short distal bristle and a very long proximal one. Fig. 50.—Scolecithrix pacifica, n. sp. (a) Female, lateral, X31. (bd) Anterior maxilliped 140. (c¢) Fifth foot X195. (d) Max- illa X83, parts as in fig. 29c. & Unknown. This specimen approaches S. porrecta closely in general char- acter, but is distinct in the length of the rami of the posterior antennae, form of the maxilla, bristle on outer margin of first joint of the outer ramus of the first foot, and in the form of the fifth feet. The anterior antennae are broken, but have prob- ably not over twenty joints. Length: Female, 2.3 mm. Occurrence: June 23, 1904, San Diego, one female. 170 University of California Publications. [ ZooLoGy 6. Scolecithrix similis T. Scott. Amallophora dubia var. similis Scott, T., 1893, p. 56, pl. 4, figs. 19-23. Scolecithrix similis Giesbrecht, 1898, p. 46. S. similis (?) Wolfenden, 1904, p. 119, pl. 9, figs. 5, 6. Fig. 31.—Scolecithriz similis. (a) Male, lateral, X31. (b) Fifth foot, male, X45. 4 First abdominal segment short, second long, twice the length of the third, which is shorter than the fourth. Right anterior antenna 18-jointed, left 23-jointed (Scott). Last two thoracic segments fused. First basal of fourth foot with a plumose inner marginal bristle; both feet of fifth pair biramous (fig. 310). Length: Male, 2.6 mm. Occurrence: San Diego, June 23, 1904. The antennae of the single specimen were broken, but the | rd form of the abdomen and fifth feet warrant one in identifying it with Scott’s species, at least provisionally. Fam. CENTROPAGIDAE. Centropagidae Giesbrecht, 1892, p. 58; 1898, p. 52. 2 Head always distinct from thorax; rostrum with two, usu- ally soft, filaments, sometimes plumose. Anterior antennae as in the Calanidae, but the second joint is more often divided into two parts, never less than twenty-three joints. Outer ramus of posterior antennae at least 24 as long as the inner. The suc- ceeding four pairs of appendages as in Calanus and like forms. In the three anterior ones are found peculiarities (in the Hete- * a yy wohtee ee wl awe ene Ee a a o@ ——————————OoUEE— OOO ee Vot.2] Hsterly.—Copepoda of the San Diego Region. ek rorhabdinae) : stunting of the second and third inner marginal lobes of the maxilla through lengthening of outer ramus; pre- ponderance of distal bristles of anterior maxilliped over the prox- imal. The four. anterior pairs of feet with 3-jointed rami; but in Temora the number of joints is reduced through fusion. The fifth pair of feet is like the others (inner marginal bristle of second joint of outer ramus of special form, sword-shaped, awl- or thorn- lke) or rudimentary, inner ramus 1-jointed or lacking, outer ramus 1- to 3-jointed. 4 Abdomen with five segments, anal segment rarely short- ened; genital orifice and grasping antenna on opposite sides of the body. Grasping antenna right or left, joints 19 to 21, and 22 to 23 fused. Both feet of fifth pair present, inner rami complete or reduced to absence; outer rami forming hooks or forceps. Sheght sexual differences occasionally in form of last thoracic segment and swimming feet. Sub-fam. CENTROPAGINAE. Centropagina Giesbrecht, 1892, p. 59. 2 Cephalothorax with six segments, abdomen with three ; ros- tral filaments soft. Anterior antennae (24th and 25th joints fused), mandibles and maxilla as in Calanus; the length of the distal curved bristles of the anterior maxillipeds and the heavily bristled first basal of the posterior maxillipeds is characteristic. All five pairs of feet with 3-jointed rami. & Grasping antenna on the right side; outer ramus of left fifth foot 2-jointed; the right foot with forceps. 1. Genus Centropages Kroyer. Centropages Kroyer, 1849, p. 602. Catopia Dana, 1848, p. 25; 1849, p. 280; 1852, p. 1172. Hemicalanus Dana, 1852, p. 1103. Ichthyophorba Lilljeborg, 1853, p. 184. Tiaptomus Lubbock, 1857, p. 403. Ichthyophorba Claus, 1863, p. 198. Centropages Brady, 1883, p. 81. Centropages Giesbrecht, 1892, pp. 59, 303, 731; 1898, p. 53. Centropages Wheeler, 1899, p. 172. Centropages Thompson and Scott, 1903, p. 247, pl. 1, figs. 19-25. 2 Head separate from thorax, fourth thoracic segment from fifth. Abdomen with three segments, genital segment asymmet- 172 University of California Publications. | ZooLocy rical. Anterior antennae 24-jointed; joints 24 and 25 fused. Outer ramus of posterior antennae 7-jointed and almost 1144 times as long as the inner ramus. The distal bristles of the ante- rior maxillipeds sickle-shaped, with spinous feathering, and much longer and thicker than the proximal bristles. First basal of the posterior maxillipeds with strongly protruding lobes, both the middle ones set with bristles, which have a spinous feathering; inner ramus well developed, 5-jointed. Rami of swimming feet usually 3-jointed, but the inner ramus is exceptionally 2-jointed. First basal with bristle on inner border in first to fourth feet, second basal thus equipped in the first pair. First basal of fifth pair without inner marginal bristle; inner marginal bristle of second joint of outer ramus thorn-like and fused with the joint (fig. 32c.) 4 Sexual peculiarities in the form of the abdomen, right anterior antenna, and fifth pair of feet. The abdomen is com- posed of five segments; anal segment in most species very short ; genital opening on the left. Right anterior antenna a grasping organ. Joints 19-21 and 22-23 fused, geniculation between the 18th and 19th. Inner marginal bristles lacking on outer ramus of left fifth foot, joints 2 and 3 fused. The outer ramus of the right foot 3-jointed, both distal joints metamorphosed into a for- ceps, one blade of which is the terminal joint, while the other is the thickened inner marginal bristle of the second joint. 1. Centropages bradyi Wheeler. Centropages violaceus Brady, 1883, p. 83, pl. 37, figs. 1-14. Centropages bradyi Wheeler, 1899, p. 174, fig. 12. 2 Second joint of outer ramus of fifth foot with a stout smooth spine (fig. 32c). Sides of inflated genital segment with- out spines or knob-shaped projections. Furea symmetrical, with a peculiar short, truneated, peg-shaped projection (fig. 320) between insertions of the two outer bristles. (Wheeler, 1899). 4 Joint 17 of right anterior antenna with smooth anterior border, not serrate; joints 19 and 20 fused, separated from joint 21; joint 18 with accessory series of teeth on lower surface (Wheeler, 1899). Vou.2] Esterly.—Copepoda of the San Diego Region. a3 Coloration: Opaque, with a large purplish spot in middle of body. Length: Female, thorax, 1.6 mm.: abdomen ? Occurrence: June 10, 1904, one female. : Fig. 32.—Centropages bradyi. (a) Female, thorax, lateral, X45. (b) Abdomen, ventral, after Wheeler 1899. (c) Fifth foot x83. SW Wheeler, 1899, p. 174, does not mention the spine-like pro- tuberance on the dorsal surface of the first segment of the ceph- alothorax in the female, but since the other characters as given by him (especially the furea) agree with the San Diego speci- men, I have not made a new species of the latter. This agrees in possessing the dorsal spine, with C. dorsispinatus (Thompson, 1903, p. 247, pl. 1, figs. 19-25), but differs widely in other respects. Sub-fam. TEMORINAE, Temorina Giesbrecht, 1892, p. 60. 2 Cephalothorax with five segments; fourth and fifth thor- acic segments fused; rostral filaments soft, sometimes plumose. Anterior antennae 23- or 24-jointed; the second joint is either not divided, or, if it is divided into two parts, the proximal! portion is fused with the first joint. Outer ramus of posterior antennae 7-jointed, and, with the following four appendages, is 174 University of California Publications. | ZOOLOGY like those of the Calanidae. The first four pairs of feet usually with 3-jointed rami, in which, however, the two proximal Joints may be fused; inner ramus absent in fifth pair, or small and 1-jointed ; the outer ramus is 1- to 3-jointed. 4 Grasping antenna usually the right; distally from the geniculation, the nineteenth and twenty-first and twenty-second and twenty-third joints are fused; sexual peculiarities often in the swimming feet as well as in the form of the body, anterior antennae and fifth pair of feet. 1. Genus Pleuromamma Giesbrecht. Diaptomus Lubbock, 1856, p. 27. Pleuromma Claus, 1863, p. 195. Pleuromma Brady, 1883, p. 45. Pleuromma Giesbrecht, 1892, pp. 61, 347, 757. Pleuromma Dahl, 1893, p. 105. Pleuromma Wheeler, 1899, p. 176. Pleuromamma Giesbrecht, 1898, p. 108. Easily recognizable by a dark-pigmented knob on the right or left side of the first thoracic segment (figs. 33a, 34a). Furea at most twice as long as broad. Rami of the first to fourth pairs of feet 3-jointed, first joint of outer ramus of third pair with a deep notch in the outer border; terminal bristle of outer ramus of third pair short and bent outward; first joint of inner ramus of second pair with hooks on inner border, on right and left foot in the female, usually on one side in male. Fifth pair in female rudimentary, 2- to 4-jointed, in male 5-jointed on each side, without forceps. Grasping antenna of male on right or left side. Abdomen of female with three segments; of male with five, sometimes asymmetrical. 1. Pleuromamma abdominalis Lubbock. Diaptomus abdominale Lubbock, 1856, p. 28, pl. 10, figs. 1-8. Pleuromma abdominale Claus, 1863, p. 197, pl. 5, figs. 1-6, 13, 14; pl. 6, fig. 1-10. Pleuromma abdominale Brady, 1883, p. 46, pl. 11, figs. 1-13. Pleuromma abdominale Giesbrecht, 1892, pp. 347, 357, pl. 5, fig. 8; pl. 32, figs. 3, 5, 13, 22, 25-30; pl. 33, figs. 43, 44, 48, 49, 52. Pleuromamma abdominalis Giesbrecht, 1898, p. 109. Vou.2] Ksterly—Copepoda of the San Diego Region. AS: ® Pigment knob on right or left side; proximal joint of first antenna with several smaller and two larger (one straight and one curved) teeth on anterior border. Fifth pair of feet 4- jointed, with three apical bristles (fig. 330). eee Fig. 33.—Pleuromamma abdominalis and P. gracilis. (a) P.a., female, lateral, X31. (0b) P.a., fifth foot X45. (c) P.g., fifth foot 195. ¢ Pigment knob, genital opening and hooks on inner border of first joint of inner ramus of second foot, on left side. Proxi- mal joint of anterior antennae with small teeth only, grasping antenna on right side. Abdomen symmetrical. End joint of left fifth foot broadened. ~ Coloration: Transparent, except for a small amount of red in the mouth region. Length: Female, about 2.4-3 mm.; male, 3.3 mm. Occurrence: A common species, both summer and winter, but males are very infrequently found; one male was taken June 2, 1904. 2. Pleuromamma gracilis Claus. Pleuwromma gracile Claus, 1863, p. 197, pl. 5, figs. 7-11. Pleuromma abdominale (in part) Brady, 1883, p. 47, pl. 2, figs. UENMGe joll, Bil, aero, Way Wee Pleuromma gracile Giesbrecht, 1892, pp. 347, 357, pl. 5, fig. 7; pl. 32, figs. 6, 18-20; pl. 33, fig. 41-47. Pleuromamma gracilis Giesbrecht, 1898, p. 110. 2 Pigment knob on right side; anterior border of anterior antennae with only small teeth; fifth pair of feet 2-jointed, three prongs at the end (fig. 33c). 176 University of California Publications. [ ZooLoGy & Pigment knob on right side; abdomen symmetrical; ante- rior antennae as in female as regards armature; grasping antenna on left side; first joint of inner ramus of second foot with hooks only on right side; third and fourth feet as in female. Coloration: As in P. abdominalis. Length: Both sexes, 1-2 mm. Occurrence: More abundant than P. abdominalis, but males are rare. Brady, 1883, p. 47, considers that P. gracilis Claus is an immature form of P. abdominalis Claus, but Giesbrecht, 1892, does not favor this view, and the San Diego specimens of P. gra- cilis present such differences when compared with P. abdominalis that there can be no doubt of the distinctness of the species. The forms represented by P. gracilis are without doubt mature, since females are often found with attached spermatophores. 3. Pleuromamma xiphias Giesbrecht. Pleuromma sxiphias Giesbrecht, 1889, p. 6; 1892, pp. 347, 367, pl. 32, fig. 14; pl. 33, figs. 42, 45, 50. Pleuromamma wxiphias Giesbrecht, 1898, p. 110. Fig. 34.—Pleuromamma sxiphias. (a) Female, lateral, X18. (b) Head of female, lateral, X48. (c) Second basal, and proximal joints of rami of second foot, x48. Vou.2] Esterly—Copepoda of the San Diego Region. A by Gri Allied to P. abdominalis, but the front of the head anterior to the rostrum is prolonged into a prominent process (fig. 340). Coloration: As in the other species. Length: Female, 4.1-4.5 mm. Occurrence: San Diego, July 31, 1903, one female; June 23, 1904, one female; taken also December 23, 1903, on ‘‘Banks.’’ The occurrence of the male is uncertain; large male animals have been taken with the characteristic shape of xiphias, but I cannot say definitely whether they belong to this species or not. 2. Genus Metridia Boeck. Metridia Boeck, 1864, p. 13. Paracalanus Brady and Robertson, 1878, p. 126. Metridia Giesbrecht, 1892, pp. 61, 339, 749; 1897, p. 254; 1898, p. 105. Metridia Dahl, 1894a, p. 10. Metridia Wheeler, 1899, p. 175. Metridia Wolfenden, 1904, p. 125. (See also T. Scott, 1893, p. 42, pl. 3, figs. 8-20.) Closely allied to Pleuromamma, but is without the lateral pig- ment knob. Terminal bristle of outer ramus of third pair of normal form; swimming feet of the male (especially the second pair) corresponding with those of the female. Furea 2 to 5 times as long as broad. 1. Metridia lucens Boeck. Metridia lucens Boeck, 1864, p. 14. Paracalanus hibernicus Brady and Robertson, 1873, p. 126. pl. 8, figs. 1-3. Metridia armata Brady, 1878, p. 42. Metridia hibernica Giesbrecht, 1892, pp. 340, 357, pl. 33, figs. 2, 12; 16, 22, 28,36, 39. Metridia lucens Dahl, 1894, p. 11. Metridia lucens Giesbrecht, 1898, p. 106. 2 Cephalothorax 124 times as long as the abdomen; lateral angles of fifth thoracic segment slightly pointed. Genital seg- ment somewhat shorter than the two last abdominal segments together, the anal segment about 34 as long as the preceding. Furea shorter than the last abdominal segment and twice as long as broad. The anterior antennae reach back hardly to posterior margin of the genital segment. Terminal bristle of end joint of outer ramus of fourth foot lttle over 14 as long 178 University of California Publications. | ZooLoGy as the joint. Fifth foot 3-jointed and with three rather long bristles on end joint (fig. 35c). © a Fig. 35.—Metridia lucens and M. boeckii. (a) M.l., female, lateral, 20. (b) M.1., second basal and first joint of inner ramus of second foot of female to show hooks, K195. (c) M.L, fifth foot of female X195. (d) M.b., fifth foot of female 195. 4 Grasping antenna on right side. Fifth foot: second joint of outer ramus of left foot without, first joint of outer ramus of right foot with a long, thorn-lke bristle. Length: Female, 3.2 mm. Occurrence: Very common, summer and winter. 2. Metridia boeckii Giesbrecht. Metridia boeckii Giesbrecht, 1889, p. 5; 1892, pp. 340, 346, pl. 33, figs. 8, 19, 31, 37; 1898, p. 107. ; 2 Like M. lucens, but furca is as long as the fifth abdominal segment, and twice as long as broad. Anterior antennae reach a little beyond the posterior border of the thorax. Fifth foot with four joints (fig. 35d). & Unknown. Length: Female, 2.5 mm. Occurrence: One or two in catches with MW. lucens. i re a a ~ Te) vou.2] Esterly—Copepoda of the San Diego Region. 1 It should be noted that not a male specimen of Metridia has been taken in any eatch, so far as I have examined them, and rather particular attention has been paid to this point. Sub-fam. LucIcurTmuNas. Leuckartiina Giesbrecht, 1892, p. 62. 2 Cephalothorax with five segments, fourth and fifth thoracic segments fused, rostral filaments thin and usually soft; abdomen with four segments, symmetrical. The second joint of the ante- rior antenna is divided, and the twenty-fourth joint is separate from the twenty-fifth. Outer ramus of posterior antennae 8- jointed, the four following appendages like those in the Cala- nidae. The first four pairs of feet almost always with 3-jointed rami, the fifth pair like the preceding ones and that of Centro- pages, with 3-jointed outer ramus and 2- to 3-jointed inner ramus. & (Known only in Lucicutia.) Grasping antenna the left ; distal to the geniculation the nineteenth to twenty-first, and twenty-second and twenty-third joints are fused; fifth pair of feet without forceps, basals 2-jointed, the right with 2-, the left with 3-jointed rami; no other sexual differences except in form of body. 1. Genus Lucicutia Giesbrecht. Leuckartia Claus, 1863, p. 182. Leuckartia (in part) Brady, 18838, p. 50. Leuckartia Giesbrecht, 1892, pp. 62, 358; 1895, p. 25 Lucicutia Giesbrecht, 1898, p. 110. Lucicutia Steuer, 1904, p. 596. Lucicutia Wolfenden, 1904, p. 121. oO Head broad; furca symmetrical. First lobe on outer border of maxilla with five bristles. © Five segments in cephalothorax, abdomen with four, sym- metrical. Rostral filaments slender, situated on a papilla. Pos- terior antennae like those of Centropages, but with eight joints in outer ramus. Blade of mandible weakly built, outer ramus bent rather far proximally. Outer border lobes of maxilla with only five bristles; inner border lobes well developed, the proximal one, however, with weak masticatory bristles. Inner ramus 2- jointed, articulated with basal; outer ramus large, oval. Distal 180 University of California Publications. ‘| ZooLowy bristles of maxillipeds little longer than the proximal; bristles of outer border of posterior maxilliped slender, without hairs. Outer rami of the five pairs of feet 3-jointed; inner ramus of first pair 2-jointed (second and third joints fused), of second to fifth pairs 3-jointed; first basal with bristle on inner margin in second to fourth pairs, second basal in first pair with inner marginal bristle, and sometimes with a tube-like process. The bristle on the inner margin of the second joint of the outer ramus of the fifth pair has the form of a curved awl (fig. 360). 4 Sexual peculiarities in the form of the abdomen, posterior antennae and fifth pairs of feet. Abdomen with five segments, genital opening on right side. The left antenna is a grasping organ, geniculating between joints 18 and 19; joints 19 to 21, 22 and 23 fused. Fifth pair of feet with 2-jointed basals; rami of left foot 2-jointed, of right 3-jointed. The distal joint of the outer ramus of the right foot is hooked and may be moved toward the proximal joints. 1. Lucicutia flavicornis Claus. Leuckartia flavicornis Claus, 1863, p. 183, pl. 32, fig. 17. Leuckartia flavicornis Giesbrecht, 1892, p. 358, pl. 5, fig. 4; pl. 19, figs. 2, 3, 15, 17, 21, 23, 29, 38; pl. 38, fig. 38, 40. Lucicutia flavicornis Giesbrecht, 1898, p. 111. ri. VS —=— Fig. 36.—Lucicutia flavicornis. (a) Female, lateral, X18. (b) Fifth foot, female, X83. Ri., inner ramus. (c¢) Outer margin of outer ramus of third foot 140. Q Anal segment shorter than the preceding; second terminal bristle of furea thick, twice as long as abdomen. The anterior an- Vou.2] Hsterly—Copepoda of the San Diego Region. 181 tennae reach beyond middle of the furea, joint 19 as long as tenth to twelfth, inclusive. Second basal of maxilla with four bristles. Inner ramus of first pair of feet 3-jointed, with eight bristles; inner ramus of fifth pair reaches almost to the distal border of the second joint of the outer ramus; first joint of outer ramus much shorter than the third, which is twice as long as the term- inal bristle. & Terminal portion of grasping antenna (joints 19-25) some- what !onger than joints 14-18. Inner ramus of right foot of fifth pair straight, with five bristles, which are at the end of the terminal joint. Coloration: Transparent, with light yellowish pigment in various locations. The San Diego specimens showed this to a very small extent. Length: Female, 1.6 mm.; male a little less. Occurrence: June 8, 1904, one male, one female; June 10, 1904, one male (?). Sub-fam. HETERORHABDINAE. Heterochaetina Giesbrecht, 1892, p. 63. Cephalothorax with five segments; fourth fused with fifth thoracic segment; rostral filaments slender, sometimes plumose ; last thoracic segment in some cases with pointed lateral angles. Abdomen with three or four segments, not always symmetrical. Second joint of anterior antennae divided, the two terminal joints usually distinet. Second joint of outer ramus of posterior antennae divided into two, so that there are as a result eight joints in the ramus (which, however, may be reduced by fusions). Blade of mandible with few teeth, inner ramus small, sometimes lacking. Inner ramus and both distal lobes of inner margin of maxilla small, occasionally absent; outer ramus always present, and usually much lengthened. Anterior maxilliped elongate, lobes small, the proximal ones usually rudimentary; bristles of distal lobes, and usually those of the inner ramus, almost always long, thick and hooked. The four anterior pairs of feet with 3-jointed rami, fifth pair like the others, rami almost without exception 3-jointed. 4 Sexual differences in form of body, anterior antennae, fifth pair of feet, seldom in structure of mouth parts. Grasping an- 182 University of California Publications. [ ZooLoGy tenna usually the left; first and second joints fused, as well as the nineteenth to twenty-first, twenty-second and twenty-third (or twenty-second to twenty-fifth) ; fifth pair of feet with 3- jointed outer and 1- to 3-jointed inner ramus; forceps incomplete or absent. 1. Genus Heterorhabdus Giesbrecht. Heterochaeta Claus, 1863, p. 180. Heterochaeta Brady, 1883, p. 48. Heterochaeta Giesbrecht, 1892, pp. 64, 372, 745; 1895, p. 259. Heterochaeta, Aurivillius, 1899, p. 38, figs. 4, 5. Heterorhabdus Giesbrecht, 1898, p. 113. Heterorhabdus Wolfenden, 1904, p. 124. 2 Cephalothorax with five segments, rostral filaments soft, situated on a papilla. Abdomen with four segments, left half of furea not articulating with anal segment, larger than the right and with much longer bristles. Rami of posterior antennae about equal in length, outer ramus with eight bristles. Anterior maxil- liped straight, terminal portion and proximal lobes with their bristles strongly suppressed, while the distal lobes are provided with strong hooked bristles. Posterior maxillipeds distinguished by shortness of the bristles on the inner ramus and by the length and thickness of one bristle on the inner margin of the first basal joint. All the feet have 3-jointed rami; inner marginal bristle of the first basal in pairs one to four, and of the distal basal joint in the first pair, well-developed and plumose. Terminal joint of outer ramus of third pair usually different in form from that joint in the other pairs, being broad and oval (fig. 38d). The inner marginal bristle of the second joint of the outer ramus of the fifth pair is thickened and sword-shaped (fig. 38e). 4 Sexual peculiarities in form of posterior portion of body, left anterior antenna and fifth pair of feet. Abdomen with five segments, genital opening on right side. Grasping antenna slender. Fifth pair of feet with 3-jointed rami; terminal joints of both outer rami hooked; processes on the distal basal joint. KEY TO SPECIES. Abdomen 4-segmented, fifth foot symmetrical.................- 2 Abdomen 5-segmented, fifth foot asymmetrical................ 5 -1. Third joint of outer ramus of third foot of same form as in second and fourth pairs) (fies 400). ar etait H. longicornis ee Pett ee ee ee VoL.2] Hsterly—Copepoda of the San Diego Region. 183 1. This joint in third pair broad and oval, terminal bristle short- Med (OA SEO) imeers aa ctersie ssh lcr sce si ctw lens ouckelere velo enersts eo ters lo are eels fel 2 2. First basal of posterior maxilliped with a very long, heavy bristle in the middle of the inner border; rostral papilla with 2 [oonini; (i, BiWMeso60cc0s0ccns00end suo5ad0asGon abe H. spinifrons 2. As above, but rostral papilla without point (figs. 38), ¢)........ 3 3.2 Inner marginal bristle of second joint of inner ramus of fifth foot much shorter and more slender than those of the third joint; first joint of outer ramus with thorn-like inner marginal Jorenisiiel, 25 Iivorisay stovare (ites GE). 66 oacnaocehecccsneaease H. clausi 3.9 Inner marginal of second joint of inner ramus of fifth foot but little shorter than those of third joint; first joint of outer ramus without inner marginal bristles............... H. papilliger 1. Heterorhabdus spinifrons Claus. Heterochaeta spinifrons Claus, 1863, p. 182, pl. 32, figs. 8-9, 14, 16. Heterochaeta spinifrons Giesbrecht, 1892, pp. 372, 382, pl. 10, figs. 1, 3, 11, 16, 19, 31; pl. 39; figs. 42, 43, 51, 52, 54. Heterorhabdus spinifrons Giesbrecht, 1898, p. 114. Fig. 37.—Heterorhabdus spinifrons. Female. (a) Head, lateral, X83. (6) Posterior maxilliped 167. 8.1, first basal. The papilla on front of head ends in a sharp point; anterior antennae reach beyond the end of the furea by the last four or five joints. The fourth lobe of the anterior maxilliped has two long, thick bristles, and a small, slender one which is hardly 14 as long as the other two; the fifth lobe has two bristles, one of which is longer and thicker than the other. A spine-like bristle at the end of the inner margin of the first basal of the posterior maxil- liped is 14 the length of the long bristle in the middle of the margin (fig. 370). Hooks at the end of both outer rami of the 184 University of Californa Publications. | ZooLocy fifth foot of the male relatively longer than in H. papilliger, the left over twice as long as the first and second joints of the outer ramus. Coloration; Transparent and colorless. Length: Female, 3.4 mm. Occurrence: June 23, 1904, one female. 2. Heterorhabdus papilliger Claus. Heterochaeta papilligera Claus, 1863, p. 182, pl. 3, figs. 10-13, 15. Heterochaeta papilligera Giesbrecht, 1892, pp. 372, 382, pl. 20, figs. 4, 7, 10, 15, 17, 23, 35, 36; pl. 39, figs. 40, 53. Heterorhabdus papilliger Giesbrecht, 1898, p. 114. 7 ---Ve2, pS Fig. 58.—Heterorhabdus papilliger. (a) Female, lateral, X31. (6b) Head, female, lateral, X83. (c) Head, female, dorsal, X83. (d) Outer ramus of third foot of male X83. (e) Fifth foot of female 140. fRe.2, second joint of outer ramus. (f) Right mandibular blade of male X83. (g) Anterior maxil- liped, female, X83. ne VoL.2] Esterly.—Copepoda of the San Diego Region. 185 Papilla on front of head elongated but not ending in a point (figs. 380, c). Anterior antennae when at the sides of the body reach about to the end of the furea. Fourth lobe of the anterior maxillipeds (fig. 389) with two long, thick bristles and a shorter, thinner one, which is over half as long as the others; fifth lobe with two bristles of about equal length and thickness. On the end of the inner border of the first basal of the posterior maxilliped is a spine-like bristle, which is hardly one-eighth as long as the bristle in the middle of the margin, and the latter one is almost twice as long as the second basal (ef. fig. 37b). Hooks at the end of both outer rami of the fifth foot in the male relatively shorter than in spinifrons; the left little longer than the first and second joints of the outer ramus together. Coloration: As in spinifrons. Length: Female, 2.2 mm.; males slightly smaller. Occurrence: A few specimens of both sexes were taken during May and June, 1904. 3. Heterorhabdus clausi Giesbrecht. Heterochacta claus Giesbrecht, 1889, p. 2; 1892, pp. 372, 382, pl. 20, fig. 2, 28, 37, 38. Heterorhabdus clausi Giesbrecht, 1898, p. 115. Fig. 39.—Heterorhabdus clausi. Fifth foot, male, 82. 186 University of California Publications. [ ZooLoGy Like papilliger, but the anterior antennae reach somewhat beyond the end of the furea. Inner ramus of anterior maxilli- peds with longer bristles; inner marginal bristle of first and second joints of inner ramus of fifth foot in the female short and slender, that of the first joint of the outer ramus thick and . hooked; second basal of the right fifth foot in the male with a long lamellar process, the second joint of the outer ramus with a shorter projection on the inner border, third relatively longer, especially on the left side. Length: Male, 2-2.5 mm. Occurrence: San Diego, July 22, 1903, one male; June 23, 1904, one male. 4. Heterorhabdus longicornis Giesbrecht. Heterochaeta longicornis Giesbrecht, 1889, p. 2; 1892, pp. 373, 383, pl. 20, figs. 14, 21, 25, 26; pl. 39, fig. 44. Heterorhabdus longicornis Giesbrecht, 1898, p. 116. Heterorhabdus zetesios Wolfenden, 1902, p. 367. Heterorhabdus longicornis (male) Wolfenden, 1904, p. 124, pl. 9, fig. 34. Fig. 40.—Heterorhabdus longicornis. Male. (a) Third foot 140. (6b) Right fifth foot X140. (c) Left fifth foot «140. 2 Anterior antennae reach beyond the end of the furca for the last eight or nine joints; inner ramus of maxilla with five bristles, first and second inner marginal lobes relatively long; anterior maxillipeds with a greater number of bristles, but with a en Vou.2] Hsterly—Copepoda of the San Diego Region. 187 less strongly developed hooked bristles than in the other species; inner ramus clearly with three joints, and with seven long bristles; bristles of first basal of the posterior maxillipeds and third joint of outer ramus of third swimming foot of usual form. Inner marginal bristle of second joint of outer ramus of fifth pair more slender, and inner marginals of first and second joints thicker than in the other species, distal border of second joint of outer ramus of ordinary form. 4 Like female in structure of maxillipeds and terminal joint of outer ramus of third and fourth swimming feet. Fifth foot (figs. 406, c): right with stiff upright process on second basal (Inner margin), covered with stiff spines, second joint of outer ramus with a projection having four teeth at end. Length: Male, 3 mm. Occurrence: San Diego, June 23, 1904, one male. 2. Genus Augaptilus. Hemicalanus (in part) Claus, 1863, p. 176. Augaptilus Giesbrecht, 1889, p. 3; 1892, pp. 65, 400, 724; 1898, p- 120. (See also T. Scott, 1893, p. 36, pl. 2, figs. 25-37; Steuer, 1904, p- 597.) 2 Cephalothorax composed of five segments; rostral filaments short and sometimes feathered. Abdomen with 3 segments, geni- tal segment usually not wholly symmetrical. Anterior antennae 25-jointed, outer ramus of posterior antennae rarely longer than the inner ramus. Mandibular blade with two teeth (mandible sometimes uniramous and with a stylet-like blade). Inner ramus of maxilla lacking; both maxillipeds with reduced proximal lobes and peculiarly equipped bristles (ef. fig. 410). Feet with spines on outer border of outer rami reduced in part, the third bristle on the inner border of the last joint of the outer ramus of the fifth foot not elongated, the inner marginal bristle of the middle joint awl-shaped or lacking; both rami 3-, rarely 2-jointed. & Sexual differences in the form of the abdomen, anterior antennae and fifth feet. Abdomen with 5 segments, genital open- ing right or left. The right or left anterior antenna may be the grasping organ. Rami of both feet of fifth pair 3-jointed (fig. 41c, d). 188 University of California Publications. [ ZooLocy 1. Augaptilus longicaudatus Claus. Hemicalanus longicaudatus Claus, 1863, p. 179, pl. 29, fig. 3. Augaptilus longicaudatus Giesbrecht, 1892, p. 414, pl. 27, fig. 31; pl. 28, figs. 11, 19, 23, 31, 35, 38; pl. 2, fig. 22; pl. 39, figs. 37, 48; 1898, p. 123. Augaptilus longicaudatus Scott, 1894, p. 34, pl. 1, figs. 24-26; pl. 2, fig. 5. Fig. 41.—Augaptilus longicaudatus. Male. (a) Dorsal X18. (b) An- terior maxilliped X83. (c) Right fifth foot X83. (d) Left fifth foot x83. 2 Genital segment not entirely symmetrical, twice as long as both the following segments together; furca as long as the anal segment, and about 5 times as long as broad. Anterior antennae longer than trunk by about the last 6 joints. Inner ramus of posterior antennae 14 longer than the outer ramus; first and sec- ond joints of outer ramus not fused; mandible uniramous. Anterior maxilliped: First and second lobes lacking, third with 1 bristle, fourth and fifth with 2, sixth with 1. First basal of posterior maxilliped with 0, 0, 1, 2 bristles. Length of first and second basals end inner ramus as 7:6:5. Outer ramus of fifth foot 2-jointed. he a i ees a Vot.2] Hsterly.—Copepoda of the San Diego Region. 189 4&4 Grasping antenna on left. Fifth foot, fig. 41¢, d. Coloration: Transparent, without pigment. Length: Male, 3.39 mm. Occurrence: June 10, 1904, 1 male. 3. Genus Arietellus Giesbrecht. Arietellus Giesbrecht, 1892, pp. 66, 415. Rhinealanus (part) T. Scott, 1893, p. 31. Arietellus Giesbrecht, 1898, p. 124. Last two thoracic segments fused, elongated into a strong spine on each side (fig. 42a), front with wedge-shaped process, rostral filaments slender. Abdomen of female with 4 segments, symmetrical; furea, and appendages with long, richly plumose bristles. Anterior antennae of female and the right one of male at most 20-jomted, joints 1 and 2, 21-25 fused; grasping antenna on the left, 19-jointed, terminal portion 2-jointed. Inner ramus of postericr antenna straight, longer than outer; mandible uniramous, inner ramus lacking; inner ramus and third inner marginal lobe of maxilla lacking, outer ramus long and characteristic. Anterior and posterior maxillipeds as in Augaptilus except in appendages of bristles (fig. 42c). Rami of first to fourth feet 3-jointed; fifth foot of female (fig. 426) 3-jointed, basals 2-, outer ramus 1-jointed, inner ramus rudi- mentary. Fifth foot of male without forceps, basals 2-, outer ramus 3-jointed, inner ramus 1-jointed. 1. Arietellus setosus Giesbrecht. Arietellus setosus Giesbrecht, 1892, p. 415, pl. 29, figs. 1, 3-7, 9-13; pl. 39, figs. 34-36; 1897, p. 254; 1898, p. 124. With the characters of the genus. Coloration: Terminal expansions of plumose fureal bristles red, the remaining portion black. Body orange red, bristles on posterior antennae and mouth parts, deep red. Leneth: 5.5 mm. Occurrence: One female was taken at San Diego, Dee. 22, 1903. 190 University of California Publications. [ ZooLocy Fig. 42.—Arietellus setosus. Female. (a) Dorsal X9. (b) Fifth foot X83. (c) Distal portion of one of the bristles of the an- terior maxilliped «140. 4. Genus Phyllopus Brady. Phyllopus Brady, 1885, p. 78. Phyllopus Giesbrecht, 1892, pp. 66, 419; 1898, p. 124. Phyllopus Wolfenden, 1904, p. 124. 2 Last thoracic segment not entirely symmetrical; abdomen with 4 segments, genital segment asymmetrical. Anterior antennae with 24 joints. Inner ramus of posterior antennae about half as long as the outer ramus of 8 joints. Blade of mandible strong, with four teeth. Anterior maxillipeds elon- gated, posterior with short, broad first basal. First to fourth pairs of feet with 3-jointed rami, second basal with inner mar- 7 a... oe oO tst—‘_aOS Vou.2] Esterly—Copepoda of the San Diego Region. Nit einal bristle in first pair and with outer marginal in first and fourth. Fifth pair with basal of two joints and 3-jointed outer ramus. Inner ramus lacking, inner marginal bristle of middle joint of outer ramus thick and long; terminal joint shortened, its distal margin toothed (fig. 430). é Like female except in structure of anterior antennae and fifth feet. Abdomen with 5 segments. Left anterior antenna 20-jointed, geniculating between joints 17 and 18. Fifth feet each with 2 basals, and 3-jointed outer rami, the right foot has a rudimentary inner ramus, broad and without spines. Second basal of each foot with a long, slender plumose bristle. 1. Phyllopus bidentatus Brady. Phyllopus bidentatus Brady, 1883, p. 78, pl. 5, figs. 7-16. Phyllopus bidentatus Giesbrecht, 1892, p. 419, pl. 18, figs. 25-33; pl. 38, fig. 35; 1898, p. 124. Phyllopus bidentatus & Wolfenden, 1904, p. 124, pl. 9, fig. 16. Fig. 43.—Phyllopus bidentatus. Female. (a) Lateral X18. (b) Fifth foot X195. With the generic characters. Both Giesbrecht (1892) and Wolfenden (1904) state that the ‘‘bidentate’’ lateral portion of the last thoracic segment does not exist as in Brady’s descrip- tion. The San Diego specimen agrees with the description of the two former authors. The male of the species is described by Wolfenden as cited, and the above description is taken from him. Coloration: Transparent, without pigment. 192 University of Califorma Publications. — [ZooLoey Length: Female, 2.2 mm. Occurrence: San Diego, May 31, 1904, one female. Fam. CANDACIIDAE. Candacidae Giesbrecht, 1892, p. 67. Candaciidae Giesbrecht, 1898, p. 126. 2 Cephalothorax with 5 segments, rostrum absent, abdomen with 3 segments. In posterior antennae the second basal and first joint of inner ramus is fused, outer ramus slender, end joints shortened. Blade of mandible with few teeth. Second lobe of inner margin of maxilla very long, third and fourth absent. Anterior maxilliped without lobes, bristles on distal portions sickle-shaped and hooked. Posterior maxilliped as in Calanus but small and weak. Inner ramus of anterior pairs of feet 2- jointed; fifth pair rudimentary. & Genital orifice on left; grasping antenna the left, seven- teenth and eighteenth and nineteenth and twentieth joints fused ; fifth foot without inner ramus, the left 4-jointed, the right 3- jointed ending in a forceps or bristle. 1. Genus Candacia Dana. Candacia Dana, 1846, p. 184. Ifionyx Kroyer, 1848-49, p. 601. Candace Dana, 1849,\p. 279; 1852, p. 1109. Candace Lubbock, 1856, p. 29. Candace Claus, 1863, p. 189. Candace Streets, 1877, p. 139. Candace Brady, 1883, p. 66 Candace Thompson, 1888), p. 148. Candace Giesbrecht, 1892, pp. 67, 423, 729. Candace Wheeler, 1899, p. 177. Candacia Giesbrecht, 1898, p. 126. 2 Fourth and fifth thoracic segments fused; front of head rectangular, lateral angles of last thoracic segment pointed; abdomen with 3 segments, genital segment often asymmetrical. Anterior antennae 23- or 24-jointed, proximal segments thick- ened, anterior border toothed. Rami of posterior antennae short, outer ramus slender, second joint elongated, terminal enes very short. Basal of mandible large, rami short, blade with 2 teeth. Anterior maxilliped elongate, without lobes, dis- Vou.2] Esterly—Copepoda of the San Diego Region. 193 tal bristles strong, sickle-shaped; posterior maxilliped small and weak, second basal and inner ramus suppressed. First to fourth pairs of feet with 3-jointed outer rami, inner rami relatively small, 2-jointed; first basal with inner marginal bristle in second and third pairs. Outer border of outer ramus toothed, fifth pair stunted, 3-jointed on each side. 4 Last thoracic segment more often asymmetrical, the point on the right side noticeable for form, size, and color. Abdomen with 5 segments, genital segment often asymmetrical with out- growths on the right side. Right anterior antenna a grasping organ. Fifth foot on right side 3-jointed, on left side 4-jointed ; the right foot terminates in a forceps or bristle. KEY TO SPECIES. 1. Terminal bristle of outer ramus of third foot with outward bent point, at least as long as the distance between the distal spines of the outer border of ther joubt (to. 44.0). --le ae-- 2 1. This bristle much shorter than the designated portion of the outer loomsler’ (Gite, AIC c do eeaogo ouae one co cop bos au0ddd so euacaddS 2. Genital segment of female longer than broad........... C. pectinata 2. Genital segment of female broader than long, male not known.. Sain A ET IE ee aoc CAPE finer en Sec. ein lap ease ese eet ot aie C. bipinnata 3. The thick proximal portion of the anterior antennae is 7-jointed (Cie anata cieaste ets cle Sr oeaieaber Yexsheiei cin dnejele.e (= mallet # "equals 4 Ze PRES) MOKWMOM, G-jOTNGER sects cer ciel ire 2-6 2 wits wee ene Seg a= oe 5) 4. Terminal joint of fifth foot of female without bristles on inner border; joint of grasping antenna proximal to geniculation with deep teeth on anterior border. Fifth foot of male (fig. AGC tay is orators ioe as er meeieg as cata atels wloks olen he leenshet “Asie = C. curta 4. Terminal joint of fifth foot of female with three bristles, apical teeth slender and sharp. Teeth on grasping antenna fine; genital segment with flat outgrowth on right side; (fig. 47b).. An aL TER OP ee eG. Groth CAC RT RS CRC CE ONC On CR ROTOR LO C. aethiopica 1. Candacia pectinata Brady. Candace pectinata Brady, 1878, p. 49; 1883, p. 67, pl. 30, figs. 1-15. Candace pectinata Giesbrecht, 1892, pp. 424, 439, pl. 4, fig. 3; pl. 91, figs. 2, 12; pl. 22, figs. 9, 17, 18, 31, 43-46; pl. 39, figs. 1, 21, 22, 24, 25. Candacia pectinata Giesbrecht, 1898, p. 128. Candace pectinata Wheeler, 1899, p. 177, fig. 15. Genital and following segment in female asymmetrical, the latter protruding posteriorly; last thoracic segment in the male 194 University of California Publications. [ ZooLocy asymmetrical. Anterior antennae with 23 joints, pectinate part of joints of grasping antenna deeply toothed, the segments on either side of the articulation suppressed. Proximal hooked bristles of second basal of anterior maxilliped as thick and almost as long as the distal ones. Terminal joint of fifth foot of female long and claw-like (fig. 440), without inner marginal bristle; right fifth foot of male with forceps. Fig. 44.—Candacia pectinata. (a) Last thoracic segment and first and second segments of abdomen, male, X45. (b) Fifth foot of female X83. (c) Third joint of outer ramus of third foot of male X49. (d) Fourth foot of female X83. Coloration: Rather transparent, lateral prolongations of last thoracic segments, genital orifice, rami and bristles of feet and mouth parts, joints 18 and 19 of grasping antenna, usually a blackish brown. Length: Females average 2 mm., males 1.9 mm. Oceurrence: Rather common; both sexes are taken in sum- mer and winter. Se Vou.2] Esterly—Copepoda of the San Diego Region. 195 2. Candacia bipinnata Giesbrecht. Candace bipirnata Giesbrecht, 1889, p. 5; 1892, pp. 424, 439; pl. 22, fig. 20; pl. 39, figs. 27, 29. Candacia bipinnata Giesbrecht, 1898, p. 129. Fig. 45.—Candacia bipinnata. (a) Cephalothorax and genital segment of female, dorsal, X31. (b) First eight joints of anterior antennae of female X83. (c) Last thoracic segment and abdomen of female X20. Q Like C. pectinata, but genital segment (fig. 45a) is broad and has a wing-like expansion on each side. § Unknown. Coloration: Much as in C. pectinata. Leneth: Female, 2.6 mm. Oceurrence: Taken usually with C. pectinata, but in fewer numbers. 196 University of California Publications. [ ZooLocy 83. Candacia curta Dana. Candace curta Dana, 1849, p. 279; 1852, p. 1116; 1855, pl. 78, figs. 6 a-d. Candace curta Giesbrecht, 1892, pp. 424, 439, pl. 21, fig. 15; pl. 22, figs. 12, 24; pl. 39, figs. 8-10, 12. Candacia curta Giesbrecht, 1898, p. 129. c Fig. 46.—Candacia curta. Male. (a) Last thoracic segment, and genital segment, lateral, X83. (b) Same, dorsal, X31. (c) Fifth foot X83. Right foot at right of figure. Allied to C. pectinata, but right side of genital segment in female has a ventral projection; fifth foot of female with two heavy teeth on the end, and one on the inner border. Proximal joint of inner ramus of first foot with but two inner marginal bristles. Coloration: As in preceding species, with very slight varia- tions. Length: Male, 1.5 mm. Giesbrecht gives 2.4-2.65 mm. Occurrence: San Diego, Jan. 4, 1904, one male. 4. Candacia aethiopica Dana. Candace ethiopica Dana, 1848, p. 23. Candace melanopus Claus, 1863, p. 191, pl. 33. Candace ethiopica Giesbrecht, 1892, pp. 424, 439, pl. 4, fig. 13, pl. 21, figs. 1, 9; pl. 22, figs. 1, 6, 13, 14, 32, 40-42; pl. 39, HOS ual als: Candacia aethiopica Giesbrecht, 1898, p. 128. Vou.2] Esterly—Copepoda of the San Diego Region. 197 Genital segment of female slightly asymmetrical, with a process on the left side; last thoracic segment of male asymmet- rical. Anterior antennae 23-jointed, denticulation of the geni- culating joints of grasping antenna fine, joints proximal and distal to the geniculation long and slender. Proximal hooked bristle of the second basal of the anterior maxilliped as thick and almost as long as the distal (fig. 472). Terminal joint of fifth foot of female with one tooth on the outer margin, 3 apical teeth, and 3 bristles on the inner border; right fifth foot of male with a forceps (fig. 47g). Fig. 47.—Candacia aethiopica. (a) Female, dorsal, 165. (b) Last thoracic segment, and genital segment of male, dorsal, X60. (c) Abdomen of female, lateral, X37., (d) Second and third joints of outer ramus of third foot, female, 60. (e) Fifth foot of female X130. (f) Left fifth foot of male x130. (g) Forceps of right fifth foot of male X130. (h) Anterior maxilliped of female 130. Coloration: Dorsal surface of cephalothorax, excepting an- terior portion of head and the last thoracic segment, black brown, 198 University of California Publications. [ ZooLocy distinguishing the species at once. Appendages colored about as in the other forms. Length: Female 2.9 mm, male 2-2.5 mm. Occurrence: Several males and females were taken October 20, 1904. Fam. PONTELLIDAE. Pontellidae Giesbrecht, 1892, p. 68; 1898, p. 131. 2 Head and thorax distinct, fourth thoracic segment usually fused with fifth. Rostrum forked, usually ending in two very strong prongs; rarely absent. Eyes large, sometimes with one or two pairs of cuticular lenses and one unpaired lens. Anterior antennae 16- to 24-jointed, the two terminal joints always fused. Second basal and first joint of inner ramus fused, terminal joints of outer ramus shortened. Mandible on the whole as in the Centropagidae. First basal of maxilla large, -second basal and rami relatively small. Anterior maxillipeds as in the Centro- pagidae, long, hooked bristles on distal portion and commonly on the proximal. First basal of posterior maxillipeds large with long bristles on lobed inner border, second basal and inner ramus relatively small. Inner ramus of four anterior pairs of feet or second to fourth, 2-jointed; fifth pair rudimentary, outer ramus 1-jointed (rarely 2-jointed), inner ramus 1-jointed or lacking. 3 Distinct from female in form of abdomen, anterior antennae and fifth pair of feet, at times also in form of eyes, rostrum and last thoracic segment. Genital orifice on left side, erasping antenna on right, middle joints much or slightly broadened; joints 19 and 21 and 22 to 25 fused. Fifth pair of feet rarely with rudiment of inner ramus; forceps of right foot incomplete or very powerful. Sub-fam. PONTELLINAE. Pontellina Giesbrecht, 1892, p. 68. 2 Cephalothorax with five or six segments; last thoracic seg- ment ends in one (seldom two) sharp points on each side and is at times asymmetrical. Rostrum ends in two strong chitinous prongs or in two filaments. One pair of cuticular lenses is occasionally found on the dorsal side, seldom two pairs, ventral Vou.2] EKsterly—Copepoda of the San Diego Region. 199 eye strongly protruding (fig. 48c). Abdomen with from one to three segments, never symmetrical. Anterior antennae 16- to 24-jointed, at least two terminal joints fused, usually also a number of proximal joints. Posterior antennae with reduced number of terminal joints in outer ramus which is often more slender and thinner than the inner ramus. Mandible as a whole as in Centropages, blade with at least five teeth. Maxilla with relatively large proximal basal, second inner marginal lobe large, second basal, rami and first outer marginal lobe accord- ingly relatively smaller. Anterior maxilliped as in Centropages, with very strong hooked bristles; posterior maxilliped short, first basal with indented or folded inner margin, set with long, strong bristles; inner ramus 3- to 5-jointed, bristles short. Outer ramus of four anterior feet 3-jointed, inner ramus of second to fourth pairs or of all 2-jointed. Basal of fifth pair 2-jointed, inner and outer rami usually 1-jointed; outer ramus seldom 2-jointed. & Sexual peculiarities in form of body, more often in eyes, anterior antennae and fifth foot. Last thoracic segment as a rule asymmetrical, right posterior angle more strongly de- veloped; abdomen with 5 segments, in cases with asymmetrical processes on right side. Right anterior antenna with broadened middie joints; beyond the geniculation either the nineteenth and twenty-first joints only are fused (besides the twenty-fourth and twenty-fifth) or also the twenty-second and twenty-third: fifth foot without inner ramus (perhaps a rudiment on left foot), four jointed on each side, the right foot with forceps. 1. Genus Labidocera Lubbock. Pontella (part) Dana, 1846, p. 184; 1848, p. 26; 1849, p. 280. Pontellina (in part) Dana, 1852, p. 1135. Labidocera (sub-genus) Lubbock, 1853a, p. 25; 18530, p. 202. Pontella Claus, 1863, p. 207; 1893, p. 233. Pontella Brady, 1878, p. 73; 1883, p. 87. Pontella Thompson, 1887, p. 34. Labidocera Giesbrecht, 1889, p. 7; 1892, pp. 70, 444, 746; 1897, p- 254; 1898, p. 132. Labidocera T. Scott, 1893, p. 82. Labidocera Wheeler, 1899, p. 178. 200 University of California Publications. [ ZooLocy Head usually without hooks on side; one pair of dorsal eye lenses, larger in the male than in the female; rostral hooks strongly chitinized. Cephalothorax of 5 segments, ending in points laterally, more strongly developed on right side in the male Abdomen of female with 2 or 3 segments, of male with 5, sometimes asymmetrical in the female. Anterior antennae of female 23-jointed; terminal section of grasping antenna (the right) of male 4-jointed. Mandibular blade with 5-7 hooked, pointed teeth. Second basal of maxilla bent toward outside, about twice as long as the second lobe of the inner margin; anterior maxilliped stunted and provided with strong hooked bristles especially on the distal half; posterior maxilliped with 4-jointed inner ramus. Inner ramus of swimming feet 2-jointed, outer ramus 3-jointed; fifth foot of female on each side with 2-jointed basal portion, rami 1-jointed, though the inner ramus may be rudimentary. Fifth foot of male 4-jointed on each side, right without inner ramus, left at times with rudi mentary inner ramus. The two terminal joints of the right foot form a powerful forceps. 1. Labidocera trispinosa n.sp. 2 Cephalothorax symmetrical, evenly rounded in front; erest, and hooks on side of head absent; rostrum bifid, very long; last thoracic segment produced on each side into sharp points (cf. fig. 48a. left side). Abdomen with 3 segments, genital longer than the last two, asymmetrical, with a prominent blunt wing-like process on the right side (fig. 48d); middle abdominal segment with a knob-like projection on the left in front. Furea symmetrical, about 3 times as long as_ broad. Anterior antennae extending back to the posterior border of fourth thoracic segment. Fifth pair of feet symmetrical (fig. 489), outer ramus ending in two teeth, the inner one twice as long as the outer; inner ramus about one-half as long as the outer, articulating with basal; outer ramus longer than the first and second basals together. & Eye lenses larger, and more nearly contiguous (fig. 48a). Last thoracic segment on right side with a long slender spine, a eee Vou.2] Hsterly—Copepoda of the San Diego Region. 201 curving dorsally, (fig. 48), a shorter straight dorsal spine, and a very short spine directed toward median line. On the left side the thorax is about as in the female. Abdomen with five segments, genital segment in some cases slightly asymmetrical ; orifice on right side; middle segment as long as the first two, and longer than the last two. Anterior antennae reaching to base of furca; joints 16 and 17 about of equal length; teeth on joint 18 directed toward distal end of antenna, those on the next joint larger and straight (fig. 481). Fifth foot, fig. 48e, f. A 18, 19.21. Fig. 48.—Labidocera trispinosa, n. sp. (a) Male, dorsal, X18. (b) Lat- eral portion of last thoracic segment from right side X60. (c) Head of male, lateral, x60. (d) Abdomen of female, dorsal, X60. (e) Left fifth foot of male. (f) Right fifth foot of male. (g) Fifth foot of female. (h) Joints 17, 18, 19 to 21, 22, of grasping antenna of male, 160. 202 University of California Publications. [ ZOOLOGY This species is distinct from any of the seventeen described species of Labidocera in the form of the last thoracic segment of the male, and the genital segment of the female. It approaches L. lubbocki Giesbrecht and L. brunescens Giesbrecht, more closely than any others, but differs distinctly from them in the above-named features as well as in the structure of the fifth feet of the sexes, ete. Coloration: Rather transparent, intestinal contents light green, thorax and abdomen yellowish with green tinge in places. Length: Female, 1.6 mm. Male, 1.7-2.2 mm. Occurrence: May 24, 1904, one female. June 16, 1904, four males. five females. Sub-fam. PARAPONTELLINAE. Parapontellina Giesbrecht, 1892, p. 73. Cephalothorax usually with five, seldom with six segments, rostral filaments slender or lacking, last thoracic segment with rounded or pointed sides. Eyes without dorsal chitin lenses. 2 Abdomen usually with three, seldom two segments, at times asymmetrical. Anterior antennae 17- to 19-jointed; sev- eral proximal joints fused in addition to the terminal points. Second basal of mandible elongate, usually eylindrical, blade narrow with from five to seven teeth. Maxilla elongate, lobes slightly protruding and not articulating; rami more often stunted; the entire second basal may be absent. Distal hooked bristles of anterior maxillipeds long and strong, seldom so on proximal portion of the appendage. Posterior maxilliped as in the Pontellinae but inner ramus is only 1- or 2-jointed. Outer ramus of anterior four pairs of feet 3-jointed, inner ramus of second to fourth or of all 2-jointed. Fifth pair stunted, basal 1- or 2-jointed, outer ramus 1-jointed usually claw-like and with- out inner ramus; not always symmetrical. & Sexual peculiarities in form of body, anterior antennae and fifth foot. Last thoracic segment and abdomen at times asymmetrical as in many Pontellinae; abdomen with five seg- ments. The right grasping antenna simulated in some respects in the left; the right antenna with but little broadened middle joints and differences in the segmentation of the joints in the eS ae eg A i NE A ma ee Vou.2] Esterly—Copepoda of the San Diego Region. 203 proximal portion; joints nineteen and twenty-one and twenty- two and twenty-five fused. Right fifth foot 3- to 4-jointed, forceps not complete, the left 3-jointed, seldom with rudiment of Inner ramus. 1. Genus Acartia Dana. Acartia Dana, 1846, p. 183; 1852, p. 118. Dias Brady, 1883, p. 72. Dias Lilljeborg, 1853. Dias Claus, 1863, p. 191. Dias Brady, 1883, p. 72. Acartia Thompson, 1888a, p. 149; 1888b, p. 141. Acartia Giesbrecht, 1892, pp. 75, 506, 721; 1898, p. 150. Acartia Dahl, 1894ce, p. 13. Acartia Wheeler, 1899, p. 182. Fifth thoracic segment and abdomen of male symmetrical ; latter with shortened anal segment. Antennae of female with seventeen segments, of the same diameter throughout the length; grasping antenna of male with very slightly thickened middle joints. Outer ramus of posterior antenna much shorter than the inner; second joint of inner elongated, first joint with nine bristles on the inner border. Outer ramus of mandible articu- lates in the middle of the margin of the second basal. First outer marginal lobe of maxilla with long bristles, outer ramus rudimentary, its place supplied by two bristles. Proximal lobes of anterior maxillipeds well developed, with long bristles; pos- terior maxilliped with four joints. Inner ramus of first pair of feet with two joints, fifth pair of female without inner ramus, with long outer terminal bristle on second basal. ° Cephalothorax with five segments, last two fused.- Pos- terior antenna very slender, second basal joint fused with prox- imal joint of inner ramus, mandible with weak blade, which has seven teeth. Posterior maxilliped without outer marginal bristies, and with three inner marginals on third joimt. Outer ramus of first to fourth pairs of feet 3-jointed, first basal without bristles, second with rather long outer marginal bristle in fourth pair. The very much stunted fifth pair (fig. 49d) consists on each side of two or three joints; the end joint (outer ramus) is a thick stylet-shaped bristle, and on the outer border of the second basal is a slender feathered bristle. 204 University of California Publications. [ ZooLoGy & Sexual differences in form of body, anterior antennae and fifth pair of feet. Abdomen with five segments, genital orifice on left side; the fourth segment and furea shortened. The right anterior antenna is a grasping organ and joints 19-21 and 22-25 are fused. The fifth pair of feet (fig. 49c) consists of a common middle part and a right foot of four joints, a left of three, each uniramous. The right especially has the form of a claw, which, because of a process on the joint preceding the terminal, becomes an incomplete forceps. 1. Acartia tonsa Dana. Acartia tonsa Dana, 1848, p. 26. Acartia tonsa Giesbrecht, 1892, pp. 508, 522; pl. 30, figs. 7, 24, 34; pl. 43, figs. 6, 10; 1898, p. 154. a) ; Fig. 49.—Acartia tonsu. (a) Female, dorsal, X45. (b) Abdomen of male, dorsal, X55. (c) Fifth foot of male X138. Dz., right foot. (d) Fifth foot of female 138. ex. b Rostral filaments present, last thoracic segment rounded laterally ; abdomen of male (fig. 495) with spines on the second segment; anal segment with lateral hairs. Anterior antennae of female without thorns, not reaching to the posterior border of 6 ee, > ~ coe clkckoooooeoosc vetoes aqecog6 *Sapphirina 206 University of California Publications. | ZooLocy 5. Abdomen with two segments, not broadened................. Corina 6. Eye lenses separated by at least their diameter; the last two oat | 13. 13. 14. 14. 15. thoracic segments without lateral sharp prolongations..... Copilia? . Eye lenses placed close together, last two segments of the anterior portion of the body prolonged into lateral pointed Processes’ (fis: sO, GAD) re arccoayshn wicks mumieltic oreteleters ora re reons *Corycaeus . Entire body much flattened, furca very long and stylet-like. .Copiliag . Body of various shapes but more rounded; if at times depressed, Mever leatelvce) 5.1 "ire ls ratio ete rae le a ee a ee 8 . Outer ramus of first foot 1-jointed, postero-lateral angles of fourth segment of body prolonged into processes (fig. 54a).... a eR a AP PUN A rie nur hrhnd RU RN ate Sets She *Clytemnestra » Outer ramusof first. foot 2 to'3-joImbed « % joes foe wim aya whet atau os 4 0M 9 . Outer ramus of posterior antenna 1-jointed; furca very short, each ramus with a very long bristle twice as long as the body at least; rami of furea and the two bristles fused in the median line; remaining furcal bristles stunted................. Aegisthus . . Outer ramus of posterior antenna 3-jointed; furea short, rami separate (fig. 52c), each with a long bristle, at least as long as the body, and at least twice as long as the other bristles GLUES eB) eh a as at ayege sachets erste tere aftetet teat atarones ratelaaee *Microsetella . Outer ramus of posterior antenna lacking; furea longer than LONG, AMT HOV ALAO 212 sects ojsurhs tale laipo re Sis ete oo eeim eae steerete eters 10 . Anterior and posterior maxillipeds alike in structure, both with LORE SPINOUS iPLISGLABn cre ot aler ‘ Price; 125 | ERRATA. Page 240, line 21, for Pl. II read Pl. XI. Page 250, line 11, for Cellariide read Cellulariide. Page 274, last line, first word, for possesed read possessed. Page 285, line 5 from bottom of page, read 84 for 93. UNIVERSITY OF CALIFORNIA PUBLICATIONS ZOOLOGY Vol. 2, No. 5, pp. 235-322, Pls. 4-16 December 9, 1905 CONTRIBUTIONS FROM THE LABORATORY OF THE MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO. Ve NON-INCRUSTING CHILOSTOMATOUS BRYOZOA OF THE WEST COAST OF NORTH AMERICA. BY ALICE ROBERTSON. The bryozoa of the west coast of North America constitute a fauna practically unknown to science. From time to time during the last half century some scattering work has been done, the earliest being by Trask (757), who deseribed and figured a number of species from San Francisco Bay. Later Gabb and Horn (’62), in connection with their work on the Geological Survey of California, identified a number of species of bryozoa, both recent and fossil; and Hincks (’82 and ’84) reported on a large collection from Queen Charlotte Islands. Still later, the present writer (’00) reported on the bryozoa of the Harriman Alaska Expedition, having previously published on the Ento- procts of San Francisco Bay (’99). Besides the work of these investigators, mention should be made of that of Conrad (755) and of Fewkes (’89), both of whom added to our knowledge of the bryozoa of this coast. The following pages constitute the first of a contemplated series of papers on the bryozoa of the Pacific coast of North America, and include about half of the sub-order Chilostomata, comprising, in the main, those Chilostomes which grow as free, flexible colonies. The inclusion of the adherent genera Aetea 236 — University of California Publications. | ZooLocy and Eucratea is somewhat arbitrary since they are more or less inerusting. In reality the forms treated comprise the first seven families of the Chilostomata as given by Hincks, and these with the exception of the two genera mentioned above are erect, flexi- ble and non-incrusting. Even these two genera send off erect free branches, and are thus partially non-incrusting. The bryozoa here identified comprise only a small part of the ma- terial on hand, the greater portion of which was collected at San Pedro and San Diego in 1901, 1902, and 1903 on dredging ex- peditions conducted by the Zoological Department of the Uni- versity of California; another portion was dredged in Puget Sound during the summers of 1903 and 1904 by the Department of Zoology of the University of Washington, and kindly placed at my disposal by Professor Trevor Kincaid. In the endeavor to make this work as useful as possible to the genera! student as well as to the expert, original figures are given of all the species mentioned. These include a habit sketch which is a photograph wherever possible, and one or more other figures giving details and variations. It is thought best even though a species is cosmopolitan in its range, and even though it has already been sufficiently well represented for diagnostic purposes, to refigure it here; and this for two reasons: first, bryozoan literature, consisting as it does, except in two or three instances, of detached papers scattered through scientifie jour- nals is not readily accessible to the general student. Second, the number of cosmopolitan species, or of species easily identi- fied, is surprisingly small, while the number new to science is correspondingly large; hence as far as illustration at least is concerned, it is desired to make this work as complete as pos- sible for the forms found on the Pacifie Coast. The diagnoses are somewhat lengthy descriptions, with references to figures wherever clearness seemed to demand it. Further, in order to assist in the identification of species, simple keys have been de- vised for the families, and where more than one genus or species: occur under a family, separate keys are given for each. The keys for species cover only those treated in this paper and are intended only for the bryozoa of this coast. Since Hincks’ (’80) VoL. 2] Robertson.—Non-Incrusting Bryozoa. 237 monograph upon the British Polyzoa is the most complete and exhaustive of any recent work, and its method is generally fol- lowed by other workers, it is here adopted as the basis of classi- fication, although his conclusions are not always accepted. Free use is made of the diagnostic definitions given by that author, especially for families and genera, supplementing and amending where necessary for greater accuracy, or in order to include forms peculiar to the region. Synonymy is based upon that of Hineks, further supplemented by Miss Jelly’s invaluable Synonymie Catalogue (’&9). No attempt has been made at re- vision of the classification. It were useless to undertake such a task except after careful study and comparison of all large collections. It is needless here to enter into an exhaustive discussion of the bryozoa as a class. Such can be found in any of the more recent treatises on zoology, and the reader desirous of obtaining a full, clear, and delightfully written article on the group, is referred especially to that by Dr. Sidney F. Harmer (’96). For the sake of clearness and convenient reference, however, a few definitions of terms used in this paper are given herewith. Bryozoa and Polyzoa are synonymous terms. These are colonial animals, and the technical term used to designate the colony as a whole is zoarium. An illustration of a bryozoan colony or zoarium may be found in any of the habit sketches, especially the photographs (Pl. XVI). The units of which a zoarium is composed consist of the zowcia and their contained polypides. A zocecium is a chamber or sac, in which the poly- pide, consisting of a digestive canal and a circlet of tentacles, is lodged. It may be ealeareous and opaque, or semi-calecareous or chitinous and transparent. The contents of a zowcium whose walls are transparent are easily made out. Below is given a figure of a zocecium of Beania mirabilis with its contained poly- pide in a state of retraction folded within it. The zoccium (z@.)is seen to be a sae or bag whose front or ventral face is bordered with a number of spinous processes, some erect (é. sp.), others curved (c. sp.). Within the zoccium is the polypide (pd.) consisting of a bent tube, the intestinal canal, having 238 University of California Publications. [ ZooLocy a cirelet of tentacles (tent.) around one extremity. Various regions of the tubular portion have specific names. In the middle of the membranous floor from which the tentacles arise is the mouth (m.). This opens into a short tube known Fig. 1—Beania mirabilis. A zowcium and its inclosed polypide shown in profile view. c. sp. crossed spines; car. val. cardiac valve; é. sp. erect spine; i. intestine; m. mouth; @s. wsophagus; or. orifice; par. mus. parietal muscle fibres; pd. polypide; ph. pharynx; py. val. pyloric valve; r. f. root fibre; re. mus. retractor muscle; st. stomach; tent. tentacles; tent. sh. tentacle-sheath; 2a. zoccium. as the pharynx (ph.) which is really a portion of the cesophagus (ws.); this in turn opens by a narrow valve, the cardiac valve (car. val.) into a stomach (st.). The stomach is a bag of a yellow or brown color due to the gland cells in its walls; it opens into the intestine (7.), by the so-called pyloric valve (py. val.). Visible above the tentacles when the polypide is retracted is a delicate membrane, the tentacle-sheath (tent. sh.). Near the distal end where the tentacle-sheath approaches the orifice (or.), a few muscle fibres may be seen on each side of the sheath ex- tending to the walls of the zowcium. These are part of the parieto-vaginal muscles which assist in retracting the tentacular VoL. 2] Robertson.—Non-Incrusting Bryozoa. 239 sheath (par. mus.). Likewise, extending from the base of the tentacles to the basal wall of the zocecium another band of re- tractor muscles is visible whose contraction draws the polypide within the zoccium (re. mus.). Fig. 2 represents a zocecium of Fig. 2—Bowerbankia imbricata. A zoccium and inclosed polypide. a. anus; giz. gizzard; int. intestine; m. mouth; ws. esophagus; py. val. pyloric valve; st. stomach; tent. tentacle. Bowerbankia imbricata, in which the polypide is expanded. Here the cesophagus (@s.) is stretched out to its fullest extent, the mouth (m.) being at the upper margin, and the tentacles outside the zoecium. This polypide possesses a region between the wsophagus and stomach known as the gizzard lined with large cells (giz.). The intestine (int.) is much elongated, the anus (a.) reaching almost to the summit of the body wall. The tentacles are commonly arranged in the form of a bell, but have 240 University of California Publications. [ ZOOLOGY the power of independent motion as shown in this figure drawn from a living specimen. Many zoaria grow erect and free, and if caleareous, their branches frequently possess flexible joints at definite points along the length (PI. V, fig. 14, j.). The zocia included between these joints form an internode, the number of zocecia in an internode being rather definite for any given species. In Menipea ternata e.g., there are, as a rule, three zocecia in an internode (fig. 14). From the lower zocecia smal] fibres are given off known as rootlets, because they serve to anchor the colony (fig. 14, 7.). Zocecia, especially if calcareous, often possess appendages of various kinds which serve as diag- nostic marks. There are also certain well marked regions to which special names are applied: Aperture—the chitinous front wall of the zoecium. This may oceupy a part of the front wall only as in Pl. IV, fig. 2, ap.; or almost the whole of it as in Pl. X, fig. 50, ap.). Area—the caleareous wall inclosing the whole or part of the aperture. (Pl. XV, fig. 88, a.). Avicularium—an appendage of the zowcium more or less resembling a bird’s head. This may be sessile as in Pl. V, fig. 14, av., or pedunculated as in Pl. II, fig. 60, av. Epistome—a ciliated lobe which overhangs the mouth, and is present only in the Phylactolemata or fresh-water bryozoa. Internode—the zoecia included between the flexible joints of a branch or stem (PI. V, fig. 14). Introvert—the thin cuticle at the anterior end of the polypide which may be retracted into the interior of the zowcium. Joint, or articulation—a non-calcified portion of the wall of the zoe- cium permitting more or less flexibility in the stem or branch (EE Vi, fie. mla9>). Lophophore—the membranous floor or rim surrounding the mouth of the polypide from which the tentacles spring. Oecia or Ovicell—synonymous terms for the chamber above the zoe- cium in which the embryo develops (Pl. V, fig. 14, @.). Operculum—the chitinous lip by which the orifice of the zowcium is closed (Pl. IV, fig. 2, and Pl. XIV, fig. 86, op.). When the aperture is entirely membranous, the operculum is inconspicuous. (RIES fio 50) Orifice—the opening at the summit of the aperture through which the polypide emerges. (Text fig. 1, or.). Scutum—a modified spine overhanging the aperture (fig. 14, sc.), often being large and flabellate (Pl. VI, fig. 19, sc.). Stolon—a creeping tubular stem from which the individuals of a colony grow. It is not found in the Chilostomata, but is characteristic of many of the Ctenostomata. Vor. 2] Robertson.—Non-Incrusting Bryozoa. 241 Spine—a jointed or unjointed process found on the margin of the aperture (fig. 1 of the text, sp.). Tentacle-sheath—the delicate membrane of the introvert which incloses the tentacles when the polypide is retracted. (Text fig. 1, tent sh.) Vibraculum—a chitinous seta of varying length, depending on the species, extending from a chamber on the dorsal side of the zocecium (PI. IX, figs. 41, 45, v.). S Vibracular chamber—a chamber on the dorsal side of the zocwcium in some of the Cellulariide, resembling an avicularium, from which extend a hairlike process called the vibraculum, and the rootlet (Pl. IX, figs. 42, 46, v. ch. and v.). For purposes of orientation, it must be explained that the front or ventral side of a zocecium is that on which the aperture with its operculum occurs; the side opposite is the dorsal. The top of a zocecium, and the ends of the growing tips of branches or colony are referred to as the distal or anterior end of zocecium, or extremity of branch or colony; the lower part or end nearest the root or point of origin, as the proximal or posterior extremity. Thirty-four species and one subspecies belonging to thirteen genera are here recognized. Of the genera, Stirparia has as yet been reported only from Australia; Synnotum only from the Adriatic; and Corynoporella only from Greenland. Of the whole list, thirteen species and one subspecies are new, while twenty species and one subspecies are restricted to the Pacific Coast.. The ovicells of Aetea anguina are here described and figured for the first time, as are the avicularia and ovicells of Stirparia, and the ovicells of Corynoporella. LIST OF SPECIES TREATED. Aetea anguina Linneus. Bugula pacifica Robertson. Aetea truncata Landsborough. Bugula flabellata Thompson. Eucratea chelata Linneus. Bugula pugeti sp. nov. Gemellaria loricata Linnveus. Bugula curvirostrata sp. nov. Menipea ternata Ellis and Solander. Bugula longirostrata sp. nov. Menipea gracilis Busk. Bugula laxa sp. nov. Menipea occidentalis Trask. Beania mirabilis Johnston. Menipea occidentalis catilinensis Beania longispinosa sp. nov. subsp. nov. Stirparia ciliata sp. nov. Menipea erecta Robertson. Stirparia occidentalis sp. nov. Menipea pribilofi sp. nov. Stirparia californica sp. nov. Scrupocellaria californica Trask. Corynoporella spinosa sp. nov. Scrupocellaria varians Hinecks. Synnotum aviculare Pieper. Scrupocellaria diegensis sp. nov. Cellaria borealis Busk. Caberea ellisi Fleming. Cellaria mandibulata Hincks., Bugula neritina Linneus. Cellaria diffusa sp. nov. Bugula murrayana Johnston. Tlustra lichenoides Robertson. Bugula californica sp. nov. Flustra membranacea-truncata Smitt. 242 University of California Publications. [ ZooLocy PHoyLtuM Mo.uuuscoirpa Milne-Edwards. Class BRYOZOA Ehrenberg. Sub-Class Ectoprocta Nitsche. Colonial bryozoa with anal orifice outside the lophophore; a well developed introvert, and a spacious eclome. Order GYMNOLASMATA Allman. Polypide destitute of an epistome; lophophore circular. Sub-Order I. CuHmostomata Busk. Gymnolemata with caleareous or chitinous zocecia whose orifice is closed by a movable chitinous lip or ‘‘operculum’’; ova usually matured in globular omcia situated above the orifice of some of the zowcia; vibracula or avicularia, or both, fre- quently present. Of the fifteen or more families included by Hincks in the Chilostomata, all but the first seven, viz., the Membraniporide, Microporide, Cribrilinide, Microporellide, Porinida, Myrio- zoidw, Escharide, are omitted here. These are incrusting, or if erect, are for the most part non-flexible in habit of growth. Below is given a key to the first seven families of the Chilostomes which are, for the most part, erect and flexible in habit of growth. KEY TO THE FIRST SEVEN FAMILIES OF THE CHILOSTOMATA. 1, Golemy \eree pings (£2 a. Faye bc eee ee ee ee ee ee oe 2 i Colonyaerect: =e er er or Pera ees eee! Se 3 2. Colony sending up erect branches from a pane just below the 2M O =) al AF < eee RoR ES ee SO oe a ee ent Eucratiide 2. Colony not sending up erect branches ee Be ne se eee eee Aeteidz 3. Colony articulated: 3 j. 6 ea ee er ees 4 3s. Colony mot ventiieulate diye: 05 ee ie oe es ene ee ae eee 5 4. Zoecia multiserial, arranged round an imaginary axis....... Cellariide A$ ZO CCL A OVS CY NRL ace eee eee es ee ee Cellulariide Shey Aclenoh Mirna dsweeegeoh lovee stor lop Kc eee: ee ee 6 5.) AOWGIa NO’ Wack GO: eC kre oe eee eae 7 6.: Zoweia, with: aviculariay 3-23 ee eee Notamiide 6:5 ZO C2 Glah swat hob you vil Gel ret ye eee Eucratiide 7. Zoecia biserial, avicularia pedunculated —.............22...... Bicellariide 7. Zowcia multiserial, avicularia sessile —........2.2 Flustride VoL. 2] Robertson.—Non-Incrusting Bryozoa. 243 Aeteidae Hincks. Zoarium composed of creeping branches more or less ad- herent to the substratum, often growing in free tufts adherent only part of their length. Zowcia uniserial, arising from each other in a tubular prolongation of greater or less length. Aperture terminal, orifice at its summit. The definitions of family Aeteide, of genus Aetea, and of species Aetea anguina are here much changed from those given by Hincks, whose diagnoses are based upon the erroneous notion which that observer entertained concerning the true nature of the Aetean zoecium. The adherent portion of the zocecium he © regarded as a stolon which together with the circle of sete in which the tentacle-sheath terminates (Smitt, 67) seemed, he thought, to relate Aetea to the Ctenostomes, and to place it in the position of a form transitional between these and the Chilos- tomes. Whatever may be the relationship between these sub- orders, Aetea cannot be said to reveal it, since as will be shown in the discussion under the species A. anguina, the adherent 99 ‘*stolonie portion’’ so-called, is not a stolon, but an important part of the zocecium. Aetea Lamouroux. Aetea Lamouroux, 1812. Anguinaria, Johnston, 1847. Aetea, Hincks, 1880. Aetea, Jullien, 1888. Zoarium adherent to the substratum. Each zow@ciuwm par- tially adherent, partially erect, the erect portion carrying at its distal extremity the membranous aperture with its operculum. KEY TO THE SPECIES OF AETEA. 1. Tubular portion of zoecium ringed, upper third spatulate, bent........ SEN 5 sos Sa eas Os SOT pa Me eR RD SB EE, ERS A. anguina 1. Tubular portion of zowcium not ringed, upper extremity not bent, Dovite exci cl Cla iY, UlTy Cal ie eee cere eee eeee eC ILE Leann es A. truncata 244 University of California Publications. [ZooLocy 1. Aetea anguina (Linneus) Lamouroux. Pl. IV. Figs. 1, 2, 3, and 4. Sertularia anguina Linn., 1758, ed. 10, p. 816. Cellularia anguina, Pallas, 1766, p. 78. Corallina anguiformis Ellis, 1767, Ger. ed., p. 50, Pl. XXII, figs. (as One DE Cellularia anguina, Ellis, 1767, p. 434, Pl. 19, fig. 10. Cellaria anguina, Ell. and Solander, 1786, p. 26. Aetea anguina, Lamouroux, 1812, Vol. III, p. 184. Anguinaria spatulata, Johnston, 1847, ed. II, p. 290, Pl. L, figs. iy. Be ; Aetea anguina, Busk, 1852, pt. I, Pl. XV, fig. 1. Aetea anguina forma spatulata, Smitt, 1867, p. 280, Pl. XVI, figs. 2, 4. a Aetea anguina, Hincks, 1880, p. 4, Pl. I, figs. 4, 5. Zoarium composed of creeping branches consisting of a single series of zoccia growing irregularly over a stem, frond, or other substratum (PI. IV, figs. 1, 2). Branches arising at right angles to the zowcia from each side of the creeping por- tion. Zoacia composed of a creeping posterior portion (fig. 2, ad.), and an anterior, erect, tubular portion (tu.). The pos- terior creeping portion of the zowcium narrow and tubular where it arises from the neighboring zocecium (ad.) gradually widening anteriorly or toward the growing point (gr.), then turning upward almost at a right angle and becoming erect (tu.), the remainder creeping on to give rise to a new zocecium. The erect portion is tubular, the lower two-thirds being minutely ringed, the upper one-third being somewhat inflated, and in many cases bent forward thus forming the so-called snake’s head (s. h.). At the base of the tubular portion and anterior to it, a septum (sep.) separates the zocecium from the one next following. The membranous aperture (mem. ap.) is situated on the ventral side of the widened upper third of the erect tube and at its upper edge is the movable lip or operculum (op.). The dorsal surface of the widened upper end of the tube is minutely granulated. The polypide is simple in structure, very minute, and when retracted is drawn downward into the hori- zontal, adherent portion of the zoccium (fig. 3, pd.). The tentacle-sheath terminates above in a circle of sete which are od VoL. 2] Robertson.—Non-Incrusting Bryozoa. 245 everted during the expansion of the polypide. Owcia globular, membranous, situated at the upper extremity on the ventral side of the zocecium just below the operculum (fig. 4, @.). As Smitt (’67), Waters (’79), and Jullien (’88) have already shown, there has been much misconception in regard to the zocecium of Actea. Busk (749) considered the erect, tubular portion alone to be the zowcium, and in this error was followed by Hineks (’80) who regards the erect tube as the zoccium, the horizontal portion as a stolon, which he compares with the stolon of the Ctenostomes. Jullien, apparently unaware of the observations of Smitt or Waters on this point, criticizes the statement of Hincks as to the stolonic nature of the adherent portion. This he considers the true zocecium, since into it he finds the polypide withdraws. itself on retraction and on its walls the retractor muscle fibers are inserted; while the upright tubular portion he regards a peristome. Smitt and Waters had previously shown, however, that the polypide inhabits the creeping portion, and that this is in no sense a stolon. Later, Waters (’96) has shown that in the species known as Aetea anguina forma recta, the ovary is situated in the creeping part, thus affording conclusive proof of the zocecial character of this so-ealled ‘‘stolon.’’ In the colonies of this species found on the California coast (figs. 1, 2), the creeping and erect portions are continuous. The polypide when expanded occupies the upright tube, but on retraction retreats into the creeping portion almost completely, only the tips of the tentacles and the long, delicate sheath with its muscles remaining in the tube (fig. 3). In this figure, part of the upright tube is represented as broken and the polypide is not in a state of complete contraction, but the insertion of the retractor muscles on the wall of the creeping part (mus.), and the presence of the ovary (ov.) on the adnate ventral wall are shown. From the evidence presented it seems to be estab- lished that the zocecium of Aetea consists of both the creeping and the erect portions and not merely of either one of these. In our specimens of Aetea the ocecia are abundant. As shown in fig. 4, each consists of a membranous bag situated on 246 University of California Publications. [ ZooLocy the ventral side of the zocecium below the operculum but exterior to the aperture, and contains an ovum in the early cleavage stage (@.). In all cases in which the ocecium is present the tubular part of the zocecium is distinctly curved, as shown in fig. 4, as if affording protection to the delicate ocecium and its contents. This is the first instance, so far as known, of the de- lineation of the occium of Aetea anguina; Waters has figured the owcium of the so-called variety recta in which it is on the dorsal side of the erect portion. The difference of location of the ocecia in these two forms is probably sufficient to separate them into distinct species. There is a possibility that the species found here is an un- described one peculiar to this coast, but the owcium of Aetea anguina not having been hitherto known, and no other distine- tions being apparent between our form and that found in for- eign localities, this identification must stand for the present. The distribution of Aetea anguina is world wide, and this fact adds to the probability that we have here the older well known form. This species is abundant at San Pedro and San Diego, grow- ing over kelp, hydroid stems, shells, and other bryozoa. 2. Aetea truncata (Landsborough) Busk. Pen tissi to, 6: Anguinaria truncata Landsborough, 1852, p. 228, Pl. XVI, figs. 57, 57*. Salpigia Hassall Coppin, 1848, p. 273, Pl. X, fig. 3. Aetea truncata, Busk, 1852, p. 31. Aetea truncata, Smitt, 1865, Pl. II, figs. 5-14; Pl. ILI, figs. 1-8. Aetea truncata, Smitt, 1867, 279 and 295, Pl. XVI, fig. 1. Aetea truncata, Hincks, 1880, p. 8, Pl. I, figs. 8-11; Pl. II, fig. 3. Zoarium growing irregularly over the substratum. Zowcia rather widely separated, the posterior creeping portion fre- quently lengthening into a long slender fiber (Pl. IV, fig. 5 ad.) ; the tubular erect portion varying in length (tw.), the membranous aperture (ap.) occupying on an average a little more than one-third of its length; slightly wider at the top VoL. 2] Robertson.—Non-Incrusting Bryozoa. 247 than at the point of union with the adherent portion; truncate, granular, no part of the zocecium annulated as is the erect portion of the zocecium of the preceding species. Oacia not known. No colonies of the branching form described by Hincks occur in our material, the simple form of Aetea truncata alone pre- vailing. The creeping portion of the zowcium is rather closely adherent to the substratum so that it is not easily removed. The polypide resembles that of Aetea anguina, being of simple struc- ture and when retracted is drawn almost completely into the ereeping part. This species was obtained in considerable quantity at La Jolla growing over the older stems of a kelp hold-fast. Eucratiidae Hincks. Zoarium forming slender, branching, phytoid tufts. Zowcia uniserial, or in two series placed back to back; expanding from the base upwards, with a terminal or subterminal and usually oblique aperture. Neither avicularian nor vibracular append- ages known. Ocacia globose. KEY TO GENERA AND SPECIES. IE MColomyaCRCe Pill Oyen ee i eee Eucratea chelata AEC OLOT AOE CI UN cna got eee tes ee ee dee ae an eee Gemellaria loricata Eucratea Lamouroux. Cellularia (part) Pallas, 1766. Cellaria (part), Ellis and Solander, 1786. Eucratea Lamouroux, 1812. Catenaria (part), d’Orbigny, 1850. Eucratea, Johnston, 1847. Scruparia, Busk, 1852. Eucratea, Hincks, 1880. Zoarium composed of creeping adherent branches and of erect free shoots. Both creeping and erect branches composed of zocecia arranged uniserially, the erect branches arising from the ventral wall of the zowcia. Zowcia prolonged into a tube of greater or less length. Aperture large. 248 University of California Publications. - |ZooLocy 3. Eucratea chelata (Linneus) Lamouroux. PL Vy figs: 7 859: Sertularia chelata Linn., 1758, ed. 10, p. 816. Cellularia chelata, Pallas, 1766, p. 77. Bull’s Horn Coralline Ellis, 1767, p. 42, Pl. XXII, figs. b, B. Cellaria chelata, Ellis and Solander, 1786, p. 25. Eucratea chelata, Johnston, 1847, ed. II, p. 288, fig. 64. Catenaria chelata, d’Orbigny, 1850, Vol. V, p. 43. Scruparia chelata, Busk, 1852, pt. I, p. 29, Pl. XVII, fig. 2. Eucratea chelata, Smitt, 1867, pp. 281 and 301, Pl. XVI, figs. 7-9. Eucratea chelata, Hincks, 1880, p. 11, Pl. ILI, figs. 9, 10. Zoarium ‘composed of ereeping branches consisting of a single series of zocecia growing irregularly over the substratum and adherent to it, and of erect branches (PI. V, figs. 7 and 8). Creeping branches (cr. br.) arising from the sides of the ad- herent zocecia; the erect, free branches (e. br.), from a point on the zocecial wall just below the aperture. Zoawcia narrowed pos- teriorly into a tube, widening gradually anteriorly; the polypide even in contraction remaining in the anterior portion of the zocecium. Aperture (ap.) oval, surrounded by a thin, raised, unarmed margin; operculum at its summit (op.). Oacia mitriform, somewhat pointed above, with a keel down the center, borne on an imperfectly developed zowcium arising just below the aperture (fig. 9, w.). Tentacle-sheath terminating above in a circle of setz which are everted during expansion of the poly- pide. The presence of sete both in Eucratea and Aetea is given only on the authority of others. Few of our specimens of Eucratca possessed ocecia, and none perfect ones. Figure 9 is taken from Hincks. Although not abundant at any one point, Hucratea chelata has been found at several localities on the coast of southern California. Gemellaria Savigny. Gemellaria Savigny, 1811. Notamia Farre, 1837. Gemellaria, Hineks, 1880. Zoarium erect, branching dichotomous, each branch given aff trom the sides of the zowcia close to their upper extremity. Vo. 2] Robertson.—Non-Incrusting Bryozoa. 249 Zoecia joined back to back, each pair arising from the anterior extremity of the preceding pair. Aperture large, sloping slightly upward. Oacia? 4. Gemellaria loricata (Linneus) Savigny. Pl. V, figs. 10, 11. Sertularia loricata Linn., 1758, ed. 10, p. 815. Cellularia loricata, Pallas, 1766, p. 64. Coat of Mail Coralline Ellis, 1767, p. 40, Pl. XXI, figs. b, B. Cellaria loricata, Ell. & Sol., 1786, p. 24. Notamia loriculata, Farre, 1837, p. 413, Pl. XX VII, figs. 6-9. Gemellaria loriculata, Savigny, 1811, Egypt. Polyp. Gemellaria loriculata, Van Beneden, 1844, p. 33, Pl. V, figs. 1-7. Gemellaria loricata, Johnston, 1847, ed. II, p. 293, Pl XLVI, figs. 12, 13. Gemellaria loriculata, d’Orbigny, 1850, p. 46. Gemellaria loricata, Busk, 1852, pt. I, p. 34, Pl. XLV, figs. 5, 6. Gemellaria loricata, Smitt, 1867, pp. 286 and 324, Pl. XVII, fig. 54. Gemellaria loricata, Hineks, 1880, p. 18, Pl. ITI, figs. 1-4. Zoarium forming densely bushy masses, 50 or 75 mm. in height, made up of numerous long, straight branches; branching dichotomous (Pl. V, fig. 10). Zoacia narrowed below, widen- ing above; aperture (fig. 11, ap.) oval, occupying about half the front, sloping somewhat toward the top, with a thin, raised, un- armed margin; operculum (op.) semicircular, at the anterior edge of the aperture. Colony attached by numerous rootlets. Main stem of colony thicker below. This species is found only on the northern shores. It is abundant at Orea, Prince Willams Sound, where it was taken from shore rocks at low tide; at Juneau, it was dredged at 20 fathoms. Hinecks reports it from Queen Charlotte Islands. Cellulariidae Busk. Escharide (part) Johnston, 1847. Cabereade Busk, 1852. Cellulariade Busk, 1852. Cellularie Smitt, 1867. Cellulartide, Hincks, 1880. Zoarium erect, branching dichotomous. Zowcia in two or more series, closely united in the same plane; avicularia and 250 University of California Publications. [ ZooLocy vibracula, or avicularia only, almost universally present, sessile. This is a well marked group all of whose members have a strong family resemblance. The walls of the zoccia are strongly calcified, being generally variously adorned with spines and sessile avicularia, and many with waving vibracula. In all the embryos mature in globose, hood-like owcia. This family is well represented in our fauna, there being at least six species of Menipea, four of which are new; and three of Scrupocellaria, all of which are peculiar to this region. Caberea is represented by but one species, the northern C. ellisi. KEY TO THE GENERA OF CELLARIID. Zocecia three or more in an internode without vibracula........ Menipea Zowcia many in an internode, vibracula present ....................------..-------- Vibracular chamber not covering dorsal surface of zoccium .......... ... ce he A To ac St a ec eg Re ee as CR Scrupocellaria Vibracular chamber directed obliquely and covering the dorsal sur- face of the zocecium ................. Baa Re bree «EES ed re Caberea De bo bo Menipea Lamouroux. Pl. V, fig. 16. Menipea Lamouroux, 1816. Cellularia, Johnston, 1847. Cellarina, Van Beneden, 1849. Emma, Busk, 1852. Menipea, Busk, 1852. Cellularia, Smitt, 1867. Menipea, Hincks, 1880. Zoacia oblong, widest above, attenuated below, often elon- gated downward; lateral avicularia generally present, sometimes wanting; frontal avicularia generally present on the zocecium at the bifureation of a branch. No vibracula. Ow@cia present; large, globose. Root fibers characteristically located, being found in two positions: first, on the front, or on the lateral wall of a zocecium just below or to one side of the aperture (fig. 16, 7. d. and r.) ; second, in a definite dorso-lateral chamber situated just above the lateral avicularium (7. ch. and u. r.). In certain mem- bers of this genus, different functions are performed by these differently located fibers. Those found to one side, or below the aperture invariably turn downward and serve to anchor the VoL, 2] Robertson.—Non-Incrusting Bryozoa. 251 colony (7.); those arising in the dorso-lateral chambers may assist In anchoring the colony when they occur on zoccia in the lower part of the colony; usually, however, they turn upward, and twining about the other portions of the colony, function as tendrils (uw. 7.). Waters (’96) calls attention to the diagnostic importance of the root fibres, their position being fairly constant so that it may be used as a specific character. Of the Menipeas of this region, M. ternata and M. gracilis have root chambers in identical posi- tions and root fibers performing identical functions. The other three species differ from these two and from each other in the number and shape of the root chambers and in the function of the root fibers arising from them. KEY TO THE SPECIES OF MENIPHA. ibe Zomcia commonly: threes am ameinGermoc eyes see eee eee 2 Te, PACs ON, waneet ay? Tira Ewa, TAKEN) eee ee Serer seee ne eeeeroreeees M. erecta Pee SPUMCS a tWiOmOlM GNC Cg arate ee er seer ee cect 2 ee ee een asevacan te eoee 3 PA STOMMES MIO ME mt ele tai OW O Ty GEC Cee sees ree oe eee ees nee 5 3. Scutum half way down on the margin of the aperture —__................. a 3. Scutum two-thirds of the way down on the margin of the aperture...... 6 AS CUE TITS 10 eas PO = ly eeee ee eae ee score ees eee M. ternata A Seutumelaraesy tabelabe 22s: anne OMA kes ee M. gracilis Oe MBS CUE ULI STAM @ eee ne crete 2k se ee Ne PEL Aes ani oned Meda USSF 6 DOSS G Ul UT aa CLUaVAliGl © Cl lapeseee ace een eee ee een et eee cr eee ee ee if 6. Scutum half way down on the margin of the aperture_....M. pribilofi 6. Scutum two-thirds of the way down on the margin of the aperture...... AS Pirie eae ne ae RR cena Ee Se BR er aes Na Se eoe ke M. occidentalis 7. Seutum two-thirds of the way down on the margin of the aperture...... BEI oe _--.------...---... MM. occidentalis subspecies catalinensis 5. Menipea ternata (Ellis and Solander) Busk. TEI We akegss WU, dG), eS als) WOR ABI NOLS sites, alyie Cellaria ternata Ell. and Sol., 1786, p. 30. Cellularia ternata, Johnston, 1847, p. 335, Pl. LIX, figs. 1, 2. Cellarina gracilis, Van Beneden, 1849, p. 67, Pl. X, figs. 1, 2. Menipea ternata, Busk, 1852, pt. I, p. 21, Pl. XX, figs. 3-5. Cellularia ternata, Smitt, 1867, p. 282, Pl. XVI, figs. 10-26. Menipea ternata, Hincks, 1880, p. 26, Pl. VI, figs. 1-4. Zoarium consisting of loosely spreading, straggling branches, ‘or sometimes of rather large dense masses, frequently 30 bo Ol bo University of California Publications. [ ZooLocy to 35 mm. high and from 60 to 80 mm. broad (Pl. V, figs. 12, 13). Branching dichotomous. Internodes consisting of three zocecia, as a rule, although there are many instances of five or seven zoccia to an internode. The longer internodes are generally terminal and bear the ovicells. Joints light colored, arising in distinct tubes (fig. 14, j.). Zoacia elongated, narrowed below; aperture oval (ap.), occupying less than half the front, with two or three spines, one on the inner margin just above the operculum, and one or two on the upper, outer margin; zoccium at bifureation of branch having sometimes one, sometimes two rather long spines at its summit. Towards extremity of branches the spines increase in length, often very considerably. Scutwm varying in shape from a mere spine to a somewhat broad flabel- late process (sc.). Lateral avicularia (av.) large, and promi- nent, generally developed on all the zocecia of an internode except the one at the bifurcation of a branch; sometimes. how- ever, missing. Frontal avicularia (fr. av.) on the zocecium at bifurcation only; small, raised, with mandible directed for- ward, often obliquely to right or left. Oacia smooth, somewhat elobose, reaching not quite to the lower margin of the next upper zocecium (oe.). From a flat disk in front or to one side of the aperture of many of the lower zoccia in a colony, root fibers (7.) extend downward, assisting in anchoring the colony to the sub- stratum; from zocecia higher in the colony other upward ex- tending fibres arise in a dorso-lateral chamber (7. ch.) which is slightly raised and which projects from the zocecial wall. M. ternata is widely distributed, being found on Channel Rocks, and San Juan Island in Puget Sound; and at Dillons Beach, Lime Point, Mendocino, and Lands End on the coast of California. It is also reported by Hincks from Queen Charlotte Islands. In the species from these different localities considerable variation occurs. As a rule, the colonies from California grow in more compact masses, and the triads of zocecia forming the internodes are shorter than are those of the Puget Sound species. Thus the length of an internode in the Puget Sound species varies between 1,100 and 1,500 p», and that of the species found VoL. 2 | Robertson.—Non-Incrusting Bryozoa. 253 farther south, slightly less. In the main points, however, the Menipeas from these various localities strongly resemble each other and in no points more constantly than in the position and form of the chambers from which the root and climbing fibres spring. 6. Menipea gracilis Busk. Pl. VI, figs. 18, 19, 20, 21. Menipea gracilis Busk, 1881. Menipea ternata, forma gracilis, Smitt, 1867. Zoarium forming a loose, tangled mass due to the great num- ber of tendril-like fibres which twine around neighboring branches. (PI. VI, fig. 18.) Branching dichotomous; internodes consisting of three zocecia, except terminal internodes and those possessing ovicells, which may have five, seven, or nine zoceeia; internodes of three zowcia only, often very long, ranging from 1,600 to 1,900 » in length. Zoawcia enlongated, aperture large. (fig. 19). Zocecia with two or three spines, depending upon the presence or absence of the lateral avicularia. If the latter are absent there will usually be two spines on the outer margin; if present, then but one (sp.). There is always a spine on the inner margin of the aperture just above the seutum. Zocecium at bifurcation possesses one or two spines, more often one, at its summit. Scutwm (sc.) large, flabellate, projecting outward and arching over the aperture in well developed specimens, pro- jecting beyond the outer edge of the zoecium. Lateral avicularia frequently absent, when present often minute. Frontal avi- cularia (fr. av.) oceur on the zocecium at the bifurcation of the branch, but these too, are frequently absent; on ocecial inter- nodes,. however, they are often found on each zocecium, situated a little to one side of the aperture and so close to the owcium as to seem perched upon its upper margin (fig. 20, av.). Oacia (oe.) high, rounded, with striations radiating from a thin place at the base of the owcial wall. Rootlets originating at two places. Those which anchor the colony springing, as a rule, from a disk to one side and below the aperture (fig. 19, r. d.) ; the others, more tendril-like, and for the most part ex- 254 University of Califorma Publications. | ZOOLOGY tending upward, arising from a definite chamber just above the lateral avicularium, the chamber projecting somewhat and being not quite circular especially as seen from the dorsal side (fig. PA Gh): This species is abundant at Orea, Prince Williams Sound; reported by Hineks from Cumshewa Harbor, Queen Charlotte Island. 7. Menipea occidentalis Trask. Pl. VI, figs. 22, 23, 24, 25. Menipea occidentalis Trask, 1857, p. 113, PISA, hip. Menipea compacta Hincks, 1882, vol. 10, p. 461. Menipea compacta Hincks, 1884, vol. 13, p. 208, Pl. LX, fig. 8. Zoarium forming bushy tufts from 15 to 50 mm. in height, attached by a large number of root fibres. Root fibres not de- veloped throughout the colony, this species being in no sense a climber. Branching extremely regular and characteristic (fig. 22), each tuft or frond consisting of a main rib or primary branch (pr. br.) from which secondary branches arise alter- nately (sec. br.), these again giving off tertiary branches (ter. br.). Internodes consisting of three zocecia (fig. 23). Joints (j.) yellow or brown arising from definite chambers on each side of the most anterior zocecium of an internode. Zoawcia elongated, narrowed below; aperture (ap.) occupying about half the front. surrounded by six jointed spines, sometimes by five or seven; two arising on the margin of the upper half of the aperture opposite each other and meeting across it (c. sp.) ; two other longer ones extending from the upper margin of the aperture (u. sp.), and between these two and the two crossed spines, two other opposite flaring spines (fl. sp.) ; on the terminal zocecia these flaring spines often grow very long so that each zocecium is then bordered anteriorly with a bristlng array of four long spines. Scutwm (sc.) sometimes a mere spine, sometimes broader than a spine, arising on the lower half of the inner margin of the aperture. Avicularia large, lateral, sessile, found typically on all the zocecia except the one at the bifurcation of a branch. Frontal avicularia wanting. Oa@cia usually developed on the tertiary branches, large, globose, covering the zocecial wall below VoL. 2] Robertson.—Non-Incrusting Bryozoa. van the apertures of the zocecia just above (fig. 24, oe.), their front walls perforated by a small number of pores. Rootlets of one sort only, extending downward, serving to anchor the colony; arising in root chambers just above the lateral avicularia (fig. 25, r. ch.) ; the root chambers situated on the lower zocecia only, and never appearing unless a root fibre is developed; rounded, projecting dorsally and laterally, the root fibre springing from the lower side and extending downward keeping close to the branch and running parallel with the other root fibres of the branch, thus forming a sort of cable or rope for the attachment of the colony. This species was first described by Dr. John Trask (757), but his description was apparently unknown to Hincks (’82), who redescribed it under the name of M. compacta. It is per- haps the most abundant species of bryozoa on the shores of San Francisco Bay and Golden Gate Straits. It grows between tide marks on rocks and sea weed forming bushy tufts of a white, yellowish or dirty grey color, depending on its age. It is quite easily recognized by its excessive spininess. It is found from Queen Charlotte Islands to San Diego, but is most abundant above Point Concepcion. South of that point it is noticeably searce on the shore and in dredgings, and the colonies are small and delicate. 8. Menipea occidentalis catalinensis. Subsp. nov. Pl. VII, figs. 26, 27. South of Point Concepcion, notably at Santa Catalina and San Pedro, M. occidentalis shows considerable variation. The internodes more often consist of five or seven zocecia, and the seutum which in the type form is a mere spine, or at most is slightly flabellate, in the subspecies catalinensis is decidedly fan-shaped, the edge being divided and extended into five, six, or more spinous processes, making it so large as to cover the lower half of the aperture (fig. 26, sc.). One or both of the spines that meet over the upper part of the aperture may be bifid (c. sp.). The bifid spines and the large and much divided scutum are very noticeable features and constitute the chief 256 University of California Publications. | ZOOLOGY differences between the subspecies catalinensis, and the typical M. occidentalis. While these variations from the type occur rather constantly in specimens from the south, affecting as they do such variable appendages as spines and scuta, they are not considered of sufficient importance to establish a new species. The drawing (fig. 26) gives the impression, unfortunately, that the individual zoccia of the subspecies catalinensis are larger than those of the type M. occidentalis, but it was unintentionally made on a larger scale of magnification than was that of figure 23. 9. Menipea erecta Robertson. PI Val, fies..28) 29573031" Menipea erecta Robertson, 1900, p. 317, Pl. XTX, figs. 1, 2. Scrupocellaria scabra, Robertson, 1900, p. 318, Pl. XIX, figs. 3, 4. Zoarium composed of numerous stiff, dichotomously divided branches from 20 to 25 mm. in length (fig. 28). Inter- nodes consisting of three, five, or seven zocecia, the number in- ereasing toward the extremities of the branches; articulations extending through the zoccia just below the aperture, some- times including its lower border (fig. 29, art.). Zoacia biserial, alternate, narrowed below ; aperture oblong, occupying more than half the front; margin raised, crenulate, with one or two blunt spines at the upper outer angle; scutuwm (sc.) a flattened spine, sometimes growing broad and bifid. Lateral avicularia fre- quently wanting (fig. 30), sometimes feebly developed (fig. 31), and again rather large (fig. 29). When avicularia are present there is usually but one spine at the upper, outer angle, although this is not the invariable rule. Frontal avicularia (fr. av.) gen- erally present on each zocecium, though sometimes lacking; when present, they are situated to one side of the aperture, at the base of the scutum of the adjacent zocecium. Oacia large, glo- bose, more or less striated. Root fibres mainly upon the lower zocecia. springing directly from the zocecial wall and extending downward (fig. 30, 7.). Examination of material from several localities, especially of that from Puget Sound, leads me to unite the species for- VoL. 2 | Robertson.—Non-Incrusting Bryozoa. 257 merly identified as Scrupocellaria scabra Van Beneden with Menipea erecta Robertson (700). This species is obtained from three localities, Kadiak and Sitka, Alaska, and San Juan Island, Puget Sound. The range of variation is considerable, but is not greater than frequently occurs in a species found in localities so remote, and consists mainly in the presence or absence of spines or avicularia. The specimens from Sitka, for example (fig. 30), show irregular development of frontal avicularia with entire absence of lateral avicularia. The colonies from Kadiak (fig. 29) show frequent absence of lateral avicularia and their replacement with spines; while a few zoecia from San Juan specimens (fig. 31) show an entire absence of spines and considerable variation in the size of the lateral avicularia. 10. Menipea pribilofi sp. nov. Pl. VII, figs. 32, 33; Pl. VIII, fig. 34. Zoartum forming a compact mass 25-50 mm. high, attached by a large number of root fibres; the upward tendril-like fibres being very slightly developed (Pl. VII, fig. 32.). Branching dichotomous, main branches long, possessing a number of shorter inward curving secondary branches. Jnternodes consisting of three zocecia except the ocecial internodes which generally con- sist of five; separated by dark colored chitinous joints. Zowcia relatively short and stout, broad at the top, attenuated below, aperture occupying less than half the front surface, with three spines on its upper margin; the bifureating zocecium may have but two spines, but it is frequently found bristling with three or four. Scutum (Pl. VIII, fig. 34, sc.) simple, often a mere spinous process; when better developed, it broadens at the free extremity. Spines and secuta distinctly jointed. Lateral avi- cularia generally present, large (lat. av.). Frontal avicularia (fr. av.) few, when present found only on the zocecium at the bifureation of a branch, large, raised, beak set obliquely, some- times transversely to the length of the zocecium. Owcia globose, smooth (Pl. VII, fig. 33, 0e.). Rootlets in two positions. Those 258 University of California Publications. [ ZooLoGy lower in the colony develop on the front surface of the lower zocecia, below and to one side of the aperture. These rootlets invariably proceed straight downwards (Pl. VIII, fig. 34, r.). Often on a zocecium in the same internode, and on zocecia at higher levels, just above the lateral avicularia, other rootlets eurving upward, develop from a circular chamber slightly raised above the zocecial wall (wv. 7.).. These upward growing fibres are of much less frequent occurrence than in M. ternata and M. gracilis, and the chambers from which they proceed are of much simpler construction. On the upper half of the colony neither fibres nor chambers are found. In the habit of growth, greater compactness, greater develop- ment of spines, and scarcity of tendril-like fibres, this species resembles M. occidentalis rather than M. ternata. Menipea pribilofi is known only from the shores of Alaska, and the islands of Bering Sea, having been obtained in consid- erable quantity from St. Paul Island, Pribilof Islands; and in small quantity from Homer, Unalaska, and Yakutat, Alaska. Serupocellaria Van Beneden. Cellularia Pallas, 1766. Cellaria, Ell. and Sol., 1786. Scrupocellaria Van Beneden, 1844. Cellularia, Johnston, 1847. Canda, Busk, 1852. Scrupocellaria, Hincks, 1880. Zoarium jointed. Zowcia numerous in each internode, rhom- boid; aperture with or without scutum; a sessile aviculariwm at the upper, outer, lateral angle; a vibraculum at the lower outer angle, and generally a sessible avicularium on the front surface of each zocecium. KEY TO THE SPECIES OF SCRUPOCELLARIA. i. Vibracular chamber tonpey ery) Z0ccerimae serene See eee 2 1. Vibracular chamber not on every z0@¢ium <2 2 3 2. Vibraculum slightly longer than a zoccium ...........-..--.--...-- S. varians 2. Vibraculum as long\as three: zomeia ee S. diegensis 3. Vibraculum not as long as a zocecium ..............2--.2-20------ S. californica VoL. 2] Robertson.—Non-Incrusting Bryozoa. 259 11. Serupocellaria californica Trask. Pl. VIII, figs. 35, 36, 36a, 37. Scrupocellaria californica Trask, 1857, p. 114, Pl. 4, fig. 2. Scrupocellaria brevisetis (?) Hincks, 1882, p. 462. Zoarium growing in large compact tufts, somewhat coarse in appearance. (PI. VIII, fig. 36a.) Branching dichotomous, inter- nodes consisting of five, seven, or nine zowecia. Zowcia shghtly attenuated below, aperture occupying more than half the front (fig. 36). Two, often three spines on the upper, outer margin and one on the inner just above the seutum. Zocecium at the bifureation (z@. bi.) of a branch with one short spine at its apex: two spines below on one side and one spine on the other just above the secutum. Scutwm (sc.) often spine-like, sometimes broadened below, and always curved with a downward slope. Lateral avicularia generally present, when absent, an extra spine may develop in that place, making the third on the outer edge of the margin (sp.). Avicularia vary in size, but unlike those of the following species, the fluctuation in size lies in the greater or less relative proportion of the muscular part (lat. av.). Fron- tal avicularia (fr. av.) present on each zocecium just below the aperture. Vibracular cell frequently lacking, and not visible from the front surface except on the zocecium at the bifurcation of a branch (v. c.). Vibracula, when present, found only on the zocecia of the lower part of the colony where root fibres are given off (r.). This is usually true except of the zocecium at the bifurcation of a branch where vibracular cells are generally present irrespective of the formation of a root fibre. Vibracu- lum shorter than a zocecium; vibracular chamber much like a lat- eral avicularium in form (fig. 35, v. ch.). The groove, lying in the part corresponding to the lower mandible of an avicularium, extending transversely across the dorsal surface of a zocecium (gr.), the short stout vibraculum much like the elongated upper mandible of an avicularium. Oacia sparingly developed, found only on internodes at the extremity of the branches, three or four in an internode, smooth (fig. 37, oe.). Rootlets abundant on the lower internodes of a colony where vibracular cells abound with their short vibracula (fig. 36, 7.). 260 University of Californa Publications. | ZooLocy This species first figured and described by Trask (757) from San Francisco Bay is probably the S. brevisetis, Hincks (’82) from Queen Charlotte Islands, of which only a short description without figure is given. Trask found neither vibracula nor vibracular chambers, but they are abundant, as I have observed, on the lower part of the colony, while almost entirely lacking on the upper portion. S. californica is distributed at various points along the coast of California; it grows luxuriantly at Dillons Beach and at many places on San Francisco Bay, and occurs in small quantity below Point Concepcion. 12. Serupocellaria varians Hincks. Pl VOW, fies: 38,39- Blo XV1, fig. 95: Scrupocellaria varians Hincks, 1882, p. 461, Pl. XTX, figs. 1-le. Zoarium forming bushy tufts 12-25 mm. in height. (PI. XVI, fig. 95.) Branching dichotorhous, internodes consist- ing of a variable number of zocecia, those in the lower part of the colony containing usually five or seven zoccia, those at a higher level, nine or eleven. Joints yellow. Zowcia biserial, alternate, slightly narrower below; aperture oval, occupying more than half the front; scutwm sometimes a mere spine, some- times bifid or trifid, usually inclined downward (Pl. VIII, fig. 38) ; two spines on the upper margin, one of them just above the seutum, the other opposite. Both lateral and frontal avicularia developed on each zocecium. Lateral avicularia of different pro- portions, the greater number being much elongated, extending upward and outward, with a long, thin, pointed, curved beak (c. av.) ; frontal avicularia just below or slightly to one side of the aperture; sessile, raised, with mandible directed transversely. Vibracular chamber triangular, the apex visible from the front (fig. 39, v. ch.) ; vibraculum (v.) longer than a zocecium, and the groove into which it falls, extending transversely across the zocecium (gr.). Owcia (fig. 38, oe.), smooth, developed on each zocecium of the terminal internodes. Rootlets springing from the base of the vibracular chamber (fig. 39, 7.). VoL. 2 | Robertson.—Non-Incrusting Bryozoa. 261 Perhaps the most striking feature of this species is the pecu- liar size and prominence of many of the lateral avicularia. This is brought about by the great extension of the beak and is not due to increase of the muscular portion (mus.) ; thus, in fig. 39 (¢. av.) the mandible (man.) is seen to be longer than the muscular part, and to be more than half as long as the whole appendage. In the shape of its vibracular chamber and in the unusually large size of its lateral avicularia, this species resembles S. cali- fornica, but as has been said, the increase of size of the avicularia in the two eases is due to increase of different parts of the organ. In other respects, S. varians and S. californica are very unlike, e.g., in habit, in shape of seuta, and more especially in abund- ance of the vibracular chambers and length of the vibracula. Although this species has been obtained in small quantities on the California coast as far south as San Pedro and La Jolla, it is characteristically a more northern form. It was first described by Hineks from Queen Charlotte Islands, and it is abundant in Puget Sound, both on Channel Rocks, and at San Juan. Material obtained in the south was dredged at 32 fathoms, while in Puget Sound this species is a shore form. 13. Serupocellaria diegensis, sp. nov. Pl. IX, figs. 41, 42, 43, 44; Pl. XVI, fig. 96. Zoarium forming a coarse bushy mass often 50 mm. in height (Pl. XVI, fig. 96). Branching dichotomous, internodes consisting of a variable number of zocecia, nine. thirteen, seventeen or more, especially in the terminal internodes. Joints somewhat inconspicuous, often occurring as high on the zocecium as the lower margin of the aperture. Zoacia biserial, the two rows inclined at an angle so that the internode is keeled, and in cross section is triangular; rather short and broad, shghtly narrowed below (Pl. IX, fig. 41); aperture oval, with a wide margin, and occupying more than half the front surface. Scu- tum (sc.) in older zocecia large, covering more than half the aperture, thickened on the inner surface and raised on a ped- unele. Spines three, four, or five, the one immediately above the seutum, and frequently the one opposite, bifid (67. sp.). In 262 University of California Publications. [ ZOOLOGY well developed zocecia there are three spines on the outer mar- gin, the lower of these frequently bifid, the second often quite long, and the third, or uppermost one shorter and sometimes missing; two spines on the inner margin, the lower generally bifid, the upper frequently lacking or broken. Lateral avicularia of moderate size, usually found on each zoccium. A series of sessile, frontal avicularia (f7. av.) extends between the zocecia, each avicularium situated usually at the base of the spine on the inner margin nearest the scutum. These frequently stand out prominently, the beak opening upward, so that the zoccia frequently have the appearance of being flanked on each side by an avicularium, the one on the inner side being slightly smaller than that on the outer. The zocecium at the bifureation of a branch usually bears a raised avicularium of extraordinary size, with elongated beak directed obliquely, sometimes to the right, sometimes to the left (67. av.). Vibracular chamber dorsal, large, its length equalling half that of the zocecium on which it is placed (fig. 42, v. ch.) ; vibraculum long, two and a half times as long as the individual zowcia (fig. 41, v.). The rootlet arises from near the base of the vibracular chamber, toward the outer side (figs. 42 and 43, r.) The rootlets are developed only on the lower zocecia of a colony; in the upper zoccia the place of the rootlet is marked by a pore (fig. 42, p.) Oacia numerous, large, reaching to the margin of the aperture of the zocecium next above, and sometimes covering it (fig. 41, oe.) ; somewhat flattened and marked by numerous pores. This species grows in large bunches, and being very calear- eous, and having many long vibracula, spines and other appen- dages which catch and hold debris, it is remarkably coarse and dirty in appearance and rough to the touch. It strongly re- sembles Caberea, and the vibracular cells while not as large as those of that genus, yet show affinities to them in size, and in the length and slope of the groove. This species likewise shows certain resemblances to S. cervicornis, Busk (752), but its habit is very different and it does not possess the peculiar scutum for which the latter is named. S. diegensis is extremely abundant at San Diego on the rocks at Ballast Point, on floats at Coronado, and wherever it can get ~~ VoL. 2 | Robertson.—Non-Incrusting Bryozoa. 263 a foothold. It seems to be almost the most abundant. bryozoan of the region; also found at many points in the vicinity of San Pedro. Less abundant northward, being found only in small quantities in San Francisco Bay and not obtained north of this locality. Caberea Lamouroux. Caberea Lamouroux, 1816. Flustra, (part), Johnston, 1847. Cellularia (part), Johnston, 1847. Caberea, Busk, 1852. Canda, d’Orbigny, 1850. Caberea, Smitt, 1867. Caberea, Hincks, 1880. Zoarium not articulated. Zoacia in two or more series, sub- quadrangular, or ovate, with a very large aperture. Sessile, frontal avicularia on the side and front of zocecia; lateral avicu- laria minute. Vibracular cells very large, placed in two rows, stretching obliquely downwards across the back of the zocecia, which they almost cover, to the median lne, notched above and traversed through a great portion of their length by a shallow groove. Vibracula usually toothed on one side. This genus is easily distinguished from others of this family by the peculiar shape and size of the vibracular cell as shown in fig. 46. It is not common in our collections, being repre- sented by only a small part of a colony of one species. 14. Caberea ellisi Fleming. Pl. VIII, fig. 40; Pl. IX, figs. 45, 46. Flustra ellisii Fleming, Mem. Wern. Soe. ii, p. 251, Pl. 17, figs. 1-3. Flustra setacea Johnston, 1847, ed. Il, p. 346. Cellularia hookeri, Busk, 1852, pt. I, p. 37, Pl. XX XVIII, fig. 2. Caberea ellisii, Smitt, 1867, pp. 287 and 327, Pl. XVII, figs. 55, 56. Caberea ellisii, Hincks, 1880, p. 59, Pl. VIII, figs. 6-8. Zoarium fan-shaped, dichotomously branched ; branches thick, widening upward (Pl. VIII, fig. 40). Zowcia in two or four rows, short, subquadrangular (Pl. IX, fig. 45); aperture elliptical, occupying nearly the whole of the front, with a broad minutely eranular margin, sloping outwards; marginal zocecia with two stout spines above on the outer side, and one on the inner; intermediate zocecia with one spine on each side. Lateral avicu- 264 University of California Publications. | ZOOLOGY laria (lat. av.) small and inconspicuous, placed a little below the top of the zowcium, with a rounded mandible; frontal avicu- laria (fr. av.) raised, two below the aperture, placed one on each side, or sometimes only one; mandible rounded, directed downwards. Vibracula very long, serrate (v.). Owcia flattened, frequently with a depressed, smooth, semicircular space in front from which fine striae radiate to the margin. footlets arise from the side of the vibracular chamber, the two on opposite sides uniting in the median dorsal line with each other and with those below, forming a thick keel-like cable along the dorsal sur- face of the branches (fig. 46, 7.). At the base of the colony the threads become free and attach themselves independently to particles of sand or to whatever forms the substratum. A small quantity of C. ellisi was dredged in 20 fathoms at Juneau. It is reported also from Queen Charlotte Islands. Bicellariidae Busk. Bicellariade Busk, 1852. Bicellariee Smitt, 1867. Bicellariide, Hincks, 1880. Zoarium erect, with or without articulated stem or pedunele, or composed of a number of zocecia connected by tubular pro- cesses. Zowcia rather loosely united in one, two, or more series, or disjunct, boat-shaped or sub-tubular; aperture occupying a large proportion of the front. Avicularia, when present, capi- tate, pedunculated, and articulated. Owcia either pedunculated or sessile. The diagnosis of the family Bicellartidw as given by Hincks is here revised in order to include the genus Stirparia Goldstein. This genus is fairly abundant on this coast, and unlike the other members of the family possesses a jointed stem of varying length surmounted by delicate feathery tufts of zoccia. KEY TO THE GENERA OF BICELLARIIDAS. COW O ray ae LCC ex os reese eee ee ce ee Goloniy:. ereeprin ce 526 eee a i ence eee Gollomy,-wumsiialle dy Bee ac ee Gollomiy; stevie aa a ee eee FioceGlan HUIS err | cone sears 5 eee en ee ee ee Corynoporella Fomeia. i= wOr Mla SO rey eee cease eee ree ee eae nate ee Bugula PO CGA WWUSCTUE UN See aN rer ee ee See ee Stirparia Zoewcia uni- or multiserial, with tubular connections ................ Beania H® Co O1 bo OUR 99 69 PO DO ~ l — Vou. 2] Robertson.—Non-Incrusting Bryozoa. 265 Bugula Oken. Cellularia Pallas, 1766. Bugula Oken, 1815. Cellularia, Johnston, 1847. Avicella Van Beneden, 1848. Avicella Van Beneden, 1849. Ornithopora d’Orbigny, 1851. Ornithoporina d’Orbigny, 1851. Bugula, Busk, 1852. Bugula, Hincks, 1880. Zoarivum erect, branched. Zoacia boat-shaped, or sub-quad- rangular, elongate, united in two or more series; aperture occu- pying a large proportion, sometimes the whole of the front, not turned upward or oblique. Avicularia in the form of a bird’s head, and jointed, usually one to each zowcium. Oacia sessile at summit of aperture, or pedunculate upon the side of the aperture. This genus is abundant in this region, not only in the number of species but also in the luxuriance of growth, and is remark- able for the large size which some of the species attain. Nine species are found, all of which are new except three; one, Bugula murrayand, is circumpolar, and another, B. neritina, is ecosmo- politan in range. KEY TO SPECIES OF BUGULA. Ji SUT OSCOMEY So ONISK Sta IES SES ew Lo ane 8 ae oe Bc epee ae J Ne 2 The see OTE, ATO LES (ENE ESSA eA ae ee ed en Ue een 4 2. Avicularia, if present, at summit of aperture -............... B. neritina Zee ASYM aT mete a SOMO Ap CL GUE Cyst ee rence ge ne ee B. pacifica 2. Avicularia between summit and base of aperture =... 0... 3 3. Avicularia much elongated, half as long as zowcium, slightly more than half way between summit and base of aperture Re ee os RE SN, Aon IRAE STi ee NE B. longirostrata 3. Avicularia not elongated, less than half the length of a zowcium, just half way between base and summit of aperture....B. californica Ape AVLGTU RABY ap TSE ier teens 8 eR he eet ged ea EAs a RD 5 cI a)a Nv CVU: Tose Bese] OYS(2) 01 ee ke, A a eee eo 2 B. laxa DaeAWwAeUl aria at baser Oh aOerbiut esse eee eee ee -B. murrayana 5. Avicularia between base and summit of aperture _.............00...... 6 6. Avicularia less than half way from base and summit of aperture........ BS eee eas oa. eee a ase ee eee eb. flabellata 6. Avicularia near base of aperture, beak much curved _... poe SSE At at Bnad AO Seema BA Soang Mares Pete EN (Oe. tS Oe B. curvirostrata 266 University of California Publications. [ ZOOLOGY 15. Bugula neritina (Linneus) Oken. Pl. IX, fig. 47; Pl. XVI, fig. 97. Sertularia neritina Linneus, 1758, ed. X, p. 38. Cellularia neritina, Pallas, 1766, p. 67. Snail-bearing Coralline Ellis, 1767, p. 40, Pl. 19, figs. a, A. Cellaria neritina, Ell. and Sol., 1786, p. 22. Bugula neritina, Oken, 1815, Ab. 2, p. 89. Acamarchis neritina, Lamouroux, 1816, p. 58, Pl. 3, fig. 2. Cellularia neritina, Johnston, 1847, p. 340, Pl. 60, figs. 3, 4. Acamarchis neritina, d’Orbigny, 1850-52, p. 324. Bugula neritina, Busk, 1852, p. 44, Pl. XLITI, figs. 1-6. Bugula neritina, Waters, 1887, p. 91, Pl. IV, figs. 3, 15. Zoarium consisting of large bushy tufts 75 to 100 mm. in height, brown or reddish brown, often tinged with purple. (Pl. XVI, fig. 97.) Branching dichotomous. Zoecia biserial, quadrangular, truneate above; aperture occupying more than two-thirds of the front; a short spine or denticle at the summit of the sides of the zowcium (PI. IX, fig. 47, den.). Owcia very conspicuous and numerous (0e.); globose, attached to the inner anterior angle of the zocwcium by a short peduncle. Avicularia none. Rootlets forming a tuft at the base of the colony. This species is extremely abundant, especially on the southern coast of California. It is not only found on rocks, floats, kelp, ete., along the shore, but it is frequently taken by the dredge, and great brown masses are constantly cast on the beach by the waves. Its most northern limit is Monterey Bay, where it is obtained in small quantities. 16. Bugula murrayana (Johnston) Busk. Pl. X, fig. 48; Pl. XVI, figs. 98, 99, ? Eschara hispida Pallas, 1766, p. 49. Flustra murrayana Johnston, 1847, ed. Il, p. 347, Pl. LXITI, figs. 5, 6. Avicella multispina Van Beneden, 1848, Vol. 15, pt. I, p. 76, figs. (he $e ? Ornithopora dilatata d’Orbigny, 1850-52, p. 323. Bugula murrayana, Busk, 1852, p. 46, Pl. LIX, figs. 1, 2. Bugula murrayana, Smitt, 1867, p. 291, Pl. XVIII, figs. 19-27. Bugula murrayana, Hincks, 1880, p. 92, Pl. XIV, figs. 2-9. Zoarium composed of bushy tufts from 25-50 mm. in height, sometimes consisting of masses of Flustra-like fronds 50 or VoL, 2] Robertson.—Non-Incrusting Bryozoa. 267 more mm. in diameter (Pl. XVI, fig. 98), or of much elon- gated, narrow, strap-like branches (fig. 99). Zoawcia multi- serial, the branches having from three or four, to twelve or four- teen rows; alternate, oblong, slightly narrower below, truncate above (Pl. X, fig. 48); aperture occupying more than half the front; an erect spine at each upper outer angle (sp.); a varying number of marginal spines, two to five on the inner margin, one to three on the outer, curving inward; occasionally in the ab- sence of an avicularium, a spine at the bottom of the aperture, curving upward (ba. sp.). Avicularia pedunculate, those on the marginal zowcia, when present, much larger than those on the intermediate zoccia; often absent from both marginal and intermediate zocecia. Oacia (oe.) large, subglobose, marked by radiating lines. Rootlets given off from marginal zocecia near base of colony. B. murrayana is found at Kadiak, Orea, Pribilof Islands, Alaska, and in Puget Sound, but does not extend farther south. The specimens from Orea, Prince Williams Sound, are ro- bust, often possessing a large number of zocecia in a series form- ing broad strap-like segments or branches (Pl. XVI, fig. 98). Those from Kadiak and Puget Sound are more delicate in ap- pearance, forming smaller colonies (fig. 99). Those dredged at ten fathoms in Puget Sound do not possess more than six or eight rows of zocecia in a series. There is great range of variation in the number of spines, and in the number and size of avicularia, among the specimens from the different localities. A consid- erable quantity of material from Pribilof Islands possesses the large marginal avicularia, while most of the material from other localities lacks them. 17. Bugula californica sp. nov. Pl. X, fig. 49; Pl. XVI, fig. 100. Zoarium consisting of a number of masses whose branches have a distinetly spiral arrangement about a central axis, some- times forming a coarse growth 75 mm. in height and from 25 to 50 mm. in diameter in the broadest place (Pl. XVI, fig. 100). 268 University of California Publications. [ ZooLoGy Each branch composed of many strap-like branchlets consisting of two series of zowcia, dichotomously divided to form a fan or frond. Zowcia narrow below, elongated, bearing three spines at the distal extremity, one stout and long (Pl. X, fig. 49, sp.), extending in a direction parallel with the branch, longer than the other two, a continuation of the wall of the zocwcium; two (sp.) at the top of the aperture projecting forward at right angles to the front surface. Aperture occupying more than two- thirds of the front of the zowcium. Avicularia large, pedun- culate, situated near the top or not more than half the length of the zocecium below the top. Oacia large, high, globose, extend- ine above the lower margin of the aperture of the next upper zocecium. Rootlets developed on lower zocecia, springing from front or ventral surface of zocecium below the aperture. The zoccia in this species are usually arranged biserially, but sometimes, especially near the point where the branch is about to divide it may contain for a short distance three or four rows of zocecia in the series (fig. 49). B. californica is most nearly allied to B. turbinata, but the zocecia are somewhat shorter and stouter; the avicularia are of much the same form and size, but are placed lower down on the zocecium; and the spines are constantly three instead of two, and of a different character from those of B. turbinata. The ocecia also differ, those of B. californica lacking the prominent process or peak described for B. turbinata. Colonies of large size oecur abundantly at Dillons Beach, California; B. californica is also found, but of less luxuriant erowth at Lands End, San Francisco Bay, and at Pacific Grove, Monterey Bay. 18. Bugula pacifica Robertson. Pl. X, fig. 50; Pl. XVI, fig. 101. Bugula purpurotincta, Robertson, 00, p. 320. Bugula pacifica Robertson, 700, p. 321. Zoarium consisting of large bushy tufts having a somewhat spiral growth, often 75 mm. in height (Pl. XVI, fig. 101) ; col- onies frequently united by a sort of cable formed by union of VoL, 2] Robertson.—Non-Incrusting Bryozoa. 269 the root fibres. More delicate in appearance than the preceding species, often tinted purple. Zowcia elongated, biserial, armed at the distal extremity with three spines, the outer one lone (300 »), projecting outward (Pl. X, fig. 50, sp.) ; the other two shorter (100 »), being continuations of the margin of the aper- ture (sp.’); Aperture occupying almost the whole of the front of the zoecium. Avicularia large, pedunculate, at the base and to the outer side of the aperture. Oq@cia remarkably small, not rising more than 60 or 80 » above the zoccium, while the em- bryo (emb.), in many cases measuring 200 p, extends down- ward into the upper part of the zoccium. Rootlets numerous. extending from the lowest zocecia of a colony. This species was formerly (’00) somewhat. tentatively iden- tified as B. purpurotincta Norman, but after more extended study of specimens from many different localities it is unques- tionably a new species. As formerly suggested by the writer, it is given the specific name pacifica, since it seems to be charac- teristic of the Pacific Coast. It ranges from Pribilof Islands, Bering Sea, to San Francisco Bay, the climax of growth. both in quantity and size being reached at Yakutat, Alaska. The purple color of the specimens obtained at this place is partic- ularly noticeable. This, as I have said, resides in part in the tissue lining the zoccia, and in part in the degenerated poly- * and is quickly lost after the material pides or ‘‘brown bodies,’ has been placed in alcohol. From one locality in Puget Sound where the species was obtained in abundance, it possessed a dis- tinctly greenish hue, while at many other places it is white or colored slightly yellow. The extreme shallowness and flatness of the omwcia are remarkable characters but are not peculiar to this species being found in at least one other species from this coast, Bugula longirostrata. The ovum matures in the upper part of the zocecium rather than in the occium, the em- bryo frequently blocking the mouth of the former. In all cases, where an embryo has formed, the polypide has degenerated into a ‘‘brown (purple) body.’’? Unlike those species which possess ocecia elevated above the mouth or orifice of the zowecium, func- tional polypides and embryos cannot exist simultaneously in 270 University of California Publications. | ZooLocy the same zocecium, and probably on account of the purely me- chanical obstruction caused by the embryo. The distribution of B. pacifica extends from Pribilof Islands, Bering Sea, to San Francisco Bay. It has been obtained from St. Paul, Pribilof Islands; is very abundant at Yakutat, and is found in smaller quantity at Orea, Prince William Sound, Alaska; abundant, but colonies smaller, at Sidney, opposite Port Orchard Navy Yard, Puget Sound, less abundant on Chan- nel Rocks, Puget Sound; fine colonies obtained from rocks at Dillons Beach, California; smaller quantity obtained from Lime Point and Fort Point, San Francisco Bay, California. 19. Bugula flabellata J. V. Thompson. Pl) axe dies SoZ: Bird’s Head Coralline Ellis, 1767, Ger. ed., p. 109, Pl. XXXVIII, Olt. Cellularia avicularia (part) Pallas, 1766, p. 68. Flustra avicularis, Johnston, 1847, p. 346, Pl. LXIII, figs. 3, 4. Avicularia flabellata Thompson, 1847, MS. Brit. Mus.; Gray, Brit. Mus. Radiata, p. 106. Avicella avicularia Van Beneden, 1848, p. 75. Ornithoporina avicularia, d’Orbigny, 1850, p. 322. Bugula flabellata, Busk, 1852, p. 44, Pls. LI, LILI. Bugula avicularia forma 2, B. flabellata, Smitt, 1867, pp. 290, 345. Bugula flabellata Thompson, 1868, Pl. VI, fig. 9. Bugula flabellata, Norman, 1868, Pl. VI, fig. 9. Bugula flabellata, Smitt, 1871-72, pt. I, p. 18, Pl. V, figs. 48-52. Bugula flustroides, Verrill, 1879, p. 52. Bugula flabellata, Hincks, 1880, p. 80, Pl. XI, figs. 1-3. not Bugula flabellata, Robertson, 1900, p. 431. Zoarium from 12-25 mm. in height, consisting of a number of fan-shaped fronds divided into narrow branches (fig. 51). Branching dichotomous, occurring where a branch reaches a width of six or seven rows of zoccia. Zowcia elongated, of about equal length throughout, aperture occupying the whole or almost the whole of the front; usually two spines placed one above the other at each upper angle, the upper spines stiff and flaring, the lower often of great length and frequently crossed in front; on marginal zocecia there are generally three spines on the outer side and two on the inner (fig. 52). Avicularia less than half VoL, 2] Robertson.—Non-Incrusting Bryozoa. 271 way from the top of each zocecium; those on the intermediate zocecia smaller than those on the margin; marginal avicularia of medium size, point of mandible bent abruptly downward. Oawcia small, globular, smooth, well elevated above the top of the zocecium, with a double line forming a band on the oral rim; opening wide. This species differs slightly from Hincks’ description of B. flabellata, in the number of spines, the English species appar- ently having but two on each side of the zowcium. It agrees, however, with the figure given by Busk in which he represents three spines on the outer side of the marginal zocecia. In other respects the English and American species seem to be identical. Bugula flabellata has been obtained in small quantity on the piles in San Diego Bay. 20. Bugula pugeti sp. nov. PIX, figs. 53,54; Pl. Xl, fig: 55. Bugula flabellata, Robertson, 1900, p. 321. Zoarium composed of numerous small somewhat spiral tufts from 10 to 25 mm. in height (fig. 53). Zowcia multiserial, each branch consisting of from four to seven rows (fig. 54); long, narrowed slightly below; aperture occupying two- thirds or more of the front; three spines on the marginal zocecia, two on the outer side (ou. sp.), one on the inner (in sp.) ; inter- mediate zocecia with two spines only; the middle of the anterior edge of each zocecium occupied by a round knob or process whose significance is unknown (k.). Avicularia (av.) found only on the marginal zoweia, large, beak somewhat curved, sit- uated half way from the top, or just below the middle of the zocecium. Oacia none. Embryos (Pl. XI, fig. 55 emb.) develop- ing in the anterior part of the zocecium, and no additional ocecium being formed. Embryos abundant in the older parts of the colony, and those zocecia possessing well developed embryos con- taining no polypide. Rootlets extending from the lower zocecia of the colony, forming a cluster at its base and attaching it to shells, rocks, or sea weed. 272 University of California Publications. [ ZooLocy This species resembles B. flabellata in some respects, and was previously so identified (’00), but it is characterized by a total lack of ocecia and is believed to be an undescribed species. A negative character is, to be sure, an unsound basis, in most cases, upon which to establish species, but there can be no doubt in this instance, that owcia are not developed. A well matured colony gives many branches in which the life cycle can be read. At the anterior edge of a branch the younger polypides are found in all stages of development. In the zoccia just posterior to these, full grown polypides appear and the genital products are abundant. In still older zowcia, a young embryo appears close to the anterior edge of the orifice of each zocecium, and the polypide in such a zocecium begins to degenerate. In older zocecia, nothing is left of the polypide but a brown body, while the embryos are well developed, almost ready to hatch (Pl. XI, fig. 55, emb.; b. b.) ; in still older zowcia, regenerating polypides (re. pd.) are found in zoccia containing no embryos and from which the larvae, presumably, have escaped. The lack of the ovicell and the development of the embryo entirely within the zocecium, are not the only characters which separate this form from B. flabellata. The shape of the avicularia, and the number and position of the spines are sufficient and constant specific differences. This species has been obtained mainly in Puget Sound. It is abundant on Channel Rocks at extreme low tide, and has been dredged at San Juan Island, Puget Sound. Qt ieee Univ. CaL.PuB.Zoo1 Vou.IL. | ROBERTSON] FLATE XIV ve aH wits oad ° FHOTU-LITHL& SP ROBERTSON, DEL o MTA: = ; 7 — oy im PLATE XV. Fig. 87.—Cellaria mandibulata Hincks. Two zoecia, young stage show- ing lateral crenulated border (er. bd.). Fig. 88.—C. mandibulata. Several zowcia at an older stage, showing operculum (op.) with its four denticles (d.), the immersed owcia above, their orifices (oe. or.) only visible, and an avicularium with its large man- dible (man.). The crenulated border (cr. bor.) has formed above and he- low the zoccia, thus surrounding them. X 50. Fig. 89.—C. mandibulata. Three zowcia at a still older stage to show inerease of caleareous wall and final form of omecial orifice (0e. or.) re- sembling a small avicularium. X 50. Fig. 90.—Cellaria diffusa sp. nov. A few zowcia (¢@.) to show shape, oecial orifice (oe. or.) with the lower margin developed into a mucro (m.), and the avicularia (av.). X 50. Fig. 91—Flustra lichenoides Robertson. A few zocwcia enlarged to show erect spines (é. sp.), operculum (op.), and omcia (0e.). X 50. Fig. 92.—F. lichenoides. The dorsal surface of a few zowcia showing rcot fibres springing from the distal angle of each zowcium. lig. 93.—Flustra membranaceo-truncata Smitt. Habit sketch. Fig. 94.—F. membranaceo-truncata. A few zoecia showing arrange- ment, immersed oewcia (0e.), and avicularium of simple structure in place of zoecium (av.). X 50. [320] Univ. CAL.PuBL.ZO0L.VoLII. | ROBERTSON] PLATE XV PHOTOLITALE SR ROBBRTSON DEL. Photographs to show habit of growth. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. o PLATE XVI. 95.—Scerupocellaria varians Hincks. — 96.—Scrupocellaria diegensis Robertson. ane HARRY BEAL TORREY _. BERKELEY. THE UNIVERSITY PRESS PRICE: $0. 25 ‘UNIVERSITY. OF CALIFORNIA PUBLICATIONS ZOOLOGY.—W.-E; Ritter, ‘Editor. Price per volume $3: 50. ~ Volume 1 completed. Volume’II {in progress): : “Introduction. A.General Statement of the Ideas and the. Present. Aims and Status of the Marine Biological Association of ae Diego, by: Wm. E. Ritter: No. 1. The Hydroids of the San Diego Region, ae Harry Beal Se } In Pages 43, text figures 23. =|) one- > No. 2. The Ctenophores~ of the San Diego Region, by Harry ‘Beal eres Torrey... Pages 6, Plate.1. is $2480" x ae No. 3. The Pelagic Tunicata of the San Diego Region, Sa veRR BE the Lar-.. vacea, by Wm. E. Ritter. Pages 62,° text Pees 23, Plates 2.) Price, z 65 No. 4. The Pelagic Canetoda of the Sai Diego Region, by G: 0. Estery, Pages 120, text figures’ 62. Price, . No. 5. Non-Incrusting Chilostomatous Bryozoa of the West Gaast of North “America, by Alice Robertson. Pages 88, Plates 13. Price, ~~ 1.00, No. 6. Differentiation.in Hydroid Colonies and the Problem of Senes- In. cence; by Harry Beal Torrey. Pages 10, text-figures 4. one cover, — No. 7. The-Behavior of Corymorpha, by Harry Beal ie ee Pages 8, | Price text figures 5. £3 eRe f - x The following series in Graeco-Roman Archaéology, Egyptian Archaeology, Ameri- can Archaeology and Ethnology and Anthropological Memoirs are publications from the» Department of Anthropology: GRAECO-ROMAN ARCHAEOLOGY. Vol. 1... The Tebtunis Papyri, Part 1. Edited by Bernard P. Grenfell, Arthur S. Hunt, and J.-Gilbart aktek rages 5 Plates 3, 1903 : Price, sb OO8es Vol. 2. The Tebtuinis Papen Part 2 (in prepaianans EGYPTIAN ARCHAEOLOGY. Vol, 1. The Hearst Medical Papyrus. Edited by G..A. Reisner and A.M. Lythgoe (in press). AMERICAN ARCHAEOLOGY AND ETHNOLOGY. Vol. 2. No. 1. The Exploration of the Potter Creek Cave, by William’ J. ‘Sy Sinclair. Pages 27, Plates 14, April, 1904... Price; .40-. No. 2. The Fanguases of the Coast of California South of San © ~~ Francisco, by A. L. Kroeber. Pages 72, June, 1904. Price, - .60 No. 3:: Types of Indian Culture in Swengciee by A. L. Kroeber. Pages 22, June, 1904. : ‘ a Price, . .25° ‘3 No. 4. Basket Designs of the Indians of AGaieners California, ~. by A..L. Kroeber. Pages 60, Plates 7, January, 1905. Price, ~ .75 Vol. 3. -The Morphology of the reps. Fanguage, by gs Earle Goddard. Pages 344, June, 1905. A> Price, 3.50 ANTHROPOLOGICAL MEMOIRS, Vol. I. Explorations in Peru, by Max Uhle (ih preparation). No. 1.. The Ruins of. Moche: No. 2. Huamachuco, Chincha, Ica. No. 3. The Inca Buildings of the ane: of Pisco. UNIVERSITY OF CALIFORNIA PUBLICATIONS ZOOLOGY Vol. 2, No. 6, pp. 323-332, Figs. 1-4 December 13, 1905 CONTRIBUTIONS FROM THE LABORATORY OF THE MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO. Vb DIFFERENTIATION IN HYDROID COLONIES AND THE PROBLEM OF SENESCENCE. BY HARRY BEAL TORREY. An examination of Pacific Coast hydroids during the past few years (vid. Torrey, :02, :04) has convinced me that many if not all species change their structural type with age. Stems straight proximally may become sinuous distally. Branches which alternate during the early stages of colonial development may later originate in pairs. Length and annulation of hydro- theeal pedicels, size, proportions and.ornamentation of hydro- theeae may similarly vary with the distance from the base of stem or branch. One hydroid in particular exhibited marked changes in structure with the growth of the colony and appeared to be especially favorable material for the investigation of cer- tain questions of differentiation. This species was described some months ago as Clytia bakeri' (Torrey, :04, p. 16). The stems, usually unbranched, rise from a ereeping hydrorhiza to the height of 20 or 30 mm. For a few millimeters (2 to 5) each stem is free of polyps and is ecom- pletely annulated. The annuli, short at the base, increase grad- ually in length until they become, at the first hydranth pedicel, 1The Hydroids of the San Diego Region. Univ. Cal. Publ. Zoology, I, No. 1, 1904. 324 University of California Publications. [ ZcoLoGy two to four times as long as broad. They ultimately grade into the internodal segments of the stem, each giving rise at its distal end to one hydrotheeal pedicel. The pedicels themselves vary in length and annulation from the base of the stem toward the tip. The first pedicel may exhibit 4 to 7 annul. The last hydro- theca may possess no pedicel at all, or one of but a single an- nulus. The number to a pedicel varies with much general uni- formity from one of these extremes to the other. From base to tip, then, each stem presents two main serial changes. (1) Below the hydranth region, the cauline annuli erow progressively longer. (2) Within the hydranth region, the number of annuli to a pedicel progressively decreases. There is a strong suggestion in these facts that they depend on changes in the internal physiological conditions of growth. Since polyps are readily regenerated where stems are sectioned, a method at once presents itself for determining the differentiation at a given level of the stem at a given time. The question to answer which the investigation was first un- dertaken was: Will the stem at a given level tend to regenerate the structural type which it originally produced there; or will it produce instead, a structural type characteristic of another region? For example, will a cut through the portion of the stem where each hydrothecal pedicel has five annuli initiate the re- generation of a pedicel with five or with one annulus, the number characteristic of the distal region ? The facts obtained from the experiments fall into three cate- gories, according as the regeneration followed (1) a cut through the distal portion of the stem, (2) a cut through the middle por- tion, (3) heteromorphically from the cut basal end. In all, 16 operations were performed. Of these, 7 were unsue- cessful. Among the remaining 9, the cut in 5 had been made through the distal region, 3 through the middle region, and 6 had regenerated heteromorphic stalks. The small number of cases involved is offset by the clearness of the results. The record of experiments in the first category is as follows: I—July 26, No. 2. Stem with 24 hydranths and terminal bud. Enumerating from base to tip, \Vou.2] Torrey.—Hydroid Differentiation and Senescence. 325 Pedicels 1, 2 with 3 annuli Pedicels 3-8 with 4 annuli Pedicels 9-11 with 3 annuli Pedicels 13-17 with 1 annulus Pedicel 18 with 0 annuli Pedicel 23 with 0 annuli Pedicel 24 with 1 annulus 10am. Cut off terminal bud, 24, 23, and part of 22. July 28,10 a.m. New hydranth No. 22 regenerated within old eup, on a short stalk from which hydrotheca is separated by a single constriction and is therefore sessile. Result: regeneration is according to the local, not the basal structural type. Il.—July 26, No. 3. Stem with 17 hydranths and terminal bud. Pedicel 1 with 3 annuli Pedicel 2 with 3 annuli Pedicel 3 with 4 annuli Pedicel 7 with 1 annulus Pedicel 9 with 1 annulus Pedicel 11 with 0 annuli Pedicel 13 with 0 annuli Pedicel 15 with 0 annuli Pedicel 17 with not even nodal constriction 12 m. Removed terminal bud, 17, 16, and part of 15. July 27, 9:15 a.m. No. 16 has regenerated as a bud on short stalk; no constriction whatever between latter and hydrotheca (fig. 1). Though there are two cut surfaces, but one bud has been formed, in place of the originally proximal individual. No. 16 developed later into a hydranth, the skeleton remaining in the same condition except for a slight general thickening. Result: regeneration is according to the local, not the basal structural type. III.—Aueust 5, No. 4. Stem with 15 hydranths and terminal bud. Pedicels 1-4 with 2 annuli Pedicels 5-11 with 1 annulus Pedicel 12 with 0 annuli Pedicel 13 with 3 annuli Pedicels 14-15 with 1 annulus Pedicel 16 with 0 annuli 326 University of California Publications. [ ZooLocy 8:45 a.m. Removed terminal bud, 15, 14, and 13. August 7, 9:45 a.m. Nos. 13 and 14 have regenerated, each with a single annulus in the pedicel. August 8, 2 p.m. Stem segment with bud of 15 has ap- peared; single annulus below hydrotheca. Result: all the regenerated parts accord with the structural type characteristic of the distal region (fig. 2). IV.—August 5, No. 5. Stem with 17 hydranths. Pedicel 1 with 8 annuli Pedicel 2 with 6 annuli Pedicel 3 with 4 annuli Pedicel 4 with 3 annuli 3 Pedicel 5 with 2 annuli Pedicel 6 with 4 annuli Pedicel 7 with 3 annuli Pedicels 8-9 with 2 annuli Pedicels 10-11 with 3 annuli Pedicels 12-15 with 2 annuli Pedicel 16 with 1 annulus Pedicel 17 with 0 annuli 8:50 a.m. Removed 17 and 16. August 11. No. 16 has regenerated. Pedicel with 1 (pos- sibly 2) annulus and 16-18 tentacles. Result: this stem was somewhat irregular with regard to the number of annuli in successive pedicels. The regenerated ped- icel, however, corresponds with the local structural type. V.—August 5. No. 6. Stem with 13 hydranths and terminal bud. Pedicel 1 with 6 annuli Pedicel 2 with 4 annuli Pedicel 3 with 5 annuli Pedicel 4 with 4 annuli Pedicel 5 with 3 annuli Pedicel 6 with 4 annuli Pedicel 7 with 3 annuli Pedicel 8 with 1 annulus Pedicel 9 with 3 annuli Pedicel 10 with 2 annuli Pedicels 11-14 with 1 annulus 8:55 a.m. Removed terminal bud, 13 and 12. Vou. 2] Torrey.—Hydroid Differentiation and Senescence. pal August 8. Nos. 12 and 13 regenerated, each with 1 annulus in pedicel. Result: regeneration according to the local, not the basal structural type. Further experiments of this sort are unnecessary to show con- Figs. 1, 2. Normal regeneration from distal wound. Fig. 3. Normal regeneration from middle region of stem. Fig. 4. Heteromorphic regeneration from base of stem. The arrow indicates in each case the level of the cut. 328 University of California Publications. [ ZooLocy clusively that, under the conditions of the experiments’ regen- eration from cuts through the distal region of the stem produces structures of the local, 2.e., distal type. Experiments belonging to the second category; cuts made through the middle region of the stem. I.—August 5, No. 8. Stem with 17 hydranths and terminal bud. Pedicel 1 with 5 annuli Pedicels 2—5 with 4 annuli Pedicel 6 with 6 annuli Pedicel 7 with 3 annuli Pedicel 8 with 5 annuli Pedicel 9 with 2 annuli Pedicel 10 with 4 annuli Pedicel 11 with 1 Pedicel 12 with 2 Pedicel 13 with 1 Pedicel 14 with 3 annuli Pedicel 15 with 1 annulus 9 1 1 annulus annuli annulus Pedicel 16 with Pedicel 17 with Pedicel 18 with annuli annulus annulus 10 am. Cut stem between 6 and 7. August 8, 3:30 p.m. No. 7 has regenerated without annuli immediately below the hydrotheca. Result: regeneration according to the distal, not the local type. I].—Auegust 5, No. 9. Stem with 17 hydranths. Pedicel 1 with 5 annuli Pedicels 2—3 with 4 annuli Pedicel 4 with 7 annuli Pedicels 5-6 with 4 annuli Pedicels 7-9 with 3 annuli Pedicels 10-11 with 2 annuli Pedicel 12 with 3 annuli Pedicel 138-17 with 1 annulus 11 am. Cut stem between 4 and 5. +The stems under observation were removed from the hydrorhiza and rested on the bottom of flat glass dishes containing about 300 ¢.c. of sea- water. None of the colonies took food during the course of the experiment beyond what they could get from the original supply of water, which was not changed. There is no reason to suppose that the results depend upon the conditions to which the stems were subjected, for several observations of regeneration in nature indicated that results are essentially similar in the two cases. Vou.2} Torrey.—Hydroid Differentiation and Senescence. 320 August 8, 9:45 am. Hydrotheea of No. 5 with 1 annulus immediately below it, supported on a segment of stem with an annulus next the cut (fig. 3). Result: regeneration according to the distal, not the local type. IlI.—Aueust 5, No. 10. Stem with 20 hydranths and ter- minal bud. Pedicel 1 with 5 annuli Pedicel 2 with 0 annuli Pedicel 3 with 0 annuli Pedicel 4 with 7 annuli Pedicel 5 with 5 annuli Pedicel 6 with 7 annuli Pedicels 7-9 with 3 annuli Pedicels 10-21 with 1 annulus 11:10 am. Cut stem between 6 and 7. August 7, 10:30 am. No. 7 has regenerated in a manner practically identical with that of the preceding experiment. Result: regeneration according to the distal, not the local type. The three cases in the second category seem to show that the mid regions of the stems no longer possess the capacity to pro- duce the type of structure originally arising there, but that they do produce a type of structure peculiar to the latest formed portion of the stem. During their growth, the stems appear to have changed their character throughout their length. The facts belonging to the third category concern the cases of heteromorphosis from the proximal cut end of the stem. I.—Aueust 5, No. 4 (see above). August 8, 10 a.m. Heteromorphic stem. August 11. Heteromorphie stem with terminal hydranth. Immediately below the hydrotheca are 3 annuli. The rest of the stem is similar in all respects to the heteromorphic stem in fie. 4. IIl.— August 5, No. 8 (see above). August 11. Heteromorphic stem similar in all respects to preceding (1), with the exception that development had stopped before a hydranth had appeared. 330 University of California Publications. [ ZooLocy III.— August 5, No. 7. Stem with 11 hydranths and termi- nal bud. Pedicels 1-2 with 6 annuli Pedicel 3 with 2 annuli Pedicel 4 with 5 annuli Pedicel 5 with 3 annuli Pedicel 6 with 1 annulus Pedicel 7 with 2 annuli Pedicels 8-11 with 1 annulus August 8, 3:30 p.m. Heteromorphie stem with three annular basal segments, a long intermediate segment and one annulus immediately below the terminal bud. IV.—August 5, No. 10 (see above). August 8, 4:30 p.m. Heteromorphie stem differing but slightly from preceding (III), surmounted by a hydrotheea (fig. 4). Latter small, and diaphragm not apparent. Hydranth at- tached to wall by a number of amoeboid strands which are com- mon along the stem but are seldom met with in hydrotheca. V.—August 5, No. 9 (see above). August 8, 4:15 p.m. Heteromorphie stem with 3 basal an- nuli, a long stem segment and a poorly defined annulus imme- diately below the hydrotheca. Shoulder process on side of long stem segment carrying growing stalk of normal type. VI.—August 5, No. 5 (see above). August 11. Heteromorphic shoot with terminal hydrotheca. Three basal annuli and a very long non-annulated stem segment which passes directly into the hydrotheea, without an interven- ing constriction or annulus. Disregarding for a moment the stalk, the number of annuli occurring immediately below the hydrotheca is typical of the latest formed (distal) region of the normal stem in 4 of the above 6 cases. In the other 2 cases, the number is larger than the typical distal number on the same stem but is smaller than the proximal number on one of the stems (II). It exceeds the proximal number on the other stem which, however, is excep- tionally small (1). The tendency to develop according to the Vou. 2] Torrey —Hydroid Differentiation and Senescence. 331 distal type is therefore conspicuous so far as the pedicels are concerned. With respect to the stalks, it may be said that they begin always with one or two short segments characteristic of the earliest formed portion of the normal stem. ‘These segments may even be shorter than the parent segment next the wound. It would seem that in this particular the regenerating stem de- velops according to the embryonic type. But the duration of this type of development is so short, lasting through the forma- tion of one or two segments only, that it closely resembles what has already been seen in the figures of regular regenerations in the lower regions of the hydranth-bearing zone, viz., that the new structures are almost invariably initiated by the for- mation of an annulus or part of one. General considerations. In seeking an explanation which shall simplify as well as summarize the results presented in the foregoing paragraphs, I think we must pass by any hypothesis which rests solely upon a basis of morphological determinants. That regeneration at a given level may not reproduce the struc- tural type characteristic of that level, while it does reproduce the type characteristic of a later level of the stem is a fact that is hardly simplified by the assumption of a residual germ plasm. So, too, does it seem improbable that the structural type is the result of a functional balance between an organism possess- ing an unmodified regenerative capacity and the conditions sur- rounding it. This view would necessitate a change in the envi- ronment between the time a polyp first appears and the time it is regenerated in a somewhat different form. But no such change is evident. The polyps half way up a stem were sub- jected during their development to external conditions essen- tially identical with those which surrounded the developing indi- viduals distal to yet differing from them. The facts, however, appear to give strong support to the view that the stem, instead of retaining unmodified its regener- ative capacity, actually loses with age its ability to produce structures which formerly characterized it; and that this is owing to a modification of conditions within the organism, which 332 University of California Publications. [ ZooLocy govern its behavior without being necessarily a part of it. These conditions are probably chemical in nature, intimately con- cerned with the metabolism. A destruction or addition of sub- stance or substances in the course of the development is readily conceivable as the efficient cause of the structural modification. The relation between internal and external conditions is under consideration. The resemblance of the phenomena of colonial differentiation in OC. bakeri to the phenomena of senescence is so strong as to suggest a similar interpretation for both. The experiments will be continued. September 1, 1905. UNIVERSITY OF CALIFORNIA PUBLICATIONS ZOOLOGY Vol. 2, No. 7, pp. 333-340, Figs. 1-5 December 13, 1905 CONTRIBUTIONS FROM THE LABORATORY OF THE MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO. Vit: THE BEHAVIOR OF CORYMORPHA. BY HARRY BEAL TORREY. In a former paper ( :04),1 it was shown that Corymorpha pos- sesses an unusually wide range of activities for a hydroid. It responds to mechanical and thermal stimuli, to chemieal stimuli that produce their effect as mechanical irritants, not as odorous substances, and to gravity. In the movements of the stem, it resembles such naked forms as Hydra, Clava and Hydractinia. The stem is everywhere sensitive to mechanical stimuli, bending from side to side or shortening under their influence. Such re- actions are due to the contractions of longitudinal muscle fibers which are situated in the usual fashion deep in the ectoderm. A lengthening of the stem may be caused wholly or in part by the circular endodermal musculature. The long proximal tentacles, about twenty-four in number, may respond singly to direct mechanical stimulation or to a stimulus applied either to another tentacle or a distant portion of the stem. In all eases of effective stimulation, direct or indi- rect, they bend invariably toward, never away from the mouth. The reaction is rapid; the recovery, which ends in the resumption of the expanded condition, is slow. The initial reaction of the distal tentacles to all effective stimuli is, on the other hand, always away from the mouth; * Biological Studies on Corymorpha. I. C. palma and Environment. Jour. Exp. Zoology. I, No. 3, 1904. 334 University of California Publications. [ ZooLoGy after which, strong irregular movements toward and away from the mouth appear. The proboscis, which carries the distal ten- tacles to the number of about fifty in a crown around its summit, may react to mechanical stimuli, directly or indirectly applied, by bending in the direction of the stimulus. These reactions of tentacles and proboscis constitute an effi- cient prehensile mechanism. A small object which happens to stimulate a proximal tentacle on its oral side is at once swept toward the mouth. The distal tentacles, which may be carried toward it by the bending of the proboscis, then move outward, meeting and transporting it by a subsequent inward movement to the mouth. All the motions thus far deseribed are due to muscular con- tractions. Locomotion is produced, as in Hydra, by the activi- ties of amoeboid cells at the base of the stem. And the circu- lation of fluids in the coenosareal canals is accomplished by the usual ciliary action, supplemented by the expansions and con- tractions of the proboscis and stem. Aside from these types of motion, the stem exhibits a marked geotropism, assuming when at rest a vertical position. This orientation does not appear to be dependent in any way upon museular activity. The behavior not only of the stem as a whole but of pieces of one-eighth or one-tenth its length from the base or various other regions, indicates that the stem is everywhere sensitive to the stimulus of gravity and furnished with an effi- cient mechanism for bringing about a response. This mechanism is expressed, I believe, in the large, highly vacuolated cells of the endodermiec axis which forms the core of the stem. Changes of orientation, according to this view, are produced by relative changes in the turgidity of such cells on opposites sides of the stem. Whether the orientation is to be reckoned as a contrac- tion phenomenon, though this is probable, cannot be said defi- nitely at this time. It is indeed a fact that the axial cells may not only decrease in size, as when the stem is shortened, but also increase in size, as when the stem is lengthened without loss in diameter. + a ae Oe capt a cn ema Vow. 2] Torrey.—The Behavior of Corymorpha. 335 Two conclusions which were formerly held, later experiments have shown to have been founded on data which were mislead- ing owing to the conditions of experimentation. The first was that a change in the polarity of a region is accompanied by a change in the reactions of the axial cells in this region. The second was that, regardless of the point at which it is supported, whether proximally or distally, the stem would orient itself ver- tically, distal end uppermost. These results were obtained on individuals which had been kept in the laboratory during the warm days of summer for a week or ten days, and did not be- have with the constancy or the precision which characterize the actions of the individuals observed last March. The latter were used for experimentation immediately upon their capture, and were kept under conditions which permitted a vigorous, healthy existence. With regard to the first conclusion, it may be said that while such a change of polarity as heteromorphosis of the proximal end of a stem segment would be accompanied by an upturning of this end, the result would not be achieved by a change in the reactions of the axial cells in this region. The essential factor lies rather in the relation of the region in question to the sub- stratum. In studying the regeneration of Corymorpha, the ob- servation was frequently made that from pieces resting on the floor of the aquarium, cut from the distal half of a stem, U- shaped figures would be formed, fastened to the floor by the loop, the two arms extending vertically upward, each crowned with sets of developing tentacles. By the side of these hetero- morphic pieces were many others fastened to the substrate by one end, which possessed an. incipient holdfast, the other end developing tentacles. In both cases, the behavior of the axial cells was constant; the cells nearer the center of the earth were relatively larger than those on the other side of the stem. The proximal ends of the pieces which developed holdfasts remained lowermost merely because they were adhesive and clung to the substrate, which the distal ends could not do. This fact appeared distinctly in the results of my recent experiments bearing directly upon the second conclusion stated 336 University of California Publications. | ZooLoGy above. I had found previously that when stems relieved of hydranths and the weight of sand grains adhering to the hold- fasts were supported at their distal ends, they would assume the normal vertical orientation, proximal end down. Last March, the experiment was repeated many times, with special precau- tions against errors. Fresh animals were used, and their distal ends were fastened by a single loop to a rigid thread. The sup- port thus afforded was secure and ample for various movements. In every case, without regard for the amount of holdfast pres- ent, the unsupported proximal end travelled wpward and came to rest only when the stem had reached a position approximately vertical, but wpside down. Comparing this result with the movements of stems attached by the proximal end, either resting on or hanging from the sub- strate, it is obvious that no reversal of behavior in the axial cells need be assumed. The variable factor is connected with the point of support. By the foregoing observations, Corymorpha seems to be re- moved from the category of animals for the elucidation of whose behavior none of the familiar mechanical explanations of geo- tropism seem to apply. Its geotropic reactions appear now to accord with the theory which Davenport formulated with refer- ence to the geotropism of free-swimming organisms. There is a difference between the resistance encountered by the stem as it moves upward (friction plus weight) and the resistance it en- counters when it moves downward (friction less weight). This difference is expressed in the stem by a tension on its upper side when it is inclined in any degree from the vertical. There is no tension on the lower side of the stem unless it be hanging downward at some angle; in which ease it never equals the ten- sion on the upper side except when the stem is vertical, its posi- tion of rest. The evidence formerly presented in favor of the view that the axial endoderm cells, and not muscles, govern the geo- tropie orientation of the stem, has been strengthened to some extent by a further experiment. In my original experiments, euts were made at frequent intervals half way through the stem . — —— “or VoL. 2] Torrey—The Behavior of Corymorpha. 337 on one side, or alternately on both sides. The stem was then laid on the floor of the aquarium, a cut side uppermost. In one to two hours, the stem had assumed an erect position. The cuts were intended to destroy the effectiveness of the muscles of the stem, leaving the column of axial endoderm cells intact. The muscles as a whole were weakened by the cuts. And the slowness of the reaction, coupled with the fact that the orientation was accomplished before the wounds closed, strengthened the view that muscles were not concerned in the reaction. Other facts supporting the same conclusion have since been obtained. A piece was removed from a stem by two cuts which 1 2 3 formed with each other an obtuse angle (fig. 1). On the cut faces, the axial endoderm was exposed. The rounded surface representing the original stem wall was still covered with ecto- derm, provided with longitudinal muscles. Figs. 2 and 3 were drawn from this piece when it was in contraction and expansion respectively. It will be noticed that in the contracted condition (fig. 2), the course of the longitudinal muscle fibers is no longer 338 University of California Publications. | ZooLoGy straight; the angle made by the cut surfaces is greater than it was at first; there is a tension factor on the side of the stem opposite the layer of muscles which tends to draw the remotest edges of the muscle layer toward each other against the pull of the muscles themselves. In the expanded condition (fig. 3), though the piece lengthens, it bends strongly toward the cut surfaces. The latter now form an angle greater than 200 de- grees. At the upper end of the figure, the tip of the piece has curled sharply over toward the wound. The tension is still more apparent here than in fig. 2. On the assumption that the axial endoderm cells have lessened their volume on the side to- ward the wound relatively to the volume of the axial cells on the side away from the wound, the configurations shown in figs. 2 and 3 are intelligible. The axial cells themselves did not push out between the edges of the wound but gave every indication of being under restraint. f 5 When long, thin, oblique slices are cut from a stem as in fig. 4, it follows inevitably that the narrow wedge-shaped ends curl toward the eut surface, as in fig. 5. The relatively de- creased turgidity of the axial cells next the wound appears to account for this condition also. In the light of these facts, it becomes clear that the axial endoderm cells under certain conditions are capable, by chang- VoL. 2] Torrey—The Behavior of Corymorpha. 339 ing their turgidity, of producing movements in the stem com- parable with those which are due to stimulation by gravity, though the evidence does not demonstrate that gravity itself affects them, directly or indirectly. But they are active, not passive elements, which increases the probability that they may be concerned in the geotropic response. If it be true, as seems probable, that the axial cells do govern the geotropie response, Corymorpha stands alone among the met- azoa in possessing a tropic mechanism distinet from the body musculature. In a recent paper, Holmes (:05) has shown that Ranatra exhibits uncommonly clear cut reactions to light. Here the tropic mechanism involves the same sense organ, the same musculature and to some extent at least the same nervous ele- ments which serve in responses to other classes of stimuli, in- ternal and external. Though Ranatra is an unusually favorable object for the analysis of the tropic mechanism, the confusion of various factors operating over the same lines renders an ade- quate analysis impossible at present. The presence of a geo- tropic mechanism distinct from the musculature in Corymorpha simplifies the problem. And I am led to believe, further, that the simply organized nervous system of the hydroid need play no part in the geotropie reaction. The axial endoderm cells probably change their volume under a given tension which is apphed directly to them. Each cell may be considered a unit, not necessarily dependent in its actions upon its neighbors, act- ing with them only in so far as they may be subjected simulta- neously to similar stimulation. The response of Corymorpha to gravity is strikingly similar to the negative geotropism of the caulicles of plant seedlings, which is unquestionably dependent upon volume changes in the stem cells. The latter are again strikingly similar in structure to the axial cells of Corymorpha. There appears to be in the hydroid mechanism no more opportunity for a pleasure-pain type of reaction than among the plants; and there is no sign of it in the geotropic movements of the stem. Neither are there signs of trials and errors. The movement of the stem is very eradual, very definite, very direct. I know of no animal which 340 University of California Publications. | ZOOLOGY: more closely approximates the plant in structure and tropic re- sponse. If the behavior of the one be explicable on the basis of direct reactions to stimuli, of the reflex type, I do not see how the behavior of the other can be excluded from a similar inter- pretation. UNIVERSITY ‘OF CALIFORNIA ‘PUBLICATIONS a, ZOOLOGY. —W. E, Ritter, . Editor.. Price per volume $3. 50. _Volume » I completed... Volume II (in progress): Introduction. A General Statementof the Idéas and the Present Aine Pre Status of the Marine Price Association of his DIES; by zi Wm. E. Ritter. No. 1. The Hydroids of the San Diego. Region, ‘by pany. Beal POrEey f Riee fe ze Pages 43, text figures 23. be ae Be. » No. 2, The Ctenophores of the San Diego Sel us by Harry Beal | RHEE: : “Torrey. Pages 6, Plate 1.- _ J No. 3. The Pelagic Tunicata of the San Diego Region, excepting the Lar. ~--wacea, by Wm.-E. seed ae oeee 62, text Sa 23, Plats 2: bs : Price, " Oo. No; 4. The Pelagic Gorspods of ie San Diego Region, by C. 0. Estey Pages 120, text figures 62. Price, No. 5. Non-Incrusting Chilostomatous Bryozoa of the West Gaist of North | America, by Alice Robertson. Pages 88, Plates 13..Price, 1.00 No. 6; Differentiation in Hydroid Colonies and the Problem of Senes- In~ cence, by Harry Beal Torrey. . Pages 10, text figures 4, - one cover, at tee No. 7. The Behavior of Corymorpha, by Harry Beal Torrey. Pages 8, | Price text figures 5. eae Ce No. 8. Dinoflagellata of the San Diego Region.—I. On Heterodinium, a New Genus of. the Peridinidae, by Charles Atwood ae Pages 26, text figures 2, Plates 3. Price, . The following series in Graeco-Roman Archaeology, Egyptian Archaeology, Ameri- — can Archaeology and Ethnology and Anthropological Memoirs are publications from Hie Department of Anthropology: GRAECO-ROMAN ARCHAEOLOGY. Vol. 1. The Tebtunis Papyri, Part I. Edited by Bernard P. Grenfell, Arthur S. Hunt,-and J. Gilbart ocak Pages 690, Plates 3, 1903 . Price, $16.00 Vol. 2. The Tebiunts Piya: Part 2 (in wrepardtiony. EGYPTIAN ARCHAEOLOGY. Vol..1.* The Hearst Medical Papyrus. Edited by G. A. Reisner and A.M, — Lythgoe (in press). : AMERICAN ARCHAEOLOGY AND -ETHNOLOGY, - Vol. 2. No. 1. The Exploration of the Potter Creek Cave, by William J.- ras Sinclair. Pages 27, Plates 14, April, 1904 . Price,- .40° No. 2. The Languages of- the Coast of California South of San Francisco, by A. L. Kroeber. Pages 72, June, 1904, Price, -60 No. 3. Types of Indian Culture in California, by A. L. Kroeber. Pages 22, June, 1904. : 2 ‘ Price, ~ .25 No. 4. -Basket Designs of the Indians of Northwestern California, by A. L. Kroeber. _Pages 60, Plates 7, January, 1905. -Price,...75 Nol. 3. The Morphology of the St Language, ae Bite Earle Goddard. Pages 344, June, 1905. Price, 3.50 ANTHROPOLOGICAL MEMOIRS. Vol. 1. Explorations in Peru, by Max Uhle (in preparation). No. 1. The Ruins of Moche. No. 2... Huamachuco, Chincha, Ica. ae No. 3. The Inca Buildings of the Valley of Pisco. - UNIVERSITY OF CALIFORNIA PUBLICATIONS ZOOLOGY Vol. 2, No. 8, pp. 341-368, Pls. 17-19 January 6, 1906 CONTRIBUTIONS FROM THE LABORATORY OF THE MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO. Vili. DINOFLAGELLATA OF THE SAN DIEGO REGION.—I. ON HETERODINIUM, A NEW GENUS OF THE PERIDINIDAE BY CHARLES ATWOOD KOFOID. The investigations of the plankton of the Pacific at the San Diego Station during the past three years has brought to heht a number of species belonging to the family Peridinidae which do not conform to any known genus. They all present in com- mon a number of characters which eall for the establishment of a new genus for the reception of the several species represented. To this genus also belong seven species described by Murray and Whitting (799) from the tropical Atlantic as members of the genus Peridinum to wit: Peridinium hindmarchu, P. milneri, P. blackmant, P. leiorhynchum, P. trirostre, P. tripos, and P. doma. Gonyaulax triacantha Jorg. is also to be referred to this genus. All of these species except P. trirostre and Gonyaulax triacantha have been seen by me and a fuller discussion of their structure will appear elsewhere. In the following pages the brief discussion of each is based upon Murray and Whitting’s figures. I am indebted to Miss E. J. Rigden, assistant in the summer of 1904 at the San Diego Marine Biological Station, for some of the sketches utilized in the plates accompanying this paper and also for the skill and thoroughness of her examination of the pO ee Byatt si gy tae Pg ; AY ACT STi 1Sitypy f f - % 342 University of California Publications. [ ZOOLOGY plankton which brought to light the most of the species here dis- cussed. The types and cotypes of the species here deseribed for the first time are in the collections of the University of California. Heterodinium gen. nov. The form of the theca resembles that of Peridinium in the presence of a median or somewhat postmedian girdle which en- circles the theca at its greatest diameter and in two antapical horns, always directed posteriorly or nearly so. The posterior margin or lst of the transverse furrow is suppressed or feebly developed in comparison with the anterior one, especially at its distal end, while the anterior one, as if in compensation, is often excessively developed. The suture lines are demonstrated with great difficulty and some uncertainty in most of the species. The plates are as follows: three apicals, one left interealary, six premedians, seven postmedians, one furrow plate, and three ( ?) antapieals, as shown in the accompanying text figures. On the ventral face about midway between the apical and flagellar pores is a small pit or pore-lke area in the mid ventral suture. An actual opening in this area has not been demon- strated. In the suppression of the posterior border of the girdle, in the number and arrangement of the plates and in the presence of the ventral pit on the epitheea Heterodinium differs from Peridinium, although in form and general appearance species of the two genera strongly resemble each other. In some species and possibly generally in the genus there is a decided asymmetry to the theea brought about by a torsion of the body on the main axis in clockwise direction, looking from the posterior toward the anterior end. This is especially notice- able in the scoop-shaped forms such as H. scrippst. DETAILED DESCRIPTION. The following is a more detailed description of the characters found in the genus. The theca is expanded in the equatorial region, with more or less well marked dorso-ventral flattening and ventral excavation near the flagellar pore. It is spheroidal, Von. 2] Kofoid—On Heterodinium. 343 ellipsoidal, elongated, rotund, flattened, or even scoop-shaped. The length always exceeds either diameter, and the transdiam- eter at the girdle equals and more often exceeds the dorso- ventral one. The greatest transdiameter is usually at the girdle but in some species the epitheca or hypotheca may exhibit a slightly greater diameter. The greatest dorso-ventral diameter is at the left of the flagellar pore. The epitheca is usually not contracted to an apical horn though in some species a short horn is present, and in others the elongated epitheca tapers gradually from the girdle to the apical pore with more or less coneavity of the lateral margins. The anterior end of the epitheca is-more often broadly rounded, being dome-shaped in the rotund species and like the end of an ellipsoid or even seoop-shaped in the flattened species. In some instances, as in H. blackmani, the epitheca is rotund at the girdle but flattened distally. The altitude of the epitheca is usually less than the transdiameter and exceeds it in only a few eases as in H. blackmani and H. hindmarchi. The ventral face of the epitheca is flattened and somewhat excavated, slightly in rotund species, more deeply and. exten- sively even to the lateral margins in the dorso-ventrally com- pressed forms. The mid ventral face is marked by the slightly sinuous suture ridge which runs from the flagellar pore to the apex and, bears midway a pit or pore-like area, a characteristic structure in the genus. This varies greatly in distinetness and in the breadth of the widened smooth suture ridge in which it is placed. The apical pore is inclined, even as much as 10° in some species, to the right and is usually well defined though rarely protuberant. The hypotheca is usually subequal to the epitheca, being longer in H. milnerit and H. sphaeroideum, and shorter in H. blackmani. The posterior end may or may not show a bifurea- tion into antapical horns. It may be broadly rounded, dome- shaped and without any antapical differentiations as in H. sphaeroideum and H. doma, with mere spinules with or without lists as in H. milneri and H. murrayi, with slight median bifur- cation as in H. whittingae or with typical elongated antapicals 344 University of California Publications. | ZOOLOGY as in H. blackmani. In the form of hypotheca Heterodiniwm thus exhibits a development parallel to that found in the allied genus Peridinium. The ventral face of the hypotheca is chan- neled by the longitudinal furrow plate which in many species extends anteriorly so that it indents the epitheca above the flagellar pore. The girdle is usually submedian in position, though preme- dian in some species as for example in H. milneri, or postmedian in others as in H. scrippsi. The girdle is much more oblique in the flattened than in the rotund forms. In H. whittingae its plane is inclined ventro-posteriorly at an angle of 45° to the axis. The girdle in all species thus far observed forms a de- scending right spiral with a displacement accelerated distally and amounting to 1-3 times the width of the furrow. The most characteristic feature of the girdle is its incompleteness distally and the absence or slight development of the posterior ridge. The furrow is bounded anteriorly by a heavy overhanging ridge which in species thus far observed is not a ribbed fin or lst but a heavy projection of the thecal wall. The posterior border is formed by a less salient ridge which becomes less prominent dis- tally and often diverges more widely from the anterior ridge towards its distal end . The feature of a more or less deficient posterior margin of the girdle is a constant character in all species thus far observed save the imperfectly known H. sphae- roideum which has, however, the ventral plates at least (the dorsal ones are not known) of the genus. The transverse furrow is indented in the thecal wall and the flagellar pore is found at its proximal end. The theeal wall is made up of discrete plates, which, however, are much less easily separated and much less clearly defined than they are in other genera of the family, as for example in Peridi- nium. The sutures are marked by flattened ridges or bands or smooth tracts in which the cleavage line may be traced in some cases in young individuals. The suture bands often have a secondary reticulation of minute polygons on their surface and are best seen on a deep focus. They are differentiated on the inner as well as the outer thecal surface. The epitheca consists VoL. 2] Kofoid.—On Heterodinium. ~ 345 of ten plates (figs. A and B) arranged as follows: three apicals about the apical pore, of which one (1) is a wide dorsal plate covering the dorsal half of the anterior end, and the other two (2,3) are ventral and are separated from each other by the mid- ventral suture which runs from the apical to the flagellar pore. Between the dorsal and the two ventral plates are lateral sutures which in subgenera Platydinium and Euheterodinium become very heavy and much more prominent than any other sutures Fig. A.—Ventral view of Heterodinium scrippsi showing thecal plates; apicals, 1-3; premedians, 5-10; left intercalary, 4; postmedians, 11- 17; antapicals, 18-20. x 450. Fig. B.—Dorsal view of same. X 450. of the theea. In some eases as in H. scrippsi and H. blackman these lateral ridges are doubled, suggesting a narrow compressed plate, but in the more rotund species they are not doubled and there is no suggestion of the presence of such a plate. I there- fore conclude that they are merely doubled margins and are not to be regarded as the margins of degenerate plates. Anterior to the girdle is a premedian series (figs. A, B, 5—10) of 6 plates of which two are ventral (5, 10), two dorsal (7, 8), 346 University of California Publications. | ZOOLOGY and one each right (9) and left (6). They are not as a rule symmetrically placed because of the considerable irregularity in their size and especially because of the disturbing effect of the left intercalary plate (4) which is found between premedians 5, 6, 7 (or 6 and 7) and apicals 1 and 3. This interealary plate is often small and in compressed species is not easily found. It is shown clearly in H. blackmani, but is merely suggested or not shown at all in the rest of Murray and Whitting’s (’99) species. Its demonstration in all species carefully examined by me leads me to expect it in others especially since they usually show the tilting of the apical pore to the right, the slight shoulder on the left marginal outline and the asymmetrical arrangement of the premedians which attend its presence in species in which it has been demonstrated. It is greatly reduced in size in flattened species such as H. whittingae, and it is certainly possible that this plate may be entirely suppressed in some of the flattened species of the subgenus Platydinium, though no conclusive evi- dence to that effect is at hand. The ventral face of the epitheea of the flattened species is formed by the two ventral-apical plates (2, 3) and the two ventral-premedians (5, 10) but in the more rotund forms the lateral-premedians (6,9) are partially exposed in a ventral view. The left interealary in the flattened species is dorsal in position, but in the rotund forms as H. murrayi and H. doma it appears on the left shoulder in the ventral view, and may be shifted dorsalwards so that it does not touch the ventral premedian 5. The girdle plate has the form of a trough-like band as in Peridinium and appears to be variously subdivided by reticular ridges some of which may be suture lines. The plates of the hypotheea are less clearly defined than those of the epitheca, especially on the ventral face. There are seven postmedians (11-17) adjacent to the girdle, 3 dorsal (13-15) and 4 ventral (11, 12, 16, 17) one of which (16) appears to extend to the tip of the right antapical without subdivision into anterior postmedian and posterior antapical moieties, ex- cept possibly in H. sphaeroideum. The separation of the adja- eent postmedian (17) from the ventral median antapical is VoL. 2] Kofoid.—On Heterodiniwm. 347 often feebly expressed. The antapical series includes 3 plates, a single dorsal (18), a median ventral (19), and a left ventral (20). The distal end of the posterior list of the girdle usually descends on the suture between the right ventral and right latero- ventral postmedians and continues to the tip of the right antap- ical horn. The boundaries of the plates on the right ventral face of the hypotheeca are subject to much obscurity and considerable varia- tion in loeation. The relations of the midventral plate (19) to ( are subject to considerable variation in connection with the vary- the adjacent postmedian (17) and to the longitudinal furrow ing degrees of development of the posterior list of the girdle. The longitudinal furrow is subject to considerable variation in length. It is relatively short in some species as in H. rigdenae, H. sphaeroideum, and H. hindmarchi where it is less than two- thirds the length of the hypotheea. In other forms as H. tri- rostre and H. murrayi it reaches the antapical border. It is without high membranous lists as a rule though one appears in HH. murray. The thecal wall is thin and hyaline and universally, except in apparently young individuals, reticulate with more or less irregular polygons formed by thickened ridges on the outer sur- face. In some species, as in H. scrippsi, and perhaps in certain stages of growth of other species these polygonal boundaries be- come so prominent as to obscure suture lines, as may be seen in Murray and Whitting’s (’99) figures of H. trirostre, H. murrayt, and H. hindmarchi. These reticulations are often quite regular as in H. blackmani and along the lateral margins of the epitheea of H. scrippsi, or very irregular as on the ventral and dorsal faces of H. scrippsi and in H. trirostre ; they may be relatively large as in H. murrayi or small as in H. sphaeroideum, with very heavy ridges as in H. trirostre or but faintly outlined as in H. milneri, or forming but a delicate network as in H. blackman. In young individuals they may be entirely lacking and the presumption is that in gen- eral, individuals with partially or feebly developed reticulations have not as yet reached the stage of completed formation of the 348 University of Californa Publications. [ ZooLoGy theea. The reticulations are found also on the girdle plate and on the girdle lists and along suture lines there are frequently rows of smaller polygons. In but a few cases as in H. murrayi and H. doma are enclosing ridges so thickened by the filling in of the angles as to leave a subcireular central area. Each reticu- lation has typically one centrally located pore. In some species with coarse reticulations there are several pores in a single area, and frequently in all species there are minor irregularities in the number and position of the pores. Small polygons fre- quently lack the pores. The reticulation is evidently formed on the outer surface of the theeal wall by plasma which is extruded through the pores, for the polygons bear a definite relation to the arrangements and distribution of the pores. The protoplasmic contents of the theca are usually hyaline and colorless, and often only partially fill the interior of the theea. The nucleus has the usual ellipsoidal form with beaded chromatin reticulum and lies near the center of the protoplasmic mass not far from the flagellar pore. It is small and is found with difficulty. No instances of diffuse reddish coloration often seen in Peridinitum and Pyrophacus have been noted as yet in Heterodinium. Chromatophores are entirely absent in some in- stances, in others they are massed in spheroidal chromospheres of pale greenish yellow or deep cadmium orange color. In some eases the chromatophores are peripheral in location and of va- rious forms. Vacuoles and pusules of varying form and distribu- tion have been observed in the cell contents. The dimensions of observed species are like those of Peridi- nium. The largest species thus far recorded appears to be H. blackmani which has a length of 225y, and transverse and dorso- ventral diameters of 1354 and 160» respectively; the smallest appears to be H. sphaeroideum with a length of only 42u, and transdiameter of 39x. The distribution of this genus from species thus far pub- lished appears to be limited, in the main, to warmer seas as shown in the following table: VoL. 2] Kofoid—On Heterodinium. 349 Species Latitude Temperature C. H. blackmani 7°-380° N. 25°-27° H. doma 34°-39° N. 16.1°-18.9° H. hindmarchi 19°—39° N. 15..6°=27 .2° H. trirostre 262 IN: Dhan H. leiorhynchum 19°—39° N. 15.5°-24.5° H. murrayi 14°-28° N. 20°-25.5° H. milneri 29°-31° N. 26.9°-27.2° H. sphaeroideum ier ea San Diego ee Bly arcane B45 (1 ANTE 14.6°-22.5 H. inaequale H. triacantha 55°=8il> N. Their vertical distribution is not known. At San Diego no individuals have been taken in the many surface catches of the tow nets made during the past few years. They have been found only in the vertical catches in from 165 to 40 fathoms to the surface. Murray and Whitting’s (’99) species were all appar- ently from plankton collected by filtermg water from ship’s pumps and therefore taken some 2-3 fathoms below the surface. The absence of chromatophores or their aggregation in chromo- spheres observed in individuals taken at San Diego is suggestive of occurrence in deep water with diminished light. The extreme hyalinity of some species is also indicative of a deeper habitat. The excessive development of the reticulum on the theca, and the asymmetry are evidently adaptations for flotation, on the one hand by inerease of friction surface, which is at least doubled in the more rugose forms, and on the other by giving a spiral course to any passive descent of the organism due to gravity, and thus prolonging its existence in the upper strata. This is a genus of somewhat aberrant structure and is repre- sented by relatively very few individuals in comparison with those of Ceratium and Peridimum. I regard it as a degenerate form unable to maintain itself at the surface and for some reason deficient in reproductive vigor. In comparison with the number of individuals observed the number of species is large. The known species are all well defined and observations on difterent individuals do not indicate as yet any noticeable intergradation. 350 University of Californa Publications. [ ZooLocy The nearest allies of this genus are plainly in the family Peridinidae, though it shows no marked structural affinities to any particular genus. The form cycle found in its species re- sembles that of Peridinium but its thecal plates are entirely different. The midventral diamond-shaped plate of the epitheca so characteristic of Peridinium is entirely lacking in Heterodi- nium, unless indeed the slight midventral expansion on the suture line be taken to represent a degenerate midventral which seems improbable. The excessive development of reticulations on the surface of the theca approaches that in Protoceratium but this genus appears to lack the midventral pit on the epitheca, and has a narrow transverse furrow which is complete distally. Its plates (see Schutt (’96)) are not known and it may prove to have a closer relationship to Heterodinitum when these are definitely determined. The midventral pit on the epitheea of Heterodinium resembles the so-called ‘‘pore’’ in Poroceratium gravidum (Gourret) but bears a different relation to the thecal plates and is probably not a homologous structure. In Poroceratium the ‘‘pore’’ hes near the middorso-ventral line in the middle of the dorsal and ventral apical plates, whereas in Heterodinium it lies in the suture be- tween the two ventral apicals. The generic distinctness of Heterodinium is thus beyond ques- tion and it belongs with Ceratium, Peridinium and Protoceratium in the sub-family Ceratiinae. SYNOPTIC KEY TO THE SPECIES OF HETERODINIUM. Sphaerodinium subgen. nov. Body spheroidal, antapical horns not present or only slightly developed as spines. Epitheca rotund without stout lateral sutures. ft; With nosamntapreall horns: Or (Sp Une St sete cae eee sgvtzeteacs 2 ie SWithantapicall’ spines yas oss es eee ees eee See ieee oe 3 2. Outline smooth, sutures faint, reticulations minute..H. sphaeroideum 2. Outline subangular, sutures prominent, reticulations coarse..H. doma 3. No apical horn, thecal markings faint S22 e ee H. milneri 3. Short apical horn, thecal reticulation prominent .............. H. murrayi Bp) slong apical) horns sects ene eee ee eee H. triacantha VOL. 2] Kofoid.—On Heterodinium. 351 Euheterodinium subgen. nev. Epitheea dorso-ventrally compressed, with straight, convex or concave sloping lateral margins which are usually thickened and have doubled ridges between the lateral margins of the apical plates. Antapical horns well developed. Girdle not very oblique. 1. Epitheca with strongly convex sides, apex broadly rounded _.... ee Ae Sat Neeson eae ore sea ane asec eats eect ten ES ANOEG UAE 1. Epitheca with straight or concave sides, apex not broadly rounded... 2 2. Epitheca low, its altitude about one-half the transdiameter -............ Be Pattie = Maen gn 8 Mes Rn dasa nee ides sect s eomaswtae ean aet kesh Ate edad H. rigdenae 2. Epitheca high, tapering, nearly equal to the transdiameter _............ 3 3h TUGHE Gale gonkel lomut wince Neel eae ae ae ce eee H. trirostre See Wehteambant cele no tbc cate Cee terpenes ee ea ee eet ee ee ee 4 4. Reticulations very coarse, scantily developed H. leiorhynchum 4. Reticulations subregular, very delicate ..............-..--.-- H. blackmani 4. Reticulations medium sized, very heavy _.....................- H. hindmarchi Platydinium subgen. nov. Epitheca dorso-ventrally compressed and hollowed out ven- trally, scoop-shaped. Lateral margins convex, not contracted to an apical horn. Girdle very oblique dorso-ventrally. Antapieal horns present. i, A\ratigyjorioal Innes): CWNAETAO ENING 5 eee cee eee eee H. scrippsi He A\inireyanicewll INOS; COMET ECG oe see occas ee cere H. whittingae Heterodinium sphaeroideum sp. nov. Pl. 3, fig. 15. A minute symmetrical species of spheroidal form without apical or antapical horns. The body is spheroidal or broadly ellipsoidal, the length 1.1 transdiameters. Dorso-ventral diam- eter equal to transdiameter. Epitheca a low dome, its altitude 0.4 transdiameters. Hypotheca exceeding the epitheea, elon- gated hemispherical, its altitude 0.6 transdiameters, with broadly rounded symmetrical antapex. Girdle premedian, transverse furrow indented, posterodexiotropic with very slight displace- ment scarcely 0.2 its width, its anterior and posterior lists equal 352 University of California Publications. [ ZOOLOGY and the latter not deflected distally, both formed by sharp pro- jecting ridges of the theeal wall. Longitudinal furrow short, its length less than 0.5 distance to the postmargin, broad and shallow, its distal two-thirds enlarged. Theceal plates imperfectly known. Ventral plates of typical number and arrangement except that the right ventral post- median (16) is not continued to the postmargin but appears to be divided into postmedian and antapical moieties. Suture lines faint, bordered by smooth structureless zones. No prominent lateral ridges. No lists or spines. Thecal wall minutely and faintly reticulate with subregular polygons with centrally located pores. Polygons relatively very numerous. Plasma dense, heavily vacuolated, chromatophores irregular, peripherally located, greenish yellow; nucleus near flagellar pore, ellipsoidal. Dimensions :—leneth, 42; transdiameter, 391; width of fur- row, 4-5; diameter of polygons, 2-3». Taken once in vertical haul from 165 fathoms off San Diego in June. Although this organism does not have the deficient girdle found in other species of the genus its thecal plates, in so far as they are known, are those of Heterodinium. Heterodinium doma (Murr. et Whitt.). Peridinium doma Murray and Whitting (799), p. 327, Pl. 30, fig. 3. Plainly belongs to Heterodiniwm because of the clearly shown ventral pit in the central expansion of the median ventral suture. The plates are only partially shown but in the one view (ventral) given they conform to Heterodinium so far as shown. The girdle and furrows are also typical. The species is characterized by the spheroidal form, sub- median girdle, broadly rounded apex, entire absence of antap- icals, median reticulations of subregular polygons and somewhat salient suture ridges. Reported from the warm temperate Atlantie between 34°- Bo Ne VoL. 2 | Kofoid.—On Heterodinium. 353 Heterodinium milneri (Murr. et Whitt.). Peridinium Milneri Murray and Whitting (’99), p. 327, Pl. 29, figs. 3a, b. The characteristic Heterodinium structures are not clearly shown in Murray and Whitting’s figures. There is only a sug- gestion of the midventral suture of the epitheca and a markedly deficient posterior list of the transverse furrow. The ventral pit is lacking and the plates are incompletely shown. The species is characterized by its spheroidal rotund body, premedian girdle with wide displacement and considerable over- lap of the ends of the transverse furrow, wide zones along suture lines free from reticulations, and coarsely reticulated plates... It is closely related to H. murrayt. Reported from tropical Atlantic in 29°-31° N. Heterodinium murrayi nom. nov. Peridinium tripos Murray and Whitting (’99), p. 327, Pl. 30, figs. 4a, b. non P. tripos (Muller), Ehrenberg (’33), p. 272 = (Ceratiwm tripos). The specific name tripos must be rejected as it was previously introduced into the genus Peridinium by Ehrenberg’s (733) transfer of Cercaria tripos of O. F. Miiller (1786) to the genus Peridinum. As figured by Murray and Whitting (’99) this species shows almost none of the generic characters except the very deficient posterior list of the transverse furrow. The ven- tral pit is questionably figured and no trace of the plates is shown. The only evidence of the presence of the left interealary is the shifting of the apex to the right. The species is characterized by its small size, rotund body, large and few subregular polygonal reticulations with a coarse mesh which hide the sutures and cover the whole theca. The oirdle is premedian and the transverse furrow is much displaced and has considerable overhang... The apex is somewhat con- tracted and the antapicals bear two short finned spinules on the left and one on the right. The anterior list of the transverse furrow is membranous. There are no antapical horns. Reported from the tropical Atlantie in 14°-31° N. 354 University of California Publications. [ ZooLocy Heterodinium triacantha (Jirg). Gonyaulax (?) triacantha Jorgensen (799), p. 35. Ceratium hyperboreum Cleve (’00), pp. 14-15, Pl. 8, fig. 5. Gonyaulax triacantha, Paulsen (’04), pp. 21-22, fig. 5. This form appears to belong to Heterodinium by reason of the reticulated thecal wall, the midventral suture of the epitheca deflected to the left, the widened distal end of the transverse furrow, and the longer right antapical spine. There is also some indication that the distal end of the posterior list of the girdle is continued in the suture on the right side of the hy- potheca. None of the figures shows the ventral area or pit, or the theeal plates in full. In so far as they are indicated in Paul- sen’s (’04) figures, they conform to those of the genus Hetero- dinium. There are difficulties in reconciling Cleve’s (’00) figures with each other, and with those of Paulsen (’04) as well as with Jérgensen’s description, probably due to the fact, that, as Paul- sen suggests; Cleve’s figure is reversed, 7.e., it is a view of the ventral face as viewed through the body from the dorsal face. This species probably belongs in the subgenus Sphaerodimum, though it does not possess a spheroidal body. It is characterized by the absence of antapical horns and post indentation, concave sides of the epitheca, the developed apical horn, and the three antapical spines. Dimensions :—length, 72-84; transdiameter, about 50,; dorsal-ventral, about 45p. Reported from coasts of Norway and Iceland. Heterodinium inaequale sp. nov. [EAL ibs mates Sh Alo) This is a small subpentagonal species with rotund epitheca and unequal widely separated antapicals. The body in face view is subpentagonal, the two anterior mar- gins are quite convex, the left posterior nearly straight, the right slightly convex and the postmargin between bases of the antap- icals is coneave. The length is 1.2 and the dorso-ventral diam- eter 0.75 times the transdiameter. The epitheca is low dome- —— VoL. 2] Kofoid—On Heterodinium. 359 shaped, compressed dorso-ventrally, its altitude (ventral) is 0.7 transdiameter. No apical horn is differentiated and though com- pressed dorso-ventrally it is not thinned down to a sharp edge at the doubled lateral sutures. The ventral face is scarcely ex- eavated. The broad midventral suture runs from the longi- tudinal furrow to the apical pore swerving towards the left at the ventral pit. The hypotheea is rotund, its altitude (mid-dorsal) 0.6 trans- diameter. The antapicals are very unequal, the right is about one-half the length of the left and is abruptly imcurved to an acute tip. The left is not incurved and is somewhat tapering. Its length is 0.3 transdiameter. The postmargin between the antapicals is slighly concave and is 0.4 transdiameter in length. The plates are typical in number, the left intercalary being confined to the dorsal face. The dorsal premedians are very low, scarcely exceeding the girdle in width. The dorsal post- medians on the other hand are unusually long, and the posterior angle of the right one projects slightly beyond the margin. The girdle is narrow and slightly oblique (15° postero-ven- trally) to the equatorial plane. The transverse furrow is postero- dexiotropie with a displacement of its own width. It is scarcely indented, the thecal wall forming a slight anterior ridge, and a small posterior one which fades into the right antapical suture distally. The longitudinal furrow is narrow and short, 0.6 dis- tance to postmargin. The theeal wall is structureless save for scattered pores in the two individuals thus far observed. These may both be young stages and the older ones may be reticulate as are other species in the genus, but there is not the slightest evidence of reticula- tions on the thecal walls of these two individuals. The suture lines are light and faint. The midventral one on the epitheca is broad in the posterior half between the ventral pit and the flagellar pore, and the lateral sutures of the epitheca and hy- potheca are doubled and prominent. The right dorsal premedian suture is very oblique. No fins or lists were noted. The plasma is coarsely granular, chromatophores few, large, spheroidal, clustered near the center. 356 University of California Publications. [| ZOOLOGY Dimensions :—length, 116-120; transdiameter, 100u; dorso- ventral, 754; furrow, 8 in width. Taken in vertical hauls from 40—95 fathoms to surface off San Diego in May and June. This species is not closely related to any described species. Its asymmetry is noticeable but no other adaptations to flotation in the theca were found in the individuals examined. Heterodinium rigdenae sp. nov. Pl. 18, figs. 6-8. A small pentagonal Peridinium-like species with a coarse polygonal reticulum and slight obliquity of the girdle, resem- bling P. acutangulum Lemm. Body pentagonal in face view with straight or nearly straight subequal sides, broadly bifurcated posteriorly with short stout conical antapicals. Length 1.3 and dorso-ventral diameter 0.6 times the transdiameter. Epitheca without apical horn, the sides sloping in a straight line from the apical pore to the girdle, compressed dorso-ventrally and somewhat excavated in the mid- ventral region, lateral sutures prominent and doubled. Hy- potheea also compressed, the right margin concave, the left with projecting angle at the junction of postmedian and antapical plates. Ventral face excavated. The antapical horns are short, 0.2 transdiameter in length, which is nearly equal to the slightly eurved margin which separates their bases. Their ends blunt with short terminal spinule. The girdle is inclined about 15° postero-ventrally from the equatorial plane. The transverse fur- row is deeply indented, more so towards its anterior than its posterior margin. It forms a descending right spiral with dis- placement slightly exceeding its width. Its distal end is feebly developed, the posterior ridge vanishing on the ventral face. The anterior ridge is a heavy projection of the body wall, nearly twice the height of the posterior. The longitudinal furrow is narrow and shallow, dilated posteriorly and extends from the Vor. 2] Kofoid.—On Heterodinium. ay flagellar pore little more than 0.5 of the distance to the post- margin. Theeal plates of the normal type. Left intercalary confined to dorsal face. Suture lines heavy, deficient on right ventral area. Thecal wall covered with irregular polygons, mostly pen- tagonal, larger ones each with single central pore. Polygons relatively few, 41 on dorsal apical. In young individuals faint suture lines but no polygons are found. The ventral area is found as usual at the junction of the sutures on the ventral face of the epitheca. It is unusually large and contains the ante- riorly located reniform pit-like structure. Individuals thus far observed have been very hyaline and colorless or with pale greenish yellow chromatophores, reniform or irregular in shape. Dimensions:—leneth, 120-1254; transdiameter, 90-92p; dorso-ventral, 155; girdle width, 10»; polygons, 2-10p. Taken in vertical hauls from 90-100 fathoms to surface in June off San Diego. This species resembles H. h indmarcht (Murr. et Whitt.) im the type of reticulations, but differs from it in its more robust form, shorter epitheca and antapicals. Heterodinium trirostre (Murr. et Whitt.). Peridinium trirostre Murray and Whitting (’99), p. 327, Pl. 29, fig. 5. This species is shown to have the typical ventral plates of Heterodinium, the ventral pit and the deficient posterior list of the transverse furrow which is deflected posteriorly on the right antapical horn. The species is characterized by the presence of pointed an- tapical horns, the left showing a bifurcation into two apices. The epitheca is high, broadly cuneate with doubled lateral su- tures. The girdle is median, the transverse furrow being dis- placed only its own width. The reticulations are very coarse and heavy and exhibit more than the usual irregularity in form. Reported from 26° N. in the Atlantie. 358 University of California Publications. | ZooLoGy Heterodinium leiorhynchum (Murr. et Whitt.). Peridinium leiorhynchum Murray and Whitting (799), pp. 326-327, Pl. 29, figs. 2a, b. This is unquestionably a Heterodinium as it shows a mid- ventral pit upon the epitheca, the doubled lateral sutures and diminishing posterior list of the transverse furrow. The plates are very imperfectly shown though there is a suggestion of the left intercalary of the epitheca in the sight shoulder on the left epithecal margin, and in the incomplete suture lines of the latera! view. The species is characterized by the somewhat differentiated apical horn, the unequal, pointed, divergent antapicals termi- nating in spines, the rotundity at the girdle, prominent suture ridges (incomplete in figure), smooth or coarsely reticulate theeal wall, and fenestrated lists of the girdle. Reported from the warm temperate Atlantie from 20°—40° N. Heterodinium blackmani (Murr. et. Whitt.). Peridinium Blackmani Murray and Whitting (’99), pp. 327-328, Pl. 29, figs. 6a, b, ¢. This superb and clearly marked species is the only one in which the left intercalary plate is shown by Murray and Whit- ting (99). It is unquestionably a typical Heterodiniwm though these authors fail to show the ventral plates of the epitheca and the ventral pit characteristic of the genus. The remainder of the plates is almost completely shown. The prominent doubled lateral sutures, the short longitudinal furrow, and the poste- riorly deflected posterior list of the transverse furrow stamp this species as a typical Heterodiniwm. The species is characterized by the curved epitheca flaring to the greatly expanded equator, submedian girdle, divergent pointed antapicals. The suture lines are marked by prominent lists and the plates are reticulate with delicate subregular hex- agonal polygons, which exhibit a tendency to horizontal elonga- tion on the epitheca. Reported from the Caribbean Sea and tropical Atlantic from 9°-25° N. VoL. 2] Kofoid.—On Heterodinium. 359 Heterodinium hindmarchi (Murr. et. Whitt.). Peridinium Hindmarchii Murray and Whitting (799), p. 326, Pl. 29, figs. la, b. This is plainly a Heterodinium as it has the characteristic ventral pit in the central expansion of the median ventral suture of the epitheea, a short longitudinal furrow and the deficient distal posterior list of the transverse furrow. The plates are not shown but the presence of the left intercalary plate is sug- gested in the figure. The species is characterized by the long stout widely sepa- rated antapicals, elongated epitheea, and coarse reticulations. Reported from tropical Atlantic from Panama to 34° N. Heterodinium scrippsi sp. nov. Pl. 17, figs. 1-5. A large species with short antapicals, scoop-shaped epitheeca and coarse irregular reticulations. The body is subheptangular in face view, swollen at the girdle, and with shallow posterior bifureation. The length is 1.5 and the dorso-ventral diameter 0.7 times the transdiameter. The epitheca is long, and its alti- tude is about 0.8 transdiameters. The ventral face is flattened and hollowed out anteriorly and thins out laterally to the doubled suture lines on the angular margin between the apical plates. Posteriorly the epitheca flares out to meet the girdle. The lateral margins have rounded shoulders about two-thirds of the distance from the girdle to the apical pore which rises from the anterior end in a fully developed apical horn. -- Vou. 2] Kofoid—On Heterodinium. 361 Heterodinium whittingae sp. nov. TeAb ley aaweys alaleals}. A large species with very oblique girdle, elliptical outline, and shallow rounded bifureation. Body elliptical in face view with broadly rounded apical end and short incurved antapical _ horns which preserve the elliptical outline. The posterior bifur- cation extends but one-fourth of the distance to the girdle and is broadly rounded anteriorly. The body is very much compressed dorso-ventrally, forming a sharp edge at the lateral margins. The girdle is very oblique being inclined at an angle of 45° to the main axis in an antero-dorsal to postero-ventral direction. The length is 1.4 transdiameters and 3 times the distance between the greatest dorsal and ventral extensions which is found in the left half of the epitheca. The whole body is slightly twisted in a right spiral. The epitheca is very much flattened anteriorly and somewhat excavated on the ventral face, forming in fact a thin sheet which expands posteriorly as it meets the oblique girdle. The hypotheca is likewise flattened and excavated ventrally about the longitud- inal furrow. The lateral postmedian plates form a posteriorly projecting tooth on the left margin. The girdle forms a descending right spiral with slight dis- placement equalling its width. Its distal end is much widened, the posterior border becoming low and deflected posteriorly into the ventral suture of the right antapical horn. The longitudinal furrow is short and narrow. The flagellar pore is found as an elliptical opening at its proximal end. The thecal wall is thin, delicate and hyaline with hght suture ridges except in the case of the lateral sutures between the apicals which are doubled and heavy, as are also the lateral sutures on the hypotheca. The plates are normal, the left intercalary being restricted to the dorsal face. The surface, including that of the girdle plate, is everywhere covered with a reticulum of irreg- ular polygons, each with a single central pore. In the several specimens thus far observed the reticulum has been very hght and delicate. The polygons are relatively numerous, 97 having been recorded on the dorsal apical plate. 362 University of California Publications. | ZooLocy The plasma is exceedingly hyaline and coarsely vacuolated and its total amount is relatively very small. The nucleus is minute (12), spheroidal and centrally located, and there is one pale chromosphere of similar size and form adjacent to it. This is a large species, 180, in length, 140 in transdiameter and 60u in greatest dorso-ventral extension. Polygons 5-12, in diameter. Taken in vertical catch from 85 fathoms to the surface off San Diego in July. VoL. 2] Kofoid—On Heterodinium. 363 LITERATURE CITED. Ola7e, 1s 10 :00. Notes on some Atlantic Plankton-organisms. Kongl. Svensk. Veten.-Akad. Handl. Bd. 34, No. 1, 34, 22 pages, 8 plates. Ehrenberg, C. G. 34. Dritter Beitrag zur Erkenntniss grosser Organization in der Richtung des kleinsten Raumes. Abh. Akad. Wiss. Berlin, 1833, pp. 145-336, Taf. 1-11. Joergensen, E. 799. Protophyten und Protozoen im Plankton aus der norwegischen Westkiste. Berg. Mus. Aarb. 1899. No. VI, pp. 1-112, I-LXXXIII. 5 plates. MiullerssO rk: 1786. Animaleula Infusoria fluviatilia et marina. Pp. LVI, 365, 50 Taf. Murray, G., and Whitting, F. G. 799. New Peridiniaceae from the Atlantic. Trans. Linn. Soc., London. Botany (2), Vol. V, pp. 321-342, Tables I-IX, Pls. 27-33. Paulsen, O. :04. Plankton Investigations in the waters round Iceland in 1903. Medd. Comm. Havunderzog. Bd. I, No. 1. 41 pages, 11 fig- ures, 2 maps. Schutt, F. 796. Peridiniales. Engler und Prantl’s Nat. Pflanzenfamilien. I Teil, Abth. b, pp. 1-80, 43 figs. EXPLANATION OF PLATES. PLATE 17. 1.—Heterodinium scrippsi ventral view. X 725. ap. p., apical pore; fl. p., flagellar pore; l. f., longitudinal furrow; v. a., ventral area; v. p., ventral pit. 2.—Dorsal view of same. X 420. 3.—Oblique view of left side of same. X 420. 4.—Diagrammatie apical view of same. X 420. 5.—Reticulations adjacent to posterior list of transverse furrow. xX 2725. [364] q ? ) ’ Linty. Can ust. Zo ou. Vat.11. [ Kororo | Pate XVIL. | on Ae oe 2 « PLATE 18. Fig. 6.—Heterodinium rigdenae, ventral view. X 410. ap. p-, apical pore; in fl. p., flagellar pore; l. f., longitudinal furrow; v. a., ventral F area; v. p., ventral pit. Fig. 7.—Dorsal view of the same. X 410. Fig. 8.—View of left side of same. X 410. Fig. 9.—Ventral view H. inaequale. X 420. Abbreviations as in fig. 6. Fig. 10.—Oblique view of right side of same. X 420. [366] een EE ————————— —_= = — —_———— Se Univ. CAL-Fus.Zoou. Vout. 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