y ke : ia. = ©; ee Pika ery ae a @ eae,” e @ a E cs ey a. Pinal A LIBRARY OF THE UNIVERSITY OF CALIFORNIA. RECEIVED BY EXCHANGE - Class Pie oe AE DANS Pet OLE EXPE DITION. VOL.V A. PUBLISHED AT THE COST OF THE GOVERNMENT BY THE DIRECTION OF THE ZOOLOGICAL MUSEUM OF THE UNIVERSITY. COPENHAGEN. H. HAGERUP. PRINTED BY BIANCO LUNO ASS. 1904-1913. Sad edit ae a Contents of Vol. V a. I. H. F. E. Juneersen: Pennatulida, p. 1-95 (3 plates), 1904. II. Tu. Mortensen: Ctenophora, p. 1-95 (10 plates), 1912. III. O. Carteren: Ceriantharia, p. 1-78 (5 plates), 1912. IV. 0. Carteren: Zoantharia, p. 1-62 (7 plates), 1913. weer ane ey coe ao J ‘ iat THE DANISH INCOLF-EXPEDITION. VOL. V, PART 1. CONTENTS: HECTOR F. E. JUNGERSEN.: PENNATULIDA. PUBLISHED AT THE COST OF THE GOVERNMENT BY THE DIRECTION OF THE ZOOLOGICAL MUSEUM OF THE UNIVERSITY. LS BRARY} O° THE f UNIVERSITY COPENHAGEN. H. HAGERUP. PRINTED BY BIANCO LUNO. 1904. THE DANISH INGOLF-EXPEDITION. VOLUME V. 1. PEAR ATULIDA. BY HECTOR F. E. JUNGERSEN. WITH 3 PLATES, 1 CHART AND 3 FIGURES IN THE TEXT. RPS OF THE e UNIVERSITY | OF : SSALIFORNIAA COPENHAGEN. PRINTED BY BIANCO LUNO. 1904. MEA EOL peste yer ge ts L: z CONTENTS. Pennatulida. Pag. Pag. dutroauctory remarks. . . . «<0 /sis)s\ ceed eee ee I. Funiculina quadrangularis (Pall) .......... 49. Remarks on the systematic classification of the Pennatulids 4. PEE ELOtORUMCIe COIL i's oe eSeiar st ss sais 8i ba oot 51. List of the northern Pennatulids............---- 10. PRIMER CITETED POO lop) clos (bso Vaire-a eo bsaceilne gue 51. Fam, Pennatulidee KOll) .sncevesueeeee a ne ses II. Protoptilum carpenteri K6ll. ............, 51. Pennatula Lams :ss ii teeneeeinna ease ss +s II —_ CROBRONE ICON oo 5 driers, o fas enetie ng. foe 55. _ Pennatulaaculeata Kor. Dan. ...........-- II. _ deriticulatins 1. :Sp..6%%.46 8 ces @ 005. 59- — phosphorea L.var. candida Marsh.&Fowl. 14. PAS CHOpUNeRN V ETEN aS ay hee Soe wel pasts 62. ied MIRE ANID IU re ess (6s o's ss 8 eae Distichoptilum gracile Verr. ........645.5. 62. See AOE SGD al os le cio 58 6) soos + 18. Fam, Anthoptilides KOM. (6.50 ei. 8 2 3 8 8 8 ee 65. Fam. Virgularide KG6ll, emend............... 24. PIPED ELSI HOON a ievally, peas hiay ete Claus. 0. oie a gaia 65. Sie MEER IANER Speed sala 5. 5 'ocs 6 Gas ee oe 8 8. 24. Anthoptilnm grandiflorum (Verr.).......... 66. Virgularia mirabilis (O.F.M.) ............ 25. _ rahe 2: Be) | eae ne a ae 67. — PMlatiscns NOM, NOV, 2 665 tis ke ee 33. Fam. Kophobelemnonide K6ll. ..........-+42- 68. MRI MERRIE Da 1G lg S ols Gnigepe ince @ os a ems 3% 37: Kophobelemnon Asbjorns.........-..+-+-+06- 68. Stylatula (Diibenia) elegans Kor.Dan........ 38. Kophobelemnon stelliferum (O. F.M.)........ 68. Fam. Pavonaride Jungersen .............+.. 39. Fam, Umbellulidee K6ll.. 0 5. eee ee tee ee 74. PTUMTER OMI gion aa hi Sante iat tne os ecko ool bas. 39. WaT RIRIIEIRCRG BEA Sehr. Siew oak are. ef tigrielie, 9 © ok alee 's 74. Pavonaria finmarchica (M.Sars)........... 39. Winwermia Hndahlt KG... sei we ba 75- MM ATIICRRIA TINO a Silos otis ie arg Wasa t 6s 68k aes (ie 44. — CCHEING (Epp hr s.s ens ee ees woe a eee 79. Halipteris christii (Kor.& Dan.) ........... 45. | General review of the occurrence and distribution of the Fam. Funiculinide K6lL, emend.............. 49. MR GeTt: PONNALINOS 51s. sca) bps use =~ oye Sines See 87. CI EEG TL Ge BC Siem gp prone at air Se ke a a Bein SMRMERMEDILO o's) 0.0 ocx acters bie Vi eater gees al ‘on 39. al eig '9nb! 6 ws 93- 42ANAD pone sone. iret} anahiot. me : ayer Kary Be missin Z Po to tee iv Same ‘ayes Rhy ar : HO wie * ‘ ewe ree . ; - aise Z 7 wt * #2 RY =) ia ey * f yi at a ae ae ae ee butte Ath is sala BL wheats Pennatulida. By Hector F. E. Jungersen. n the present treatise an account is given, not only of the Pennatulids brought home by the «Ingolf», | but also of material in later collections from within the territory of the Ingolf-Expedition and from the adjoining regions which were investigated afterwards. Thus it comprehends the results from the surveying cruises of the «Diana» at Iceland and the Faeroe Islands, from the expedition to East- Greenland by Messrs. Amdrup and Hartz, from the participation of cand.mag. Ad. Jensen in the investigation of the «Michael Sars» in the North Atlantic during the summer of 1902 under the direction of Dr. Hjort, and from the cruise of the «Thor», the steamer for the international investi- gations of the sea, during the summer of 1903 under the direction of Dr. Joh. Schmidt; finally it includes also the material sent to our museum from Mr. Jéusson, district physician on the Vestman- Islands, and by Mr. Miiller, agent in Thorshavn. Our knowledge of the Pennatulids rests in several respects on a rather slight footing; this holds good also for the apparently so well examined Scandinavian forms, in spite of the numerous and beautiful treatises which have appeared from Norwegian scientists. I have tried personally therefore to examine all the forms that have hitherto been described from the coast-regions of Scandinavia, as well as the forms from the territory adjoining the one which was investigated by the «Ingolf». For this purpose I have examined the collections in Christiania, Bergen and Stockholm; from the - museums in these towns I have received later all that I wished to examine more closely and to compare with our own material here in Copenhagen; for this great liberality I beg to offer my best thanks to Professors R. Collett and Hj. Théel, and to Conservators Dr. A. Appelldf and Mr. J. Grieg. I have further had the opportunity at the British Museum of bringing into the comparison part of the material of the Challenger-Expedition; on the other hand, I have searched in vain in several English museums for the Pennatulids from the expeditions of the «Porcupine», the «Triton», and the «Knight-Errant». The result of my endeavours has thus been a revision of the Pennatulid- fauna of a large sea-territory, viz. the Polar Sea between Europe and Greenland, the sea to the West of Greenland, and the northern part of the Atlantic down to 55° Lat.N. and to the meridian of Cape Farewell. This revision, as will be seen, has led to a somewhat new conception of several species hitherto established; further, several forms have been added, of whose occurrence within territory so far north we have hitherto known nothing, and could know nothing; of undescribed species only two have been added. The Ingolf-Expedition. V. 1. 2 PENNATULIDA. I have followed K6lliker in regard to the terminology; some of his terms have been used from much older times, and are based on the external |resemblance to a feather, especially conspicuous in forms like Pennatula, Pteroeides etc. .The stem of the colony is accordingly divided into the shaft, rhachis, carrying all the individuals, and the peduncle, which is naked and under natural conditions sunk into the bottom of the sea; polyp is the name used only for the individuals posses- sing the entire equipment of an octocoral, tentacles etc, zooid for the dwarf individuals wanting sexual organs and tentacles (only in Umdéellula do they carry one tentacle), The individual is said to be provided with a calyx, when the basal part of its body’) is stiff so as not to be retractile, but the upper part of the polyp with mouth and tentacles can be retracted and hidden in it. The upper edge of the calyx may be more or less marked, in the former case it is provided with lobes, up to eight in number; the axial (dorsal) side of the individuals is turned towards the stem, the abaxial (ventral) away from the stem. Wings, ale or pinne, is the name of the oblique series of polyps placed transversely on the stem, where the polyps are mutually coalesced. I have differed only in one point from Ko6lliker; what he de- scribes as the ventral side of the colony, I call the dorsal, and vice-versa (except in Renzliwm see below); this I shall try to explain more closely. In the great majority of Pennatulids the colony, as is well known, is constructed bilaterally. The plane that divides the stem longitudinally Fig. 1. Diagram, part of a Penna- tulid-rhachis. V—D indicates the direction of the middle plane. 1—8 polyps in one transverse series; the figures give the age of these polyps, into two corresponding parts, has quite naturally always been interpreted as a dorso-ventral one (V—DZ in the annexed figure); but whether the side of the stem in the figure denoted by J, is to be called the dorsal (po- no. I being the eldest one; the arrows indicate that the polyps accordingly appear and are developed to either side away from the naked sterior), or the ventral (anterior), has always been decided quite arbitrarily. The earlier authors — with the exception of Lamarck — have generally called it the dorsal, or posterior side. K6lliker, in his monograph (p. 5), declares it to be the ventral for no other reason than that it is «am streak D—D. zweckmassigsten». Later, all authors have adopted the decision of Kdlliker although some of them (e.g. Verrill, Koren and Danielssen) had earlier used the contrary desig- nation. The feature distinguishing this side of the colony, is first and foremost that new polyps develop on either side, away from the median line; thus, the polyps placed nearest to this line are the oldest in each transverse series; further, the median line itself is always «sterile» in so far as polyps are never formed in it; only rarely is it covered with zooids (for instance Kophobelemnon); generally along the median line there is a more or less narrow «naked» streak, increasing in breadth down towards the peduncle. The opposite surface of the stem (V in the figure) is characterized by the fact that the new polyps are developed from both sides in the direction towards the median line; accordingly, the 1) By the body of the polyp is understood the part of the polyp projecting from the surface of the stem; in reality some portion of each individual issuing from the stem is inclosed in the latter. PENNATULIDA. youngest polyps in each transverse series are those nearest to this line; the median line and its more immediate surroundings is often also «sterile» of polyps; when this is the case the naked streak, how- ever, is narrower here than on the opposite surface, with the exception of the lower part towards the peduncle (Revzzlla, however, is a remark- able exception). Thus the two median surfaces of the stem are gener- ally easily distinguished‘). Making the median line our starting-point we might say that the surface D shows a «centrifugal» development of the polyps, the surface V a «centripetal» one; so far we might well abandon the terms ventral and dorsal side, and instead of them use centrifugal and centripetal side; these latter names, however, are somewhat clumsy in descriptions, as is also the case, I think, with the terms proposed by Bourne?): prorhachis (for Y) and metarhachis (for V); dorsal and ventral side are, and will always be, the most handy names, if only a rational choice were agreed upon. In my choice I have started from the following consideration: the stem of the sea-pens, rhachis + peduncle, is the direct product of the individual developed from the egg (the «. This may be seen in every specimen of Renzlla; but it has been further proved by the examination of its development by Wilson. From his examination it is also seen that the naked side of the Renilla corresponds to the ventral or sulcar side of the primary polyp; but Wilson has not seen either that it corresponds to Kolliker’s «dorsal side» in the other Pennatulids; I have, however, proved this fact in my treatise, quoted above, on Pennatula phosphorea. Nevertheless the matter does not seem to be understood yet with regard to Renilla; Delage and Hérouard (Traité de Zoologie concréte, T. 2, 2, Ig01) take the same erroneous view as KOlliker, so does Moroff; whether it is also the view of Bourne is not distinctly seen, but from his endeavours to derive Rendlla from an Umbellula-like original form I am inclined to think so. To prevent any misconception in future I figure here a Renilla and a Pennatula, seen from the same — homologous — side, the side I designate as the dorsal one. Remarks on the systematic classification of the Pennatulids. Recent authors have generally adopted the system set forth by K6lliker in the Challenger Report vol. I, 1880. According to this, the order Pennatulida is divided into four groups (sections), in the first three of which the colony has a bilateral formation; when the sub-sections are omitted this system is as follows: I. Pennatulee. Fam. Stylatulide. Stylatula Vert. Diibenia Kor. & Dan. Acanthoptilum Koll. Fam. Pteroetdide. | Pteroeides Herkl. Godeffroya Koll. Sarcophyllum Koll. Il. Spicate. | Fam. Pennatulide. Fam. Funiculinide. | Pennatula Yam. Funiculina Yam. Letoptilum Verr. Halipteris Kéli. Ptilosarcus Gray. Fam. Stachyptilide. Halisceptrum Herkl. Stachyptilum Koll. Fam. Virgularide. Fam. Anthoptilide. Virgularia Lam. Anthoptilum Koll. Scytalium Herkl. Fam. Kophobelemnonide. Pavonaria Koll. Kophobelemnon Asbjorns. PENNATULIDA. 5 Sclerobelemnon Koll. HT Reniilew: Bathyptilum Koll. Fam. Umbellulide. Umbellula Yam. Fam. Protocaulide. Protocaulon Kol. Fam. Renillide. Renilla Yam. IV. Veretillec. Cladiscus Kor. Dan. Fam. Cavernularide. Fam. Protoptilide. Cavernularia Val. Protoptilum Koll. Stylobelemnon Koll. Lygomorpha Kor. Dan Fam. Lituaride. Microptilum Koll. Lituaria Val. Trichoptilum Koll. Veretillum Cuv. Leptoptilum Kil. Policella Gray. Scleroptilum Koll. Clavella Gray. Studer (Versuch eines Syst. der Alcyonaria. Arch. Naturgesch. 53. Jahrg. 1. 1887, p.22, and Challenger Report vol. 31, p. XX VII) has followed Koren & Danielssen?), and like these authors added a fifth to the four groups of K6lliker: Géndulee with fam. Géudulide, genus Géndul Kor. Dan. but has otherwise restricted himself to the inserting of the later established genera into the families of KGlliker (Sczophyllum Verr. in fam. Pennatulide, Svava Kor. Dan. in Stylatulide, Gunneria Kor. Dan. and Dzstichoptilum Verr. in Protoptilide). Later (Note prélim. etc. Camp. de l’Hirondelle 1890) he has inserted into the fam. Pterocidide the new genus Gyrophylium Stud. Kiikenthal (Diagnosen neuer Alcyonarien aus der Ausb. d. Deutschen Tiefseeexpedition. Zool. Anz. 25. Bd., 1902, p. 302) has added still another (the sixth) group: Verticilladee: fam. Chunnellide, genera Chunella Kiikenth. and Amphianthus Kkth. Bourne (Anthozoa in the treatise on Zoology edited by Ray Lankester, part II, 1900) almost without any alteration — at all events, without any essential alteration — has followed Kdlliker; it has to be remarked, however, that he regards the two families of Kélliker Vir- gularide and Stylatulide as subfamilies of one family Virgularide; Pennatulide and Pteroeidide Koll. are regarded also as subfamilies; and the family Géndulide is placed beside, and in the same group with these families; finally also Fumiculinide and Bathyptilide are degraded to subfamilies of one family, as are also the two families of Veretilline. The system has been more highly modified in Delage & Hérouard (Traité de Zoologie concréte T. 2, Partie 2, 1902); here there are five groups: 1) Frondina (Renilla), 2) Umbellina (Umbellula), 3) Juncina (Spicate +-Veretillee of Kélliker), 4) Pennina (= Pennatulee of KGlliker), 5) Acauline (Géndul). The extent of the families remains otherwise almost unaltered; the family Protocaulide of KOlliker, however, has been placed under the family Kopho- belemnonine D. & H., and the genus Dewtocaulon Marsh. & Fowler has been included in the same family; the genus Cladzscus Kor. Dan. has been removed to the family Protoptiline; the genus Svava Kor. Dan. to the family Virgularine; several genera of the same family, as Zygus Herkl, Radicipes Stearns, as also some of the genera established by Gray in the family Pexwnatuline are also enumerated. t) Nye Alcyonider etc. 1884. 6 PENNATULIDA. I cannot regard the grouping of Delage & Hérouard as an improvement on the system of Kélliker, and most of the genera, found in these authors but not in those before mentioned, will have to be abandoned. Nor can I, for my part, regard the system of Kolliker as satisfactory; the characters, by which the division into groups has been made, hardly always make the line of separation in a natural way; when, for instance the first group Pexnatulee is made to contain the forms in which the individuals of the polyp-series are coalesced to form wings, this feature is seen not to be of absolute validity: in the genus Virgularia, species are found in which the wings are partly dissolved into free individuals, or, more properly, in which only some individuals are coalesced, and only to quite a slight extent (Virgul. bromleyi K. and Virg. cladiscus mihi); the same is found in the genus Sty/atula (in the sub- genus Dzibenza). On the other hand, forms are found in the group Sfzcaze, in which a coalescing of some individuals in an oblique series takes place, in which, accordingly, there is as distinct a beginning of wings as in the mentioned Virgudarie; this is seen in Amthoptilum grandifiorum Verr., while in another species of the same genus, A. murrayi K. the polyps are quite separate, and partly even not arranged in regular oblique series. A one-sided prominence, given to this single character, will thus easily separate closely related forms into two groups; but when other structures are also taken into consideration, I think that the four groups of Kélliker may be kept — at least provisionally — but, to be sure, with altered contents. If we knew more of the development of the colony in a larger number of Pennatulids than is the case now, the mutual relations of the forms might certainly be seen more clearly, and then, perhaps, the grouping might be different. For the present, it is exceedingly difficult’ to reach a sure classification; difficulties occur on all points, especially in limiting off the species. Most of the features that have generally been used as specific characters prove to be more or less useless: the number, grouping, size of the polyps and zooids often change considerably during growth, as also all the relative sizes of the parts of the polyps and the stem, which are moreover exceedingly influenced by the degree of contraction. Nor does the anatomical structure give many holds for the classification, as it is upon the whole too uniform; the spicules also give but slight help: species of the same genus may quite lack spicules or be abundantly provided with them (for instance, within the genus Uméellula); nor is the form and size of the spicules often of much assistance; in specimens of the same species both the number and the size may be very varying (for instance, in Funiculina quadrangularis, Kophobelemnon stelliferum). Amongst the better characters, I reckon the presence or absence of a calyx on the polyps, and the form of the calyx. For the delimitation of genera and families, these features along with the grouping of the polyps and zooids in the developed colony will for the present be of most value. I am not in a position to undertake a complete reformation of the system of the Pennatulids on the basis of the material I have hitherto had for examination, but on several points I shall be able to revise, with greater certainty than has before been possible, certain parts of the system of Kélliker and the later additions to, and alterations of, this system; on other points I shall have to be content with giving suggestions more or less positive. The more particular reasons for the position of the forms dealt with in the following section, will be given there; in this place I shall only give some conclusions with regard to these forms. PENNATULIDA. Y In the group Pennatulee, the genus Halisceptrum Herkl. will have to be removed from the family Pennatulide to the family Virgularide; it is quite obvious that this genus is closely related to the genus Virgularia itself; perhaps it might even be embodied into it (see later under Virgu/aria); further the genera of the family S¢y/atulide show so close a resemblance to Virguwlaria, that they can scarcely be regarded as forming a separate family; they are certainly more closely related to the genus Virgularia than this genus is to the genera Scytalium Herkl. and Pavonaria KOll.; on the other hand, I should be inclined to separate these latter genera into a separate family (or, at all events, a sub- family) characterized — in contradistinction to Virgularia and the Stylatulids (with the exception of Acanthoptilum) — among other things by the fact that the sexual organs are only developed in the fully formed polyps of the older wings, and that new polyps, at any rate in the younger forms, bud out on the upper, older part of the rhachis; in this family Pavonaride mihi must further be included a member of the group Sfrcate, viz. the genus Halipteris KOll., although its polyps are not united into wings; but in most other features it shows the greatest resemblance to Pavonaria K6ll., much more than to Funiculina, with which it has been placed by KGlliker. The genus Svava Kor. Dan. which Studer has referred to the Stylatulids, Delage & Hérouard to their Vergularine, will have to be dropped as a genus; it is only a species of the genus Virgularia; Lygus, which in Del. & Hér. is given as a separate genus, is synonymous with V7rgwlaria; and Radicifes Stearns (Proc. U.S. Nat. Mus. Vol. 6, 1883, p.96, Pl. VII) is no sea-pen at all, but a Gorgonid (a species of Strophogorgia Perc. Wr.); the «genera» of Gray: Ptilella, Phosphorella, Crispella (the family Pennatuline Del. & Hér.), and Argentella (of their Pteroecideidine) have already, and certainly rightly, been condemned by K6lliker in his monograph. Further, the genus Stachyptilum K6ll., in my opinion, is to be included in the family Pennatulide. In the group Sficate of Kélliker (= Umbellina + Juncina — the family Veretilline in Del. & Hér.) the greatest alterations will have to be made. In the first place the whole family Protocaulide must be done away with: two of its genera, Protocaulon KOll. and Deutocaulon Marsh. & Fowl., because they are young stages of Virgularta-species, Cladiscus Kor. Dan., because it is only a (wrongly inter- preted) species of Virgularia, in reality identical with Svava Kor. Dan. (for further particulars see under Virgularia mirabilis and V. cladiscus mihi). Consequently, all inquiry may be omitted with regard to the question whether «Cladiscus» is to be included in Protoptilide or Protocaulide, and whether Protocaulon and Deutocaulon, as has been done by Del. & Hér., may be placed under the family Kophobclemnonine. Again, most members of the family Protoptilide must be dropped as separate genera. Lygo- morpha Kor. Dan. is a young form of Halipteris (for further particulars see under H. christiz); Microp- tilum KON. of Pavonaria; Leptoptilum KOll. and Trichoptilum K6ll. are young forms of Funiculina (see under this latter); Ganmneria Kor. Dan., which was inserted not only by the authors of the genus, but also by Studer and Delage & Hér,, is a wrongly determined Kophobelemnon (see under K. stelli- Serum); further, the genus Scleroptilum K6ll. must be separated from this family; its polyps, in spite of their dense provision of spicules, want a calyx, and it can only be by an oversight of Kdlliker that it has been placed in a family especially characterized by him as provided with a calyx. Finally we have only left Protoptilum KO6ll. and the later added Distichoptilum Verrill. These two genera, at all 8 PENNATULIDA. events, have the common feature that their polyps are provided with a well developed calyx, the abaxial part of which is fully formed, often with calyx-teeth, while its axial side is more or less con- nected with the stem; young stages of Protoptilum, or such simple species as Prot. carpenteri (if it is not only a young stage also) resemble Dzest:choptilum very closely; the arrangement of the zooids, however, is quite different, as Profoptilum is provided with numerous dorsal zooids, and moreover with zooids everywhere on the rhachis between the polyps, while Dzstichoptilwm has only lateral zooids, two for each polyp. The other forms of the group Sfzcate in which the polyp has developed a calyx: Famniculina, Halipteris, and Stachyptilum, seem to me to show no close relation, so that they certainly cannot — as has been done by Bourne — be placed in one family. On the contrary, the genus Stachyptilum K@ll. seems to me to be altogether a stranger in this group; as far as I can see from the description and figures of Kélliker, I think it must be referred to the first group, Pennatulee, and be placed there in the family Pennatulide; the whole form of the colony and the arrangement of polyps and zooids is as in Pennatula: the polyps are placed quite regularly in oblique series, but the members of one series are not mutually coalesced; the dorsal surface of the stem is covered with zooids, as are also the lateral and ventral intervals between the polyps; apart from a slighter development of the points of the calyx it looks like a Pennatula with free polyps. The genus Funzculina occupies a quite isolated position by wanting real zooids; only the quite young polyps appear temporarily as zooids; by and by they increase in size, get fully developed ten- tacles, form sexual organs, and become perfect polyps; the arrangement and growth of the individuals is not regular, although they, as it were, tend towards a regular arrangement in transverse series. From all these facts, /wnzculina seems to me to occupy an especially primitive place below all known sea-pens, and, at all events, a sufficiently peculiar place for it to form a separate family in which no other genus can be included for the present. Haldpterts, which has been placed as its nearest ally, since K6lliker in his monograph had made this arrangement, is in reality far removed from it; the resemblances found are common to most long and slender sea-pens. In Halifieris, there is as strong a contrast between zooids and polyps as in any other sea-pen; in the calyx of its polyps the abaxial side is far more developed than the axial one where teeth are quite wanting in the calyx. In reality, I think that the original reference of the typical species H. christit to «Virgularia» has more nearly approached the correct thing; it is, as stated above, a Pavonarid; as in /avonaria the calyx of the polyp is provided with two abaxial teeth; the only essential difference is that the polyps do not coalesce to form real wings. With regard to the genera with polyps without calyx, Scleroptilwm Koll. shows the simplest form of colony; as it shows no immediate relation to the other known genera it must, I presume, form a separate family Scleroptilide’). The genus Axthoptilum must still form the type of a family Axthopiiiide; but in this family 1) The genus was established by KdOlliker (Chall. Rep. Vol. I, p.30, Pl. VII, fig. 29) for the species S. grandifiorum and durissinum, both from considerable depths in the Pacific off Japan; Verrill has later found another species S. gracé/e, of a length up to 1 foot, in the depths of the Atlantic to the east of North America (Cape Hatteras) (Rep. Comm. Fish and Fisheries for 1883 (1885) p. 510 [8], PL III, fig.6, and Am. Journ. Sc. Vol. 28, 1884, p. 219); the description in the latter place speaks of «ventral» zooids, and such zooids, according to Kélliker, ought to be wanting. PENNATULIDA. 9 the genus Benthoptilum*) of Verrill may probably also be included; further, I think that the two families Kophobelemnonide and Umbellulide are to be kept unaltered. Umbellula I regard — contrary to Delage and Hérouard — as a particularly specialized form, related to Kophobelemnon; an Umbelliula might be thought to be derived from a Kophobelemnon-like form in which the polyps in the greater part of the rhachis were checked so as to be kept in the zooid-stage; the zooids in Umbellula are somewhat more developed than ordinary zooids, in so far as they are generally provided with one tentacle; and a certain resemblance to Kophobelemnon is undeniably seen in such Umbellula-species as U. gtintheri K. in which the polyp-bearing part of the rhachis is lengthened in a club-shaped manner. The radial structure seen in certain Umébellula-species (for instance U. encrinus), is, at all events, a secondary development, having plainly arisen from an original bilateral formation, as is sufficiently shown by the younger stages. To Umbellulide, the Chunnelide Kkth. are no doubt related; in these latter also the radial arrangement of the polyps is plainly a secondary one; the bilateral formation may unmistakably be traced in the arrangement of the zooids, but may also be seen in that of the polyps. I think it superfluous to form a separate main group for this family. According to this, the section Sfzcate will have to be divided into two subsections: A. The polyps with calyx. Fam. Funiculide (Funiculina Lam.). » Protoptilide (Protoptilum, Distichoptilum). B. The polyps without calyx. Fam. Scleroptilide (Scleroptilum). » Anthoptilide (Anthoptilum, Benthoptilum). » Kophobelemnonide (Kophobelemnon, Sclerobelemnon, Bathyptilum). » Ombellulde (Umbellula). » Chunellide (Chunella, Amphianthus). The sections Renillee and Veretillee must probably still remain. Even if Renilla in its development, shows a distinct connection with a common primitive Pennatulid, it is so peculiar when fully grown, that it does not agree with other groups. This holds good to a smaller degree with regard to Veretillee; the radial arrangement of polyps and zooids in these forms, however, is a peculiarity justifying a separation, at least provisionally. I think it probable that this radial arrange- ment will meantime prove to be of secondary origin, since a bilateral formation may be traced in the inner structure of the stem in Veretzl/wm (comp. Kélliker Monogr. p. 430). It is rather remarkable that no young stage of this group is as yet known, although it is represented, in the Mediterranean for instance, by two rather common genera: Veretillwm and Stylobelemnon. Both these groups: Renillee and Veretillee, 1 must regard as very specialized and anything but primitive Pennatulids. With regard, finally, to the group Gindulee (Acaulina Del. & Heér.), it must be quite omitted; it has been established only on the «genus» Géndul, which is nothing but a mistaken fragment of Pavonaria finmarchica (see under this species). 1) &. sertum Verrill, from the depths of the Atlantic to the east of North America (Cape Hatteras)..Rep. Comm. Fish etc. 1883, p. 510 [8], Pl. II, fig. 4, and Am. Journ. Sc. (3), Vol. 29, 1885, p. 149, note. The Ingolf-Expedition. V. 1. 10 PENNATULIDA. List of the northern Pennatulids. By northern Pennatulids are here meant the species known from the Davis Straits and Baffins Bay, the Greenland Sea, and the part of the Atlantic north of 55° N. Lat. and east of the meridian of Cape Farewell. The reduction made in the previous section of formerly established genera, will appear clearly in a list of the Pennatulids from this territory, which has been the special object of my investigations; to this is further to be added that I have found it necessary to unite together several of the earlier established species. Notwithstanding the fact that several forms have now been added from within this geographical territory, which were not hitherto known to occur there, and among these two new species, the total number of species will be seen to be diminished to about two thirds of the number which might be enumerated from previous publications. The following list A contains the earlier established species, the list B the species as I now think they should be. In the latter list, the species marked with an asterisk have been brought home by the Ingolf-Expedition itself. A. B. 1. Pennatula phosphorea \,. : = 1. Pennatula phosphorea lu. 2. » distorta Kor. Dan. 3. > aculeata Kor. Dan. Af noe aculeata Kor. Dan. 4. » grandis Ehrenberg Ce » grandis Ehb. As it See prolifera un. sp. 5. Virgularia affints Kor. Dan. 5. Virgularia affints Kor. Dan. 6. > mirabilis (Ly) 6. » mirabilis (1,.) 7. Cladiscus gracilis Kor. Dan. 8. » lovent =» » 9. > kélltkert » > = 7. *Virgularia cladiscus nom. n. 10. Svava glacialis ee 11. Deutocaulon hystricis Marsh. & Fowl. 12. Dzubenia abyssicola Kor. Dan. 13. > elegans M25 ED = 8. Stylatula (Diibenia) elegans Kor. Dan. 14. > borealis » 15. Pavonaria finmarchica (M. Sars) ; : 4 paren = 9. *Pavonaria finmarchica (M. Sars) 16. Géndul mirabilis Kor, Dan. 17. Halipterts christit (Kor. Dan.) 18. ZLygomorpha sarst » » ene rae ; : ; : = 10. Halipteris christi (Kor. Dan.) 19. Stichoptilum arcticum Grieg 20. Protoptilum tortum » PENNATULIDA. Il 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33- Protoptilum lofotense Kor. Dan. » mohni » » > carinatum » armatum » » Funiculina quadrangularts (Pall.) Kophobelemnon stelliferum (O. F. M.) > abyssorum Kor.Dan. > moebit Pes = > leuckartit Koll. Gunnerta mirabilis Kor. Dan. Bathyptilum carpentert Koll. Umbellula encrinus (1,.) > gracilis Marsh. II. 12. 13. 14. 15. 16. 17. 18. 19. 20. ai. Protoptilwm thomsont Koll. *Protoptilum carpentert Koll. * > denticulatum un. sp. *Distichoptilum gracile Vert. *Funiculina quadrangularts (Pall.) *Anthoptilum grandifiorum (Vert.) *Anthopttlum murrayt Koll. *Kophobelemnon stelliferum (O. F. M.) Bathyptilum carpentert Koll. *Umbellula encrinus (1,.) * » lindahii Koll. Fam. Pennatulide KOll. Pennatula Lam. Pennatula aculeata Kor. & Dan. PL I, fig. 1. Pennatula aculeata Kor. Dan. 1858. Forhandl. i Vidensk. Selsk. i Christiania 1858, p. 25. > Kor. Dan. Fauna littor. Norvegice. III. 1877, p.86. Tab. XI, figs. 8—o. » » » distorta var. aculeata Kor. Dan. phosphorea var. aculeata KOll. Marshall. >» >» > americana Moroff. Octoc. 1902, p. 381. Nye Alcyon. Gorgon. etc. 1883, p.24. Tab. XI, figs. 5—10. Monogr. p. 134 and 366. Penn. «Triton», 1883, p.123. Tab. XXI, figs. 4, 5, 7; Tab. XXII. The | 46> | 45 > | 38 » 38 >» | 40> | 30> | 35> |] ..... | 14 > Wumber of wings .................. | 23/24 19/20 20/19 17/17 17/17 21/21 14/ 14 14/15 13/13 teres 8/8 Length of a well-developed wing.... | 2omm | 22mm | ygmm | 24mm | 24mm | grmm_ yqmm | grmm | y5mm 21mm | 8mm Breadth of such a wing above the base. 3» 3.26 )ca3 21 3 3% [25 > | 22% | 2» 2 > |2%6—3 » 5 > Number of polyps in such a wing... 8 be) 10 9 9 9 7-8 98.157, g-II 3—4 Length of the largest spines (zooids). | 3mm) 2mm | 3mm | 28mm|/ 3mm | 2mm | 3mm | 3mm | 2-2mm | 33:5 mel c, Imm | | | | None of the first ten specimens show any distinct terminal polyp; some (Nr. 1, 4, 5, and 7), on the other hand, have a distinct terminal zooid, in Nr. 1 with a spiniform, large, but distinctly two- pointed calyx; in some, rudimentary wings with reduced polyps are found at the apex; in some distinct top-zooids. Nr.10 is a torn-off pen, not quite complete below. In the specimen Nr. 11 (from St. 166), on the contrary, the terminal polyp and terminal zooid are well developed; in spite of its slight size it may already be recognized quite distinctly as belonging to the species aculeata; a few of the zooids are not only larger than the others already, but have also the long calyx-tooth distinctly marked; moreover, the specimen is of the same dark, violet-red colour as the ten specimens from St. 167. Pennatula aculeata is originally known from the western coast of Norway; it is here (see Grieg 30, p.9) said to go farther north than P. phosphorea, viz. to the Lofotens, and to greater depths (400 fathoms); it seems not to have been found farther south than Askevold (Séndfjord). Later it has been found by the English expeditions of the «Porcupine», «Triton», and «Knight Errant» at the following places in the Atlantic: west of Brittany (48° 26’ N. Lat. 9° 44’ W. Long., 358 fathoms); north- west of Rona (the Hebrides) (59° 4o' N. Lat. 7° 13' W. Long., 556 fathoms, and 59° 37’ N. Lat. 7° 19' W. Long., 520 fathoms); further, by the Prince of Monaco near the Azores (38° 34' 30” N. Lat. 30° 43' 30” W. Long., 1287 M.); by the French expedition of the «Caudan» in the Bay of Gascony (46° 28' N. Lat. 7° W. Long.*), 1710 M.); by the «Travailleur»>, west of the Spanish peninsula (P. Fischer, 23, p. 38). To these loca- lities are to be added as new ones: south of the Vestman-Islands, 63° 5’ N. Lat. 20° 7' W. Long., 557 M. (the «Thor», St. 167, "/, 1903), and 52° 57’ N. Lat. 19° 58’ W. Long., depth 956 M. (the «Thor», St. 166, ‘4/, 1903). The species is not known from the North Sea nor the adjoining seas, the Skager Rak, Kattegat etc, nor from the Mediterranean. On the American side of the Atlantic it is very common, and is hitherto known from the Gulf of St. Lawrence as its northernmost locality (Whiteaves), to off Chesapeake Bay. To this occurrence on the American coast is to be added the new locality, St. 25 of the «Ingolf», Davis Straits, 63° 30! N. Lat. 54°25’ W. Long., 582 fathoms, by which observation its northern limit on the American side has been carried considerably farther north. According to the above, this species is assuredly to be regarded as a purely North-Atlantic one; it is not known south of the Azores, nor on the American side farther south than 33° N. Lat. (Agassiz: Three Cruises of the 1) Reckoned from the meridian of Paris. 14 PENNATULIDA. Blake, vol. II, p. 142. Bull. Mus. Comp. Zool. vol. 15); it seems to be limited essentially to the margins of this part of the Atlantic and to the slope into great depths. As to its bathymetric range the lowest depth is 30 fathoms, at Norway, but upon the whole it occurs in greater depths; on the American side, where, as mentioned, it is very common, it is still found at 1255 fathoms (Verrill: Rep. U.S. Fish. Comm. 1883 (1885) p. 509), but it is most abundant between 150 and 300 fathoms. Whilst the very closely allied Penn. phosphorea has not yet been found in America, a Penn. americana has been described by Moroff. According to the description, it is beyond all doubt, however, P. aculeata. M.’s specimen is also from Massachusetts (see Zool. Anz. 1902 and Zool. Jb. 17. Bd., 1902). Pennatula phosphorea L. var. candida Marsh. & Fowl. A. Milnes Marshall and G. H. Fowler: Report on the Pennatulida dredged by H. M.S. «Porcupine». Trans. Roy. Soc. Edinb. vol. 23, part 2, 1887, p. 456, Pl. XXXI; Pl. XXXII, Fig. 3. This form has been founded by Marshall and Fowler lc. on three fragments taken by the «Porcupine» in 1869 at 62° 1’ N. Lat. 5° 191 W. Long. Our Zoological Museum has obtained 17 specimens from the Vestman Islands, 7 from the physician Thorsteinn Jénsson, who, according to «Skyrsla um hid islenzka natturufreedisfélag drid 1897—98», p.12, has also given a specimen to the Natural History collection at Reykjavik; 8 specimens from the cruise of the «Diana» in 1900 (A. C. Johansen); r specimen from rgor (R. Horring), and 1 from the collection of the «Thor» in 1903. The species is peculiar by its being perfectly white in spirit, almost all the spicules being colourless; the animal, when living, may be of a slight reddish colour (according to Herring), and in a single specimen the red colour is still distinctly seen (see below). Otherwise it agrees in several features with Penn. phosphorea var. angustifolia K6ll. (Monogr. p. 130). The pinnules are slender, and the calices of the polyps are long and conspicuous, provided with 8 long teeth formed by spicules; the number of pinnules on either side 17—24 (often differing somewhat on the two sides of the rhachis; comp. below). The tentacles of the polyps carry spicules along the dorsal side of the stem. The large spicules, as elsewhere in P. phosphorea, are fusiform, and thus only conspicuously triangular at the ends; in section, they are somewhat round; the smaller ones, on the other hand, are altogether triangular. The largest polyp-spicules measured are 1'072™ long, and o'072™" thick; the smallest are 0°04—0'048"" long, but the general length of the spicules of the tentacles is o7096—o-128™™. The specimens in hand show the following particular features: Nr. I 2 3 4 5 6|7|8|o zo | tr | 12 | 33 | 1 15 16 17 t g Total length in mm. || 37 65 65 65 | 66.) 9892 he te Fe So Length of stalk.. » 15 on 30 90; -| 320k BO eee hae salt as 35 40 | 40 4) | cag Nias Number of wings... |12/19| 22/21 | 19/19 |20/18 17/19 22/23/22/22 18/17 19/18 19/20| 23/24 2t/ar 20/21 | 22/21 20/20)25/23| 18/18 Number of polyps in one of the large wings 6 |9—10|8—r10] 9 II 9 10 9 8 |8—g| 1o—12 Io |g9—11|8—1r| 9 10 |8—r10 } | Length of a wing in | | ps Ba ea eieeeele te 10 15 II 13 14 12 II 16 14 | 17 16 | II'5 | 19 Rela es 12 13 PENNATULIDA. 15 The specimen Nr.1 is distinguished by having rather a strong red tinge, so that it approaches somewhat the common phosphorea; both polyps and zooids contain red spicules in considerable num- bers; the polyps are red, but with white calyx-teeth and white lateral edges; the edges of each wing are therefore white, and the red surface is streaked with narrow white lines marking the boundaries between the coalesced polyps. A quite similar distribution of the colours, only with a far deeper red, is not uncommon in typical phosphorea specimens from the Kattegat. Further, this specimen has kept both terminal polyp and terminal zooid, as is also the case with the specimens Nr. 4 and Nr.g. In the specimens Nr. 3, 7, 12, 15, 16, and 17 the quite distinct terminal zooid is preserved in its full development, but the terminal polyp is transformed into a zooid, either of equal size with the terminal zooid (Nr. 15), somewhat smaller than this (Nr. 12), not much smaller (Nr. 3) — or quite atrophied (Nr. 7, 16, 17). Although the specimens in question, with the exception of Nr. 1, are large and must be considered as fully developed, the features mentioned are quite uncommonly distinct. The other specimens show much altered features at the upper end of the rhachis; some specimens (e. g. Nr. 6) show very beautifully a number (4) of large «top-zooids», amongst which, probably, both the terminal polyp and the terminal zooid are to be sought. In the specimens Nr. 2 and Nr. 7, the peculiarity is found that one of the dorsal zooids near the middle of the edge of the rhachis is developed as a soli- tary polyp, 7 and fully 4™™ long respectively. The specimen Nr.14 approaches somewhat the «lanci- foliate» form‘). Most specimens contain ripe sexual organs, and some seem to contain larve. Occurrence. This light-coloured form has been taken by the «Porcupine», as mentioned, on 62° 1' N. Lat. 5° 19! W. Long., accordingly E. of the Faeroe Isles, at a depth of 114 (English) fathoms. All the specimens here mentioned are from the Vestman-Islands; with regard to Nr. 2, 5, 8, 10, II, 13, and 14 there is no information as to the depth or the bottom; but to judge from the attached particles it must have been black Basalt-mud; Nr.g has been taken at 50—6o0 fathoms, E. of the Vestman-Islands, 2'/, miles E.N.E. of Storhdfdi; the others have been taken (the «Diana», 1900, St. 41) on «clay-bottom», at a depth of 68—7o fathoms; Nr. 17, the «Thor», 1903, St. 169, 63° 24’5’ N. Lat. 20° 1’ W. Long., depth 120—158 m., ooze. All the specimens have been taken during the summer months (July, August). This pale or white variety probably occurs on restricted localities within the wide extended area of the species, but with regard to the more particular conditions under which it is produced and thrives, we have no information which could render any certain statement possible. The question can scarcely be of a casual albinism, as all the specimens known from the Vestman-Islands belong to the same pale variety, and in its red forms Penn. phosphorea is altogether unknown hitherto from any locality at or round Iceland or the Faeroe Isles. Other discoveries of white forms of Penn. phosphorea are, so far as I know, only referred to by two authors. K6lliker describes a white specimen in his Monograph, p. 133; it belongs to the Museum in Copenhagen, and I have thus been able to compare it with the form from the Vestman-Islands; it is very different from this latter, and K6lliker with good reason has determined it as a subvariety alba of Penn. ph. var. lancifolia. Unfortunately, we 1) When Marshall and Fowler give 4:2: as the greatest length of a pinnula in their specimens, this is surely due to a misprint; such a length is scarcely found in any form of P. phosphorea, and the figures on Plates 31 and 32 give also a length of only ca. 14—15™™, 16 PENNATULIDA. have no information at all as to the place where the specimen was found. Esper’s Pennatula alba which K6lliker has referred to his var. angustifolia, closely resembles, to judge from the figure, 22, pl. VI, the specimens in hand from the Vestman-Islands; their colour is said to pass almost impercep- tibly into a reddish one; but we have no information as to the place where these specimens of Esper were found. Verrill mentions a white specimen of Pennatula aculeata (Amer. Journ. Sc., vol. 23, p. 310, and Report... «Blake» p. 3) taken by the «Fish-Hawk» at st. 1025, depths 216 fathoms; he says of it: «This is doubtless only an albino»; it appears to have been taken together with numerous specimens of the common red form of Penn. aculeata (comp. Am. J. Sc. 23, p. 315). Remarks. According to what has hitherto been known, it must be regarded as hardly pro- bable that Pexnatula phosphorea should be found outside the Atlantic. Studer, however, in his preli- minary report of the «Albatross» Expedition (p.55) has given Penn. phosphorea as from the Pacific, from o° 19' N. Lat. go° 34’ W. Long., 331 fathoms, and «Pennatula distorta, Dan. Kor, var. pacifica, n. var.», from 1° 7’N. Lat. 81° 4’ W. Long. Of the former, it is said that the (single) specimen was not to be distinguished from the Atlantic (i.e. the typical) phosphorea I, As to Penn. distorta Kor. and Dan., this species cannot be maintained, in my opinion; it is nothing but Penn. phosphorea with unusually narrow and (casually?) distorted wings; and the Penn. distorta var. aculeata described by the same authors, is obviously only «distorted» specimens of the species Penn. aculeata. And with regard to Studer’s «Penn. distorta, pacifica>, his remark that the colour is whitish yellow, and that the calyx- spicules appear to be longer and more conspicuous, seems to me to indicate that another species must be in question; from the Asiatic side of the Pacific several Pennatula-species are known, among others just such forms with narrow leaves in which yellow or reddish yellow is prevalent in the colour, and where the type of spicules, moreover, is different from that of phosphorea; also the depth, 1740 fathoms, must be regarded as somewhat great for the supposed species. Th. Moroff (Zool. Anzeiger 1902, p- 579, and Zool. Jahrb. Abth. Syst. 17 Bd., 1902, p. 380) has established a P. phosphorea var. longispinosa from Japan (I specimen); unfortunately, his figures show only that the specimen is a Pennatula; but some remarks in the description would indicate that Ahosphorea is not in question (for instance the broad streak between the «ventral» zooids). Pennatula grandis Ehrb. Pennatula grandis Ehrenberg. Die Corallenthiere des rothen Meeres, 1834, S. 66. > borealis M. Sars. Fauna litt. Norvegie, I, 1846, p.17, Pl. II. Ptilella borealis Gray. Catalogue of Sea-Pens, 1870, p. 21. This splendid species has not been taken by the «Ingolf» itself; but during one of our sojourns at Thorshavn I received from Mr. Finsen, apothecary, a specimen taken on «the bank south west of the Faeroe Isles at a depth of go fathoms»; later, 19 specimens from the Faeroe Isles have been sent to our Museum by the agent Mr. Miiller, taken on the fishing bank east of Fuglo, ca. 150 fathoms. With the exception of one individual (Nr. 20) all the specimens are large and well-developed. I can add nothing essentially new to the former descriptions (by Sars, Kélliker, Koren & Danielssen, PENNATULIDA. 17 Verrill, a.0.) of this species which can scarcely be confounded with any other. I shall only make a few observations on the spicules, as these have not been described in particular by the earlier authors. The spicules are of a marked triangular type; in the larger types especially the polyp-spicules (i. e. of the calices, the upper part with the tentacles containing no spicules) the three ridges which constitute the spicule show their free edges to be distinctly thickened, whilst the sides are hollowed; in the red spicules, therefore, the colour is deeper at these thick edges. The polyp- spicules measure 0.144—0.897™ long, and o.o16—o,048™" broad. The colourless part of the peduncle is provided with short, coarse spicules 0.088—o.120™ long, the thickest ones being 0.024™™ thick; these spicules are of a very slight reddish colour; the deep-red spicules from the band-shaped edge of the bulb are 0.096—0.200"" long, and o.016—0.032™" broad. The specimens in hand from the Faeroe Isles show the following particular features: Nr. I | 2 3 | 4. | 5: 6 Cs 8. g. | 10. | Bie kad b4. [Ae | Thee] TGS] 37. 18. Ig. | 20. | ] Totallength inmm. || 445 | 445 | 435 435 433 | 430 430 | 425 j-440 420 | 410 | 400 | 398 | 395 | 390 | 380 | 370 | 315 | 210 | 107 Lengthof peduncle, | | to the edge of the | | bulb .... in mm. | 125 | 110 | go | 130 124 110 | 108 | 110 | 125 | 110 | 82 | 92 | ror | 80 | go | 105 | svael 30 Diam. of bulb » 32 | 34 35 | 32 30 | 30 | 29 ma | 2 | 30. | a4 | eel 6 20 | 23 23 7 Number of polyps | | in a well-devel- | | loped, arge wing | 83 | 73 | 70 | G74 7b 70 173, | 68 | 82 | 78 | 67 |. 72.) 78 | Sr)... | G2) Ba |... | 25 Number of wings. /39/40\42/44 44/41 37/ nine 37/35|37/ 38 36/35|37/40\39/39|41/41/42/4x/89/39|37/37|34/34|39/39| (34/32) 39/31) 18/17 Length of a large | | | | | wing.... inmm. | | 45 | 44 | 35 4r | 23 | 38 | 43 | 45 | 47 | Lieu, 9 Greatest breadth of | | | AGE ry in mm 23 | 30 |... 26 Sg: ise jee [az |' 24°} x7 | : een Ss Length of polyp- | | | | calyx ...inmm. eet ti roe} an res BS 3—5/3—4| 5 | 5 |4-6| 5 ee fl | 4 The specimens Nr. 18 and 19 are defective; in Nr. 18 the lowermost wings are wanting, the tissues of the bulb and the peduncle having been rubbed away; Nr. 19 is broken off below the bulb; the rhachis, which is complete, is ca. 70™™ long. Nr. 20 is the smallest specimen of this species I have ever seen amongst the large number I have had the opportunity to examine in our Museum and in Bergen and Christiania; Verrill mentions that he has had several young specimens and one very young, but he gives no measurements nor any further information. For the sake of comparison, I may just mention that the two specimens of M. Sars, upon which the species P. borealis was established, were of a respective length of 420 (16") and ca. 810" (31"), with 37 and 57 pairs of wings; in other Norwegian specimens measured by Koren and Danielssen, the length is given as 250—780™" with 30—7o pairs of wings and 48—100 polyps, whilst Verrill mentions a specimen of 530™ in length with 36 pairs of wings, as one of the largest American specimens. The colour in some specimens is brownish or dark brownish-red (especially so in Nr. 20, partly in Nr. 18 and 19), but most frequently of a beautiful red or reddish yellow. Occurrence. The species was only known hitherto from the fjords and coast-regions of Norway, the eastern coast of North America where it is common on the fishing banks off New The Ingolf-Expedition. Vr. 3 18 PENNATULIDA. Foundland, Nova Scotia, and the northernmost of the United states (as far as off Nantucket Island and Martha’s Vineyard). To these localities are now to be added the banks round the Faeroe Isles. It has been taken at depths from s50—1255 fathoms’). Pennatula prolifera un. sp. Pl. II, figs. 15—24. From two stations in Davis Straits, 24 and 36, we have obtained a Pennatula-species in very large numbers, from the former station about 4o specimens, from the latter upwards of a hundred; but all the specimens are young ones, several of them as small and apparently as young as the youngest known stages of Pennatula phosphorea’); even the largest and most developed specimen is still little developed, as will be shown more particularly below, so far as the pen’s provision of polyps is concerned. It is a very difficult thing therefore to characterize the species; or, to speak more cor- rectly, it is impossible to diagnose it briefly by such features, as are elsewhere used in distinguishing species within the genus Pennatula. That the species is a new one not hitherto described, will be obvious, I think, from the following description, in which I shall give an account of the peculiar mode of propagation in this species which has led me to give it the specific name frolifera. All the specimens were white, with the exception of two from St. 36, which were light-red; but these also have now turned as white as the others in spirit. The largest specimen (from St. 24) has to some degree the penniform appearance (figs. 15, 16). The total length is 35™", in which the rhachis accounts for 18™™, measured from the terminal zooid (see below) to the lowermost polyp-bud, the peduncle 15!/.™™; the breadth of the pen is 4.5™™ (the polyps being pressed in against the stem); the rhachis itself is ca. 1.5™™ broad, the terminal bulb of the peduncle is 2.5™™ long and ca. 1.5™™ broad. On either side of the rhachis are eight distinct wings and below these four rudimentary wings. The calyx-mouth of the polyps is provided with 8 long and strong spines supported by long spicules. The form of the calyx, as is ordinarily the case in the genus, is somewhat different accord- ing to the degree of retraction; when the polyps are quite retracted, the calyx-teeth are closed and the whole calyx has a lengthened oval form reminding one of a narrow grain of barley. The longest calices are 5™™ long and ca. 1™ broad. The tentacles are on the aboral side provided with longi- tudinally arranged spicules, both on the stem and the lateral branches. With regard to the arrangement of the polyps it has first to be pointed out that no terminal polyp is present: the pen ends above in two polyps placed (almost) side by side and equally developed; between their bases the upper end of the rhachis itself is found, formed by a large zooid (with two calyx-points). Each of these two uppermost polyps forms a wing (I). This uppermost pair of wings is separated from the next pair (II) by a somewhat long part of the rhachis, this latter pair again by a somewhat shorter distance from the third pair (III), whilst the following pairs (IV—VIII) follow one another more closely. 1) Verrill: Rep. U S. Fish. Comm. 1883, p. 509. 2) Figured by Jungersen, 1888, Pl. V, figs. 1—2. PENNATULIDA. 19 Apart from the uppermost wing the other wings contain more than one polyp, as at all events a rudiment of one or two polyps more is found in each wing to the ventral side of the oldest one. In the uppermost pair of wings but one (II) the polyp of the left side (1) carries a small protuberance (*), which I take to be the first trace of individual Nr.2; the right wing has a distinct individual 2, but still in the zooid-stage; in wing III, polyp 2 is large and well-developed on either side of the colony, but nothing is seen of 3; in wing IV, polyp 2 is still larger, and 3 is present as a zooid; in wings V—VII there are also 3 individuals, but 2 is smaller than in the preceding wings, and decreases in development below, while 3 is still only a zooid in all of them; in wing VIII there are only two polyps, of which Nr. 2 is only a zooid. The four rudimentary wings (IX—XII) consist, each of them — more or less distinctly — ot two individuals, both still zooids. Of zooids, there are 2 longitudinal rows dorsally, one on either side, separated by a bare streak (i.e. devoid of individuals), about 0.57" broad below, narrowing above (a similar naked streak runs along the rhachis between the polyps of the ventral side); in each row an indication of a grouping into 3—4 zooids for each wing may be traced. The series is already indicated below at the smallest rudiments of wings. — Between the uppermost wing and the next the series is very open, and only one zooid is found at each of the uppermost polyps (I). The zooids look like small scales supported by spicules. At the upper end of the rhachis the rows are completed by a median, con- siderably larger, single top-zooid, Z, provided with two distinct calyx-points'). Of lateral zooids, one seems to be begun immediately above each wing, with the exception of the two uppermost pairs of wings. ‘These rudiments, however, are very small, and difficult to determine with certainty. The spiculation is copious and dense; the longest spicules are found on the polyps, the next on the zooids and rhachis where they are mostly arranged longitudinally; there are also many transverse, rather long and thin, spicules on the peduncle, intermingled lower down with longitudinal, short, oval ones; towards the terminal bulb the spiculation diminishes to open, longitudinal rows of short, oval spicules, and the bulb itself has no spicules, and is therefore transparent, so that the interior longitudinal septum is seen (but no terminal pore; I think upon the whole that the existence of such a pore in Pennatula normally is very doubtful). The spicules (especially with regard to the long ones) are of quite a different type from that predominant in Penn. phosphorea. The fact is that all the spicules (excepting the oval ones in the peduncle) are distinctly and sharply triangular, recalling mostly those of Penn. grandis. The longest measured are 0.784™" long, 0.04™™ broad; the short and thick ones are respectively 0.160™™ and 0.04— o.112™, and 0.032™; the small oval spicules of the peduncle are 0.064—0.088™™ long, and 0.016™™ broad. The calcareous axis terminates above, bent to the right, opposite to the third uppermost wing, and below, also curved, just above the terminal bulb. The sexual organs are fully developed; they may be seen shining through the po- lyps without any further preparation, but still more distinctly by clearing with oil of cloves; in 1) In this specimen the top-zooid has by chance been wedged in between the two uppermost polyps, so that it is only partly seen in the figure, but both from the dorsal and the ventral side. 2) Comp. Jungersen, 1888, p. 161. a 20 PENNATULIDA. all the more developed polyps, from the uppermost to the two lowest, large eggs (or larve) are seen; in the rhachis, on the other hand, no sexual products are found. In the other specimens, about 150, that I have been able to examine, there is found, a so to speak continuous series of stages, from the one just described down to a stage where the colony is provided with only one pair of wings (I). Representatives of these stages may be found figured on Pl. I], figs. 17—24. One specimen (St. 24) of a total length of 29™™, is provided with six distinct pairs of wings, and below these with tiny rudiments of two pairs more. Each wing of the uppermost pair (I) consist of three individuals: one fully developed polyp, one somewhat smaller, and one still a zooid; the next wing (II) shows the same development. The wings III—V still consist of three individuals, but two are in the zooid-stage, and the most ventral, youngest one, is already very small in IV, in V both are very small; VI consists of only two individuals, a small polyp, and a small zooid; both the rudiments (VII, VIII) consist of a larger and a quite small zooid. Another specimen (St. 24), of 25™, shows 5 pairs of wings and 2 pairs of rudiments; all the wings consist of two individuals, the two uppermost of a developed polyp and one little developed, (wing I on the right, however, has three individuals, being provided with a small zooid), the others of a polyp and a zooid, both decreasing in size. from above downwards; the rudiments consist of two zooids. Other smaller specimens of 21™™, also show 5 pairs of wings and 2 or 3 pairs of rudiments; others again (25—16™™) 4 pairs of wings, 3 pairs of rudiments, others (17—13™™) 3 pairs of wings, 3 pairs of rudiments; others again 2 pairs of wings and 2 pairs of rudiments, Pl. II, figs. 17,18; in all these specimens each wing consists of two individuals, a polyp and a zooid. Finally, we have several specimens with only one pair of wings, and below one or two pairs of rudiments, and some specimens with only one pairs of wings and no rudiments below (PI. II, figs. 19, 20). Each wing in these latter, however, consists also of two individuals, a developed polyp and a small zooid. The proportion between the rhachis and the peduncle as to length is somewhat varying; some- times the former, sometimes the latter is the longer, but often they are about equal in length. Devel- oped sexual organs (sometimes larvae) are seen in most polyps, at all events in the developed ones; in quite small (10o™™ long) specimens indeed with only one pair of wings and 2 pairs of rudiments, I have seen large and developed sexual organs in one or both of the large polyps. Even the smallest specimen, of 6™™, represented in figs. 19, 20, shows (by clearing with glycerine) the sexual organs, though not yet ripe. All the specimens have the two rows of zooids (z), but their number in each row is very varying, increasing with the number of wings; in the specimens with only one pair of wings only one or two zooids are found in each row. ‘The small lateral zooid above each wing is only seen in the tolerably well developed colonies. A common feature in all the specimens is further, that the stem contains a calcareous axis, and that the rhachis terminates above in a large zooid (Z). This series of stages therefore is evidently a developmental series, in which the last-mentioned specimens with only one pair of wings are the youngest. That this developmental series belongs to a species of the genus Pennatula is undoubtedly a fact; but the rather obvious supposition that the young stages of the pale variety of Penn. phosphorea var. candida before mentioned might be in question, is inadmissible on account of the difference in the type of the spicules, and a closer comparison with PENNATULIDA. 21 the developmental stages known of P. phosphorea renders it quite untenable. Penn. phosphorea always shows more numerous and more advanced polyps in the single wings, at stages where the number of wings is as in specimens of the present new species, and the colony is upon the whole more developed at corresponding sizes; on the other hand, the young colonies of P. phosphorea show no ripe sexual organs. These features, however, might be thought to be influenced by the conditions of life at such great depths as the one from which the new form has been obtained. The most con- clusive thing to my mind, however, is the great difference in the construction of the upper end of the rhachis. In young colonies of Penn. phosphorea the rhachis always terminates above in a terminal polyp, and on the dorsal calyx-base of this there is a large terminal zooid which completes the rows of dorsal zooids'). Both terminal polyp and terminal zooid are found even in well-developed colonies with complete pen-form (for instance, in specimens of more than 35™", with 12 pairs of wings, the largest of which are provided with 5—6 polyps; but more rarely in still larger specimens); in the new species however, no stage not even the youngest, shows the least indication of a terminal polyp: the rhachis ends as stated in a latge zooid, and the two nearest polyps (or wings) are equally developed and placed at almost the same height; thus the younger stages especially have a dichotomous appea- rance, markedly different from that of Penn. phosphorea. According to what has been known hitherto, the primary individual, developing directly from the larva which arises from the egg, is in all Pennatulids a polyp with tentacles (see my treatise pp. 162, 173); as this polyp grows greatly in length, its lowermost part below the calyx forms the stem (rhachis -+ peduncle); on this lengthened part the other individuals, polyps and zooids, appear gradually (in some genera zooids are also formed on the polyps). During the growth of the colony the terminal polyp may later disappear; this is probably the case in most of the genera of sea-pens, and it is the rule in Penn. phosphorea. Where the disappearance of the terminal polyp is not due to a mere displacement (as in Kophobelemnon), it may be due either to an abortion or a transforma- tion into a zooid. In the genus Pteroeides the latter seems to be the case; when the colony has grown to a certain size, the end Of the axis, at all events, carries a very large zooid instead of a ter- minal polyp, (see Kélliker, Monograph, p. 356, my previous treatise, p. 172, and v. Koch 2, p. 398). In Penn. phosphorea, the end of the rhachis of large colonies carries generally several very large zooids «top-zooids»; amongst these presumably, are both the original terminal zooid and — by a retrograde transformation — the terminal polyp and some of the uppermost wing-polyps (comp. Jungersen p. 168, and v. Koch). With these facts in view we return to our new Pennatula-species. It may be main- tained with a probability, bordering on certainty, that this species like P. phosphorea, has also had a stage not only with a terminal polyp, but also with a terminal zooid on its calyx-base. — I shall only recall the fact that such a distant form as Renzlla agrees completely in this respect with P. phosphorea. How, then, has the terminal polyp disappeared here? 1) If a displacement had been the cause, one of the two uppermost polyps would have to be regarded as the terminal polyp, and the large zooid of the end of the axis would represent the terminal zooid. This supposition, however, is inadmissible, partly because this large zooid is not placed in the same way as the real terminal zooid in P. phosphorea and Renilla, on the dorsal calyx-base of either 1) Comp. my previous treatise p. 168, and Marshall & Fowler, Penn. «Porcupine», Pl. XXXII, figs.5 and 7. 22 PENNATULIDA. of the polyps in question, partly because both the upper polyps (I) always (or, at all events, with very rare exceptions) distinctly appear as lateral polyps, i.e. wing-polyps, a polyp Nr. 2 sometimes even a Nr.3 being found at their base on the ventral side of the colony (whether these latter ever grow to their full size is of no consequence in this connection); in Penn. phosphorea (and Renilla) the term- inal polyp is always single, and continues to be so. 2) The large zooid on the end of the rhachis might be interpreted as the terminal zooid, the terminal polyp itself being aborted. This hypothesis would require so early an abortion of the terminal polyp, that we know of no parallel in any other Pennatulid whatever. 3) The terminal polyp might be transformed into the large zooid, which ought, in this case, to be denoted as the «top-zooid». This interpretation would be met by a difficulty quite similar to the one in supposition ‘Nr. 2, and further by the fact that the original terminal zooid must have dis- appeared at the same time — that is to say, at a very early stage. This seems little probable, when it is remembered that v. Koch has in one case found the real terminal zooid to be preserved in a specimen of P. phosphorea, in which the terminal polyp seemed to have disappeared, a full grown specimen with 38 pairs of wings (comp. v. Koch, 2, p. 397, fig. V). Although, in spite of the difficulties indicated, the possibility is scarcely quite rejected that one of the two latter suppositions, or both, may apply to this species, I believe I have observations which show quite a different interpretation to be the correct one, and make the others superfluous. The fact is that amongst the many specimens from St. 36 no less than 18 individuals, and 3 individuals amongst those from St. 24, show that this species is capable of transverse fission. By this fission only the upper end of the rhachis with the uppermost pair of wings (I) is separated. The separation always take place close above the second uppermost pair of wings (II), and before it is quite accomplished the part of the rhachis situated between the two uppermost pairs of wings evidently stretches longitudinally; by this means, the part prepared for separation becomes similar to the the youngest known stages of development described above; compare figs. 19, 20 with the top of figs. 21 and 22. A clearing with glycerine or oil of cloves shows, however, that all the specimens in hand which are in the act of transverse fission, contain no calcareous axis in the top-part destined for separation; this part contains only the continuation of the longitudinal partition which farther down in the rhachis contains the calcareous axis. This axis itself terminates always immediately below the point of separation, generally with a hook turned to the right but this may also assume a somewhat different form (comp. fig. 22). This want of an interior calcareous axis is literally the only difference that can be pointed out between the top and one of the youngest independent colonies. The specimens showing the transverse fission belong to the large and middle-sized; the total length is between ca. 17 and 25™"; the number of wings is 4 pairs with 2 or 3 pairs of rudiments, and up to 7 pairs and 3 pairs of rudiments; each wing is generally provided with two individuals, of which Nr. 2 is almost always a zooid; yet the largest specimen, with 7 pairs of wings, shows three individuals in wings II and III: a large polyp, a quite small polyp and a zooid. Developed sexual organs are found in some of the specimens, but in several others, e. g. the one represented in fig. 21, 1 have not been able to find them. The process of transverse fission itself is seen in somewhat different stages; where the constriction of the rhachis is not deep, the top is least lengthened (comp. fig. 23); where the constriction PENNATULIDA. 23 is so deep, that the top is only connected with the other part of the colony by a quite thin string, the top (on the dead specimen) is hanging down like a «broken flower» along the stem of the colony, and then its rhachis is always prolonged stalk-fashion, most often bent into the shape of an S, so that the top gets a still more independent appearance (comp. fig. 21). Unfortunately, I have had no oppor- tunity of observing the separation itself, the further fate of the separated top, nor that of the remaining part of the colony. That I am not here guilty of a misinterpretation of chance mutilation is certain; to be sure, casual injuries are not wanting in many of the specimens in hand, but they have quite a different appearance; here the spicules on both sides of the constriction are placed quite regularly and in such a way as to be evidently arranged for the purpose; broken spicules are never seen as in injuries; moreover, the facts that the constriction always takes place at the same spot, that it is found in different stages and in a somewhat large number of specimens, and further that the top grows in length and emancipates itself, as it were, tend without a doubt to show that this species really is able to separate off the top. Of course it may be doubted whether this is a normal process; but I think it an obvious conclusion to take it to be normal, and to regard the phenomenon as a kind of propagation. The whole individualized character of the top, ready for separation, seems to me to indicate that this part will later live independently as a small colony, develop an interior calcareous axis‘), continue to grow and increase its number of individuals. The rest of the colony, the «mother- colony», will then probably form the large zooid at the place of constriction. One single specimen (fig. 24), without any constriction, seems to me of some interest in this connection; in that place of the rhachis where the constriction is seen in the other specimens, we have here a large, single and median zooid (Z') projecting in a polyp-like manner and interrupting the two rows of smaller dorsal zooids; it is provided with two lateral calyx-points in quite the same manner as the zooid which terminates the rhachis above between the two wings I (Z). The interpretation of this feature might be that the large top-zooid had been formed exceptionally before the constriction, this having not taken place or having been delayed for some reason or other. The specimen in question is otherwise of a good size (27™™), has 7 pairs of wings and 3 pairs of rudiments and contains developed sexual organs (even larvze); the calcareous axis reaches only to the height of the pair of wings III, that is to say, to a tolerably great distance from the spot where the large zooid is formed. My conclusion is accordingly, that all the mentioned features of the transverse fission observed, tend to show that this species may justly be called Pennatula prolifera. We should then have found a mode of propagation within the Octactinia somewhat reminding one of that known in the Poly- actinia, for instance in Goniactinia prolifera (M. Sars), and the Fungie; but the question, with regard to this sea-pen, is of a transverse fission of a colony. Several facts will be more intelligible viewed in this light. In the first place, the circumstance will now be readily understood why the species was taken in great numbers at both places where it was found, and that, in spite of this, it is only represented by colonies of a youthful character. It is obvious that the colonies need only a slight degree of development to be capable of transverse fission 1) It might, however, be possible that an interior calcareous axis might come up, before the separation took place; the contrary cannot be said to be proved by the specimens in hand. 24 PENNATULIDA. (as in Gontactinia, it is only young individuals that divide); the new colonies separated off by the division will soon get so far as also to divide; even the mother-colony itself will soon again be able to throw off the top part, and so on. Secondly, the fact will be explained that in the whole series of development found, no specimen, however small, is provided with terminal polyp and terminal zooid; the smallest colonies may possibly simply be tops of somewhat older colonies recently divided, which older colonies had before divided — perhaps more than once. Accordingly, we do not yet know the real primary polyp of this species, nor do we know the appearance of the species in its full-grown state. Whether the finished Pennatula-shape is ever reached must remain doubtful; I think, however, that the largest specimen in hand indicates that some individuals attain this form. This would not be necessary for the sake of the sexual generation, as a great number of the imperfect colonies have proved to be sexually ripe. Occurrence. The Davis Straits; St. 24, 63°6' N. Lat. 56° W. Long., (ca. 40 specimens), and St. 36, 61° so’ N. Lat. 56° 21’ W. Long. (upwards of roo specimens). The depths are 1199 fathoms and 1435 fathoms; accordingly, the species seems to be a marked deep-sea form; it has been found together with numerous other deep-sea forms; other Pennatulids found there, were specimens of Distichoptilum gracile, a series of young stages of Kopho- belemnon stelliferum, and a young stage of Uméellula lindahii. Fam. Virgularide KOll., emend. Virgularia Lam. Among the peculiar characters of this genus, I shall point out a few which have either not been noticed or only imperfectly: 1) The polyps are provided with a calyx, the edge of which may be simply circular or furnished with 8 more or less distinct small teeth without spicules; in the former case (for instance Virg. mirabilis), the edge of the calyx is not seen at all, when the polyps are most extended. I need only mention that the calyx here, as elsewhere, means that the hinder part of the polyp-body is rather stiff, so that the fore part with the tentacles may be retracted into it; thus the necessary stiffness is found here in spite of complete want of spicules. 2) The shaft (the rhachis) may be divided into three regions; (a) that of the developed wings, (b) that of the rudimentary wings, and (c) that of the stalk-zooids. (a) is the longest, upper part of the shaft; the polyps are fully formed, with tentacles etc, but generally contain no sexual products’); (b) may be rather long, and is on either side provided with close-set wings with undeveloped polyps which still want tentacles, but contain sexual organs and ripe sexual products; these wings become smaller below and gradually mere rudiments looking like low transverse ridges, and finally pass into (c); this last region is provided on either side with a single row of small zooids, which I shall call stalk-zooids. Close below these the peduncle or stalk begins; the boundary is generally indicated 1) An exception is found in Virg. affinis Kor. & Dan. (see KGlliker, Monogr. p. 198). PENNATULIDA. 25 by a slight constriction. Unless the examination is thorough, however, the peduncle seems to begin where the rudimentary wings cease. The stalk-zooids are never developed to polyps, but keep the zooid-form *). 3) New polyps are never seen in the act of formation at the ventral edge of the wings (as in Pennatula and many other genera); this holds good not only of the whole region (a), but also far down in (b); only in the smallest rudimentary wings are polyps seen to arise as elsewhere in the Pennatulids; that is to say in such a way, that the most dorsal polyp in each rudiment is the oldest, the most ventral one the youngest. One gets the notion indeed, that all the polyps belonging to a wing are begun contemporaneously?); certainly every wing, as soon as it has the character of a wing, whether its polyps are fully formed or not, is provided with the number of polyps that it has altogether. Nevertheless the polyp forming the dorsal edge of the wing continues always to be the largest, the one terminating the ventral edge the smallest, even if the difference is often very slight. 4) The calcareous axis has always a broken end above; it generally projects even with a more or less long naked end from the sarcosoma. This is a well-known and often mentioned fact; but it has been interpreted as due to injury, whilst I think it is caused by the mode of growth of the colony (for further particulars see under V. mzradziis). 5) All the species of the genus are provided with the regular radiate canals in the rhachis, pointed out by KOlliker. In all these features, and in several more, Halisceptrum agrees with Vzirgwlaria; it is quite certain therefore, that the genus 77 is to be referred to the same family as Virgularia; and it may be doubted whether it ought to be maintained as a separate genus distinct from the latter. Virgularia mirabilis (O. F. Miiller). Pennatula mirabilis O. F. Miller. Zool. Dan. Prodrom. 1776. 3074. 5S > > Zool. Danica. The Danish edition 1781 («Straa-Sofjeren»), p. 43; the Latin edition. 1788, p.11, Tab. XI. Virgularia mirabilis Lamarck. Anim. s. vertébres. 1816. T. II, p. 430. > > Kolliker. Monogr. 1872, p. 190. > Lyungmanni KOll. Monogr. p. 196. > multifora Kner. Verhdl. k.-k. zool.-bot. Ges. Wien. 1858, p. 295. 1) Kélliker has first seen these facts in Halisceptrum and in Virgularia; but in the description of what he calls «lateraler Zooidstreifen», he does not distinguish between the zooids that become partly the wing-polyps, partly the lateral zooids placed under the wings, and those that continue to be zooids and to be arranged in a long single series; these latter are the stalk-zooids. 2) In this way the fact has been interpreted by Marshall; but everybody may, by careful examination of the most rudimentary wings, see that this interpretation is not correct. M., however, was scarcely in possession of any specimen with this part of the rhachis fully preserved. Otherwise he seems to be the only author, who has especially observed the unvarying nature of the number of polyps in the wings. See: Report on the Oban Pennatulida, 1882, p. 63. This little treatise, which amongst many well-known things, contains several new interesting facts, is not found in our libraries, so that, when in 1898 I communicated my conception of the mode of growth and development in Virg. miraéilis to the Natural History Society here, I did not know that anyone had before observed the want of increase in the wings of Virgularia. The inter- pretation of M. of the mode of growth, however, is essentially different from mine (see later). The Ingolf-Expedition. V. 1. 26 PENNATULIDA. Of this species, I have had the opportunity to examine a very large number"), partly from Danish seas, partly from various Scandinavian regions and from Iceland (the Vestman Islands). I have been able to examine both well-developed and large specimens, to a length of 530™™2), and also young stages down to a size of 10™", i.e. smaller than those hitherto mentioned in the literature. In larger specimens I find the number of polyps in the developed wings very varying, from 12 to 4; specimens with three polyps in the wings or fewer are young stages which will be more particularly mentioned later on. K6lliker, it is true, in his diagnosis of the species, states the number of polyps to be 6—g, but in the more particular description, specimens occur with 10, with 11—12, and with 3; O. F. Miller gives the number as 8, Dalyell 8—10. The lateral zooids placed under each wing occur, in a corresponding manner, in varying numbers; where they are most numerous they are grouped in two transverse rows, in the young stages in a single row and in a number almost similar to that of the polyps. The peduncle is always shorter than the pen, but otherwise of very varying relative length, When K6lliker says: «Feder ungefaéhr 2'/,—3 mal langer als der Stiel», his state- ment is by no means always correct. The peduncle, here as in many other Pennatulids, may obviously be stretched very much in the living state of the colony, and in a few preserved specimens it may also be found much longer; in a specimen 350™™ long from the Kattegat I thus find it to be 250™" long (measured from the real boundary towards the rhachis at the cessation of the stalk-zooids). Of the numerous young stages examined, most, and especially all the youngest ones, belong to the museum in Stockholm. They form together a rather close series, elucidating, I ithink, the essential facts of development and growth in the Virgularia. There is still, however, a regrettable break between my youngest stage and the single primary polyp, the «, this supposition is no better founded than the one that Protopti/um KGll. is the young stage of Mrgularia(!). 32 PENNATULIDA. stage, 307" long, with wings with 3 polyps, a quite short pen with developed polyps, but a long rudi- ment-region; here also two naked axial ends project from the sarcosoma at the top, one quite short broken off, the other rather long (5—6™™) and evidently in process of decay, being plainly divided into a series of short, loosely connected pieces, like joints *). In a complete specimen, 423™" long (the peduncle 80™™), with wings with six polyps, from the Norwegian coast off Arendal, the wings of the left side are normal, whilst those of the right side, even quite down in the rudiment-region are placed in an inverted position, so that the calyx- openings and the tentacles of the polyps are turned towards the peduncle. It is well-known that the wings have a considerable power of motion, and I have often seen that they may also be turned in such a way as to make the polyps point downwards, but here the question is somewhat different; - in the present specimen, the group of lateral zooids belonging to the inverted wings is placed above the wing instead of below it, and by serial sections I have convinced myself that both zooids and polyps here turn the ventral side upward, pointing towards the top of the colony, whilst the zooids and polyps of the correctly situated wings of the left side are placed normally with the dorsal side turned upwards. The same abnormality is seen in two fragments of the upper part of the pen in a specimen with four polyps in the wings, from Bohuslan, Flatholmen, and in another fragment from the same locality, with three polyps in the wings, in which latter the abnormal side is disting- uishable as the right one, the end of the stalk being preserved. The last-mentioned specimens belong to the Museum in Stockholm. An abnormality, probably of a somewhat similar kind, is mentioned by Kolliker in a specimen of Halipteris christii (Mongr. p. 244); I have found no statement elsewhere of such features having been observed before. As to the synonymy of V. mirabilis, 1 have without any hesitation referred the V. Zyung- manni of Kélliker to this species. He has established his species on a fragment (in the Museum in Stockholm) from the Azores, and he himself expresses (Monogr. p. 197) a strong doubt as to its specific difference from V. mzradilis; after all, I think that its geographical occurrence, so far distant from what is otherwise known for mzradzlis, has been the special reason of its being established as a separate species. Later, the correctness of this species has hardly been carefully examined; it has, so far as I know, only been mentioned later by Studer as having been obtained in fragments by «l’Hirondelle» in the Bay of Gascony, without, however, any further information as to its structure, and by Whiteaves from the Gulf of St. Lawrence (Catal. Mar. Inv. East. Canada, 1901, p. 33). So far as I can discern from the careful description of this «species» by K6lliker, there is no sufficient reason for retaining it. The same can also be said for the V. multiflora Kner of the Adriatic, which K6lliker himself seems to be most inclined to regard as a local variety. Only the fact that the specimen in our Museum, in spite of its size, is provided with remarkably small polyps, could raise some doubt in my mind; the polyps, however, are just as small in a large fragment of V. mzradziis from the Vestman Islands (Th. Jénsson); but this specimen has evidently been somewhat dried. No doubt also the Virgularia sp. mentioned by P. Fischer, from the west coast of France (Loire-Inférieure, at Croizic), 1) A double calcareous axis in lérg. mirabilis has been observed before in a single specimen by Dalyell (l.c. p. 187) Pl. XLII, fig. 8. «The bones were unequal, a longer and a shorter, the longer extending so regularly, that the shorter might have been almost considered a fragment accidentally or supernaturally introduced along with it amidst the fleshy substance». PENNATULIDA. 33 is identical with V. mzradilis. It is said to resemble V. multiflora Kner, and to have its, «calicles aussi profondément séparés que ceux des V. mirabilis Miill.» (Bull. Soc. Zool. de France, vol. 14, pp. 36, 37); but a more particular description of it is wanting. With regard to the distribution, I have to add here a new locality of no small interest, viz. the Vestman Islands off Iceland. During the summer of 1900, about fifty large and small frag- ments of specimens with wings with 4—g9 polyps were taken here, also 14 young stages, most of them with 3 polyps in the wings (the «Diana», A.C. Johansen), and in the summer of r901 the Museum has received from Th. Jénsson, physician in the Vestman Islands, a large fragment (145™™ long) with (8—9) polyps in the wings. Hitherto V. miradilis was only mentioned from the coasts of Scandinavia, Den- mark and Great Britain. On the west coast of Norway it reaches as far north as Lofoten (Grieg, lc. p.11), on the British coasts it does not seem to have been found farther north than Scotland; in the Danish seas its southern boundary is given as the Great Belt and the Sound down to south of Hveen (Levinsen, Hauchs Togter, p. 399); for England, in the Channel (Falmouth and Eddystone, Marshall: Oban Penn. p.75, W. P. Marshall (sen.): Journ. Mar. Biol. Ass., vol. III. p. 335). The fact of this species having now been found at the Vestman Islands not only removes its northern boundary and considerably extends its western territory, but, when we consider that in recent years we have been able, just at the Vestman Islands, to point out several Atlantic animals known as belonging to the fauna, partly of the south-west of Europe, partly of the Azores, the probability that the « V. Zyung- manni> taken at the Azores and in the Bay of Gascony is identical with V. mzrabilis rises almost to a certainty. If my supposition that V. mwztiflora is also to be referred to mzrabilis, be correct, the Mediterranean will also belong to the territory of the species, the Adriatic at all events where V7. mzltz- Jlora has been taken in the Bay of Fiume (at the island of Veglia)'); also at Naples (Nisida) has a Virgularia been found (Kdélliker, Monogr. p. 369), but of its more particular determination nothing is as yet known; also at southern France, Cap Béarn (Lacaze Duthiers, Comptes rend. Acad. Sc. 1891, and Corallaires du Golfe du Lion, p. 359). On the American side of the Atlantic the species has not hitherto been found, according to existing statements; but according to what has been said above of V. Ljungmanni KOll. it no doubt occurs there also, since 15 specimens of V. «Lyungmannt» were taken already in 1872 at a depth of 200 fathoms in the Gulf of St. Lawrence north east of Cap des Rosiers (Whiteaves Le. p. 34). The depths at which V. miradilis has hitherto been found seem not to exceed 150—200 fathoms. At the Vestman Islands it has been taken at 98 fathoms (the «Diana» St. 41) and at 49 fathoms (the «Diana» St. 18). Virgularia cladiscus nom. nov. Cladiscus gracilis Kor.& Dan. Fauna litt. Norv. II], 1877, S. 101, Tab. IX, Fig. 13—15. > Lovent > Berg. Mus. Nye Alcyonider etc. 1883, S. 23, Tab. XI, Fig. 1—4. Virgularia tuberculata Marshall. Rep. Penn. H. M.S. «Triton» 1883, S. 129, Tab. XXXI, Fig. 1—3. Cladiscus Kollikert Kor.& Dan. Norske Nordhavs Expedition, Pennat., 1884, S.57, Tab. II, Fig. 8—13. Svava glacialis Kor.& Dan. Norske Nordh. Exp. 1884, S. 4,5, Tab. I, Fig. r—16. 1) From Trieste it is not known; at all events, it is not mentioned by Graeffe in Ubers. der Seethiere des Golfes v. Triest, IIL Coelenteraten. Arb. Zool. Inst. Wien, T. 5, 1884. The Ingolf-Expedition. V. 1. 5 34 PENNATULIDA. The calyx of the polyp distinct, with 8 small papillae or wart-shaped calyx-teeth (sometimes, however, they may be indistinct and seem to be wanting), often with 8 slight longitudinal ridges; the wings formed of groups of 3—4, up to 6 polyps, somewhat coalesced only at the base; more frequently one of the outermost polyps is quite, or almost quite, free; below each wing a series sometimes double, of 3—4 lateral zooids. The wings of the two sides are, in the younger part of the rhachis, placed almost opposite to each other, farther up they are distinctly alternating, with a distance of up to 2—3"™ between each set of wings. The dorsal radiate canals on each set of wings grouped more or less distinctly in a fan-shaped manner. Spicules are quite wanting (as in V. mirabilis). The colour of the living animal yellowish white. Of this beautiful Pennatulid the «Ingolf» has obtained four specimens at three different stations; a fifth specimen (a fragment) has been taken by the «Diana» at the Vestman Islands. Only in one specimen (Nr. 4) has the peduncle been preserved complete with its terminal bulb, in the three largest (Nr.. 1—3) the lower part of the peduncle is wanting, but the corresponding end of the calcareous axis with its fine, bent hook is preserved; as in other Virgularie, a denuded part of the axis projects above from the sarcosoma. Nr. I | oa | 3 4 5. Total‘ lengths 2s Oy a eae eee in mm. 94 85 71 32 40 The rudiment-region + the stalk-zooids .in mm, 25 20 20 10 » Number of polyps in each developed wing....... 4 3 4 3 3 Length “ota polyp 32457 iaccs ss cca eee in mm. 2 2—2,5 2 2 Number of lateral zooids under each wing....... 3—4 3 3-4 3 > In the upper part of the rhachis irregularities and variations are found with regard to the wings and their number of polyps, which is probably, at least to some extent, connected with a normal atrophy of the top part. At the top of the larger specimens, for instance, some of the upper wings are quite wanting on one side, so that only something like a «scar» is left; towards the top there may be wings with two polyps or even with one, or 3- to 4-polyp wings with one or two quite small, probably atrophied, polyps. As mentioned above, the polyps are provided with a distinct calyx, about 1™ long, with eight small soft calyx-teeth; these teeth, however, may be very indistinct or even imperceptible in some individuals of the same colony; the forepart of the polyp with extended ten- tacles is generally of the same length as, or a little longer than, the calyx; whether the fully developed polyps are able to retract this part completely into the calyx as in V. mirabilis, I must leave undecided; in the specimens before me all the fully developed polyps, at all events, are stretched out. As in other Virgulariz the rudiment-region contains the sexual organs, whilst the developed polyps are sterile. The longest colony I have been able to measure myself, is 105™™ long (a specimen taken north of Norway by the Barents-Expedition); Grieg (Norges Penn. p.20) gives a size of even 486™™ (a specimen from Fotlandsvaagen north of Bergen). As is seen from the synonyms this form has been described before, but as constituting a separate genus, or even two genera, and several species. The reason why I have given it a new specific name within the genus Virgudaria, and not retained its oldest specific name «graczlis» is that the PENNATULIDA. 35 name «Virgularia gracilis» has long ago been used by Gabb and Verrill for another, Californian, species’). I have therefore used as specific name, Koren and Danielssen’s oldest name for the genus which, I think, is quite descriptive in itself. Koren and Danielssen originally established the genus Cladzscus with the species C/. gracilis on a single, fragmentary specimen (Fauna litt. III. p.102); the genus was referred to the subfamily Funiculinee of the family of the Virgulariz in the (old) system of KOlliker; later, Kélliker (Chall. Rep. p- 35) placed this «genus» far from the Virgularie in his family Protocaulide*). This erroneous posi- tion — far from the Virgularie — is, I think, essentially due to the misinterpretation of certain features in the structure of the colony by Koren and Danielssen, which misinterpretation, among other things, led them to establish the genus Cladiscus. The fact is that in the original diagnosis of this genus, only the following essential features are mentioned viz. that «the cells are situated separately», and «the zooids ventral». (Lateral zooids are not mentioned). In the description of the single specimen of the species Ci. gracilis upon which the genus has been established, it is said of these zooids: «On the ventral surface ..... there appears a series of scattered zooids which, in the spaces between the groups of cells are more agglomerated, become larger, and form strong ventral protuberances of a very peculiar, as it were, crenulated appearance; the zooids, which are very much elongated, lying in the protuber- ances nearly in fan-like arrangement (fig. 15 a). In each protuberance there are from 20 to 25 zooids». Later (Bergens Mus. Nye Alcyonider etc. p. 23) with the help of a «new species» C/. Loveni the generic characters have been supplemented with a remark that the lower end of the axis is knob-shaped (which is not correct); on the other hand, nothing whatever is.said iu the altered diagnosis as to zooids. According to the description, the new species is not provided with the above mentioned «ventral zooids», but on the contrary with a single series of 3—4 lateral zooids below each wing. At the same time, the authors protest against KGlliker’s transferring the genus Cladiscus to the family Protocaulide, in which the calyx of the polyps is said to be wantings), justly drawing attention to the fact that the polyps are here provided with distinct «cells»; but when they then place their genus in the family Protoptiiide KOll., they are still farther in the wrong. Finally, the same authors again find in a third «species» Cl. Kéliikerd (Norwegian North-Atlantic Expedition p.57) the «ventral zooids», arranged upon the whole as in C/. gracilis, but bell-shaped and containing sexual organs(!) (comp. l.c. Pl. Il, figs. 8—13); further, two transverse rows (each of three individuals) of lateral zooids below the wings. From a close examination of the type-specimens of these three Cladiscus-species in the Bergen Museum (where I have also examined the later added specimens of Cl. Loveni, mentioned by Grieg (Lc. p. 20)) it was easy for me to see: 1) that the specimen described as Cv. gracilis has also lateral zooids, 2) that in CZ Loveni the same «ventral zooids» are found as in the two other «species», and 3) that the lateral zooids in all three species may occur in a single row, or some may be placed in such a way that the row seems double. As the number of polyps in the wings, three or four, which number varies with growth, cannot be used here as a distinguishing specific character there can be no doubt ') This species is hardly caracterized in such a way, that one would recognize it from the original description only. Comp. Kélliker, Monogr. p. 215. 2) This family (under the section Sfica/a, subsection /unciformes) consists in K6lliker of the genera Profocaulon KOIL, and C/adiscus Kor. Dan.; as both the genera must be abandoned, the family must also be dropped. Marshall & Fowler (Rep. Penn, «Porcupine» p. 462) had seen already that its place was unnatural. 3) In «Protocaulon», however, a calyx is found. See above p. 31. a 36 PENNATULIDA. whatever that their three species will have to be united into one. Further from an examination of the type-specimens of the genus Svava of Danielssen and Koren (North-Atlantic Exp. Penn. p. 45) — the species Sv. glacialis with the var. alba —I have convinced myself that this genus agrees in every respect with the «C/adiscus-species»'); here also «ventral-zooids» of the same kind and the same arrangement as in these species may easily be detected; they may always be seen — even under a magnifying glass — when the specimen is cleared in oil of cloves, or, what is often better, in bergamot. But if the same method is used with regard to any tolerably well-preserved specimen of Virgalaria mirabilis, the very same «ventral zooids» will also be found there, arranged in the same way, only in far greater numbers and in several layers. The fact is that these so-called «ventral zooids», which in some specimens of «Cladiscus» are seen so easily, in others only with difficulty unless in a clearing medium, are not at all zooids, but on the contrary the «radiate canals» described by KOlliker in Halisceptrum and Virgulartia, There can be no question therefore of maintaining genera like Cladiscus and Svava; they are one and the same species, and this species must belong to the old genus Virgularia; it is even very closely allied to the species miradilis; indeed, young specimens of this latter species with the same number of polyps in the wings can with difficulty be distinguished from Virg. cladiscus mihi; on clearing, the young V. miradilis also show radiate canals in smaller numbers and in a single layer; the polyps, however — whose degree of coalesence in V. miradilis is somewhat varying, — are never so slightly coalesced as in V. cladiscus, and distinct calyx-teeth, at all events, are not found. That the species V. cladiscus may be provided with more than four polyps in the wings is seen in a couple of specimens from the Dutch Barents-Expedition (1881) now before me, which have five polyps in the wings (whilst others specimens from the same expedition have four or three polyps in the wings), as also in a specimen from Finmark, Grélsund, and one from the Skager Rak (both belonging to the Stockholm Museum): and Grieg (lc p.11, under Svava glacials) men- tions a specimen from Trondhjem Fjord with up to six polyps. A young stage, with two polyps in the wings, two lateral zooids, 52™™ long, from the Skager Rak, is found in the Stockholm Museum. This species may also vary in the way in which the polyps of the separate wings are connected, and partly also in the degree of their coalescence (comp. Grieg l.c. p. 19). That I have also enumerated Virg. tuberculata Marshall among the synonyms will require no long explanation. From the description in Rep. Penn. «Triton» pp. 129—133 and from the figures, every one will easily recognize its identity; and moreover, this specific name has been withdrawn in the later work by Marshall and Fowler (Rep. Penn. «Porcupine», p. 464), on account of its identity with Cladiscus gracilis Kor. & Dan., and it is added that these authors find «the generic characters as given by the Norwegian authors too slight to justify a separation of the form from Virgularia>?) one must certainly allow that they are right here, although they pay no regard to the false «ventral zooids» whose nature was unknown to them. Strange to say, the same authors have retained Svava glacials, 1) The description by Danielssen and Koren of the genus Svava, agrees so well with the one given earlier of the genus C/adiscus, that it is astonishing they have not themselves seen the identity. And when it is stated for Svava that the generative organs are only developed in the lower, imperfect polyps, whilst the upper, perfect ones are sterile, it is also strange that their attention was not immediately drawn to the genus Virgu/aria, with regard to which Kélliker had long ago pointed out the same fact. 2) Marshall figures no lateral zooids in his V. ¢wéercu/ata; from his remarks on p. 132 it is seen, however, that he has noticed them, but does not venture to state them as certain. PENNATULIDA. 37 although they had a specimen before them; but they have referred it to the right place, in the family of the Virgulariz '). Distribution. The «Ingolf» has taken the species at the following localities: St. 6: 63° 43’ N. Lat, 14°34! W.Long., 90 fathoms, bottom temp. + 7° (Nr. 4); St. 59, 65° N. Lat, 11° 16’ W. Long., 310 fathoms, temp. + 0.1° (Nr. 2), both these stations near Iceland; St. 138, 63° 26’ N. Lat. 7° 56’ W. Long., 471 fathoms, temp. + 0.6° (Nr. 1, 3), near the Faeroes to the north. The «Diana» has taken Nr. 5 at the Vestman Islands at 68 fathoms. To judge from the hitherto known places, the species occurs in different fjords along the Scandinavian coast from Finmark through the Skager Rak to Bohuslan; in the deep channel of the Skager Rak it is evidently of frequent occurrence, to judge from the numerous specimens from this place in the Stockholm Museum (taken by the «Gunhild»); north of Norway it has been’taken (by the North-Atlantic Expedition and by «Barents») up to 75° 31’ N. Lat. (between Bear Island and Spitzbergen); north of Asia, west of Taimyr by the «Vega»-Expedition; by the English expeditions («Triton», «Porcupine») it has been taken north of the Hebrides and southwest of Ireland (51° 51’ N. L., 51° 51' W. L,), its most southerly occurrence hitherto known, while the Vestman Islands, about 20° W. Long., is the most westerly. It is remarkable that it may occur both in the warm and the cold area; its proper territory, however, seems to be within the warm area; within the cold area, it has only been taken at the two stations of the «Ingolf» enumerated above, and at four stations of the North-Atlantic Expedition (18, 31, 124 and 251); but all these stations are near the boundaries to the warm area, the Norwegian ones (according to the chart) even at the very border. At all events, it has not hitherto been taken in the deep and cold polar basin, properly so called. The bottom is commonly soft, clay or mud, sometimes sand (e. g. St. 6). The depth rather varying, from 40—555 fathoms, most frequently, however, over 100 fathoms. | This species is not known from the American side of the Atlantic. On the other hand, a «Cladiscuss-species is described from the Pacific, CZ Agassizii Studer (Note prélim. sur les Alcyon. «Albatross» Exp.; Bull. Mus. Comp. Zool. Vol. 25, Nr. 5, 1894, p.58) [under the family Protocaulide), of considerable size, 280", with three polyps in the wings («Albatross», St. 3424, 20° 15’ N. Lat. 106° 23' W. Long., depth 676 fathoms). That Virgularia-species of «Cladiscus»-form are also known from other places in the Pacific, is seen from the Vérg. bromled KOll. of the Challenger-Expedition, from Japan (565 fathoms, temp. + 3°.3 C, provided with spicules), and V. gracillima KOll. from New-Zealand (at small depths, 10 fathoms or thereabout). Stylatula Verrill. In Fauna litt. Norv. III, p. 92, Koren and Danielssen separated the genus Deidenza from the genus Stylatula for two Norwegian species, which at the same time they figured and described in detail, later a third species was added. These three species cannot be maintained; they are only one species, and I think the justification of the genus Dzidenza to be doubtful. To the best of my belief, Dzdenza is related to the other Stylatuda-forms (with S¢. gracilis Verr. as type) in quite the same manner as the «genus» Cladiscus is to the typical Virgudaria; here as there, the most conspicuous difference from the nearest 1) Danielssen and Koren had said nothing at all of the more particular relationships of Svava. 38 PENNATULIDA. allies is that the polyps are almost free and only few in each wing; further, the polyps in Dzibenia are said to want calyces; generally, they are only contracted — in very different degrees — and not with the circle of tentacles fully retracted; but that this circle may be completely retracted, is seen from Kor. & Dan.’s fig. 3, lc. Pl. X*), and it is also mentioned l.c. p.95; when the circle of tentacles is re- tracted the other part of the body of the polyp then forms a «calyx»; I find this feature to be quite similar in Stylatwla gracilis, in which, however, it is the rule that the individuals of the wing are retracted. Finally, the lateral zooids are arranged in a group above each wing with numerous indi- viduals, whilst in the other S¢y/atwla-forms they seem to be arranged in a single transverse row. Altogether, these features seem to be too insignificant to justify a separation from S*y/atula as an independent genus; but, as my material of typical forms of S?¢y/atuda has been limited to one specimen of SZ. gracilis, I think it better to leave the matter for future determination. Stylatula (Diibenia) elegans Dan. Kor. Virgularia elegans Dan. Videnskabsselsk. Forhdl, i Christiania 1859, S. 251. Stylatula elegans Richiardi. Monografia etc., 1869, S. 73. > » K6ll. Monogr., 1872, S.225, Tab. XVI, Fig. 137—38. Diibenia elegans Dan. Kor. Fauna litt. Norv. II, 1877, $.97, T. III, Fig. 1—7. > abysstcola » » © Ibid. $.94, Tab.X og XII, Fig. 1—3, » borealis» > N. Nordh. Exp. Penn., 1884, $.97, T. III, Fig. 1—7. > abysstcola Marshall. Rep. Triton Exp., 1883, T. XXIII, Fig. 17—21. From the Vestman-Islands we have two fragments (17 and 20™", probably of the same colony) of a «Dzbenia» with four polyps in the wings, above each of which is a group of zooids consisting of several rows; the large needles of the «calcareous plate» are 2.4™™ long, and at the thick end up to 0.240™™ broad; the smaller needles are 0.544—0.768"™ long, up to 0.048™" broad. The large needles of the calcareous plate seem to grow in thickness at the proximal end in such a way that the original long spicule is united with several of the smaller neighbouring spicules. The spicules along the aboral side of the tentacle-stem are of a length of 0.064—0.160™. The body of the polyps is without spicules, or only provided with a few. The polyps measure up to 3™™ (but the size is here, as is so often the case, very varying according to the degree of extension). I have stated above that I cannot acknowledge the three established species as separate, Neither from the descriptions, the figures, nor the specimens I have had the opportunity of examining, can I discern any other difference than in the number of polyps in the wings and the number of lateral zooids; and here where the number of polyps varies within the limits 2—6, this difference means no more than in Vrgularia, the mode of growth of the colony being the same as in that genus (even Koren and Danielssen have given up laying absolute weight on the number of polyps by making the form «smaragdina» a variety of adyssicola). Only in appearance are the specimens of 1) I find the feature to be the same in a great number of polyps in a specimen in our Museum; it is determined as D. abyssicola by Koren and Danielssen, and has been sent from Bergen Museum. PENNATULIDA. 39 «elegans» and of Koll. Monogr., 1872, S. 243, Tab. XVII, Fig. 144. Géndul mirabilis Kor.& Dan. Berg. Mus. Nye Alc. etc. 1883, S. 19, Tab. X. This Pennatulid is said to have been taken at Iceland and brought to the Paris Museum by Gaimard 1839 (KGll. Monogr. p. 243). According to its known distribution, its occurrence at Iceland was very probable, though up to the present we have had no other authority for it than the above. From the results of the «Ingolf», however, I am able to confirm the correctness of this occurrence. At one of the stations south of Iceland (St. 47) the « and the systematic group based upon it will have to disappear and be forgotten as soon as possible! Distribution. Pavonaria finmarchica has been found in the fjord-regions of Norway, from Finmark (Oxfjord) to the neighbourhood of Bergen; it seems also to extend farther south, into the Skager Rak, to Bohuslan (at Koster, Grieg, lc. p. 11); it has further been found at Iceland (Gaimard), and south of Iceland, St. 47 of the «Ingolf», 61° 32' N. Lat, 13° 40’ W. Long., 950 fathoms; next it has been found, frequently and in specimens more than one yard long‘), on the east coast of North America, especially on the fishing banks. It has been taken at depths from 60 to 980 fathoms. According to the localities where it has hitherto been found, its territory seems to be the whole northern part of the Atlantic, and there is every possible reason to expect that it will be found again in several places at Iceland (i.e. at the south coast and especially at the Vestman-Islands), and on the fishing banks round the Feroe Isles. From the Atlantic another species has been described: P. africana Studer (Ubers. Anth. Alc. «Gazelle». Berl. Monatsber. 1878, p. 672), 4 specimens of which were obtained on the west coast of Africa at 10° 12.9! N. Lat., 17° 25.5’ W. Long, depth 360 fathoms. The possibility is not excluded that we have here really the same species which has so wide a distribution farther north; the very low wings and the small number of polyps in these (4, or 5 to 6), specially pointed out by Studer, can scarcely — the number of polyps at all events cannot — be regarded as reliable specific differences from P. fin- marchica; at the top, the wings become higher and the polyp-calyx more distinct; the teeth of the polyp-calyx are not very distinct, but this is often the case in northern specimens also, and here also t) In the rich collection of P. fixzmarchica of the Christiania Museum is a specimen from Oxfjord, 5 feet long. 6* 44 PENNATULIDA. the wings may be quite low. Whether the want of an enlargement of the calcareous axis at the thickest part of the peduncle, is of any importance I must leave undecided, but I think it doubtful. The colour, to be sure, is not in exact agreement with the statements of Sars with regard to living P. finmarchica, but the difference is not so great as to make an identity of the species impossible. The genus Pavonaria also occurs, but as other species, in the Pacific. In our Museum we have representatives of two undescribed species from Japanese seas') and the Gulf of Korea. To these must be added a young stage from the coast of Korea (depth 80 fathoms); it is 305™™ long, but very slender. In spite of this considerable length it is still nearest to the A/icroptilwm-stage; the calyx-teeth are long as in the Ingolf specimens, and in many of the polyps also fused together so as to appear as one tooth. The polyps are arranged in two rows, one belonging to either side, larger (older) polyps and smaller (younger) alternating somewhat regularly, and where two polyps are placed opposite to each other their bases touch; one zooid or only a few between each two polyps on the same side, etc. Most probably it is the same species described at a still younger stage by K6l- liker as Microptilum willemoéstt (Rep. Chall. Penn. Pl. VII, fig. 27) taken south of Yedo, 34° 7' N. Lat. 138° E. Long., depth 565 fathoms. In the Riksmuseum of Stockholm is a Pavonaria (1075, "4/3 79) taken by the Vega-Expedition at Bering-Island (depth 65 fathoms); and Moroff (1. c. pp. 390, 393) describes a Pavonaria dofleint from Monterey in California, and a P. californica. Finally, we have several older notices of large Pennatulids from the American side of the Pacific (originally by Gray called « Osteocella», but he, to be sure, established the genus on the naked calcareous axes); they have been described by Stearns and Moss, and belong doubtless either to Pavonaria or Halipteris; the descriptions, however, are so imperfect, that it cannot be decided with certainty which of these closely allied genera are in question (comp. below under /alzpterts). Halipteris K6lliker. The genus Halipteris is very closely allied to Pavonaria K6ll. This may be seen, apart from the anatomical structure of the colony, its polyps and zooids (with regard to which I may refer to the monograph of Kolliker), especially in the polyp-calyx, the form of which is the same in both genera, viz. a truncate, oblique cone whose abaxial side is the longer and whose abaxial edge is provided with two calyx-points supported by spicules. The most essential difference between the two genera is only that in Halpteris, the oblique polyp-rows are not coalesced to real wings; the young stages therefore show so great a resemblance to one another as to be easily confused. To separate these two genera into two different families, as done by Ko6lliker, must be regarded as quite erroneous. 1) The remarkable «genus» LEchinopitlum described by Hudbrecht (Proc. Zool. Soc. Lond. 1885, p. 512, Pls. 30, 31) is, I think, one of the Japanese species, viz. a multilated fragment, from which, as is the case with Géndu/, the calcareous axis has been torn out. As I have not, however, had the opportunity of seeing Achinxoptilum itself, I can only advance this as a supposition, and leave it to a more particular test. PENNATULIDA. 45 Halipteris christii (Kor. & Dan). Pl. II, Figs. 30, 31, 32. Virgularia Christi Kor. Dan. Nyt Mag. for Naturvidensk. 5 Bd. 1848, S. 269; Faun. litt. Norv. II. 1856, S.g1, Tab. XII, Fig. 7—12. Norticina Christit Gray. Catal. Sea-Pens, 1870, S. 13. FHlalipterts > KGlliker. Monogr., S. 241, Fig. 146, 147. Lygomorpha Sarsii Kor. Dan. Faun. litt. Norv. III, 1877, S.99, Tab. IX, Fig. 7—12. Protoptilum tortum Grieg. Bergens Mus. Aarbog 1886, S. 13, Tab. VII, Fig. 19, 20, Tab. VIII, Fig. 1-3. Stichoptilum arcticum Grieg. Ibid. $.15, Tab. VIII, Fig. 4, 5, Tab. IX. From a fishing bank ca. 20 miles east of Fuglé, the Feeroe Isles, the Copenhagen Museum received through Agent Miiller in 1899 a fine and complete specimen, 1040" long, of which the peduncle makes only 80", and again in 1902 from the same donor an excellent, smaller specimen from the Feroe Isles, 400™ long (the peduncle 56™"). Young stages of this species differ so much from the grown form, that they have been described as independent genera and referred to quite a different family. On some of them Koren and Danielssen have formed the genus Lygomorpha, on others Grieg has formed the genus Sticho- ptilum, whilst some specimens have been referred by the same author as a new species to the genus Protoptilum K6ll.; all these «genera» were again gathered into the family Protoptilide of Kolliker. That I can now with perfect surety refer them to their right place as Halipteris christit, is due to the circumstance that I have been able to examine the respective type-specimens, and to compare them with one another, and with a large and good material of larger and full-grown specimens; upon the whole I have had before me a tolerably entire series from a length of ca.73™™ up to about four feet. The most important aid to recognizing the younger stage as Halipteris christii is the form of the calyx. The arrangement of the polyps will naturally vary; in the youngest specimens one must expect a more or less distinct arrangement in two rows, in the somewhat older ones we must expect to find new polyps on the inner side of the original rows (i.e. somewhat nearer to the ventral median line). In large specimens of Halipteris christii the polyp-calyx has a somewhat different appearance, according as the polyps are stretched out or more or less retracted, and to this is to be added that the two abaxial calyx-points may vary somewhat in size; generally they are short, one (the inner one) is often quite indistinct, which may also hold good with regard to the outer one; this however, will scarcely ever be found in all the polyps of one and the same specimen. When the polyps are much expanded the mouth of the calyx is distended, and then the two teeth may be widely separated and are easily overlooked; when, on the other hand, the polyps are quite retracted, the mouth of the calyx is drawn together; then the calyx is very similar to a grain of wheat, and in well-preserved specimens the calyx-points are always seen distinctly. They are supported by layers of spicules continuing down the calyx. The longest of these spicules I have found to be of fairly constant length — having measured them in several specimens, both large and small; on an average they are some 0.480™™ long, 0.032™" broad; the very longest I have found to be 0.496™™ long, 0.040™™ 46 PENNATULIDA. broad (with these measurements those of Kélliker: 0.4—0.7, agree very well); the spicules are often, as it were, divided longitudinally into, or composed of, several incomplete individuals; this holds good also with regard to the shorter forms, a8 also for the tentacles, Passing now to the hitherto wrongly understood young stages we shall begin with the smallest specimen, It is only 73 long (of which the peduncle makes ca, 15"), and belongs to the Bergen Museum, where it was found, together with two other larger specimens, in a glass labelled: «Protoptilum tortum Grieg; Vadeb, G,O, Sars», It is figured on PI II, fig, 30, three times the natural size; if this figure is compared with fig, 32, which represents the top of a fully formed Halipteris christ (viz, the above mentioned specimen from the Feeroe Isles of a length of 400") magnified to the same degree, the agreement in the form of the calyx will be so apparent that any further account — Ee will be unnecessary; it may be remarked, however, that the calyx-points are upon the whole somewhat more marked in the very young specimen, It is evident that Grieg has founded the deseription of his Proloptilum lortum upon the largest specimen in the same glass (143 long), and this is the specimen figured lc. Pl. VI, figs, 19—20, Pl, VIII, figs, 1-3; he does not even mention that he had two more specimens, In «Bidr, til de norske Aleyonarier» (1886) p.13 it is said of the calyx: «The edge of the cell is smooth; on the free side, however, is found a small tooth very richly provided with spicules»'), To this I shall only add that in this as in the other specimens I find two teeth, one of which is in most polyp-individuals some- what larger than the other, : The next smallest specimen I have seen is 86" long (the peduncle only 7™™) It deviates rom the former by the fact that the part of the rhachis where the polyps are developed, is much — longer, whilst on the other hand the lower part with the rudiments is extraordinarily short; the new, inner polyps are upon the whole also larger, ‘Together with a fragment 146% long’) it was labelled: «Lygomorpha Sars, Kristiansund, 80—100 fathoms, G, O. Sars coll.» Next come two specimens, respectively 106 and 145” long, the peduncle respectively 20 and ag", labelled: «Zygemorpha Sarsit, Lofoten, Risveer, G,O, Sars coll» These are the type-specimens of Koren and Danielssen of the genus and species stated on the labels) Here all the polyps are fairly well expanded, and the calyx consequently is distended; nevertheless, the calyx-points seem to be comparatively long and are considerably more conspicuous than is commonly the case in adult wpecinensa with expanded polyps; they seem also to be somewhat richer in spicules, but an examina- tion of the spicules shows the form and size characteristic of //adipferts, Nor have Koren and Danielssen quite overlooked the resemblance; in Fauna litt, III, p, 100, they say: «The genus Z. comes nearest to the //a/ipéerss, Kallikers, but in the same breath they mention a series of features which ‘) pao: «the free margin ia furnished with an extremely minute papillas, *) In thie specimen the lower part of the rhachis together with the peduncle is wanting; in the upper part of tne rlhachia the appearance ia already //aipferéy-like on account of the increase in the number of polyps, whilst the lower part reaemblea more that of the amall apecimen; on either aide of the rhachis two polyp-individuals are inverted (comp, Virgwd. miradilis above p. 42)} here, the abnormal position ia eapectally conspicuous, the long side of the calyx-cone and the two pointe of the calyx-edge being turned towarda the peduncle inatead of towards the top, as in the other individuals, 4) That my meaasureamenta are a ttle different from those in Fauna litt, Nory, p. too, is owing to the fact that Koren aud Danielaaen have paid no regard to the curves, and aa to the peduncle, have not seen its real limit; the quite small polyp-radimenta at the lower part of the rhachia have presumably been overlooked, In Aedip/er’y the peduncle in reality is alwaya very short, ; "Pa Te PENNATULIDA. 47 in their opinion, justifies the establishement of the new genus, which is placed between Hadipteris and Funiculina. From a cursory examination of the large specimens of Halipterts of a meter or more, the resemblance, to be sure, is not very conspicuous; in most polyps of large specimens the edge of the calyx seems often but little formed, as for instance is shown in the figure of Kdlliker (Monogr. Pl. XVII, 146—147). A closer examination, however, of any tolerably well-preserved specimen will always show a larger or smaller number of polyps, especially among the younger ones, for instance in the lower end of the rhachis, in which the two calyx-teeth so conspicuous in Lygomorpha are quite distinctly seen, and even rather long; and when once seen they may be traced in almost all the polyp-calyxes. In the figures of Koren and Danielssen of the adult Hadféeris in Fauna litt. Norveg. 2 part, Pl. XII (figs. 7—10) they are also given very distinctly, and in the text it is said that the calyces are of a «conical form and end upward in two points». Kolliker says (Monogr. p. 244): «Miindung ganzrandig mit einem kurzen Zahn und einer demselben entsprechenden schwachen Leiste an der unteren ventralen Seite, und swacher Andeutung eines ahnlichen Vorsprunges am gegeniiber- liegenden Rande». Of the two ventral teeth also, the one which the polyp turns toward the naked dorsal surface of the rhachis is generally the most developed. It is evidently this tooth that Grieg has remarked upon in his description of «Protoptilum tortum», whilst he has overlooked the other. All other features, in which Lygomorpha Sarsii might seem to deviate from Halpéerts christi, are only such as are a consequence of its being a young stage: the arrangement and smaller number of the polyps, the small size of the colony etc. Next we come to some (3) specimens of lengths from 234—310™™'); they were labelled «Stich- optilum arcticum» Grieg. Vads6. 30—40 fathoms. G. O. Sars». Grieg has described them in Bergens Mus. Aarb. 1886, p. 15 and p. 21, and given figures 1. c. Pl. VIII and IX; of the calyx he says p. 16: «The edge is provided with eight small, little prominent spines. The spicules are numerous in the cell, being especially found in large numbers in one side of the spines, whilst the other side is without spicules (Pl. IX, fig. 6). The spicules never project over the edge, they are mostly placed longitudin- ally, and are narrow, pointed, 0.210" long». I must suppose that the statement here should be «in the spines of one (the ventral) side»; and the fig. 6 quoted above shows the two ventral calyx-teeth which I find in the polyps of all these specimens; as in the grown specimens, one of the calyx-teeth is even the stronger one in most individuals. What has given Grieg the notion of the other six teeth is only slight undulations of the edge of the calyx; they may also often be seen in large Halpterts, and I take them to be the last traces of the equipment that must be said to be typical of the Penna- tulids. Also with regard to the arrangement and number of polyps, these «Stichoptilum», in accordance with their greater size, are nearer to the «//alipterts»-character than is «Lygomorpha». For the rest, Grieg has himself later observed the resemblance with the latter; in Ovs. Norg. Penn. (Berg. Mus. Aarb. 1891) p.22 he says: «When I enumerate these two species (viz. Protoptilum tortum and Stichop- tilum articum), it is not without some hesitation; they ought perhaps, to be enumerated as varieties of Lygomorpha Sarsit, which they resemble very closely». (The succeeding observations on the varia- 1) Between these and the preceeding, the above mentioned type-specimen of Protoptilum fortum, 143™™ long, and the «Lygomorpha»-fragment, 136™™ long, from Kristianssund, fit in with regard to development and size. 48 PENNATULIDA. tions of the calyx are partly incorrect; the calyx, when it is correctly understood, will always give good characters). Stages such as the last mentioned are immediately followed by others as for instance, the specimen 400" long from the Feeroe Isles. If we are first persuaded that the difference between «Lygomorpha» and Halipteris with regard to the teeth of the polyp-calyx, is only a difference in degree dependent on the growth, and that all other features of the polyps are the same, it will easily be seen that the other, apparently great, difference in the appearance of the colonies, the number and grouping of the polyps and the zooids, is due only to age: at the same time that the colony — as is commonly the case in the Pennatulids — grows in the lower part of the rhachis, at the boundary towards the peduncle, and adds here new polyps and zooids, a constant increase in the number of the polyps also takes place in Halipteris in the older, upper part of the rhachis; as is indicated in the young stages, new individuals grow up in the interspaces between the older ones. In this way arise the short and little regular, oblique rows peculiar to Halfpterts. The irregular arrangement seen in the younger stages is for the rest constant, solitary polyps or irregular little groups being always found between the more regular oblique rows; and apart from the lower part of the rhachis, the boundary line between the polyps of the two sides on the ventral side of the colony is, as in the young stages, only seen with some difficulty; in some places towards the top it seems to be quite wanting. The increase in the number of the zooids follows that of the polyps; Kélliker has carefully traced the mode of development of both these kinds of individuals in the lowermost part of the rhachis (Mongr. p. 245). Distribution. Hitherto, the species has been known from various Norwegian fjords, from the Varanger fjord (Hjzelm6) to the Jader; also, the young stages were found within the same territory; away from Norway it is only noted as having been found on the east coast of North England (Northumber- land, Tynemouth; Alder: Ann. Mag. Nat. Hist.; (3) vol. IX, p. 316). To these localities I may add the Dogger Bank in the North Sea’), and as a new locality also the Feeroe Isles, the fishing bank 20 miles east of Fuglé (agent Miiller). Hitherto it has only been found at depths to about 200 fathoms. From the American side of the Atlantic it is not hitherto known with full certainty. Verrill, to be sure, (Proc. U. S. Nat. Mus. vol. 2, 1879, p. 199, and later in Americ. Journ. Se. and Arts vol. 23, 1882, p. 309) mentions a Halipteris (christiz) as taken on the east coast of America on a fishing bank (Grand Bank), but nothing more definite has appeared concerning it later; if the genus be correctly determined, it is most probable that the species is also H. christ. It has probably a similar distribution to Pavonaria Jinmarchica and Pennatula grandis etc.?). We have various indications that the genus Halipteris is not restricted to the Atlantic. In «San Francisco Mining and Scientific Press» August 9 1873, and Proceed. Calif. Acad. of Sciences 1873, as also in Am. J.Sc. (3) vol.7, p.68, Stearns refers a Pennatulid (up to 66 inches = ca. 168™ long) to the Pavonaria of Cuvier, as subgenus Verrillia, the species V. Blakei. In a note added (in the last named place, p. 70) Verrill declares the form to be most closely allied to Halipteris christ (Kor. & Dan.), and accordingly Stearns has later (Americ. Naturalist vol. 16, 1882, pp. 55—56), called the t) Four specimens in the British Museum labelled: «Funiculina quadrangularis. Dogger Bank. 94. 6.24. I—4». 2) It is possible, however, that this Ha/ip/eris has later been tacitly withdrawn and placed by Verrill under Pavonaria Jinmarchica. PENNATULIDA. 49 form Halipteris blakei. The place of discovery was the Gulf of Georgia, Burrard’s Inlet (comp. also Proc. Calif. Acad. Sc. Vol. V, 1873, pl. I, pp. 7—12). From the description given I would not maintain as a certainty, however, that this species is not a Pavonaria KOll., and at all events nothing more particular can be said as to its specific difference from Halipteris christit. The description of the calcareous axis with its lower enlargement agrees with both genera. The specimen from the same locality, described and figured by Moss (Proc. Zool. Soc. Lond., 1873, p. 730), undeniably reminds one most of a Pavonaria; but neither the description nor the figure yields any reliable basis for a determin- ation. Naked calcareous axes of the same forms have been described by Gray (Catal. Sea-Pens, p. 40; Ann. Mag. Nat. Hist. (4), IX, p. 405, X, pp. 77, 406) as «Osteocella septentrionalis» from the west coast of America, and others similar as «Osteoc. cliftoni», possibly from West-Australia. Thus the genera Halipteris and Pavonaria are probably rather widely distributed in the Indo-Pacific Ocean. Fam. Funiculinide KOoll., emend. Funiculina Lam. Funiculina quadrangularis (Pall.). Pennatula quadrangularis Pallas). Elenchus Zoophyt. 1766, p. 372. Pavonaria antennina Cuvier. Régne animal. 1817, Vol. IV, p. 85. Funtiulina tetragona Lamarck. Anim. s. vert. Vol. II, 1816, P- 423. > quadrangularis KOll. Monogr. 1872, p. 256. Leptoptilum gracile K6ll. (Rep. Chall. Penn. 1880, p.27, Pl. VII, Fig. 28). > >» var. morvegicum Kor.& Dan. Bergens Mus. Nye Alcyonider. 1883, p. 29, Pl. XIII. Funiculina armata Verrill. Am. J. Se. (3) Vol. 17, 1879, p. 240 and Bull. Mus. Comp. Zool. Vol. XI, 1883, p.6, Pl. x, Fig. 4. South of Iceland the «Ingolf» has obtained a young specimen of a length of 225"; the length of the peduncle cannot be decided with certainty, as the soft parts of the lower part of the rhachis have been scraped off; otherwise the specimen is very well-preserved; the polyp-calyxes up to 3.5™™ long. In appearance it is strikingly similar to the Z7ichoptilum brunnewm of Kolliker (Rep. Chall. Penn. Pl. VIII, fig. 31), and agrees also with the Leptoptilum gracile var. norvegicum of Koren and Danielssen lc. Pl. XIII. The polyp-calyxes are richly provided with spicules, the arrangement of which quite agrees with fig. 3 of Pl. XIII of the last-mentioned work; again, the tentacles are also 1) Bohadsch is stated by KOlliker to be the author who has given the earliest description of the species in: De quisbusdam Animalibus Marinis, etc. Dresdz 1761. Here it is mentioned, chap. VI, p. ror under «Penna», as no. 3, P. rubescens (spinnis carens, tentaculis in corporis trunco positis») and p.112 as a species neglected by science, but known to fishermen who called it «Penna del pesce pavone». His specimen which was incomplete, was of a length of 2 feet 10 inches («multo vero longiorem eam fuisse non dubito»), and has probably been from the Mediterranean (Naples? from which locality he has several things). However, I have found this species mentioned as early as 1655 in Museum Wormianum p. 235: «Penne marine nomine ad me transmissum ex Norvegia corpus gracile, oblongum, cartilagineum, quadrangulum, interior substantia albicante, quodammodo extremitatem penne anserine plumis denudate referens, exterius membrana sublutea tectum. Tenuis licet sit hac penna, crassitie extremitatem penne anserine non superans, longitudine tamen eeqvat tres pedes Romanos. Nullam video habere affinitatem cum Penna marina a Gesnero, Rondeletio, Aldrovando & aliis descripta, ut quo referam vix habeam; hic interim locum obtineat, donee comodior detur». The Ingolf-Expedition. V. 1. 50 PENNATULIDA. provided with spicules along the aboral side of the stem. The polyps contain well-developed sexual products (eggs). The polyps are arranged in two alternating rows each corresponding to one edge of the quadrangular calcareous axis; between the developed polyps smaller ones are found, and between these again numerous small ones more or less in the zooid stage; in the very smallest, however, ten- tacles may be traced as small papille, and their calyxes are provided with 8 points; true zooids that remain as such, are not found. In all these features this specimen agrees with the «genus» Lef- toptilum of KOlliker. At station 18, south west of Iceland, about midway between Greenland and Ice- land, the », re 4 2,8 2,8 4 Greatest breadth of rhachis » 2 2 2 2,2 2 2 PENNATULIDA. 53 Nr. 1 is entire and shows no terminal bulb; Nr. 5 and 6 are fragments of the rhachis; in the latter, the upper end of the rhachis is undamaged; half of Nr.4 is a naked calcareous axis, and only in the lower part of the pen are some polyps preserved; Nr.7 is rather damaged, broken, and the uppermost 32™" only a naked calcareous axis. By clearing of Nr.1 and 6 in oil of cloves, quite whole, undigested shells of Zimacina are observed partly in the cesophagus, and partly in the gastric sac of several polyps; at the locality in question, Z7macina was especially numerous in the plankton. This fact, I suppose, indicates that these sea-pens, although bottom animals, get part of their food from the animal life of the surface, individuals from this part sinking to the bottom; in a similar way I have seen larve of lamellibranchs forming a prominent part of the food of Virguwlaria mirabilis from the Sound. Sexual organs were not seen on clearing; nor did an examination by serial sections show any such organs. Closer examination, on the other hand, has shown the following interesting features with regard to the inner structure. From the dorsal longitudinal canal of the rhachis, trans- verse canals issue at quite regular and somewhat short interspaces towards the periphery, where they open partly directly into the gastric sac of the dorsal zooids, partly — in the interspaces between the zooids — into a system of canals which is, otherwise, again connected with the zooids. These trans- verse canals evidently correspond to the «radiate canals» of the Virgularia (and also of other Penna- tulids); here they are characterized by being in the form of a long funnel, narrow at the dorsal main canal and widening at the peripherial end; their epithelium stains very intensely, so that they are seen with unusually sharp outlines. In each transverse section only one or two such dorsal radiate canals are generally seen. Into the ventral principal canal open radiate canals situated in the middle of the rhachis; they are much larger and wider than the dorsal ones, but run also towards the periphery; their epithelium also stains very intensely. These wide «collecting canals» connect the polyps and the other (ventral) zooids with the ventral main canal of the stem either directly, or indirectly through a network of canals with less intensely staining epithelium. The «collecting canals» are much less numerous, but much longer than the first-mentioned dorsal radiate canals, and their course is a much less horizontal one; they go upwards or downwards, and only rarely are more than two seen along-side each other. The small number of these «collecting canals», I suppose to be connected with the small number of the polyps; both features — to which may also be added that no sexual organs were observed — might favour the belief that Prot. carpenteri is only a young stage. The features given above of the « lofotense Dan. & Kor. N. North-Atlantic Exp. Penn. 1884, p.61, Pl. H, Figs. 14—20. > mohnt » » > > » > » »63, » Ill, » 1-7. > carinatum » > > > > > >» 265, » TI, » 8—11. > armatum > > > > > > » »68, » IV, » I-—7. Of a Protoptilum-species which I refer to P. thomsoni of KOlliker, I have had for examination three considerable fragments taken by the «Michael Sars» in the summer of 1902; these fragments I have been able to compare with a specimen from the Skager Rak belonging to the Riksmuseum in Stockholm, and with the material of the Bergen Museum amongst which were the type-specimens of the above named four «species» of the North-Atlantic Expedition. The description of K 6lliker is based on four considerable fragments of the rhachis; both the upper and the lower end with the whole peduncle were wanting. Only the abaxial edge of the polyp- calyx is free, and this edge is most frequently provided with four small points or teeth (to judge from the figures 221—222, Pl. XXIV, these teeth are as often wanting or slightly marked); the axial side passes into the coenenchyma, and downwards towards the peduncle the calyxes pass upon the whole so very gradually into the rhachis, that their length cannot be determined exactly; the projecting part itself, however, measures hardly more than 1.5—2™™; the calyx is richly provided with calcareous spicules 0.36—1.05™" long and 0.027—0.066™" broad. The zooids have an «incomplete» calyx supported by calcareous needles which form a small point. The polyps are provided with a row of spicuies on the aboral side of the stem of the tentacles (0.11—o.19™ long, 0.0o1—o.04™ broad). According to Kolliker the arrangement of the polyps is difficult to define precisely; it may evidently be described as two-rowed, i.e. two rows of polyps on either side of the median line of the colony (this line, however, is only to be determined with difficulty on the ventral side); but K. prefers to describe the arrangement as consisting of short oblique rows of three polyps on either side, being of opinion that each of the dorsal polyps forms a group with the two lower polyps placed more ventrally; these oblique rows would then have to be designated as ascending towards the dorsal side, whilst the oblique rows (or the individuals of the wings) in the Pennatulids generally ascend towards the ventral side. The zooids are found in large numbers, and everywhere on the rhachis between the polyp- calyxes, with the exception of a streak along the dorsal surface of the rhachis. The colour is pink, varying in intensity in the different specimens. In the Scandinavian specimens before me the calyx of the polyps is rather varying with regard to the degree to which it projects from the rhachis; this holds good with regard to different specimens as well as to different places in one specimen; but the axial side to the very mouth is always incorporated into the ccenenchyma. Thus the measurement of the length and breadth of the calyx may give very different results; it is especially difficult to determine the lower end of the calyx. The edge of the calyx is also varying, being often quite without teeth, often with one large and several smaller and indistinct; the most frequent number seems to be three, and these variations 56 ; PENNATULIDA. may be seen in the same colony, even in neighbouring polyps; the calyx-edge of the zooids is just as indistinctly furnished with projections; most frequently, it seems to me, without such. The tentacles are provided with a distinct streak of spicules; otherwise the retractile parts of the polyps are naked, whilst the calyx is furnished with long spicules, placed densely and numerously on the abaxial side. The size of the spicules agrees upon the whole fairly well with the measurements given by’ Kolliker. The arrangement of the polyps presents some variation, but the differences are evidently connected with the different age and development of the colonies; also, the arrangement shown by Kdlliker for the type-specimens of 'the species Pr. thomsoni is represented here. A common feature in all the specimens is, that along the dorsal median line of the pen’ runs a distinct, sometimes rather broad streak devoid of polyps and zooids (m, figs. 4, 5, 6); it is broadest in the lower part of the pen towards the peduncle; this streak is uncoloured (or yellowish white) like the peduncle. A similar streak, more or less distinct, also runs along the opposite, ventral surface (# figs. 6, 7, 8); it begins below at the peduncle where it is almost as broad as the dorsal one, but it very soon becomes quite narrow, often only seen as a very fine line, in some specimens even interrupted in several places by polyps placed quite medianly. Otherwise the whole rhachis is covered with polyps and zooids; the latter occur in very large numbers and fill all the interspaces between the polyps. On the lowermost part of the pen (figs. 6, 8) undeveloped polyps together with zooids form a long, wedge- shaped streak on either side, between the two naked streaks mentioned above. In the most developed specimens the polyps may be said to be arranged in three or four longitudinal rows on either side; but if we follow K6lliker, and take into consideration the oblique rows ascending towards the dorsal surface, each of these contains generally four, five (or in a single place six) polyps (comp. fig. 7); in reality, the arrangement on either side is nearest quincuncial. Less developed specimens show two longitudinal rows on either side with two or three polyps in each oblique rows; the least developed show something between one and two rows, i. e. they approach the arrangement that may be found in P. carpenterz; in these specimens there seem to be transverse groups of three polyps: one belongs to the right side, one to the left whilst the middle one is apparently placed in the ventral median line («Pr. lofotense>). In places where the ventral median line is only indicated (it may be very difficult or in some places quite impossible to see it) the real condition is that a polyp has been added alternately on the right and the left side inside the one belonging to the original first row (the dorsal polyps) nearer to, or, as it were, in the ventral median line; oblique rows cannot be said to be marked here as in the others; or, if they are present, they must be growing in the opposite direction and alternating in such a way, that opposite to an oblique row of two polyps on one side there is a solitary polyp on the other side. In the table below, of all the speci- mens which have hiherto been described and which I think must be referred to the same species, some of these features have been indicated and some measurements given. The statement of the «total length» means here only the length of the specimen in question, as no specimen is complete. This species probably grows to as greath a length as such forms as Halipferis, Pavonaria or Funtculina. rd by, fs 7 PENNATULIDA. 57 5: Il. I. 2. 4. Killiker’s 6. cP Nr. «P. lofo-| «P. 3. | «Pr. ar- typeof |« » 2 5 Length of the spicules of the RMPROR To tie a'ss 6 in mm. 0.560 0.445 0.142 0.36 0.222 0.992 0.192 | 0.592 —0.893 | — 0.625 —0.893 —I.05 —1.023 —0.992 | —0.960 Breadth » » » » » 0.020 0.027 0.016 0.027 0.041 0.032 0.032 | 0.032 —0.035 | —0.045 —0.046 —0,066 —0.055 —0.064 —0.056 | —0.048 Length of the spicules of the Remtacles 3... .5:.. in mm. 0.290 0.268 Were 1) O.1I. 0.136 “yee 0.118 Sivie 0.192 | 0.128 ; —o0.318 —0.19 —o.128 —0.192 i Breadth » » » » » 0.020 MOR Pes 1) 0.01 0.014 .... | 0.0080 | -.... | oo16 | 0016 4 —0.027 —0.04 —0,032 Length of the polyp-calyx » 3-5 2A 1 A—5 3 1.52 3-5 | 3—4 | 3—4 |4(2—5)| 2—4 | 2-4 Breadth » ee » Boe Gabe 2 I Rete ae sey 2 1.5—2| 1.8—2 1.8 Number of calyx-teeth........ 3 2—5 3 2—3 | (most often) 4 3 3(0—2) 3 3(0—2) | 3(0—5) | 3(0—4) Number of longitudinal series | POR fcr ios cp es wie gals 1(—2) 2 2 2 2 3 3 3 3 3 3(—4) Number of polyps in each ob- : UC oe ae 1(—2)2)| 2—3 22) 2-3 iz 3(—4) 3 3-4 4 4-5 5 The most complete specimen is Nr. 11 from the Skager Rak, in which the whole peduncle is preserved; the upper end, however, is broken off3). The four specimens Nr. 3, 7, 8, and ro are of especial interest because their upper end is intact; this shows that any supposition that the genus Protoptilwm was constructed like the Virgularie, is unwarranted; when the top is so often wanting, and the calcareous axis appears naked for a greater or less distance, this in Protoptilum is with tolerable certainty only due to mutilation, generally caused by the fishing apparatus. In the four mentioned specimens the thachis terminates above as a longer or shorter process (fig.4 /), bent towards the ventral side in a more or less hook-like way; this process is covered with zooids to the point; the point itself may be occupied by a zooid, which, however, does not appear to be larger or upon the whole built differently from the others. No stress can be put on the differences with regard to the size of the spicules; in the first place, I have by no means always measured exactly the largest and the smallest specimens, and secondly, the measurements of the specimens of the North-Atlantic Expedition and of the type- specimen of the species P. thomsoni, have been made by Danielssen and by KéOlliker, and these measurements I have not been able to control; besides, spicules are upon the whole often very varying both in length and thickness in different individuals of the same species of Pennatulids. As to the colour of polyps and zooids, and consequently of the rhachis as a whole, it is also t) Is said to be «as in the other species». 2) The oblique series are here most easily seen to be such, when viewed in a direction opposite to that in the other specimens, viz. from the dorsal polyp upwards towards the ventral median line. 3) This specimen was in the Riksmuseum in Stockholm labelled as «Pr. gunhilde», but the name has never been published (comp. Grieg. Ovs. Norges Penn. p. 21). The Ingolf-Expedition. V. r. 8 58 PENNATULIDA. rather variable, but is almost always different shades of red; only Nr. 6 «P*. carinatum» is stated by Danielssen to have been straw-coloured, the calyxes almost white. «For several specimens the colour of the calyxes of the polyps is stated to have been of a brick-colour; now, however, they are almost quite bleached or only pink; exceptions in this respect are formed by the Swedish specimen Nr. 11 and one of those taken by the «Michael Sars» (Nr. 10), both being still of an intense brick- red colour. By a careful comparison of all the Scandinavian specimens I have not been able to find any difference that might justify a division into several species. The four species established by Danielssen on the material of the North-Atlantic Expedition: Pr. lofotense, armatum, carinatum, and mohni, are each based on only one single specimen; in other words, each specimen found has been made a separate species, although the localities might have advised some caution; scarcely anybody will be able to find reliable specific characters in the (otherwise very careful) descriptions of Danielssen, no more than in his figures, and the specific diagnoses he himself gives, must be rejected by all, who have had any experience with regard to the Pennatulids. The fact is that the differences really found in these four specimens are individual ones, either such as are due to different development and growth (as the number and particular arrangement of polyps and zooids), or such as owe their origin to a castial degree of retraction (for instance, the features that have evidently given rise to the specific name carznatum; further, that the polyp-calyxes may be narrower, more or less projecting and marked off from the rhachis etc). That it is injudicious to seek characters for the division of species in such features is immediately seen, when new specimens are found. In this respect I may refer to the fact that the two specimens found by Dr. Appelléf (Nr.3 and 7 of the table), when judged in the same way as has been done with those of the North-Atlantic Expedition, would of necessity have to be established as two new species, neither of them agreeing exactly with any of the species of Danielssen, but being more or less closely related to each of them and being, besides, provided with something new. Grieg, to be sure, has seen this (at all events with regard to one specimen, comp. Norges Penn. p. 21), but he has also seen that it would be dangerous to create new species; the more so since the Swedish specimen (Nr. 11 of the table), which Grieg has also had the opportunity of seeing, might as well lay claim to the same title, as it cannot be said absolutely that it «seems to be quite similar to» the type-specimen of «Pr. mohni>; only with the specimens of Appelléf in mind could there be from Grieg’s standpoint any question of referring it to «Pr. mohni>. But when it is understood that «Pr. mohni> may assume so many different shapes, the necessary consequence will be the incorporation of the three other forms of Danielssen into the same species. I am therefore quite persuaded that every one, who can now inspect and compare all the Scandinavian specimens I have had at my disposal, will arrive at the same view with regard to their specific identity. Strange to say, no comparison with the species Pr. thomsoni seems hitherto to have been attempted; at all events this species is not mentioned at all by Danielssen or Grieg; nevertheless, the agreement between the Sandinavian forms and this one is in all essential features so close, that I think myself fully justi- fied in using the specific name of Kélliker. Also, the features of the inner structure mentioned by KGlliker (Monogr. p. 371), have been found by me in the Scandinavian specimens I have examined in this respect. Thus, I find the radiate canals, to which KdOlliker especially calls our attention as PENNATULIDA. 59 being found Breadth » » LS hae kon sae ig) Sic yale RS Sergei ie i eae a pri eae ara I.5 » Breadth of rhachis measured across the mouths of two opposite polyp-calyxes. . . aN t's Length of a polyp calyx, from the base to the point of a tooth................ 4—4.5 » » > > » ite » » notch between two teeth ....... «tea SIRENS. ro ha ea a a I—I.5 >» fumnrm meciniouth of the calyx....2... 000.60. eee eee eee 1.3 > Since, according to the description by Kélliker of Protoptilum smittit, there might be reason to suppose that the specimen of the «Ingolf» was perhaps the same species, I have, by the kindness of Professor Théel, examined the type specimen of Py. smittiz. This specimen is also at a little developed stage, mainly with one row of polyps on either side; but the top is broken off (it is upon the whole rather damaged); some agreement is present in the outer appearance, but the teeth of the polyp-calyxes are broader and vary more irregularly in size and number (3—5), the zooids are 62 PENNATULIDA. much more numerous, the retractile parts of the polyps very dark-coloured, the spicules more slender and fewer, especially on the peduncle. Although several of these differences are of such a nature, that in some other Pennatulid-genera they would not be regarded as specific differences, I dare not overlook them with regard to the genus Protoptilum, our knowledge of the variation and the change in growth of this genus being still very incomplete. Accordingly I take — for the present — Py. smittit to be a young stage of another species than my Pr. denticulatum. ‘They belong both of them to the Atlantic Ocean, and the great distance between the localities, respectively ca. 36° N. Lat. 14° W. Long. (the Josephine Bank), and ca. 58° N. Lat. 4o° W. Long., need not in itself — any more than the difference in depth (228—1695 fathoms) — prove any specific difference, as may be seen from other cases (e.g. Anthoptilum grandifiorum a. 0.). Distichoptilum Verrill. The genus has been established by Verrill on one specimen of the species mentioned below, and characterized in the following way: «Slender pennatulids, with an axis through the whole length. Polyps arranged alternately, in a simple row, on each side. Calicles bilobed, appressed. Zooids three to each polyp, one in front and one on each side of each cell. Spicula abundant in the calicles, rachis, and stalk; those in the stalk are small, oblong, triquetral, interwoven». In this diagnosis, however, the following alterations have to be made: the calyxes of the lateral polyps are provided with several (6) teeth on the abaxial edge; the axial edge is toothless, and the axial side of the calyx fuses with the rhachis. The number of zooids is two at each polyp; they are placed directly above the mouth of the calyx of the polyp, one on the dorsal side of the rhachis, and one on its ventral side. The peduncle increases evenly in thickness downwards. This genus is especially interesting in that it retains as a constant feature, an arrangement of the polyps which in other Pennatulids is only found as a transitory one in young stages, and which may really be more or less distinctly traced in all the Pennatulids of which we know sufficiently young stages; it is seen most distinctly in the «Protocaulon»-stage of the Virgulariz and in Protoptilum. This simple arrangement in Déséi- choptilum is not only a constant feature, but persists throughout with the utmost regularity, which gives to the whole colony an exceedingly emphatic character of simplicity. Distichoptilum gracile Verr. (PL I, Figs. 12, 13, 14). Distichoptilum gracile Verrill. Am. Journ. of Sc. 24, 1882, p. 362, Note. » > > Bull. Mus. Comp. Zool. Harvard Coll. Vol. XI, Nr. 1, 1883, p.8, Pl. I, Fig. 1. > Verrilla Studer. Ibid. Vol. 25, Nr. 5, 1894, p. 59. At two stations, very far from each other, one in the Davis Straits, the other south of Iceland, the «Ingolf» has obtained several specimens of a Dustichoptilum which, in spite of some PENNATULIDA. 63 differences, will have to be referred, I think, to the species D. gracile of Verrill. The features in which all the specimens differ from this species are the following: the polyp-calyxes are only here and there seen with a two-lobed edge; and when this is the case it has only arisen by a closing of the mouth of the calyx round the completely retracted polyp’); in other cases — where the contraction of the retractile parts is not so complete — the abaxial edge of the mouth of the polyp-calyx shows six pointed teeth supported by spicules. Next, the polyps are not placed so close to each other, as given in the description of Verrill and in the figure l.c. 1 a; generally, the front edge of a polyp-calyx does not reach farther upward than to the lower end of the calyx placed next above it, and consequently it covers no part of this next calyx; often, especially in the upper part of the rhachis, one polyp is far from reaching to the base of the next. With regard to this point, however, it is to be remem- bered that the sarcosoma of the rhachis in all Pennatulids may be shortened by a contraction of its longitudinal musculature, so that the distance between the polyps may be altered; as in Virgularia (mirabilis), Stylatula, etc, the appearance of Dzystichoptilum may also be influenced by this fact. Further, there are found at each polyp, not three but only two zooids (each with a small calyx provided with spicules); they are placed close to the (axial) upper edge of the calyx, one on the dorsal, one on the ventral side of the rhachis, and (very nearly) of the same relative height. When Verrill states that one more zooid is placed «in front of» i. e. above the calyx, such a zooid is absolutely wanting in all the specimens of the «Ingolf»; in this respect as in all others these specimens agree with Studer’s VD. verrilliz; in Verrill’s figure also, this third zooid is not seen, and I think its existence quite problematical. Finally, a more or less distinct longitudinal stripe or furrow is found both on the dorsal and on the ventral side of the rhachis; the appearance of this furrow may however, I think, be quite dependent on the degree to which the longitudinal canals are filled, and to the casual contrac- tion of the sarcosoma. The colour is stated by Verrill to be «bright orange red, due to the spicula; end of bulb yellowish»; in the specimens of the «Ingolf» the colour is only red in the upper, abaxial part of the polyp-calyxes, otherwise it is yellowish to yellowish-white; the red colour of the calyxes extends over two only of the lateral points, whilst the other teeth contain colourless spicules (a further reason for overlooking these teeth). In all the features stated, the D. verrilli¢ of Studer agrees with the Ingolf-specimens, except that the calyx is said by Studer also to be provided with two teeth; he declares himself that his «species» is very closely allied to that of Verrill — the only differences being the greater distance between the polyps, and the greater size of his specimens — but these features are of no value as specific characters. So far as I can see, both Studer’s specimens and ours will have to be referred to the species D. gracile first described. The caleareous axis is round, in some specimens a little angular or compressed in part of the rhachis. In the upper retractile part of the polyps, spicules seem to be wanting, but they are found in the tentacles on the aboral side of the stem, not, however, to the very point; pinnule are naked. The specimens taken by the «Ingolf» show the following proportions: 1) Verrill’s fig. 1a, which, by the way, is not very detailed, shows only quite retracted polyps. 64 PENNATULIDA. Nr. 2 2: 3. 4. 5- 6. 7: 8. 9. 10. Il. er ] PGA Wen Be eS Fst oak ee eee in mm. | 289 | 227 106 104 101 98 83 75 70 47 36 Length of, peduncle \... 0.45 S445 she's » 74 | 70 32 25 35 47 4o | 33 25 22 15 Greatest breadth of peduncle .......... > i) 1.3 I 1.3 I I rt |o8—1 I I I Breadth of rhachis measured across a polyp- | COLES es a oeh< Lnig ate Sieetes tae in mm. 1,5 | 1.7 | I—1.3 I te 1.8 a, 1.8 1.8 1.8 1.5 Length of polyp-calyx.. a: 52. ces ws Gates > 3 3 3 3 3 2.3 2 2.5 2.5 2 2.5 Breadth » > » across the mouth » I I TAR Tae I bo 4 Sa tees I a 0.8 0.8 | As a matter of comparison I may note that Verrill’s specimen is 456™ long, the breadth in the middle 2™™, the length of the peduncle 1oo™™, whilst Studer says that his «species» probably reaches to a length of upwards of a metre; the single entire specimen, however, measured only 470™", Of the specimens before me Nr.6, 7, and 10 are damaged, more or less of the upper part of the calcareous axis having been denuded; the others look as if they were entire, but a more thorough examination of the upper part of the rhachis makes it doubtful whether the original upper end of the colony is preserved. With exception of the specimen Nr. 9, which I take to be abnormal, and which will be more particularly mentioned hereafter, the rhachis ends in a solitary polyp which, moreover, on its axial (dorsal) side carries a solitary and rather conspicuous large zooid (see Pl. I, fig. 13 z). It would be natural therefore to suppose these to be the terminal (i.e. the original) polyp and a terminal zooid, such as we know to occur in young /emzmatula-colonies. But so far as I can see, only six calyx-teeth are found in this apparently terminal polyp, as in the other polyps, the lateral ones; now we might, with some probability, expect eight calyx-teeth and a fully developed calyx-edge in the real terminal polyp, but the calyx-edge does not appear to be developed on the side towards the apparently terminal zooid — which, however, is difficult to decide with perfect certainty, as the polyp in question is always strongly contracted, so that the calyx-teeth join each other. At all events, the terminal zooid is placed close to the calyx-teeth of the end-polyp, whilst in Pennatula, as is well known, it is placed below at the base of the calyx of the terminal polyp. In other words, it is placed exactly where the one zooid would be found, if the apparent end-polyp is a lateral polyp. I am most inclined to suppose therefore, that the «terminal polyp» is the upper- most lateral polyp present, and the «terminal zooid» one of the two zooids of this polyp together with an adjoining small part of the rhachis, so that the real upper termination of the colony is close to this zooid. But such a termination of the rhachis can scarcely be the original one; it is probable that in all these specimens an upper part of the colony has dropped off either as a consequence of earlier mutilation or perhaps through atrophy in a somewhat similar way as in the Virgulariz. That at all events, the original features are not found at the upper end of the longest specimen but one (Nr. 2), although this end looks as in the others, may be concluded with certainty from the fact that the calcareous axis ends immediately at the «terminal zooid», cut off quite abruptly, and is there of the same thickness as farther down. With regard to the others I have not been able — even by clearing (in oil of cloves and the like) — to get a clear notion of the upper end of the calcareous axis, but it seems to be tapering, and does not appear to be broken. On the other hand, the lower end may easily be observed; it is found in the outermost point of the peduncle, and forms here a curved hook. PENNATULIDA. 65 The specimen Nr. 9 deviates from all the others in having the lateral polyps in the uppermost part of the rhachis arranged in (almost) opposite pairs; three such pairs are present (PI. I, fig. 14); between the two polyps of the uppermost pair the rhachis ends as a little knob (k) containing the upper, fine end of the calcareous axis, and provided with one ventral zooid at all events for one of the two polyps. The spicules on the calyx have a length of 0.528—o.800™™ and a breadth of 0.032—0.048"™; on the tentacles, the spicules are 0.096—0.176™ long and 0.008™™ broad. Occurrence. The specimens of the grandifiorum Verrill. Am. J. Sc. (3), Vol. 23, 1882, pp. 312, 315; Bull. Mus. Comp. Zool. Vol. XI, p.5, PL I, Fig. 6. The «Ingolf: has obtained six specimens at one station in the Davis Straits; three of these are well-preserved, two rather large, the third small; two are more or less damaged, and one specimen is quite denuded (Nr. 6); two specimens (Nr. 7, 8) have been taken at another station, both of them in bad condition. Nr. I. a 3. 4. 5: 6. 7 8. Total length. ade F AE, Fon tein. ous Gate in mm.|) 415 300 205 172 122 1801)| 180 130 Leng of peduncle hs Spike pete oop Gots sia vee » 52 41 32 Rae 23 sels 31 20 Diameter of the upper enlargement of the peduncle... » 8 5-5 3 Pee 2 ee 4 a Breadth of the naked dorsal surface of the rhachis ... > c. 4 4 3 Feld I—2 oat ez 2 Number of polyps in the wings... 05 260. 170700. Eee ee 5—7 | 5-7 | 3-5 StU 2—3 ee 3-5 | 2-3 Length of an extended polyp with the tentacles. .... in mm. 16 A bate a 5 eps 5 5 As is generally the case in the species, spicules are only found in the end of the peduncle; in one of the specimens the largest measure to 0.024—0.030™", whilst others show a size of ca. 0.012 —o.020™"; the smallest seem to be even smaller than the minimum size of 0.007™" given by KoOlliker; a regular combination of four spicules is rather common, but more irregular combinations of three or four spicules are also found. The number of polyps increases during growth, not only in the lower, boundary part of the rhachis, but also farther up in the older part, where the oblique rows here representing the wings are often seen on the ventral side of the colony to be increased by a young polyp or a bud. The colour of the polyps when living is violet, that of the stem reddish; in spirit this sea-pen loses its colour only the tentacles of the polyps remain coloured, and there it is turned brown. In several of the specimens from both stations peculiar, worm-like parasites were found, partly projecting from the mouth of the polyps; they were parasitic Copepoda of the genus Lamifpe Bruz.; evidently a distinct new species, to which I give the name of ZL. anthoptili. Anthoptilum grandiforum has hitherto only been mentioned from the American side of the Atlantic; from North America it has been known from off Guadaloupe to off Nova Scotia (42° 46! N. Lat. 63° 45’ W. Long.) (Verrill, Agassiz); from South America from 37°17’ S. Lat. 53° 12’ W. Long., south of Buenos Ayres (the «Challenger»). To these localities two new ones are to be added, showing that the species ranges over a far wider geographical territory, as well to the north as to the east, in the Atlantic. The specimens of the «Ingolf» have been obtained in the Davis Straits, partly south-west of Godthaab, at St. 25, 63° 30’ N. Lat, 54° 25’ W. Long., 582 fathoms (soft mud, somewhat like potter’s 1) The greatest breadth of the naked calcareous axis on the compressed lateral surface 2™m™, on the narrow sur- face 1.5mm, PENNATULIDA. 67 clay), partly west of Sukkertoppen, at St. 28, 65° 14’ N. Lat, 55° 42’ W. L., 420 fathoms (soft, brown-gray mud with innumerable arenaceous shells of Rhizopoda); and from the Manchester Museum our Museum has obtained a gigantic specimen’) taken near the Cape of Good Hope ( ee 4 4 ‘6 4 3-5 URINE RUMEN ere 2 a ais ow ec ew cs en dene e whe » 8 | 8—9 6 4-5 4 5 Breadth of the naked dorsal streak................... » | 2 2 | CR | 1.5—2 1.5—2 t) It measures 1350mm, the peduncle 190mm, its enlargement about 30™™ in diameter. The polyps are coalesced into groups of 5—8 individuals, and measure 17—-25™m, American specimens grow to a height of 2 feet, and a diameter of ca. 25mm (Verrill, U.S. Fish. Comm. p. 510). 2) West of Tristan @’Acunha, at 35° 41' S. Lat., 20°55’ W. Long., depth 1500 fathoms, the «Challenger» obtained an Anthopt. simplex Kéll. (a damaged specimen of a length of 4oomm), which resembles 4. grandiflorum, but with only two polyps in each row; this specimen is possibly only an A. grandiflorum the development of which has been delayed, and in which the number of polyps for some reason or other has remained small; in other regions, however, the number of polyps may be greater at a far smaller size of the colony, which is proved by the specimens of the «Ingolf», especially the smallest of a length of only 122mm; therefore I think it rather hazardous to suppose 4. simplex and A. grandifiorum to be identical. 3) I am therefore able to declare with complete certainty that Roule’s «7vichoptilum» sp. (l.c. p. 307) has nothing to do with the genus Anthoptilum, but is really, as stated on p. 50, Funiculina quadrangularis. 9* 68 PENNATULIDA. St. 47. Nr I 2 3 4 5 Bboy 8 9. | 10. | a. | 12 Total Ten gt isos oes cakes Seales in mm. 285 255 | 250 | 245 | 245 | 245 | 240 | 240 | 235 | 230 | 230 | 230 Length ‘of peduncle 2205.4. 29 ees > 43° 5)" 351) S4* 37 PRayaegan ast Sgt ge" amy Saori eae Diam. of the upper enlargement of the peduncle — » 5 5 5 5 5 5 5 4 4 5 5 4° Length, of a: poly®.. 0.5-.645 5 ot ee ae eae > 6 8 8 8 8 | 11 / 10 |8—r12|8—g9 7 8 7 Breadth of the naked dorsal streak ........ » I.5—2 2 2 2 2 2 2 2 2 Nr. 13 I4. | 15. | 162) a7. | 28) aga eae. 21. 2255) 24. OUR BORN afc 3in ets wc St areralege ino ae in mm. || 225 | 225 | 220 | 220 | 220 | 220 | 220 | 210 | 295 ; 200 | 3195] 185 Lenoth of peduncle... 5 Ser Nees > cg ey pai & § 2 hi see 32 saga ae 31 29 24 28 Diam. of the upper enlargement of the peduncle» 5 5 |4—5 5 4 5 5 4 5 55 4 4 Rength ata DOD oon cathe nee roy week > +l ee 8] 12 6 g- | IO | <3 [AO 8 6 10 Breadth of the naked dorsal streak ........ > SDH tees (a PRRs Mba: xc 22 2 Car: eee oe Nr. 25. | 26. | 27. | 28. | 29. | 30. | 31. | 32. | 33. |.34.| 35. | 36. | 37% See eS ROtal Ten Ste ces. ceases ea ee eee in mm. | 185 | 175 | 175 | 175 | 175 | 175 | 155 | 145 | 125|108| to2 | 85 80 TLengthiof pedtncle: i) 5205, 302 LE. Sas > 28} 30] 28] 26] 27] 21} 25] 26] 2a1t| 20] 20] 17 18 Diam. of the upper enlargement of the peduncle» 4 | 4 x 4 4 4 4 4 ry rane Sei 2.5| 2 Length GF a poke p35 sicca tpn el pa eee » 6 | 7 | S| 6 2 INNA oe ot ene real Wet eel BE 6 Breadth of the naked dorsal streak ........ > nee Kris sae Anthoptilum murrayi has originally been taken by the «Challenger», south of Halifax (one specimen, 510™™ long), 48° 8’ N. Lat. 63° 39 W. Long., depth 1250 fathoms; later, it has proved to be of frequent occurrence off the east coast of North America, partly at the same localities as the preceding species, in depths from 640—1362 fathoms (Verrill, Am. J. Sc. (3), vol. 28, p. 220; U.S. Fish. Comm. Rep. 1883, p. 511). On the European side of the Atlantic, it was found almost at the same time at two rather widely separated localities, viz. by the «Caudan» in the Bay of Gascony (depth 1410 metres, 45° 57'N. Lat., 6° 21’ W. Long. (Paris), one specimen, 400" long), and by the «Ingolf» south of Iceland, at four stations situated on almost the same line west to east between Greenland and the Feeroe Isles, viz. St. 83, 62° 25' N. Lat., 28° 30’ W. Long., 912 fathoms; St. 40, 62° N. Lat. 21° 36’ W. Long., 845 fathoms; St. 65, 61° 33’ N. Lat., 19° W. Long, 1089 fathoms; and St. 47, 61° 32’ N. Lat., 13° 40’ W. Long., 950 fathoms. Thus it is evident that the species belongs to the deep part of the whole northern Atlantic inside the territory of the positive bottom temperatures. Fam. Kophobelemnonide KOll. Kophobelemnon Asbjérns. Kophobelemnon stelliferum (O. F. Miller). Pennatula stellifera O.¥F. Miller. Zool. Dan. Prodr. 1776, Nr. 3076; Zool. Dan. 1788, p. 44, Pl. XXXVI. Kophobelemnon Millert Asbjornsen. Fauna litt. Norvegize II, p. 81, Pl. X, Figs. 1—8. ie ase PENNATULIDA. é 69 Kophobelemnon stelliferum K6ll. Monogr. 1872, p. 304, Pl. XXI, Figs. 179—181. > Leuckartii KOll. Ibid. p. 306, Pl. XXI, Fig. 182. > Moebit Kor.& Dan. Berg. Mus. Nye Gorgon. etc. 1883, p. 25, Pl. XII. > abyssorum Kor.& Dan. N. North-Atlantic-Exp. Penn. 1884, p. 10, Pl. IV, Figs. 17—20. > scabrum Verrill. Bull. Mus. Comp. Zool. Vol. XI, 1883, p.7, Pl. I, Fig. 5. ? > tenue > Rep. Comm. Fish and Fisheries 1883, p. 510, Pl. III, Fig. 5. Gunneria borealis Dan. & Kor. N. North-Atlantic-Exp. 1884, p. 58, Pl. IV, Figs. 8—16. Of the genus Kopfhobelemnon several species have been established, some from the Atlantic (and the different branches of that ocean), others from the Pacific. Of these species as of so many other sea-pens, the same things holds good, that the material has been too scarce to allow their determ- ination to be reliable. Here, however, only the Atlantic species are to be mentioned in detail. As wili appear from the above list of synonyms I am of the opinion that they must all be united into one species, which then receives the old specific name steliiferum given it by O. F. Miller. To this *must further be added the genus and species Gunneria borealis of Danielssen and Koren (as to this see p.72). The specimen of O. F. Miiller was taken at Drébak, the numerous specimens of Asbjérnsen at different places in the Christiania Fjord. When Kélliker worked up the genus in his monograph, he justly referred all the Scandinavian specimens at his disposal to one species, not only those origi- nating from about the same localities as the specimens of Miiller and Asbjérnsen, but also a few from somewhat great depths in the Atlantic proper, north-west of Scotland (even if he thought them to be a variety «durum» on account of the size of the spicules; later examination of another specimen from another part of the Atlantic attached this variety still more closely to the type (l.c. p.370)). At the same time, however, he established a new species A. deuckartt? on some specimens from the Mediterranean (Nizza); to be sure he declares this form to be very closely allied to the former, « 28° so’ > » 788 y > 3°.5 » » 24: 63° 06’ » 56° oo’ » » 1199 » » 2°.4 » » 28: 65° 14’ » 55° 42’ > » 420 » > 3°53 The steamer «Thor» has further added as new localities: 63° 05’ N. Lat. 20° 7’ W. Long., 557 metres (St. 167 of the «Thor», south of Iceland, '/, 1903). Here 14 specimens were obtained, of which the largest is 105™™ long with 12 polyps, the smallest 50™™ with 7 polyps. Further: 62°57’ N. Lat. 19° 58’ W. Long., 957 metres; here three specimens were taken: two small young stages with solitary primary polyp, the one only 8™™ long and very small, with one large and two quite small zooids; the other 24™™ long, rather big (ca. 2™™ broad), with a polyp-bud just appearing on the right side and with many and large zooids; the third specimen 55™™ long, damaged, with several polyps and large zooids. Apart from the coast-regions of Norway where the Vest Fjord (ca. 67—68° N. Lat.) is given as its northernmost occurrence, the species has not hitherto been known farther north than 60° 18'N. Lat, in the Feeroe Channel (the «Triton» Exp. St. 8), whilst its most westerly occurrence on the European side of the Atlantic did not extend much farther than 10° W. Long., west of Brittany (the «Porcupine» Exp. 1870; see K6lliker, Monogr. p. 370). Now its territory is seen to extend, not only to Iceland but across the Denmark Straits and quite into the Davis Straits, west of Greenland; its occurrence there further corroborates my supposition that the Kofhoselemnon occurring farther south on the east coast of America belong to the same species. Formerly, it was known from many of the Norwegian fjords and the adjoining coast-regions, viz. from the Vest Fjord and farther south through the channel of the Skager Rak to the Christiania Fjord and to Bohuslin; also from many places in the Feroe Channel; off Brittany; from the coast of Les Landes: from different places in the Mediterranean (Cape Béarn, Nice The Ingolf-Expedition. V. 1. 10 74 PENNATULIDA. Naples: «K. deackartit>); finally on the American side of the Atlantic near George’s Bank (41° 29! N. Lat., 65° 47’ W. Long., «K. scabrum»), and at several other places, not more exactly designated. Its distribu- tion, then, probably comprises the whole northern part of the Atlantic (at all events north of ca. 40°) within the territory of the positive bottom-temperatures. It must be observed, however, that a few specimens have been taken by the «Porcupine» and the «Triton» outside the warm area, but at its very border, in the peculiar narrow strip of water which stretches from the cold depths of the North At- lantic into the Fzeroe Channel towards the Wyville Thomson ridge. The question is of two localities, very close to each other: 60° 14' N. Lat. 6° 17' W. Long., and 60° 18’ N. Lat., 6° 15’ W. Long., where the respec- tive bottom-temperatures are given as + 08° C. (the «Triton», St. 8, Marsh. p. 120) and + 1.1°C. (the «Porcupine» 1869, St.57; see «Depths of the Sea» Pl. IV and p. 143); only two specimens were taken at either place. That the occurrence within the cold area must be regarded as a mere exception and is only possible so close to its border, will appear from the fact that neither the North-Atlantic Expedition nor the «Ingolf», any more than the «Michael Sars» or the «Thor», have obtained any Kophobelemnon in the really great depths of the cold area of the North Atlantic, and also from the fact that K. s¢elliferum evidently reaches its highest development in such regions as the Norwegian fjords and especially the Skager Rak where the largest specimens hitherto found have been obtained. The bathymetrical range of the species is evidently very wide: in the Skager Rak it has been taken at depths from 26—400 fathoms; at Norway from 20—400 fathoms; in the Mediterra- nean at ca. 30 fathoms, as also at greater depths; in the Atlantic itself it has upon the whole been taken at considerable depths, between 420 and 1199 fathoms, but most frequently, it seems, as small or even very small specimens; at America, finally, in depths from 980 to upwards of 2000 fathoms (the latter depth is stated by A. Agassiz in: «Three Cruises of the «Blake»», Vol. 2, 1888, p. 142, and Verrill for his two «species» gives depths from 499 to 2369 fathoms; Rep. U. S. Fish Comm., 1883, p. 510). : I shall only add further that the genus Kophobelemnon is also distributed over the Pacific, and, according to the statements in the literature, is represented there by several species; but the number of these will probably also be reduced in future; of one of these species, K. affine Stud. its author already states that it is very closely allied to K. stelliferum («Albatross», Rep. Bull. Mus. Comp. Zool. Vol. 25, Nr. 5, p.57), and the same is said by K6lliker (Chall. Rep. p. 16) of a young stage from the seas north-east of New Zealand. Fam. Umbellulide Koll. Umbellula Cuv. This genus, which was down to. the year 1871 only known from Ellis’s and Mylius’s im- perfect descriptions of the dried specimens of U. encrinus brought home from the Polar Sea in 1753 by the Jutland skipper Adrians, has been proved to have a worldwide distribution in the great depths of the sea by the expeditions of recent times, and several species have now been established. The knowledge of these species, however, leaves much to be desired with regard to accuracy; of several PENNATULIDA. 75 — I think most — of them only young stages are known, and we know from UW. encrinus that the appearance of the species changes much during growth; no single investigator has been able to com- pare to any considerable extent the many species established on more or less scarce material etc. So much seems to be certain, however, that there is a number of species with well-developed spicula- tion, and others where the spiculation is quite reduced, only the peduncle containing quite micro- scopic spicules; others again are stated to be quite devoid of spicules, but, as will be shown with regard to the two species more particularly mentioned below, not all the statements in this respect are correct. The following species have been described as devoid of spicules: U. encrinus (L.), magniflora KG6ll., “ndahlit KOll, gracilis Marsh., geniculata Stud., and (I suppose) dazrdi Verr. Apart from U. genz- culata, with regard to which I can say nothing, I believe all these species to be closely allied; but as long as the knowledge of the characteristics of the species, and their variability and changes during growth is so uncertain, I think that some stress is to be laid on the geographical occurrence; thus, I believe we may take it for granted that those of the mentioned species which have been found in the Pacific, viz. U. magniflora and geniculata (and Studer’s « U. encrinus») are specifically different from the others which belong to the Atlantic and the Arctic Oceans; and further, as the whole group of Penna- tulids in its distribution within the northern parts of the ocean has proved to be remarkably depen- dent on the bottom-temperature, there is every reason to suppose that the form which has been found well- and even enormously developed within the «cold area», is specifically different from those found in the «warm area» of the Atlantic and the adjoining seas. On the other hand, I am inclined to regard these latter U. lindahlii, gracilis, and bairdii as one and the same species. Umbellula lindahlii K6ll. Tab. III, Figs. 37—46. Umbellula miniacea Lindahl. Kgl. Sv. Vet.-Akad. Handl. Bd. 13, Nr. 3, 1874, p.12, Pl. I, II, Figs.12—19. > pallida » ibid. p. 13, Pl. III. > Lindahl Kolliker. Festschr. Phys.-Med. Gesellsch. in Wiirzburg 1875, p. Io. > gracilis Marshall. Rep. «Triton»-Penn. 1883, p. 143, Pl. XXV. > Baird Verrill. Am. Jour. Sc. (3) Vol. 28, 1884, p. 219 (Note) and U.S. Fish. Comm. Rep. 1883, (1885), p. 509, Pl.I, Figs. r—2. Two specimens of the genus Uméellula, taken by the «