Natural History Museum Library
VOYAGE OF H.M.S. CHALLENGER.
ZOOLOGY-VOL XM.
7^
%
e.O. & ru rt^
REPORT
ON THE
SCIENTIFIC RESULTS
OF THE
VOYAGE OF H.M.S. CHALLENGER
DURING THE YEARS i87 3-7 6
UNDER THE COMMAND OF
Captain GEORGE S. NARES, R.N., F.R.S.
AND THE Late
Captain FRANK TOURLE THOMSON, R.N.
PREPARED UNDER THE SUPERINTENDENCE OF
THE LATE
Sir C. WYVILLE THOMSON, Knt., F.R.S., &c.
REGIUS PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF EDINBURGH DIRECTOR OF THE CIVILIAN SCIENTIFIC STAFF ON BOARD
AND NOW OF
JOHN MURRAY
ONE OF THE NATURALISTS OF THE EXPEDITION
Puf)lt£il)eli bp of i^er jHajestp’g ^obernment
PRINTED FOR HER MAJESTY’S STATIONERY OFFICE
AND SOLD BY
LONDON LONGMANS & CO.; JOHN MURRAY; MACMILLAN & CO.; SIMPKIN, MARSHALL, & CO. TRUBNER & CO. ; E. STANFORD ; J. D. POTTER ; and KEGAN PAUL, TRENCH, & CO. EDINBURGH ADAM & CHARLES BLACK and DOUGLAS & FOULIS.
DUBLIN A. THOM & CO. AND HODGES, FIGGIS, & CO.
1885
Price Sixty Shillings.
PRINTED BY NEILL AND COMPANY, EDINBURGH, FOR HER MAJESTY’S STATIONERY OFFICE.
The Editor of the Challenger Reports will be greatly obliged to Authors sending him copies of separate papers, or references to works, in which the Challenger discoveries are referred to, or the observations of the Expedition are discussed.
This will greatly facilitate the compilation of a complete Biblio- graphy, and the discussion of the results of the Expedition, in the final Volume of the Series.
Letters and Papers should be addressed —
JOHN MURRA F,
Chau.enger Office,
J2 Queen Street,
EDINBURGH,
EDITOKIAL NOTE.
The Report on the Annelida Polych^ta, by Professor William C. MTntosh, F.R.S., &c., occupies the whole of the present volume, and forms Part XXXIV. of the Zoological Series of Reports on the Scientific Results of the Expedition.
The preparation of this valuable Report has occupied Professor MTntosh over seven years, and it will be welcomed by all naturalists as a most laborious and painstaking contribution to Science.
The Manuscript was received by me in fifteen batches, at various dates, between the 8th August 1884 and the 20th July 1885.
John Murray.
Challenger Office, 32 Queen Street, Edinburgh, \st SepUmher 1885.
CORRIGENDA ET ADDENDA.
vii, line 12, for “ Edward” read “ Edouard.”
21, “PI. IlA. figs. 8-12” should he “PI. IIa. figs. 9-11.”
27, “PL III. fig. 3” should he “PI. III. fig. 2.”
29, “PI. IIIa. figs. 10-12” shoidd he “PI. IIIa. figs. 10, 11.”
34, Aphrodita australis, for “PI. VII.” read “PI. VI.”
39, Loitmoniee producta, insert reference to “PL VI. figs. 1, 2.”
67, Lepidonotus eristatus, “ PL XI. fig. 2 ” {7iot “ fig. 3 ”), and “ PL XVII. fig. 1 ” {not “ PI. XVIII.”) 71, after Eunoa opalma” insert “PL VIIIa. figs. 9-11.”
77, for “ PL XXXII. fig. 7 ” read “ PL XXXIIa. fig. 7.”
Ill, for Polynoe platyeirnts” read Polynoe platycirrataf 135, for “ PL XXIIa.” read “ PL XIIIa.” '
170, line 9 from foot, for “ Genetyllis lutea'" read “ Genehjllis oculata.”
172, line 7 from foot, for “ Genetyllis lutea,” read “ Genetyllis oculata.” '
223, line 10 from foot) transpose and “long.”
240, in explanation of woodcut, fig. 4 “ capensis” should be pettigrewi.”
244, line 4, for Lumhriconereis capensis” read “ Notoeirrus capensis.”
299, line 6 from bottom, after “PL XXXVIII. figs. 6-8,” add “fig. 19.”
337, line 14 from bottom, after Hyalinceeia tuhicola, &c.,” add “PL XL. fig. 2.”
343, for “ Glycera tesselata ” read “ Glycera tessellata.”
343, Glycera capitata, also from Station II.; lat. 38° 10' X., long. 9° 14' W.; depth, 470 fathoms; sea- bottom, green mud.
346, line 2, for “ hremewris ” read “ hrevicirrus.”
359, line 14, /or “Station 141, &c.,” read “Station 145a, December 27, 1873; lat. 46° 41' S., long.
38° 10' E. ; depth, 310 fathoms; sea-bottom, volcanic sand.”
385, insert “ Ghtetozone, Malmgren.”
394, line 16, for antaretia” read antarctica.”
394, line 17, “west” should he “east.”
480, line 4 from foot, add “also from Christmas Harbour, Kerguelen.”
481, line 12 from foot, /or “above”’ read “below.”
481, lines 5 and 6, for “ above the ventral edge ” read “ below the dorsal arch.”
THE
VOYAGE OF H.M.S. CHALLENGEK.
ZOOLOGY.
KEPORT on. the Annelida Polych^ta collected by H.M.S. Challenger during the years 1873-76. By William C. M‘Intosh, M.D., L.R.C.S.E., LL.D., F.R.SS. L. & E., F.L.S., C.M.Z.S., Soc. Psychol. Par. Soc. Honor., &c.; Professor of Natural History in the University of St. Andrews.
INTRODUCTION.
During the organisation of the Challenger Expedition the subject of the Annelids had several times been brought under my notice by the late Dr. Owyn Jeffreys, who more than any other marine zoologist in our country had kept the department in view during various dredging expeditions in the neighbouring seas. Though unable, by reason of strict and responsible official engagements, to lend any active service to the Expedition, I had explained as fully as possible the best mode of preserving these somewhat delicate animals (Annelids) both to Professor Wyville Thomson and Mr. Murray. Towards the middle of 1877 I received intimation from Sir Wyville Thomson about the examination of this part of the collection, which he described as somewhat limited in extent. A review of the specimens, however, on their arrival proved that instead of being limited, the series was an extensive one, and reflected much credit on the scientific staff of the Expedition. This was probably due 'to the interest taken in the group by the late Dr. Rudolf von Willemoes-Sulmi, a young naturalist of great ability, whose previous acquaintance with the Annelida had been considerably extended by a trip to the Fseroes just before the equipment of the Challenger. He was chiefly occupied with the Crustacea, it is true, during the voyage, though a few notes and sketches he made on the Annelids will be specially referred to. The loss of this
(ZOOL. CHALL. EXP. PART XXXIV. 1885.) LI U
11
THE VOYAGE OF H.M.S. CHALLENGER.
accomplished and energetic young naturalist ^ was a serious blow to the zoological stafl' of the Expedition.
Mr. Murray forwarded eight pages of sketches and notes made by Dr. v. Willemoes- Suhm. The first refers to a Nematoid parasite in a prawn procured on February 24, 1874 (Antarctic Ocean), in 150 fathoms, apparently very similar to one found in a prawn in Shetland many years ago, and forwarded to Dr. Cobbold. The second is an Eteone (probably new) from the Spanish coast. Station L, lat. 41° 58' N., long. 9° 42' W., in 1125 fathoms. The third is termed by Dr. v. Willemoes-Suhm a Syllidean, but it also approaches the Alciopidse in the structure of the feet. The fourth sketch enables me to supplement the description of Dalhousia atlantica (p. 186), and to correct the remark that there is no median tentacle. Dr. v. Willemoes-Suhm did not attach any name to this form, but there can be no doubt it is the same specimen. In the fresh animal the elevated areas of the head do not seem to be so prominent, and Dr. v. Willemoes-Suhm figures the crescentic pigment-mass on each side as the posterior pair of eyes, while the smaller pair behind the large anterior eyes are not shown. In his view of the foot a long slender ventral cirrus is indicated, this being absent in the preparation. His figure of a ventral bristle is quite recognisable, and he adds a wing or guard in the fresh specimen at the tip. Moreover, the drawing of the pharyngeal region and stomach is characteristic, no armature, as already stated, being present. The tail terminates somewhat bluntly, has two median slender cirri, and two lateral jointed processes, thus agreeing with the arrangement at the anterior end. It is satisfactory that the further information derived from the sketches of the lamented naturalist does not alter in any material respect the position assigned to this novel form.
The fifth sheet represents a pen and ink sketch of the peculiar Trophonia ivyvillei (p. 366) of the natural size, and one of its bristles, under the term “ Annelide aus der Ophelien-Familie,” while it is further stated “ Fuss-stummeln und Kiemen fehlen ganzlich.” It is not remarkable that this form should have given rise to ambiguity. The sixth series of sketches are connected with Mynochele from Station 20 (p. 410). Dr. V. Willemoes-Suhm termed the form Myriochele abyssorum, “ n. sp.,” but as his figure of the hook is not minutely accurate enough for satisfactory comparison, and the figure of the body of the animal shows no new feature, the decision that it is Myriochele lieeri, Malmgren, is perhaps at present prudent.
The seventh plate represents a Protula from “ 600 fathoms,” Cape St. Vincent, Portugal. It is uncertain whether this be Protula lusitanica, from Station II., 470 fathoms, or another form. All Dr. Willemoes-Suhm states is that -it is a “ Serpuloid.” The eighth and last sheet gives an imperfect sketch of Hydroplianes, procured on August 20, 1873, off St. Paul’s Eocks. From the partial outline with pen and ink this seems
^ For an interesting memorial of E. von Willemoes-Suhm, Ph.D., vide Challenger Briefe an C. Th. E. von Siehold ; Nach dem Tode des Verfassers herausgegehen von seiner Mutter. Leipzig, 1877, and also the Narr. Chall. Exp., vol. i. p. 769, 1885.
EEPORT THE ANNELIDA.
iii
to be a remarkable form with some superficial resemblances to the Polynoidse, though it may be a larval animal belonging to a different tyj^e. The absence of minute descrip- tions, with the exception of Trophonia loyvillei, and still more of specimens, prevents anything further being advanced about the foregoing sketches.
Numerous specimens of Sternaspis occurred in the collection, but, believing with Selenka, Vejdovsky, and others, that this group lies between the Chsetopoda and Gephyrea, they were at once forwarded with a few other forms to Prof. E. Pay Lankester, who at that time intended to work up the Gephyreans.^ It was my intention to describe the Nemerteans, and, indeed, sections of the majority had been made, and an outline of the group and its literature prepared. So much work, however, had fallen to my lot since my return to St. Andrews, especially in connection with the fisheries, that with Mr. Murray’s sanction I had great pleasure in handing over the Nemerteans to my friend Prof. Hubrecht of Utrecht, in whose skilled hands the interests of science will be more than safe. I confidently look forward to the publication of the recent important researches of Ur. Hubrecht, based on the Nemerteans of this Expedition.
A few Crustacean parasites occurred on the Annelids, and it has been deemed proper to describe them along with their hosts rather than separate them by giving them over to another worker.
In order to gain a correct view of the position occupied by the group to which this Report is devoted, I have made a few notes on some of the previous voyages. These must not by any means be regarded as complete or exhaustive, but simply represent a few broad touches to aid in bringing out the relations of the series of Annelids collected by the Challenger to previous efforts in this department.
The earlier voyagers seldom included the Annelids in their collections, though it is true a ship captain brought some specimens to Pallas, and gave that author an oppor- tunity of describing certain new forms; while a few others, for iustance Adler, mention them in connection with phosphorescence. In some of these voyages the invertebrates, however, formed a prominent feature, e.g., in Phipp’s Voyage to the North Pole in H.M.S. “Racehorse.”^ Moreover, in this early expedition it is evident considerable care had been taken to secure specimens, and the use of the trawl on the northern shores of Spitzbergen is a feature of considerable interest. Three Annelids are mentioned as having been thus procured, viz., Serpula spirorbis, Serpula triquetra, and Sabella frustiilosa, the latter characterised by “ Testa solitaria libera simplici curvata; fragmentis conchaceis sabulosisque.” The attention given to zoology in this expedition is note- worthy, and in contrast, for instance, with what was done in M. Soniierat’s Voyage a la Nouvelle Guinee,® which was published shortly afterwards.
1 Vide Keport on the Gephyrea, collected by H.M.S. Challenger, by Prof. Emil Selenka of Erlangen. Zool. Chalk Exp., part xxxvi., 1885.
^ London, 1774, 4to.
^ Paris, 1776.
IV
THE VOYAGE OF H.M.S. CHALLENGEE.
The voyages in the latter part of the eighteenth century and the begioning of the nineteenth showed little improvement in this respect. Thus the cruise round the world in the ships “ King George ” and “ Queen Charlotte ” ^ gave no addition to our knowledge of this and some other invertebrate groups, though crabs, shells, and birds are mentioned and figured, and the same may be said of the French Voyage de la Perouse autour du Monde.^ Similar remarks apply to the trip to Cuba and St. Domingo by M. E. Descourtily,^ and to Baron Albert von Sack’s Voyage to Surinam,^ The cruise of H.M.S. “ Investigator ” to Australia and other parts was even less productive in this department.
Captain Tuckey’s voyage to the Zaire (usually called Congo) contains a note® by J. Cranch that a new species of Nereis was taken on a bit of floating wood, lat. 0° 21' 0" N., long. 5° 49' 37" E., together with a genus not known to him. A single species {Nereis heteropoda) also is given by Chamisso and Eysenhardt in their Voyage Round the World. ^ There can be little doubt that during Sir John Ross’ two Arctic voyages (1818 and 1829) Annelids of considerable interest must have been obtained, indeed, he mentions in his first voyage that “worms’’ were procured in the mud at the depth of 1000 fathoms. Unfortunately the collections in each case have disappeared.
In the Narrative of a Survey of the Intertropical and Western Coasts of Australia, by Captain King, the versatile talents of Dr. J, E. Gray added an Annelid to the list, viz., Leodice gigantea, Savigny,* which measured nearly five feet in length.
The presence of so acute an officer as Captain Ed. Sabine in Parry’s first voyage ® to Greenland, accounts for the mention of two species of Annelids from the “Fauna Groenlandica,’’ viz.. Poly me cirrata and Polynoe scabra. The notices of Annelids, however, at this time by navigators are brief and fragmentary, a single species, jDerhaps, only coming under observation, as for instance in Eschscholtz’s voyage from Cronstadt to St. Peter and St. Paul, in which Tomopteris oniscifoj'inis is mentioned. Even in more ambitious voyages they made a small appearance, as for example in Freycinet’s Voyage autour du Monde. In other expeditions certain groups of Vermes become prominent, as in Riippel’s Atlas zu der Reise im nbrdlichen Africa, where the Planarians and Gephyreans {Sipunctdi) are specially noticed by Leu chart. The Nemerteans, again, occupy a plate in the Voyage de 1’ Astrolabe, and reference is made to the elegant forms and rich coloration of such Annelids as Amphitrite, Serpida, Nereis, and TerehellaP
In the Voyages en Scandinavie et en Laponie,^^ considerable attention is given to
1 By Captains Portlock and Dickson, London, 1789, 4to.
^ 4 vols., Paris, 1797. ^ Voyages d’un Naturaliste, &c., Paris, 1809.
* London, 1810. ® 2 vols. 4to., London, 1814.
® Narrative of an Expedition to Explore tlie Eiver Zaire, &c., London, 1818, Appendix, p. 418.
T Berolini, 1819-1822 (?). « London, 1818-1822, p. 437.
^ Supplement to the Appendix of Captain Parry’s First Voyage for the Discovery of a N.W. Passage, &c., London, 1824, p. 239.
Frankfurt am Main, 1826.
1838-1840.
Voyage de I’Astrolabe (ZooL), MM. Quoy et Gaimard, Pari.?, 1834.
REPORT ON THE ANNELIDA.
V
the Nemerteans and Planarians, but little to the Annelids, though an atlas of fifty folio plates (many coloured) enriches the work. The groups just mentioned, on the other hand, disappear from such works as The Zoology of Captain Beechy’s Voyage to the Pacific and Behrings Island in H.M.S. “Blossom,”^ and the Voyage autour du Monde par les mers de ITnde et de Chine execute sur la corvette de I’Etat la Favourite, though many other groups are mentioned in both works.
The extensive area traversed by H.M.S. “Sulphur,” under the command of Sir E. Belcher, was barren of results in regard to the Annelids ; and even Mr. Darwin’s classic voyage in H.M.S. “Beagle” produced little in the department of the marine forms further than a few interesting Planarians, including a pelagic form in the open sea, off Fernando Noronha.
During the Antarctic voyage of Sir James Eoss, Dr. (now Sir Joseph) Hooker^ mentions that Ditrypa and another Annelid were dredged at 400 fathoms, while between 200 and 400 fathoms, off Victoria Land, Serimlce and various other Annelids were procured by the same instrument. The interest taken by this author in the group is well shown by a series of carefully coloured drawings of Annelids made during the voyage in H.M. ships “ Erebus ” and “ Terror,” and forwarded to me by Mr. Murray for examination. These include examples of Syllis and Nereis, of the Phyllodocidse and Lumbrinereidse, fragments of the Terebellidse, and a complete young form, a species like Potamilla, and various examples of the Serpulidse, amongst which Serpula narconensis, with its roseate branchial plumes, appears to be represented. Excellent drawings of Tomopteris and a Pontobdella conclude the list. That a botanist should have done so much under the circumstances merits more than a passing comment. From the published account of this voyage^ a considerable amount of information is obtained about the nature of the natural harbours at Kerguelen, the muddy bottoms of which especially abound in Annelids.
The latter are, again, absent from the account of the voyage of H.M.S. “Fly,”^ but the zoology of this expedition, it is true, comprises only four pages of the appendix. None occur in the invertebrates found during the voyage of H.M.S. “ Samarang.” ^
A change was now, however, looming in the distance, and the publication of Prof. Edward Grube’s account of the Annelids procured in the Eeise in den aussersten Norden und Osten Sibiriens,® forms one of the earlier indications of increased attention to the group. Several expeditions, however, the accounts of which were j)ublished after this date, give little information on the subject, such as the United States Exploring Expedition (C. Wilkes),^ the Voyage autour du Monde of “La Bonite,”® and
1 London, 1839, 4to. ^ Ann. and Mag. Nat. Hist, ser. i. vol. xvi. p. 238, 1845.
3 Voyage of Discovery and Research in the Southern and Antarctic Regions, London, 1847, 4 vols.
* 2 vols. 8vo., London, 1847.
^ Zoology of the Voyage of H.M.S. “ Samarang,” under Sir E. Belcher, London, 1850.
® St. Petersburg, 1851. ^ Philadelphia, 1852. ® Paris, 1852.
VI
THE VOYAGE OF H.M.S. CHALLENGER.
Macgillivray’s Narrative of the Voyage of H.M.S. “ Eattlesnake,” ^ yet in the latter the powerful help of Assistant-Surgeon T. H. Huxley was available. As an example of the scanty reference to the group in these, it may be mentioned that the only example of the “Vers” in the Zoologie of the “La Bonite” is Sagitta hipunctata. This work, however, is pre-eminent in its folio atlas of beautifully coloured steel engravings. No improvement on the foregoing in regard to the Annelids occurred in the zoology of H.M.S. “Herald,” edited by Edward Forbes;^ and the same may be said of The Last of the Arctic Voyages, by Sir E. Belcher.^
The presence of W. Stimpson in the United States Surveying Expedition to the North Pacific, Japan Sea, &c., resulted in the brief description^ of various Annelids, but the con- tribution is limited and difficult to follow from the absence of illustrative figures. In the voyage of A. S. OErsted to the West Indies and Central America, and that of H. Kroyer to South America, about a hundred Annelids were procured, and these formed the materials for Grube’s well-known Annulata CErstediana,® which only lacked figures to have been much more important. All previous expeditions, however, were eclipsed by the able report of Kinberg on the Annelids of the Swedish frigate “ Eugenie,” ® in which a thoroughly scientific grasp of this subject was taken, and the aid of a really skilful artist obtained in drawing the structural features of the animals. Unfortunately, but a fragment of the work is complete, the majority of the forms being only known by brief descriptions in the Ofversigt k. Vetensk.-Akad. Forhandl. Such a contribution marks an era in the list of voyages, and is equally creditable to Kinberg and his country.
Following close upon this publication is the treatise by Schmarda on the Turbellarians Rotifers and Annelids procured “Auf einer Reise um die Erde.” The second volume is composed for the most part of an account of the marine Annelids, with nineteen brightly coloured plates (steel engravings) and many woodcuts, and both it and the former are referred to constantly in the subsequent Report on the Annelids collected by the Challenger. The work^ forms a conspicuous landmark in the history of the subject, and though often failing in severe attention to structural details, bears evidence of much labour and perseverance.
The many voyages undertaken in our country previous to the Challenger Expedition had produced zoologically results of considerable value, though the scientific staff connected with their production both at home and abroad was often of a very limited description. We were, however, not in advance of other countries in this respect. This is boldly shown by the liberal subsidy by government which enabled the fine series of volumes (six of which were zoological) giving the results of the Austrian Novara Expedition to
1 London, 1852. ^ London, 1854. ^ London, 1855.
* Proc. Acad. Nat. Sci. Philad., 1855, &c.
® Vidensk. Meddel. f. d. nat. Foren., 1856-58.
® Kongliga svenska Fregatten Eugenics Eesa omkring jorden, &c., Stockkolm, 1858.
^ Zweite Heft, Leipzig, 1861.
REPORT ON THE ANNELIDA.
vii
surpass anything of this kind previously published in our own country. Grube took the Annelids ^ in hand, and in less than half a hundred pages and four fine steel plates carefully described them and figured the most interesting. In this department, therefore, the comparison with the Annelidan results of the Challenger is noteworthy, since both ships traversed similar seas.
Stimulated by such examples as the foregoing, as well as by the activity of the Scandinavians and Americans, and more than all by the influence of the late Sir Wyville Thomson and Dr. Gwyn Jeffreys, in conjunction with Dr. Carpenter, our own government fitted out the “Lightning” and “Porcupine,” and in both ships, notably in the final expedition of the latter (1870), very considerable additions were made to our knowledge of the Annelids, especially by the labours of Prof. Ehlers of Gottingen, who, after the death" of Edward ClajDarede, examined those frequenting deeper water than 500 fathoms in the “Lightning” and “Porcupine” expeditions of 1868 and 1869. Subsequent voyages have further extended our information in the Annelidan department, as for instance the cruise of Dr. Gwyn Jeffreys in the “Valorous” to Davis Strait, the last North Polar Expedition under Sir George Nares, the Transit of Venus Expedition to Kerguelen, the dredgings of Captain St. John in the China Sea, and those of Dr. John Murray in the “ Knight Errant ” and “ Triton ” in the North Atlantic.
The recent advances made in our knowledge of the Annelids by the expeditions of other nations have been numerous ; indeed, no well-organised exploration of the ocean now returns without representatives of the group. Amongst others that occur to me are the American expeditions in the “ Blake,” the Annelids of which are now in the skilled hands of Prof. Ehlers ; the collection made by the German exploring ship “ Gazelle,” a preliminary account of which was published by the late veteran zoologist, Prof. Grube, in 1877, and which is frequently referred to in the following pages ; the Annelids of the Swedish North Polar Expedition under Prof. Nordenskiold, as described and figured by Theel ; and those of the Norwegian North Atlantic Expedition by Hansen. In the Challenger a large number of the Annelids were procured in the trawl, and this is consonant with our own experience in British waters. The extensive surface afforded by the trawl, and the readiness with which Annelids, Nemerteans, and even such forms as Corymorpha cling to the meshes is remarkable, while perhaps it is occasionally vain to search for them with a dredge.
General Condition of the Preparations.
In reviewing the condition of the preparations forwarded for examination, it has to be stated that many, as might have been expected, are injured or have to be described from mere fragments. A few had been dried. The great depth from which others were obtained probably caused laceration or softening before they reached the surface, and in
1 Annel. Novara-Exp., Zool. Theil., Bd. ii. Atth. 3, 1868.
THE VOYAGE OF H.M.S. CHALLENGER.
viii
this respect it is well to remember that many sj)ontaneously rupture on the slightest irritation.
In regard to the preservation of the Annelids, it is unsafe to mix them with other classes, for when separation is carried out by hands that perhaps are imperfectly acquainted with the grou|), loose scales or cirri are apt to be overlooked, and are thus irretrievably lost.
One important aid in dealing with any group was entirely absent, viz., coloration. The staff on board the Challenger w^as wholly inadequate to overtake this department, yet the beauty of the marine Annelids as a whole depends on the endless variety and often gorgeous loveliness of their hues.
Methods followed in Description.
In dealing with the materials placed at my disposal, an external survey of each was made under a lens, the structure of the feet, the minute anatomy of the bristles and hooks, as well as of the body-wall and other parts, was considered. It w'as impossible, however, to do more than glance at the anatomy of the group in passing, leaving for the present, for instance, such interesting questions as the nature of the remarkably folded organ (called liver by Johannes Steen at the anterior jiart of the alimentary canal (below and at the sides of the gullet) of Terehellides, for future consideration. Little reliance was placed on the description of the bristles and hooks without accurate representa- tions, since many species come so close that it would be very difficult for one’s successors to comprehend all the details. The distinctions while reliable are fine. Moreover, the hard parts just mentioned are less liable to be altered by the spirit than the soft tissues of the animals. The remarkable modifications observed in the bristles of every foot in many of the groups, and which are so disposed that a regular gradation in form exists between those at the superior border, and those at the inferior border, afford even a more complex subject for reflection than the changes undergone by the spines of an Echinoderm.
Classification.
The large number of new forms brought within our knowledge by the Challenger would have been supposed to lead to a noteworthy change in classification, but from the first it was apparent that no new family was required. All the types fell under the groups already constituted, and which have been very satisfactorily given by Malmgren in his Annulata Polychseta.^ A careful review of these groups in connection with the arrangement and relations of the nerve-cords, and the general structure of the body-wall,
^ Jenaische Zeitsclir., Bd. xvi. p. 227, Jena, 1883.
2 Helsingfors, 1867.
REPORT ON THE ANNELIDA.
IX
in 1876^ led me to give a general support to this classification, and further experience has not as yet shown the necessity for any material change. It is true there are some forms, such as the genus Eulepis, which almost merit the distinction of a separate family, but they have only recently been discovered, and may properly be left for further investigation.
Species.
While perhaps some species might have been united, the difficulties surrounding the subject (arising chiefly from imperfect descriptions and figures of essential parts) have been considerable. It is hoped, however, that the present observations and drawings will enable subsequent observers to clear up the discrepancies. There can be no doubt, as A. S. CErsted observed, that a single accurate figure, for instance of a characteristic hook or bristle, is of greater cod sequence in certain cases than an elaborate Latin description ; moreover, experience does not altogether bear out the statement made by Hansen with regard to the Polynoidse, viz., that the scales are of greater importance in specific separa- tion than the bristles. A specimen certainly would be more easily and accurately determined with both scales and bristles present, but some, including myself, would consider a specimen of greater value with bristles and without scales, than with scales and without bristles.
No less than about two hundred and twenty new species fall to be noticed.
Food of Annelids.
In many cases the food of the Annelids has been examined, and as there can be no question (excluding surface forms) that this was obtained on the bottom of the ocean, the condition of the various types in their alimentary canals has a direct bearing on their bathymetrical distribution. Thus the almost perfect state of some organisms, for example, Foraminifera and Radiolaria, with their contained protoplasm in the digestive tracts of Annelids from great depths, leads us to conclude that in all probability they live there, and do not in all cases fall to the bottom for the nourishment of the fauna of that region. The discrimination shown by the Annelids in regard to food may readily be observed by contrasting the muddy contents of the alimentary canal with that forming the tube. Most feed on mud containing minute organisms, but others devour their neighbours, small Crustaceans, zoophytes, and sponges, while a few, such as certain Nereids and Eunicidse, are partial to Fuci and other Algse.
1 Proc. Roy. Soc. Edin., 1876-77, vol. ix. p. 372. This paper has escaped the notice of Dr. Pruvot in his Eecherches Anat. et Morphologiques sur les systeme Nerveux des Annelides Polychetes, Archives d. Zool. expe'r., 1885, No. 2,
p. 210.
(zool. chall. exp.— part XXXIV. — 1885.) LI h
X
THE VOYAGE OF H.M.S. CHALLENGER.
Tubes.
The majority of the tubes formed by the Annelids collected by the Challenger have been described along with their inhabitants, but a few remarks are necessary on certain empty tubes. A very striking form is one procured by the trawl at Station 298 (south of Valparaiso); lat. 34° 1' S., long. 73° 56' W.; depth, 2225 fathoms; bottom tem- perature 35°‘6, surface temperature 59° ; sea-bottom, blue mud. The same form comes from the neighbouring Station 299. It consists of a triangular tapering tube (PI. XLIX. figs. 8, 9) composed of very fine dark mud, and having the three ridges hispid vdth short muddy processes. On magnifying the surface between the ridges (fig. 9) peculiar trans- verse markings, which almost resemble scutes, are observed. Its occupant probably pertained to the Terebellidse or a neighbouring family, but no trace of it was found.
The tubes composed of the secretion produced by the body of the animal, such as those of Hyalincecia and Eunice, are of course independent of their surroundings, but the majority have a composite nature, viz., have either an internal lining of the secretion or an admixture, and an external investment of mud or other solid particles. On the blue mud and red clay the tubes are often almost entirely formed of these deposits. As GlohigerincB appear these are studded over the surface of the mud ; while in certain localities the discoid Foraminifera are set on edge on the surface of the mud so as to render the tube hispid. Massive tubes, almost entirely composed of Foraminifera, occurred at Station 158 (south of Australia). Small bivalves and other Mollusks are also largely used to strengthen and protect muddy tubes, and the extreme development of this method is shown in the empty tube in fig. 13 of PI. XXXIXa. from Port Jackson. The tube is quite squamous, from a close series of Molluscan valves which overlap each other around the tube.
General Kemarks.
The drawings of the first fifty-five plates were made by my niece, who patiently endeavoured to render them as life-like as possible, though it was hardly possible in all cases to represent minute structural detail. Moreover, many of the specimens were so much injured that difficulty was felt in making a satisfactory picture ; indeed, previous knowledge of their structure was necessary in this respect. Upwards of thirty plates and the woodcuts are from my own drawings, a fact which will explain the somewhat tardy a^Dpearance of the Report, which had to be carried on amidst one or two distractions. Mr. Edward Prince and Mr. John Wilson, my former students and prize- men, aided me with several of the concluding plates ; while Dr. R. Marcus Gunn, previously associated with me in Perthshire, and now one of the ophthalmic surgeons at Moorfields, illustrated his own observations on the eyes of the Alciopidse and Phyllodocidse.
REPORT ON THE ANNELIDA.
XI
It may be deemed by some an inconvenience to refer to several plates for the structural and other details of the same species, but the study of the Annelids is a matter that requires so much deliberation that the inconvenience is comparatively slight. Moreover, it was found that different treatment was required in the execution of the structural plates. In the explanations of the latter, it has not been thought necessary to go into details, since this had already been done in the text.
I have to acknowledge the unfailing courtesy of Dr. John Murray, who, after the death of Sir Wyville Thomson, became Director of the Challenger Commission, of Mr. Hoyle, M.A., M.R.C.S., Naturalist to the Commission, and Mr. Monteith of the same office, who suggested improvements in passing the work through the press. Prof Moseley also kindly forwarded a fine specimen of the Hexactinellicl Sponge {Cratero- morpha meyeri) infested by Syllis ramosa, from the Oxford Museum. It was procured off the Philippines by Capt. Chimmo of H.M.S. “Herald.”
To Prof Flower, Director of the British Museum, as well as Prof Jeffrey Bell and Mr. S. 0. Ridley of the same institution, I am much indebted for their aid in examining the Annelids in the collection. I am especially obliged to Prof Jeffrey Bell for his patient and courteous attention in this respect. Lastly, Dr. Murie’s valued aid in the Library of the Linnean Society deserves my cordial acknowledgments.
GEOGRAPHICAL AND BATHYMETRICAL DISTRIBUTION.
In the following arrangement I have adopted the regions given by Prof. Busk in his able Eeport on the Polyzoa, which regions are generally acquiesced in by most observers. They are, indeed, such as suggest themselves in a natural grouping of the oceanic areas.
In the first of these. A., the North Atlantic Eegion (see the sketch-map), a large number of forms occur, and relatively few range to other areas ; but this apparent definition in so vast a region is probably due to the comparatively unexplored condition both of it and the other oceans. "Within its limits marked diversities present them- selves, such as the comparative absence of the Amphinomidse (with the exception of Paramphinone) in the north-eastern part of the area, and their abundance in the south-western; the appearance of the Euphrosynidae between tide-marks in the southern parts, and their limitation to the deeper water in the northern. Most of the genera are cosmopolitan in their range, but the remarkable new genus BusJciella is entirely confined to the abysses of this and the South Atlantic. Many interesting extensions of the previously known range of genera have been made by the Challenger, for example, the finding of Pulepis in the West Indies, its original habitat being in the Philippines*. The cosmopolitan habits of such types as Harmothoe imhricata, Hyalinceda tubicola, Scolecolepis cirrata, and Terebellides stroemi have also been more clearly disclosed.
SUEFACE FoEMS.
Besides the purely pelagic Alciopidae, which were met with by the Challenger in the Atlantic and Antarctic Oceans, as well as more abundantly in Mid Pacific, oflF the Sandwich Islands, Tomopteridae and various larval forms occurred. Amongst the latter were young Terebellidae, about half an inch in length, which were caught in the Atlantic on the return voyage. Various larval Polygordii were found in the tow-nets near St. Vincent, Cape Verde Islands, along with Tornaria, Pteropods, and Copepods. Tomopteris, again, ranged to both Atlantic and Pacific Oceans, and was ? ccompanied by Sagitta, Copepoda, and various larval forms such as the young of Chh'odota. The
XIV
THE VOYAGE OF H.M.S. CHALLENGER.
surface-fauna of our own seas presents similar cliaracters, though such types as Alciopa are rarely met with. A remarkable feature in the surface collections of the Challenger is the occurrence of a Glycera of the ordinary marine type on the surface of an inland sea in Japan. The specimen is about an inch in length, and by no means larval, for the proboscidian armature is complete. The state of the nets perhaps may not have been such as lead to absolute reliance on this peculiar exception.
A. North Atlantic Eegion.
Station II. Off Setubal ; lat. 38° 10' S. (dredged).
Allmaniella setubalensis, n. sp. Nephthys malmgreni, Theel.
Syllis setubalensis, n. sp.
long. 9° 14' W. ; 470 fathoms; green mud
Praxilla cliallengerice, n. sp. Euthelepus setubalensis, n. sp. Protula lusitanica, n. sp.
Station III. Off Cape St. Vincent; lat. 37° 2' N,, long. 9° 14' W. ; 900 fathoms; blue mud (dredged).
Nothria conchylega, Sars.
Station VI. Off Gibraltar; lat. 36° 23' N., long. 11° 18' W. ; 1525 fathoms; Globi- gerina ooze (trawled).
Evarne tenuisetis, n. sp. Nereis longisetis, n. sp.
Maldane malmgreni, n. sp. Amphicteis gunneri, Sars.
Station 3. South of the Canaries ; lat. hard ground (dredged).
Chloenea atlantica, n. sp.
Polynoe [Robertianella) synopli- thalma, n. sp., B.
25° 45' N., long. 20° 14' W. ; 1525 fathoms ;
Halodora reynaudii, Aud. and Edw. (surface).
Dalliousia atlantica, n. sp.
Station 20, East of the Antilles; lat. 18° 56' N., long. 59° 35' W. ; 2975 fathoms; red clay (dredged).
Myriocliele heeri, Malmgren.
REPORT ON THE ANNELIDA.
XV
Station 23. Oflf Sombrero and St. Thomas,
AphrocUta intermedia, n. sp.
Eulepis challengericB, n. sp. Psammolyce occidentalis, n. sp. Macduffia honhardi, n. sp.
Nothria sonibreriana, n. sp. and var.
Station 33. Off the Bermudas ; lat. 32° mud (dredged).
Eulepis ivyvillei, n. sp.
Eunotomastus grubei, n. sp.
Station 36. Off the Bermudas ; lat. 32° (dredged).
Notopygos megalops, n. sp.
Off the Bermudas, floating at surface on a ] Amphinome rostrata, Pallas.
Between tide-marks, Bermuda.
Eurythoe paci/ica, Kinberg, D. Polynoe pustulata, n. sp.
Nereis iPerinereis) melanocephala, n. sp.
Eunice vittata, Della Chiaje, var. cirrohranchiata, n. sp. harvicensis, n. sp.
West Indies ; 390 to 450 fathoms (about).
Scolecolepis cirrata, Sars, var. 2. Chcetozone atlantica, n. sp.
Maldane atlantica, n. sp,
Ampharete sombreriana, n, sp. Melinna maculata, Webster.
L' N., long. 64° 35' W. ; 435 fathoms; coral
Placostegus assimilis, n. sp. Spirobranchus occidentalis, n. sp.
' N., long. 65° 4' W.; 30 fathoms; coral
Hipponoe gaudichaudi. And. and Edw. (surface).
f
r,
Hermodice carunculata, Pallas. Eunice sp,
Aricia platycephala, n. sp. Cirratulus assimilis, n. sp.
Terebella crassicornis, Schmarda. Pista sombreriana, n. sp.
Dasychone bairdi, n. sp,
Serpida sombreriana, n, sp.
Station 44. Off Chesapeake Bay ; lat. 37° 25' N,, long. 71° 40' W. ; 1700 fathoms ; blue mud (dredged).
Melinnopsis atlantica, n. sp.
Station 45. Off the North American coast; lat. 38° 34' N., long. 72° 10' W. ; 1240 fathoms ; blue mud (dredged).
Harmothoe benthaliana, n. sp. Nephthys phyllobranchia, n. sp. Lumbriconereis punctata, n. sp.
Eunice cerstedi, Stimpson. Spioclicetopterus sp. Praxilla occidentalis, n. sp.
XVI
THE VOYAGE OF H.M.S. CHALLENGER.
■ Station 47. Off the North American coast; lat. 41° 14' N., long. 65° 45' W. ; 1340 fathoms ; blue mud (dredged).
Laranda longa, Webster. Lumbriconereis ehlersi, n. sp. Aricia norvegica, Sars. Aricidea fragilis, Webster.
Notomastus agassizii, u. sp. Maldane sp.
Myriochele Jieeri, Mgrn., var. Thelepus sp.
Terehellides stroemi, Sars.
Station 48. Nova Scotia; lat. 43° 4' N., long. 64° 5' W. ; 51 fathoms; rock (dredged).
Eusyllis tubifex, Gosse. ' Thelepus cincinnatus, 0. Fabricius,
var. canadensis, nov.
Station 49. Off Halifax ; lat. 43° 3' N., long. 63° 39' W. ; 85 fathoms ; gravel, stones (dredged).
Euphrosyne borealis, CErsted. Lcetmonice producta, Grube, var. assimilis, nov.
Nereis pelagica, Linnaeus. Potarnilla torelli, Malmgren. Protula americana, n. sp.
Station 50. Off North American coast; lat. 42° 8' N., long. 63° 39' W. ; 1250 fathoms ; blue mud (dredged).
Chcetozone benihaliana, n. sp.
Station 63. Mid Atlantic; lat. 35° 29' N., long. 50° 53' W. ; 2750 fathoms; red clay (trawled).
Amphicteis gunneri, Sars, var. atlantica, nov.
Eupista darwini, n. sp., var., Gr. Lanassa benihaliana, n. sp.
Ehlersiella atlantica, n. sp.
Station 70. West of the Azores; lat. 38° 25' N., long. 35° 50' W. ; 1675 fathoms; Globigerina ooze (trawled).
Lcetmonice producta, Grube, var. xvilleinoesi, nov., B, C, D.
Station 73. Off the Azores ; lat. 38° 30' N., long. 31° 14' W. ; 1000 fathoms ; Pteropod ooze (dredged).
Leanira hystricis, Ehlers. | Staurocephalus atlanticus, n. sp.
REPORT ON THE ANNELIDA.
XV] 1
Station 75. Off Fayal, Azores; lat. 38° 38' N., long. 28° 28' W. ; 450 fathoms; volcanic mud (dredged).
Lepidonotus squamatus, Linn. Sigalion huskii, M‘Intosh. Hyalincecia tubicola,0. F. Mtiller, B.
Glycera tessellata, Grube. capita ta, (Ersted.
Branchiomma vesiculosum, Montagu.
^ O
Ditrypa arietina, 0. F. Muller.
Station 76. Off the Azores; lat. 38° 11' N., long. 27° 9' W. ; 900 fathoms; Pteropod ooze (dredged).
Praxilla sp.
Off St. Vincent, Cape Verde Islands.
Hermione hystrix, Savigny. Lagisca tenuisetis, n. sp.
Lagisca per acuta, n. sp. Phyllodoce sancti-vincentis, n. sp. Hesione pantherina, Eisso.
Nereis atlantica, n. sp.
Terebella sp.
Nereis [Platy nereis) dumerilii, Aud. and Edw.
Nematonereis sp.
Eunice torquata, De Quatrefages. Sahellaria{Pallasia)jolmstoni, n. sp. Loimia savignyi, n. sp.
Laonome haeckelii, n. sp.
Station 101. Off Sierra Leone ; lat. 5° 48' N., long. 14° 20' W. ; 2500 fathoms; blue mud (trawled).
Buskiella ahyssorum, n. sp., B.
Station 106. Mid Atlantic, near Equator; lat. 1° 47' N., long. 24° 26' W. ; 1850 fathoms ; Globigerina ooze (trawled)
Lagisca {Agnodice) moseleyi, n. sp. | Buskiella ahyssorum, n. sp., B.
St. Thomas, West Indies, shallow water and between tide-marks.
Hesione sp. (1). j Sabella bipunetata, Baird.
Nereis antillensis, n. sp. Dasychone ivyvillei, n. sp.
Marphysa goodsiri, n. sp.
nigro-macidata (Baird).
B. South Atlantic Eegion.
In this region the two chief centres for specimens were the coast of Brazil and the Cape. As formerly indicated, the remarkable Buskiella ahyssorum extended into the deeps (2025 fathoms) of this as well as the North Atlantic. The wide range of
(ZOOL. CHALL. EXP. PART XXXIV. 1885.) LI C
xviii THE VOYAGE OF H.M.S. CHALLENGER.
Hyalinoecia tubicola, of wliicli a large variety occurred off the South American coast, is also noteworthy. Some of the types, such as Euphione, Pista mirahilis, Eupista, and Ranzania are also peculiar.
Station 113a. OIF Fernando Noronha; lat. 3° 47' S,, long. 32° 24' 30" W.; 25 fathoms; volcanic sand and gravel (dredged).
Nereis eatoni, M‘Intosh, C.
Station 122. Off Brazil; lat. 9° 5' S., long. 34° 50' W. ; 350 fathoms; red mud (trawled).
Syllis hrasiliensis, n. sp. Nematonereis schmardce, n. sp.
Nereis [Ceratonereis) hrasiliensis, Eunice equibranchiata, n. sp.
n. sp.
Station 124. Off Brazil; lat. 10° 11' S., long. 35° 22' W. ; 1600 fathoms; red mud (dredged).
Polynoe {Robertianella) synophthalma, n. sp., A.
Station 133. Off Tristan da Cunha ; lat. 35° 41' S., long. 20° 55' W. ; 1900 fathoms; Globigerina ooze (trawled).
Lcetmonice producta, Grube, var. willemoesi, nov., A, C, D.
STx^tion 141. South of the Cape; lat. 34° 41' S., long. 18° 36' E. ; 98 fathoms; green sand (dredged).
Polyiwe cape.nsis, n. sp.
Syllis capensis, n. sp. Lumbriconereis p>ettigreioi, n. sp.
Ranzania capensis, n. sji. Prionospio capensis, n. sp. Praxilla capensis, n. sp.
Stations 141, 142. Between tide-marks. Sea Point, Cape Town, and Simon’s Bay, &c.
Euplirosyne capensis, Kinberg. Lepidonotus walilbergi, Kinberg. Eunoa capensis, n. sp.
Polynoe attenuata, n. sp.
Eulalia capensis, n. sp. Notocirrus capensis, n. sp. Nematonereis sp.
Eunice murrayi, n. sp.
Trophonia capensis, n. sp. Cirratulus capensis, Schmarda. Nicomaclie capensis, n. sp. Sabellaria (Pallasia) capensis, n. sp. Sclimardanella pterochceta, (Schmarda).
Dasyclione violacea (Schmarda).
REPOET ON THE ANNELIDA.
XIX
Station 142. South of the Cape; lat. 35° 4' S., long. 18°- 37' E. ; 150 fathoms; green sand (dredged).
EupJiione elisahethce, n. sp. | Dasychone capensis, n. sp.
Protula capensis, n, sp.
Station 320. Off South America; lat. 37° 17' S., long. 53° 52' W. ; 600 fathoms; green sand (trawled).
Eulagisca corrientis, n. sp., C. Lcenilla fusca, n. sp.
Syllis robertiance, n. sp.
Pista mirahilis, n. sp.
corrientis, n. sp. Sahella assimilis, n. sp.
Hyalinoecia tuhicola, 0. F. Muller, var.. A,
Station 322. Off South America; lat. 35° 20' S., long. 53° 42' W. ; 21 fathoms; sand and shells (trawled).
Terebella seticornis, n. sp.
Station 325. Off the coast of Buenos Ayres; lat. 36° 44' S,, long. 46° 16' W. ; 2650 fathoms ; blue mud (trawled).
Myriochele heeri, Malmgren, var. | Amphicteis sarsi, n. sp.
Eupista grubei, n. sp.
Station 333. West of Tristan da Cunha; lat. 35° 36' S., long. 21° 12' W. ; 2025 fathoms ; Globigerina ooze (trawled).
BusMella abyssorum, n. sp., A.
Station 335. North of Tristan da Cunha; lat. 32° 24' S., long. 13° 5' W. ; 1425 fathoms ; Pteropod ooze (dredged).
Nothria abranchiata, n. sp., C.
Station 344, Off Ascension; lat. 7° 54' S,, long. 14° 28' W, ; 420 fathoms; volcanic sand (dredged).
Eunice prognatha, n. sp.
XX
THE VOYAGE OF H.M.S. CHALLENGER.
O. South Indian or Kerguelen Eegion.
One of the most striking features in the collection made in this region is the large pro- portion of species pertaining to Kerguelen. The abundance of Annelids in the deep water of the land-locked bays of this island, as well as the special opportunities of the naturalists for collecting them, probably account for their number. The bottom of these still inlets is also favourable to the increase of such forms. This is also the headquarters of Lcetmonice ‘pvoducta, Grube, varieties of which, however, extend into several of the great oceans. The cosmopolitan nature of certain Annelids is shown by the occurrence of Scolecolepis cirrata and Terebellides strcemi in the same water. This region and the neighbouring sea contain also many new forms, Terebellidse being especially conspicuous.
^Station 144a. Off Marion Island; lat. 46° volcanic sand (dredged).
Lagisca antarctica, n. sp.
magellanica, n. sp., var. gruhei., nov.
Exogone heterosetosa, n. sp.
48' S., long. 37° 49' 30" E.; 69 fathoms ;
Nereis eatoni, MTntosh, B. Terehella Jiabellum, Baird.
Neottis antarctica, MTntosh, G. Serpula narconensis, Baird, G.
Station 145a. Off Prince Edward Island ; lat. 46° 41' S., long. 38° 10' E.; 310 fathoms ; volcanic sand (dredged).
Polyeunoa Icevis, n. sp. | Eunice edwardsi, n. sp.
Scalibregma injlatum, H. Eathke, var.
Station 146. East of Prince Edward Island; lat. 46° 46' S., long. 45° 31' E.; 1375 fathoms ; Globigerina ooze (trawled).
Lcetmonice producta, Grube, var. willemoesi, nov.. A, B, D.
Polynoe {Admetella) longipedata, n. sp.
Maldanella antarctica, n. sp.
Station 147. Between Prince Edward and Kerguelen Islands; lat. 46° 16' S., long. 48° 27' E.; 1600 fathoms ; Diatom ooze (trawled).
Lcetmonice producta, Grube, var. j Lagisca crosetensis, n. sp.
benthaliana, nov., E. | Petta assimilis, n. sp.
Amphicteis wyvillei, n. sp.
REPORT ON THE ANNELIDA.
XXI
Station 149. Off Kerguelen ; lat. 49° 8' S., long. 70° 12' E., &c. (dredged at and near).
LcBtmonice producta, Grube.
Evarne kerguelensis, n. sp.
Eupolynoe mollis, M'Intosb.
Hemnadion kerguelensis, n. sp.
NephtJiys trissophyllus, Grube.
Salvatoria kerguelensis, n. sp.
Eusyllis kerguelensis, n. sp.
Syllis gigantea, n. sp.
Sg)hcBrosyllis kerguelensis, n. sp.
Autolytus maclearanus, n. sp.
Nereis kerguelensis, n. sp.
Lumhriconereis kerguelensis, Grube.
Eunice magellanica, n. sp. young (Vj. Glycera kerguelensis, n. sp.
Spirorbis sp.
Scoloplos kerguelensis, n. sp. Travisia kerguelensis, n. sp. Trophoniob kerguelarum, Grube. . Brada mammillata, Grube. Scolecolepis cirrata, Sars, var. Notomastus sp. (1).
Praxilla kerguelensis, n. sp.
assimilis, n. sp. Ampharete kerguelensis, n. sp. Ampliitrite kerguelensis, M‘Intosh. Ereutho kerguelensis, n. sp.
Poly cirrus kerguelensis, n. sp. Artacama challenger ice, n. sp. Terebellides stroemi, M. Sars, var.
Between Kerguelen and Macdonald Islands ; surface.
Tomopteris carpenteri, De Quatrefages.
Station 151. Off Heard Island; lat. 52° 59' S., long. 73° 33' E.; 75 fathoms; volcanic mud (dredged).
Phyllocomus croceus, Grube.
Surface of sea between Stations 154 and 155 (Antarctic Sea). By tow-net.
Alciopa antarctica, n. sp.
Station 156. Antarctic Sea; lat. 62° 26' S., long. 95° 44' E.; 1975 fathoms; Diatom ooze (trawled).
Nothria abranchiata, n. sp., B. Grubianella antarctica, n. sp.
Ephesia antarctica, n. sp. Lecena antarctica, n. sp.
Station 157. Midway between the Antarctic region and Australia ; lat. 53° 55' S., long. 108° 35' E.; depth, 1950 fathoms ; Diatom ooze (trawled).
Lcetmonice producta, Grube, var.
ivyvillei, nov.
Nothria armandi, n. sp.
Trophonia ivyvillei, n. sp. Praxilla abyssorum, r. sp. Pista abyssicola, n. sp.
xxu
THE YOYAGE OF H.M.S. CHALLENGER.
D. The Australian Eegion.
The types in this area are in many cases peculiar and novel. It and the Indian region are distinguished by the ^irominence of the Chloeia group, and the Australian specially by the remarkable nature of the Polynoidse, e.g., Lejndonotus cristatus, Polynoe mirabilis, Polynoe platycirrata, Scalisetosus ceramensis, Eunoa iphionoides, and Eunoa ahyssorum. The latter is also the region of fine examples of the genus Thalenessa, of one of the most remarkable discoveries of the Challenger, viz., the branched Syllis, of many Eunicidee, from the large Eunice amphroditois downward, of Nicidon, the edible Palolo, and the striking Nothria willemoesii with its spinose tube. Comparatively few of the species range into the other areas if we except the ubiquitous varieties of Lcetmonice producta and one or two others.
Station 158. Considerably south of Australia; lat. 50° 1' S., long. 123° 4' E. ; 1800 fathoms ; Globigerina ooze (trawled).
HyalincBcia henthaliana, n. sp. | Gruhianella antarctica, n. sp., var.
Station 160. South of Australia; lat. 42° 42' S., long. 134° 10' E,; 2600 fathoms; red clay (trawled).
Eunoa ahyssorum, n. sp. | Polynoe ascidioides, n. sp.
Station 162. Bass Strait; lat. 39° 10' S., long. 146° 37' E. ; 38 fathoms; sand and shells (dredged).
Polynoe platycirrata, n. sp. Thalenessa oculata, n. sp. Staurocephalus australiensis, n, sp.
Eunice vittata, D. Chiaje.
pycnohrancliiata, n. sp. hassensis, n. sp.
Station 163a. Twofold Bay, Australia ; lat. 36° 59' S., long. 150° 20' E.; 150 fathoms; green mud (trawled).
Phyllodoce duplex-, n. sp. [ Sahellaria (Pallasia) giardi, n. sp.
Terebella gruhei, n. sp.
Station 163b, Off Port Jackson; lat. 33° 51' S,, long. 151° 22' W. ; 35 fathoms; hard ground (dredged).
Aphrodita australis, Baird. Thalenessa Jlmhriata, n. sp.
Eunice aphroditois, Pallas. Sahella fusca, Grube.
REPORT ON THE ANNELIDA.
xxiii
Station 166. Off New Zealand; lat. 38° 50' S., long. 169° 20' E.; 275 fathoms; Globigerina ooze (trawled).
Eunoa iphionoides, n. sp.
Station 167a. Queen Charlotte Sound, New Zealand; lat. 41° 4' S., long. 174° 19' E. ; 10 fathoms ; mud (dredged at and near).
Leanira Icevis, n. sp.
Nephthys verrillii, n. sp. Hyalinoecia tuhicola, 0. F. Muller, var. longihranchiata, nov.
Eone trijida, ii. sp.
Glycera lamelliformis, n. sp. Apomatus elisabethm, n. sp. Pomatocerus strigiceps, Morch.
Station 168. Queen Charlotte Sound, New Zealand; lat. 40° 28' S., long. 177° 43' E. ; 1100 fathoms ; blue mud (trawled).
Eumenia reticulata, n. sp. Melinna armandi, n. sp.
Maldanella neo-zealanice, n. sp. Lecena neo-zealanice, n. sp.
Lecena langerhansi, n. sp.
Station 169. Off New Zealand; lat. 37° 34' S., long. 179° 22' E. ; 700 fathoms; blue mud (trawled).
LcBtmonice producta, Grube, var.
willemoesi, nov.. A, B, C. Polynoe [Macellicephalci) mira- hilis, n. sp.
Lumbrico nereis neo-zealanice, n. sp.
Nothria tenuisetis, n. sp, minuta, n. sp. Chatozone pacijica, n. sp, Lanassa sarsi, n. sp. Terebellides sp.
Station 170. Off the Kermadec Islands; lat. 29° 55' S., long, 178° 14' W. ; 520 fathoms ; volcanic mud (trawled).
Lagisca hermadecensis, n. sp.
Nicidon balfouriana, n. sp.
Station 171. North of the Kermadec Islands; lat. 28° 33' S., long. 177° 50' W. ; 600 fathoms ; hard ground (trawled).
Polynoe magnipalpa, n. sp.
Terebella kermctdecensis, n. sp.
XXIV
THE VOYAGE OF H.M.S. CHALLENGER.
Station 172. Off Tongatabu ; lat. 20° 58' S., long. 175° 9' W. ; 18 fathoms; coral mud (dredged).
Hesione pacijica, n. sp. | Nereis {Platynereis) tongatabuensis, n. sp.
Station 174. South of Fiji Islands; lat. 19° 6' S., long. 178° 14' E. ; 140 fathoms; coral mud (dredged).
Praxilla hdllikeri, n. sp. | Te7xhellides ehlei^si, n. sp.
OIF the Fiji Islands,
Palolo vwidis, Gray.
Off Levuka, Fiji,
Eurythoe pacijica, n. sp., var. levu- Tcaensis, nov., A.
Psammolyce fijiensis, n. sp.
Station 186. Torres Strait; lat. 10° 30' S., long. 142° 18' E.; 8 fathoms; coral mud (dredged).
Lepidonotus cidstatus, Grube.
Eupompe australiensis, n. sp.
Eunice torresiensis, n. sp.
Thelepus sp.
Eunice Unhranchiata, n. sp. Hyalincecia tuhicola, 0. F. Muller, var. papuensis, nov.
Arafura Sea, south of New Guinea.
Nephthys dibrancliis, Grube.
Ne7-'eis [Platynereis) a7^afu7^ensis,n.sg.
Eunice elseyi, Baird. Protula a7’'afurensis, n. sp.
Station 192. Off Ki Island, Flores Sea; lat. 5° 49' 15" S., long. 132° 14' 15" E. ; 140 fathoms ; blue mud (dredged).
Syllis ramosa, n. sp., E.
Off Amboina (dredged).
Nothria willemoesii, n. sp.
Glycera ambome7isis, n. sp.
REPORT ON THE ANNELIDA.
XXV
Station 194a. Off Ceram; lat. 4° 31' S., loug. 129° 57' E. ; 360 fathoms; volcanic mud (trawled).
Scalisetosus ceramensis, n. sp.
Near Banda, off the Moluccas.
Chloeia fiisca, n. sp.
Admiralty Islands ; 16 to 25 fathoms.
Thalenessa digitata, n. sp.
E. Philippine or Japanese Region.
This region (E), like D, contains many species common to the shores of Hindustan, as shown by a series of beautifully coloured drawings made under the superintendence of Sir Walter Elliot of Wolfelee,^ and of the collections made by Dr. Anderson of the Imperial Museum of Calcutta. The Chloeia group is as well represented as in the former area (D), and in this respect agrees with the Bengal series. A variety of Lattmonice 'producta is common to this region and C, while the rich waters of Japan add a new species to the genus, amidst a profusion of other remarkable t}^es. The area is specially rich in new Polynoidse, such as those commensalistic in Euplectella, and in the Hexac- tinellid Sponge lodging the branched Syllis. It is also the region of Palmyra and Eupholoe, and of the large-eyed Genetyllis oculata.
Station 201. Basilan Strait, south of the Philippines; lat. 7° 3' N., long. 121° 48' E.;
82 fathoms ; stones and gravel (trawled).
Notopygos lahiatus, n. sp. ' Polynoe{Lepidonotus)iphionoidesp.?p.
Eunoa mindanavensis, n. sp. Eupholoe philippensis, n. sp.
Eunice mindanavensis, n. sp.
Station 205. Offthe Philippines; lat. 16° 42' N., long. 119° 22' E.; 1050 fathoms; blue mud (trawled).
Serptda philippensis, n. sp.
Off Hong Kong; 10 fathoms (dredged).
Dasychone orientalis, n. sp.
1 I have to thank Sir Walter Elliot for so kindly permitting me to have these for several years. Unfortunately, no specimens accompanied the drawings, and thus much less could be accomplished with them than I at first anticipated.
(ZOOL. CHALL. EXP. PART XXXIV. 1885.)
LI d
XXVI
THE VOYAGE OF H.M.S, CHALLENGEE.
Station 209. Off Zebu; lat. 10° 14' N., long. 123° 54' E.; 95 fathoms; blue mud (dredged).
Lagisca hexactinellidce, n. sp. Polynoe [Langerhansia) euplectellce, n. sp.
Syllis ramosa, M‘Intosh, D. Cirratulus zehuensis, n. sp. Artacama zehuensis, n. sp.
Sahella zehuensis, n. sp.
Station 214. South of Mindanao ; lat. 4 33 N., long. 127° 6' E.; 500 fathoms; blue
mud (trawled).
Ipliionella cimex, De Quatrefages. |
Station 232. South of Yedo, Japan ; lat. 35° green mud (dredged).
LcBtmonice japonica, n. sp.
Eunoa yedoensis, n. sp.
Leanira areolata, n. sp.
Lumhriconereis hifurcata, n. sp.
japonica, Marenzeller.
Genetyllis oculata, n. sp.
11' N., long. 139° 28' E.; 345 fathoms;
Nothria inacrohranchiata, n. sp. Maldane sarsi, Malmgren. Nicomache jap)onica, n. sp.
Praxilla lankesteri, n. sp. Amjohicteis jap>onica, n. sp.
Off Japan; lat. 33° 56' N., long. 130° 27' E.; 30 fathoms.
Ammotrypane gracile, n. sp.
Station 233a. Near Kobe, Japan; lat. 34° 38' N., long. 135° 1' E.; 50 fathoms; sand (dredged).
Palmyra' aurifera, Savigny (1). Lepidonotus gymnonotus, Maren-^ zeller.
Polynoe ocellata, n. sp.
Leanira japanensis, n. sp.
Nereis [Platynereis) kohiensis, n. sp.
Eunice Jcobiensis, n. sp. PhyllochcBtopterus claparedii, n. sp. Pista fasciata, (Grube).
Dasychone picta, n. sp.
japonica, n. sp.
ELydroides mul tispinosa, Marenzeller.
Station 233b. South of Japan; lat. 34° 18' N., long. 133° 35' E.; 15 fathoms; blue mud (trawled).
Cldoeia fiava, Pallas.
Dasyhranchus sp.
REPOET ON THE ANNELIDA.
xxvii
Station 235. South of Yedo, Japan ; lat. 34° 7' N., long. 138° 0' E.; 565 fathoms ; green mud (trawled).
Loetmonice apJiroditoides, n. sp. | Folynoella levisetosa, n. sp.
Station 236. South Japan; lat. 34° 58' N., long. 139° 29' E.; 775 fathoms; green mud (trawled).
Tlielepus marenzelleri, n. sp.
Off Yokohama, Japan ; 550 fathoms.
Lagisca yokohamiensis, n. sjd. |
Station 241. North Pacific; lat. 35° 41' N. clay (trawled).
Lcetmonice producta, var, hen- j thaliana, C. 1
Lumhriconereis heteropoda, Marenzeller.
long. 157° 42' E.; 2300 fathoms; red
Nicomache henthaliana, n. sp. Eusamytha pacijica, n. sp.
Station 244. Depths of the Pacific; lat. 35° 22' N., long. 169° 53' E.; 2900 fathoms ; red clay (trawled).
Placostegus ornatus, Sowerby.
Station 246. Mid Pacific ; lat. 36° 10' N., long. 178° 0' E.; 2050 fathoms; G-lobigerina ooze (trawled).
Melinna pacijica, n. sp.
F. North Pacific Eegion.
Comparatively few Annelids come from this region, but three out of the five are surface forms of interest (Alciopidse). The members of this family thus frequent the superficies of all the great seas, from the Arctic to the Antarctic Oceans, and from the eastern to the western shores of the Americas. Prof. Huxley’s remark that it is doubtful whether any well marked provinces of the ocean can be defined by the occurrence of purely pelagic animals thus still holds.
As to the causes which have led to the relegation of such peculiar forms as Buskiella to the remote abysses of the Atlantic, and of Lecena ahyssorum and Placostegus henthalianus to the depths of the North Pacific, various opinions may be held. We have no reliable data in support of the view which supposes that these “ primitive ” types have
XXVlll
THE VOYAGE OF H.M.S. CHALLENGER.
been gradually pressed by the more prolific and hardier shallow-water forms deeper and deeper into the ocean. In the case of Lecena and Placostegus, again, both are members of genera not unknown in comparatively shallow water, and even between tide-marks.
Station 253. Mid Pacific; lat. 38° 9' N., long. 156° 25' W.; 3125 fathoms; red clay (dredged).
Lecena ahyssorum, n. sp. | Placostegus henthalianus, n. sp.
Surface of the sea at Honolulu.
Alciopa quadrioculata, n. sp. | Alciopa sp. {V).
Greeffia oahuensis, n. sp.
G. South Pacific Eegion.
The majority of the specimens from this area come from the Strait of Magellan, the confined waters of which seem to be favourable for their development. Of characteristic forms the most striking are Aj)hrodita echidna, De Quatrefages, two species of Lagisca, Eunoa opalina, Leanira magellanica, Eunice, Nothi'ia, Plemipodus, Sarny thoq)sis, Eupista, and Euthelepus.
Station 272. Mid Pacific; lat. 3° 48' S., long. 152° 56' W.; 2600 fathoms; Radiolarian ooze (trawled).
Myriochele paeijiea, n. sp.
Off Tetuaroa Islands.
Glyeera sagittarice, n. sp.
Station 285. Pacific ; lat. (trawled).
32° 36' S., long. 137° 43' W.; 2375 fathoms; red clay Placotegus morchii, n. sp.
Station 298. Off the west coast of South America; lat. 34° 7' S., long. 73° 56' W.; 2225 fathoms ; blue mud (trawled).
Lumbriconereis ahyssorum, n. sp. Nothria pycnobranchiata, n. sp. ehlersi, n. sp.
Maldanella valparaisiensis, n. sp. Samythopsis grubei, n. sp.
Eupista darwini, n. sp., A.
REPORT ON THE ANNELIDA.
XXIX
Station 299. Off Valparaiso; lat, 33° 31' S., long. 74° 43' W.; 2160 fathoms; blue mud (trawled).
Euthelepus chilensis, n. sp.
Station 302. West of Patagouia ; lat. 42° 43' S., long. 82° 11' W.; 1450 fathoms;
Globigerina ooze (trawled).
Vermilia sp.
Station 306a. Strait of Magellan; lat. 48° 27' S., long. 74° 30' W.; 345 fathoms; blue mud (trawled).
Leanira magellanica, n. sp. | Hemipodus magellanicus, n. sp.
Station 307. Strait of Magellan ; lat. 49° 24' S., long. 74° 23' W.; 140 fathoms; blue mud (trawled).
Aplirodita echidna, De Quatrefages.
Station 308. Strait of Magellan; lat. 50° 8' S., long. 74° 41' W. ; 175 fathoms; blue mud (trawled).
Lagisca magellanica, n. sp.
darwini, n. sp. Eunice magellanica, n. sp.
Seipula narconensis, Baird, var. magellanica, nov., C.
Station 310. Strait of Magellan; lat. 51° 27' S., long. 74° 3' W.; 400 fathoms; blue mud (trawled).
Lagisca magellanica n. sp., var. murrayi, nov.
Station 311. Strait of Magellan; lat. 52° 45' S., long. 73° blue mud (trawled).
Eunoa opcdina, n. sp.
46' W.;
245 fathoms ;
Station 313. Strait of Magellan; lat. 52° 20' S., long. 67° 39' W.; 55 fathoms; sand (trawled).
Nereis patagonica, n. sp.
XXX
THE VOYAGE OF H.M.S. CHALLENGER.
BATHYMETEICAL DISTRIBUTION.
In regard to bathymetrical distribution, the greatest number of species occur in the shallow water (10 fathoms and under), probably because much work was done in this region in the pinnace, especially in rich localities, and also because the opportunities for collecting between tide-marks were fully taken advantage of ; moreover, the surface-forms are included in this division. The two regions ranging from 10 to 50 fathoms and from 50 to 100 fathoms have each about the same number of Annelids, and both are similar in respect to new forms. In the three areas just mentioned, the known species are fairly represented, though the majority are new. The number between 100 and 200 fathoms is less than the foregoing, but the proportion of new forms is much higher, and several are of considerable interest, e.g., Syllis rcanosa. In the rich region ranging from 200 to 500 fathoms, very few known forms break the long list (the second of the series) of novel Annelids. About five new genera are included, and the remarkable types are exemplified by Allmaniella, Scalisetosus, Eulepis, Genetyllis ocidata, Macdiiffia, Hemipodus, and Euthelepus. The number of species at this depth, however, may be partly due to the more frequent dredging therein. Between 500 and 600 fathoms the number falls to less than half that in the previous grouj), but the majority are new. The number found between 600 and 1000 fathoms include two known S23ecies out of a list of fourteen.
The four sjDecies occurring between 1000 and 1200 fathoms are new. Those between 1200 and 1500 fathoms are more than five times as numerous as the last, and include only five known forms, most of which, however, are found in shallow water as well as at this great depth, e.g., Eunice oerstedi, Aricia norvegica, Amphicteis gimneri, and Terehel- lides stroemi, the latter three, besides, having a very wide geograj^hical range. About the same number were procured between 1500 and 2000 fathoms. All are new, and three new genera required to be formed.
Between 2000 and 2500 fathoms the total numbers about half that just mentioned (between 1500 and 2000 fathoms), and all are new, while four new genera are present in the series. In the region between 2500 and 3000 fathoms several known forms occur, viz., LcBtmonice producta, a marked variety [henthaliana) of which, however, only is found at this dej^th, Myriochele lieeri, Amphicteis gunneri (var. atlantica), and Placo- stegus 07'iiatus. If the diagnosis be correct, the latter was first procured by Sowerby in shallow water in the Phili^Dpines. The two forms from the profound abyss of 3125 fathoms are new, though the genera are well known.
In glancing over the lists, and excluding the pelagic types, it is evident that no definite law as to the j^resence or absence of genera at particular depths, can be enunciated, though it is true that such genera as Lecena, Eupista, Euthelepus, Melin-
REPORT ON THE ANNELIDA.
XXXI
nopsis, Eusamytha, Samythojjsis, GrubianeUa.^ and others come from profound depths. But it must be remembered that much as the memorable voyage of the Challenger has added to our knowledge in this respect, still more remains to be accomj^lished in the vast oceanic areas. If the deepest abysses have produced in the present instance known genera, it is probable that further research will enlarge the areas to which the new genera are limited. It is also noteworthy in connection with the Annelids obtained from the profound abysses, that the majority are tube-dwellers, and thus are more or less protected, though at the same time this fact renders their presence in the trawl or dredge more frequent.
10 fathoms and under.
Euphrosyne capensis.
Amphinome rostrata (surface). Hermodice carunculata.
Eurythoe pacifica.
var. levukaensis.
Hipponoe gaudichaudi (surface). Hermione hystrix.
Lepidonotus wahlhergi.
cristatus.
Eunoa capensis.
Polynoe pustulata.
attenuata.
Eupompe australiensis.
Leanira IcBvis.
Nephthys atlantica.
Phyllodoce sancti-vincenfis. Eulalia capensis.
Alciopa antarctiea. j
quadrioculata. \
Alciopa sp. surface.
Halodora reynaudii. j
Greeffia oahuensis. J
Hesione assimilis.
Hesione sp.
Nereis melanocephcda.
Notocirrus eapensis.
Palolo virid.is.
Neinatonereis sp.
Eunice torresiensis. vittata, var. eirrobranchiata. aphroditois. torquata. barvicensis. tribrancliiata.
Eone trijida.
GlycerOj lamelliformis.
Aricia platycepliala.
Trophonia cape7isis.
Brada whiteavesii.
Cirratulus capensis.
Nieoinache capensis.
Sabellaria jolinstoni.
capensis.
Terebella crassicornis.
Sehmardanella pterochcefa.
Thelepus sp.
Sabella bipunetata. fusca.
Dasychone baWdi.
oidentalis.
wyvillei.
nigro-maculata.
violacea.
XXXll
THE VOYAGE OF H.M.S. CHALLENGEK.
10 to 50
Chloeia Jiava. fusca.
Notopygos megcdops.
Aphrodita australis.
Palmyra aurifera.
Lepidonotus gymnonotus.
Lagisca tenuisetis. per acuta.
Evarne herguelensis. tenuisetis.
Polynoe platycirrata. ocellata.
Thalenessa oculata.
d igitata. fi.mhriata.
Leanira japanensis.
Hesione pacifica.
Autolyt us maclearan i is.
Nereis hrasiliensis.
Jcohiensis.
Hydroides i 50 to 10<
Euphrosyne borealis.
Notopygos labiatus.
Lagisca yokolia mie nsis.
Polynoe liexactineUida.
Harmoth oe hcdiaeti.
Eupolynoe mollis.
Hermadion he i 'guelensis.
Polynoe euplectellce. capensis. ipliionoides.
Eupholoe philippensis.
Nephthys trissophyllus (and 20 fathoms).
Syllis capensis.
fathoms.
Nereis tongatabuensis. eato7ii.
Staurocephalus a ustraliensis. LumbriconeiNs hetei'opoda. Eunice hobiensis. murrayi. equibranchiata. bassensis.
Gl ycera amboinensis. Ammotrypane gracile.
Travis ia herguelensis.
Brad a man imilla ta . Phylloclicetopterus claparedii. Dasybranclius sp.
Sabellarm giardi.
Terebella seticornis.
Pista fasciata.
A rtacania ch allengei ia. Dasyclione picta.
japo7iica.
ultispmosa.
fathoms.
Syllis gigantea.
Eusyllis tubifex.
Exogone heterosetosa.
Nereis pelagica.
hergueleiisis. p>atagonica. Notocirmis capensis.
Eunice mandanavensis. Nothria ivil lem oesii.
Glycera tessellata. capitata.
Scalihagma inflatum. Ranzania capensis. Prionospiio cap>ensis.
EEPORT ON THE ANNELIDA.
xxxiii
50 to 100 fathoms — continued.
Cirratidus zehuensis.
Pmxilla ca'pensis.
Phyllocomus croceus.
Amphitrite hei'guelensis.
Thelepus cincinnatus, var. cana- densis.
Neottis antarctica.
Artacama zehuensis. Potamilla torelli.
Sahella zehuensis. Branchiomma vesieulosum. Protula americana. Serpida 7iarconensis. Ditrypa arietina.
100 to 200 fathoms.
Aphrodita eehidna.
Lcetmonice producta.
Euphione elisahethcB.
Eunoa mindanavensis.
Lagisca antarctica.
magellanica.
darwini.
Phyllodoce duplex.
Salvatoria kerguelensis.
Sphcerosyllis kerguelensis.
Syllis ramosa.
Lumhriconereis kerguelensis.
Eunice edwardsi.
pycnohrancJiiata.
Glycera kerguelensis.
Scoloplos kerguelensis.
Trophonia kerguelarum.
Spirorhis sp.
Scolecolepis cirrata.
Praxilla kollikeri.
kerguelensis.
assimilis.
Ampharete kerguelensis.
Terehella gruhei.
jiahellum (and to 50 fathoms).
Ereutho kerguelensis.
Polycirrus kerguelensis.
Terehellides strcemi, var.
Dasychone capensis.
Protula capensis.
Apomatus elisahethcB.
Serpula narconensis, var. magel- lanica.
Pomatocerus strigiceps.
200 to 500 fathoms.
Lcetmonice japonica. Aphrodita intermedia. Iphionella cimex.
Lepidonotus squamatus. Eunoa iphionoides. opalina. yedoensis.
Polyeunoa levis.
(ZOOL. CHALL. EXP. PART XXXIV. 1885.')
Lagisca magellanica, var. murrayi
var. gruhei.
Allmaniella setuhalensis. Scalisetosus ceramensis.
Eulepis wyvillei.
challengerice.
Sigalion huskii.
Psammolyee occidentalis.
Lie
XXXIV
THE VOYAGE OF H.M.S. CHALLENGEE.
200 to 500 fathoms — continued.
Leanira magellanica. areolata.
Ne'phthys malmgreni.
GenetylUs oculata.
Syllis setubalensis. brasiliensis.
Nematonereis schmardce. Lumbriconereis bifurcata.
japonica.
Eunice magellanica.
prognatha.
Macduffia bonhardi.
Nothria sombj^eriana, and var.
macrobrancliiata.
Hemipodus magellanicus.
Spirobranchus
Scolecolepis cirrata, var. ChcBtozone atlantica. Notomastus grubei. Maldane sarsi. Nicomache japonica. Maldane atlantica. Praxilla. challengerice. Ampharete sombreriana. Amphicteis japonica. Melinna macidata. Euthelepus setubalensis. Terebellides ehlersi. Protula lusitanica. Serpula sombreriana. Placostegus assimilis. occidentalis.
500 to 600 fathoms.
Aphrodita aculeata. Lostmonice jilicornis.
aphroditoides. Dasylepis equitis.
Lagisca propinqua.
kermadecensis. Eulagisca corrientis. Evarne johnstoni.
Lcenilla fusca.
Polynoe magnipalpa. Polynoella levisetosa. Syllis robertiance. Nicidon balfouriana. Nothria quadricuspis. Hyalinoecia tubicola, var. Terebella kermadecensis. Pista mirabilis. corrientis.
Sabella assimilis.
600 to
Polynoe mirabilis.
Leanira hystricis.
Staurocephalus atlanticus. Lumbriconereis neo-zealanice. Nothria tenuisetis.
conchylega.
minuta.
1000 fathoms.
ChcBtozone pacijica. Praxilla sp.
Terebella sp.
Lanassa sarsi. Thelepus marenzelleri. Ehlersiella atlantica. Terebellides sp.
REPORT ON THE ANNELIDA.
XXXV
Eumenia reticulata. Melinna armandi.
1000 to 1200 fathoms.
I Lecena neo-zealanice.
I langerhansi.
1200 to 1500 fathoms.
Harmothoe henthaliana. Polynoe synophthahna. longipedata.
Nephthys phyllobi^anchia. Laranda longa.
Lumbriconereis punctata.
ehlersi.
Eunice cerstedi (to 83 fathoms). Aricia norvegica.
Chcetozone henthaliana. Notomastus agassizii.
Maldanella antarctica.
neo-zealanim. Nicomache henthaliana. Maldane sp.
Praxilla occidentalis. Myriochele heeri. Amphicteis gunner i. Thelepus sp.
Terehellides stroemi. Serpida philippensis. Vermilia sp.
1500 to 2000 fathoms.
Chloenea atlantica.
Lcetmonice producta, var. ivyvillei (and to 50 fathoms).
Lcetmonice producta, var. wille- moesii (to 700 fathoms).
Lagisca moseleyi.
crosetensis.
Dalhousia atlantica.
Nereis longisetis.
Nothria ahranchiata. armandi.
Nothria henthaliana.
Ephesia antarctica.
Trophonia wyvillei.
Mcddane malmgreni.
Praxilla ahyssorum.
Petta assimilis.
Amphicteis sarsi.
Gruhianella antarctica, and var. Melinnopsis atlantica.
Pista ahyssicola.
Lecena antarctica.
2000 to 2500 fathoms.
Lumhriconereis ahyssorum. Nothria pycnohranchiata. ehlersi.
Maldanella valparaisiensis.
Samythopsis gruhei. Eusamytha pacijica. Melinna pacijica. Euthelepus chilensis.
Placostegus morchii.
xxxvi
THE VOYAGE OP OHALLENGEE.
2500 to 3000 fathoms.
LcBtmanioe producta^ var, hen- thaliana (to 1600 fathoms).
Eunoa abyssorum.
Polynoe ascidioides.
Buskiella abysmrum (to 1850 fathoms).
Myviochele heeru
Serpula sp.
Myriochele pacijica.
Amphicteis gunneri, var. atlantica.
sarsi.
Eupista darwini (to 2225 fathoms). grubei.
Lanobssa benthaliana^
Placostegus ornatus..
Lemia abyssorum^
3125 fathoms.
I Placostegus henthalianus^
: DESCRIPTION OF GENERA AND SPECIES.
Family EuPHEOSYisrrD^.
The Euphrosynidse, as the first group of his Amphinomea, have been so clearly indicated by Ehlers/ that at present they stand in no need of further notice. Their short ovoid form — hirsute with the characteristic bristles, rows of dorsal branchise, and the carunculated head— is diagnostic.
The Challenger was fortunate in procuring two species, for as their haunts are often littoral, they would thus seldom come under observation. A single species was found by Kinberg and Schmarda, since the Euphrosyne polyhi'anchia of the latter is undoubtedly synonmous (as Ehlers states) with Kinberg’s Eivphrosyne capensis. Grube gives none in his Annulata (Erstediana, nor in the account of the Annelids of the German exploring ship “ Gazelle,” and only one (Euphrosyne aurifei'a) in Semper s fine series from the Philippines. In the “ Novara ” collection he found the same species as Kinberg and Schmarda. One form also is noted by Marenzeller^ from southern Japan.
Euphrosyne, Savigny.
Euphrosyne capensis, Kinberg (PL II. fig. 5 ; PL I a. figs. 1-3).
Euphrosyne capensis, Kinberg, Ofversigtk. Vetensk.-Akad. ForbandL, d. 14 Januari 1857, Stockh., 1858, p. 14; and Fregatten Eugenies Resa, 1858, taf. 12, fig. 14.
Euphrosyne polyhranchia, Schmarda, Keue wirbeUose Thiere, vol. i. part 2, 1861, p. 136, taf. 32, figs. 264-287.
Body rather more than usually elongate, somewhat fusiform, and of a reddish-orange colour. The caruncle, consisting of an elevated keel and a flat band, reaches to the eighth segment, and a short and somewhat stout conical tentacle exists in front of the ridge, with a well-marked eye at each side of the latter anteriorly. A pair of much more minute and in some nearly confluent eyes occurs on the ventral surface of the narrow cephalic ridge. The arborescent branchise are purplish, with the elliptical or sub-oval tips pale reddish, and each complete row generally consists of eleven trunks. The bifid dorsal bristles are of
^ Die Borstenwiinner, i. p. 64 et. seq.
2 Denhschr. d. h. Akad. d. TFiss. Wien, Bd. xli., 1879, p. 2 (sep. Abd.).
(ZOOL. CHALL. EXP. — PART. XXXIV. — 1885.)
LI. 1
2
THE VOYAGE OF H.M.S. CHALLENGED.
two kinds — smootli and crenated in the fissure, while the bifid ventral bristles are all smooth. There are about fifty-seven segments. Length, 45 mm. ; breadth, 12 mm.
Habitat. — Under stones between tide-marks at Sea Point, near Cape Tovm, South Africa, November 1873.
The species seems to be tolerably abundant at the Cape, and it is noteworthy that it, was the only one procured by Kinberg and Schmarda in their well-known travels. It is evident that both refer to the same form, as Prof. Ehlers^ more than susj)ected, and consequently Kinberg’s name has the priority. It belongs to the first group of Ehlers, viz., those having the sides of the segments furnished with both cirri and branchiae, all the latter, moreover, being ramose. Grube describes the same species from St. Paul, in his account of the Annelids of the “Novara” Expedition.^
The body of Ewphrosyne capensis is proportionally longer, more finely spinose and more flattened than that of the British Euphrosyne foliosa. Kinberg observes that the caruncle is semiglobose anteriorly — a feature that has not been seen in these examples. Schmarda, again, describes and figures seven tentacles (Ftihler) in connection with the caruncle, but such apparently is a misapprehension. None sho'wed any trace of the rudi- mentary clavate tentacles observed by Ehlers in his Euphrosyne racemosa. In front of the puckered opening of the mouth are the two kidney-shaped pads, separated by a deep median fissure. These pads are probably of some importance as pivots during the complex actions of the buccal apparatus. A typical segment of the body consists of a convex dorsal and a flattened ventral arch. The former is bare in the middle line, but has laterally a superior division carrying a dense series of bristles and a posterior row of the branchiae, with a cirrus at the dorsal margin and another midway between the bristles and the branchiae. The latter is the longer and shows a slight constriction in the middle, but neither tapers much. The inferior division, again, presents a tuft of bristles, and interiorly and posteriorly a cirrus, which is generally more slender at the tip than either of the fore- going. All the cirri are shorter and stouter than in Eiphrosyne foliosa. The dorsal row of bristles is comparatively short when contrasted with Euphrosyne borealis, and even less boldly marked than in Euphrosyne foliosa ; while their apertures in the thick cuticle appear as if punched out. Kinberg’s figure is a very fair, though not quite accurate, representation of the serrated kind, while Schmarda’s deviates still further from nature. The curve of the tip (PI. Ia. fig. l) is less pronounced than in Euphrosyne foliosa, and the disproportion between the processes better marked. When viewed antero-posteriorly, the notches on the limbs of the fork are observed to be due to transverse grooves. At the dorsal edge as well as in the centre of the row many with a smooth fork occur (PL Ia. fig. 2), the long process being much attenuated. All the smooth bristles project considerably beyond the
1 Die Borstenwiirmer, i., 1864, p. 65.
2 Annel. Novara Expect, p. 6 (sep. Abd.).
REPORT ON THE ANNELIDA.
serrated kind. M. Claparkle^ has drawn attention to the special characters of the bristles in JEuphrosyne, viz., their extreme brittleness, their tubular structure, calcareous nature, and the entrance of air into the interior. The same features have been noticed in Chloeia.^ Schmarda also mentions the occurrence of a yellow fluid in the central canal of the present form. Nothing has been seen to verify his statement that a network of slightly curved bristles, pointed at both ends (fusiform-acerate, Bowerbank), exists at the lower border of the branchiaj. His figures very much resemble sponge-spicules. The ventral bristles (PL Ia. fig. 3) are considerably larger than the dorsal, and are terminated by a shghtly curved blunt tip, with a conical spike at the base. The internal canals from the processes join after a short course downward, and there is a slight dilatation of the cavity opposite the enlargement at the upper third of the shaft.®
The dorsal region of each segment is further supplied with a densely ramose series of branchial processes, which are eleven in number in the typical segments. Schmarda says there are twelve, but such a discrepancy is of little importance. Each is dicho- tomously branched, and the tips end in a series of elliptical or sub-oval bodies somewhat like those in Euphrosyne foliosa. The superior are the longer. So far asTan be observed in sections of the organs, the view of M. Claparede — that these structures are entirely devoid of an axial cavity, if we exclude from this term the blood-vessels — seems to be correct, especially as regards the distal branches. This author, indeed, thought that the general surface of the body exercised the function of respiration. On the other hand, it requires very little manipulation to trace the large blood-vessels from the body-cavity into the branchise and follow their branches up to (but not into) the tips of the organs. In sections of the base of the trunks a complex series of muscular fibres appear, and the cuticle and hypoderm of the entire structure are dense except distally, where the former becomes very thin.
In the structure of the body-wall this form agrees with its congeners. The nerve- cords have superiorly a firm investment which is continuous from side to side. In the hollow between them superiorly is a fascicle of muscular fibres, and below them a central granular structure. Moveover, the cords are united by a streaked isthmus interiorly. The anatomy of one region of this form is especially interesting, viz., that of the buccal apparatus. The anterior part of the structure consists of a cylindrical protrusible proboscis densely covered with cuticle. The centre of the latter in front is occupied by a large muscular and vascular, but chiefly glandular mass, the inner lining of which is throvm into bold longitudinal rugae. The vast collection of glands in this organ would indicate
1 Anu. Chetop. clu Golfe de Naples, p. 109.
^ “ Porcupine ” Annelids, Trans. Zool. Soc. Lond., vol. ix. p. 396.
® A remarkable and apparently new form in the British Museum shows the peculiarity of having along with the ordinary forked ventral bristles a dense series of long slender dotted hairs tapering to a tip which is slightly birlbous. The dorsal serrated forms are very boldly marked, and the smooth ones are long. The branchise are much branched, and the tips somewhat lanceolate. The species is fully an inch in length, and comes fioin the Pascadores Islands — Consul Swinhoe’s Collection (70, 6, 18, 13).
4
THE VOYAGE OF H.M.S. CHALLENGER.
important secerning functions. Behind is a remarkable lingual organ, which in longi- tudinal section presents much resemblance to the tongue in mammals. It is attached at the base posteriorly to a dense glistening muscular mass — separated by a layer of horizontal fibres from the complex series of radiating muscles which diverge upward to the periphery of the organ. In vertical longitudinal sections in the middle fine these fibres are separated posteriorly, by septa passing downward from the thick cuticular (chitinous) coat, into spaces that enlarge as we proceed anteriorly. In such sections most of the fibres in the posterior half are more or less vertical, though it is evident they are closely interwoven. The muscles at the tip of the tongue arch backward over the posterior part of the preceding glandular region (which forms a highly vascular cushion), and pass downward to the front of the dense glistening basal mass formerly aUuded to — the curve of the arch becoming less and less till the fibres are nearly vertical behind the cushion, or at the commencement of the smooth and dense cuticular region of the tongue. The chitinous cuticular surface of the organ is so largely developed at this part that it is as thick as the hypodermic layer. In this region also there are at least three strong plates of transverse muscular fibres extending from summit to base. If the section deviate to the lateral regions, the decussation of the fibres is extremely complex. In transverse sections, again, near the tip of the tongue, the thickness of the hypoderm on the rugae of the prelingual mass as well as in the lingual papillm is considerable. Moreover, it is apparent that the entire diameter at the base of the prelingual mass is formed on the same plan, the central region (lingua proper) being, however, most muscular. When the section passes through the thick mass of the tongue an inextricable series of crossed and interwoven fibres occurs, the base assuming a somewhat stratified appearance, since the numerous blood-vessels are situated at intervals in regular horizontal rows. Nothing can exceed the complexity of the muscular structure of the organ, which is thus eminently calculated to subserve very important functions. The tip of the tongue is highly vascular ; indeed, the organ throughout is largely supplied with blood-vessels. The dorsum of the free tip is covered by a series of elevated papillae — truncated at the tip, and with one angle pointed — composed of processes of the hypoderm sheathed in cuticle. Schmarda describes a series of small blackish horny teeth as situated on the ridges of the vault of the proboscis, but, in the present instance, nothing further than the hypodermic longi- tudinal furrows covered with cuticle have been observed. Elders found simdar ridges in Euphrosyne racemosa, but in the latter as well as in the Euphrosyne foliosa of Britain the parts are less developed than in the large African form.
Behind the tongue the alimentary region presents on its ventral aspect a large and a small fold, the former in longitudinal section often having a broad summit applied to the vault of the canal and a narrow base so that the whole resembles a pedicled mass, but it varies according to the condition of the spirit-preparations. This region is furnished with numerous transverse folds, so deep in some cases as to
REPORT ON THE ANNELIDA.
o
merit the name of lamellae (like those in the ruminant’s “ manyplies ”). The canal then bends upward (from the posterior pit) and forms a kind of pylorus, with thick walls and bold longitudinal furrows, and terminates in the intestine, into which the part freely projects like a long os uteri into a vagina. The thick rugose intestine, which agrees in structure with that in the ordinary Annelids, terminates in a dorsal anus, with two styloid processes, of the usual shape, at the sides interiorly. None of the specimens presented developed generative organs, which in the British examples from Herm are in full maturity in August. Claparede mentions that they originate at the bases of the feet.
So far as structure shows, the animal would appear to evert its lingual organ and apply it with a rasping action to the surface of the sponge, the successive pieces being crushed against the firm rugose wall behind by the hardened posterior surface of the tongue, and again subjected to the action of the deeply folded or lamellar region behind it. Thus, before the (especially soft tissues like those of sponges) food reaches the intestinal surface, it has been partially disintegrated and fitted for absorption. Minute shells of Spirorhis are of course less affected, though many are broken into fragments, and all are in a condition to give up their contents. Like the British Euphrosyne foliosa at Herm, the South African form would seem to browse on the patches of Halichondria and other sponges which abound under stones between tide-marks ; and there is little wonder therefore that there should be so many interesting homologies between its buccal apparatus and that of such Nudibranchs as Doris tuberculata and Doris johnstoni, which follow precisely similar habits.
Euphrosyne borealis, (Ersted (PI. I. figs. 2, 3 ; PI. Ia. figs. 4-6).
Euplirosyna horealis, CErsted, Gronl. Ann. Dorsibrancli., p. 18, Tab. II. figs. 23-27.
Euphrosyne horealis, Auctorum.
Habitat. — A small example apparently referable to the foregoing occurred at Station 49 (off Halifax), May 20, 1873, lat. 43° 3' N., long. 63° 39' W.; depth, 85 fathoms; bottom temperature 35°’0, surface 40°‘5 ; gravel and stones.
Observations, — The specimen agrees in external appearance with the Norwegian form — the smooth dorsal bristles projecting far beyond the branchise, and giving the back a very hirsute appearance. The serrated bifid dorsal bristles (PL Ia. fig. 4) are strongly marked ; and it is to be observed that when viewed antero-posteriorly they are also bent laterally. The ventral bristles present rather an elongated tip beyond the spur, and the edge above the latter is often marked by a series of irregularly placed serrations — two varieties being shown in PL Ia. figs. 5, 6. Each of the processes at the tip has its internal canal. Some of the stronger bristles have the terminal portion less bent than
6
THE VOYAGE OF H.M.S. CHALLENGEPv.
those represented ; and all the characters are better marked in the large examples from Greenland. The latter very well bear out Oersted’s original remark — to the effect that the branchiae are two or three lobed. In the present specimen, however, the organ next the dorsal median line is sometimes quadripartite. GErsted also observes that there is no dorsal cirrus, but near the base of the latter branchial process is an organ which seems to be the homologue of the cirrus in the other forms, and which he had overlooked. This species is entered in the American fauna by Stimpson,’ who states that it is not uncommon on muddy bottoms. It is not mentioned by Prof. Verrill in his dredgings on the coast of New England ; and no example was obtained by Mr. Whiteaves in the Gulf of St. Lawrence, though the other forms procured at this Station closely resembled those from the latter region. In this form the nerve-cords seem to be placed farther inwards than in Euphrosyne foliosa, or than in Euplirosyne capensis. The layers of the body- wall — cuticular, hypodermic, and muscular — are largely developed. The proboscis has a well-marked chitinous layer (usually thrown into numerous wrinkles in the preparations) just within its external investment in situ. In his excellent brochure on the Annelids of Denmark,^ Tauber includes this species, but he labours under a misunderstanding in comprising Hipponoe jeffreysii as a synonym.
Family Amphinomida:.
Few representatives of this family were procured by the Challenger. In Grube’s Annulata CErstediana three are noticed. Sixteen are mentioned by Kinberg, but it is doubtful how many of these came from the voyage of the frigate “Eugenie.” Four occur in Grube’s list from the ship “ Gazelle ” ; while five are included in Semper’s PhilqDpine series of the same author. Schmarda describes nine. The moderate number obtained by the Challenger probably depended on the prevalence of the common kinds amongst the shore rather than the deep-sea fauna, though the occurrence of others as surface- forms doubtless increased the list. They are chiefly southern, though the patient and successful labours of the elder Sars and those of his distinguished son have added two species to the northern fauna, viz., Euryihoe horealis Parampliinome pulcliella,^ both ranging to the British shores. Moreover, both forms occur occasionally in deep water (about 600 fathoms) while extending to comparatively shallow water, or, in the case of Euryihoe horealis, to the tidal region. Those procured by the Challenger with one exception do not pass to a greater depth than 100 fathoms, several inhabiting the littoral region. The exception, however, is marked, for Chloenea atlantica, n. sp., descends to no less a depth than 1525 fathoms.
1 Marine Invertebrata of Grand Manan, p. 36. ^ Annulata Danica, Kjobenhavn, 1879, p. 78.
3 On some Eemarbable Forms of Animal Life, &c. i., 1872, p. 45. This species was probably first described by the author from specimens procured off Shetland by Dr. Gwyn J effreys, vide Trans. Boy. Soc. Edin., vol. xxv. p. 406.
REPORT ON THE ANNELIDA.
7
Tlie members of the family have thus a varied distribution, ranging from the surface to the bottom, and from great depths to the littoral region. It has to be noted, however, that the species found at the surface have generally been on floating timber or other buoyant substances.
The Chloeia group is at present in a somewhat unsatisfactory state, even the number of eyes having been for a long time erroneously described; indeed it is only about twelve years ago that Kinberg and W. Baird made this correction. The head must thus have been superflcially examined by Savigny, Lamarck, Audouin and M. -Edwards, Eisso, Grube, Schmarda and De Quatrefages. Even in the work of the last-named author a form called Aristenia conspurcta, Sav., is admitted amongst the Amphinomacese of uncertain seat. Now the figures in Savigny’s plate (pi. ii. figs. 4^-E) clearly show that a Trophonia from the Eed Sea has been represented, and that probably from bad keeping or otherwise certain changes had caused the artist to represent branchiae. The actual structure of the dorsal and ventral bristles of the forms described up to the present date is shrouded in considerable obscurity, and though Kinberg’s plates partly improve this condition, they also show that more yet remains to be done.
The bristles throughout the Amphinomaceae have certain characters in common, viz., brittleness, calcareous composition, oleaginous contents often forming crystalline aggre- gations, and the well-marked tubular structure. In the Mediterranean Chloeia from the “Porcupine” the unusual form of a dorsal bristle with a simple tip (PI. IIa. fig. 7) occurs, but it is interesting to observe that there is a distinct bend (on the right in the figure) a little below the serrations, and that in certain examples which have no evident serrations there is a rudimentary indication of the spur.
In regard to the structure of the bristles, again, the examples of the Chloeia group that have occurred in this collection and in the “ Porcupine ” range themselves under six heads. (1) The Mediterranean Chloeia from the “Porcupine,” which has its dorsal bristles simple and serrate, and the ventral very slightly bifid. (2) Those from East Indian and Arafura Seas [Chloeia jiavoL), in which the dorsal bristles are boldly serrated and bifid (small spur), and the ventral more distinctly bifid than the former. (3) Chloeia fusca, from the region near the Moluccas, which has bifid bristles of three kinds, viz., (c<) very slender and attenuate, (6) with stout, short tips, and (c) with longer tips, serrated externally, while the ventral bristles are bifid and smooth. (4) In this and the two following a second dorsal or branchial cirrus is present, and the bifid dorsal bristles are of two kinds — (a) boldly bifid and smooth, and (6) bifid and serrated externally on the long limb. The ventral bristles are bifid and serrated internally on the long limb. The branchiae are branched. It comes from the Atlantic (Canaries). (5) From the Bermudas. In this both dorsal and ventral bristles are bifid and serrated only on the inner side of the long limb, and the tips of both are short. (6) From the Phfiippine Islands. The dorsal bristles are very strong, with smooth bifid tips. The ventral are
8
THE VOYAGE OF H.M.S. CHALLENGER.
similarly shaped, but with two serrations on the inner side of the long limb. Tbe sbgbt differences in tbe characters of tbe last mentioned series correspond with tbe close resemblances of tbe forms generally. Lirione agrees with tbe three last in ba^dng tbe branchial cirrus, but it is much shorter. There is only a single dorsal cirrus (in its usual position behind tbe bristle-tuft) in Amphinome vagans and in Eurythoe pacijica.
Chloeia, Savigny.
Chloeia jiava (Pallas) (PI. III. figs. 1 and 3 ; PI. Ia. figs. 7-9).
Aphrodita flava, Pallas, Miscell. Zool., p. 97, pi. viii. figs. 7-11.
Chloeia flava, Auctorum.
Habitat. — Two examples were trawled at Station 233b (south of Japan), May 26tb, 1875, in lat. 34° 18' N., and long. 133° 35' E., at a depth of 15 fathoms, amongst blue mud, the occurrence of Synapta in the same bottle bearing out the latter statement, though it would rather have been expected elsewhere. Surface temperature, 66° ‘3. The locality of the third specimen (a small one) is doubtfully marked “ Arafura Sea.”
The range of this form is great — from Cape Verde to Amboina, and generally throughout the Indian Ocean and Chinese waters.
Observations.— In his Miscellanea Zoologica, published in 1766, Pallas gives a very good account of the external characters of this form, one example of which Schlosser obtained for him from the shores of Bengal, whence it was fished uj) by the anchor of a Belgian trader, while the other came from Amboyna. Pallas gave it the name of Aphrodita flava, from the fancied resemblance of the colour of the bristles to the locks of Dione, the mother of Aphrodite (Venus). Gmelin then included it in his edition (1788) of Linnmus as Terebella flava.^ Bruguieres next mentioned the same form under the name of Amphinome cajoillata.^ Eisso ® afterwards described a new form [Chloeia rupjestris) from the Mediterranean, a fact which escaped the notice of some of his successors. Cuvier* then joined the two species as Amphinome jaune ou chevelue. Savigny,® again, established the genus Chloeia, in the fourth family of his Amphinomidse, entering the same species as Chloeia capillata — the name given to it by his countryman being chosen rather than the rightful one of Pallas. This author and several others state that the eyes are only two ; but there are really four, two on each side. Audouin and M. -Edwards ® followed the latter, and pointed out the serrated condition of the dorsal bristles, and the bifurcated nature of the ventral bristles. They also only knew of the single form described by Pallas. Grube likewise gave Chloeia only two eyesd and both
1 Loc. cit, p. 3114, No. 7.
2 Encycl. mdtfi., Diet, des vers, i. p. 45, No. 1, pi. lx. figs. 1-5. ^ ]^^t. Eur. merid., iv. p. 425.
^ Diet. d. Sci. Nat., t. ii. p. 71 ; and Eegne Anim., t. ii. p. 527, edit. 3, pi. ix.
® Syst. des Ann61. p. 58. ® Hist. Nat. des Annel., p. 120. ^ Earn, der Anneliden, pp. 40 and 121.
REPORT ON THE ANNELIDA.
9
he and Blainville ^ term the species Chloeia Jiava. Kinherg ^ separates the Chloeia Jlava of De Quatrefages as Thesmia Jlava, since it differs in the structure of the bristles ; and the remarks by Dr. Baird ® on the latter author’s species are important. Thus I agree with him in considering that Chloeia incerta, De Quatrefages, cannot be separated from Chloeia jlava, and that probably an error has crept into his description of the dorsal bristles of the latter. The observation of De Quatrefages that Chloeia furcigera is distinguished by having bifid bristles in both dorsal and ventral series only shows that the true nature of these organs in the group was misunderstood, since all are morphologically bifid. Finally, Prof. Grube describes the occurrence of the present form from Amboina, Salavatti, and Cape Verde, in the collection made by the German frigate “ Gazelle,” as well as in that made in the Philippines by Prof. Semper. The same author points out ^ that his Chloeia ceylonica ® is only a young form of Chloeia Jlava ; while the Chloeia pul- chella of Dr. Baird in the British Museum seems to be the same species. It was procured by H.M.S. “Herald” from Eaine Islet, North Australia. Chloeia Jlava appears, on the whole, to be an eastern type.
The larger of the two Japanese examples is less than 3 inches, and therefore does not come up to the original specimen procured by Pallas ; and this difference in size would appear to correspond with the diminution in the number of segments. Pallas mentions forty segments, whereas M. de Quatrefages says from thirty -five to thirty-seven. In the present case the segments numbered res23ectively thirty-five and thirty-eight.
The body is somewhat fusiform in outline, but the addition of the bristles gives it an ovoid appearance. The head is small, and dorsally almost wholly occupied by the tentacles and caruncle. Anteriorly are four tentacles, the inferior pair being pale in the preparation, widely separated at the base, and considerably shorter than the superior. The latter (which alone are mentioned by Pallas) arise close together in front of the caruncle, and are pale ventrally, but tinted of a rich madder-brown dorsally. Behind
them is the large rugose caruncle, which consists of an elevated, doubly crenate crest and
a wrinkled horizontal portion ; moreover, the middle of the crest is marked by an interrupted brown line. The organ, fixed to the first two segments, extends to the commencement of the fourth, and terminates in a free tapering extremity. Attached in a groove at the anterior end of the caruncle is the long brown median tentacle, which is about twice as long as the pair in front of it : like the rest it is a simple subulate process. In a line with the anterior margin of the latter, or in the smaller form decidedly in front, is a large jDigment-speck, and behind it a smaller, on each side, so that, as mentioned in the Mediterranean Chloeia,^ four is the correct number of eyes, and not two, as mentioned by Savigny and many of his successors. Kinberg ^ noticed the error,
^ Diet. d. Sci. Nat., art. “Vers.” 2 Qfversigt k. Vetensk.-Akad. Forhandl., 1867, p. 86.
® Journ. Linn. Soc. Land. (Zool.), vol. x. p. 230. ^ Annel. Fauna d. Philippinen, p. 10.
® Proc. Zool. Soc. Lond., 1874, p. 326. ® Trans. Zool. Soc. Land., vol. ix. p. 396.
7 Freg. Eugen. Resa, pi. xi. fig. 1.
(zoou CHALL. EXP. — PART XXXIV. — 1885.)
LI 2
10
THE VOYAGE OF FI.M.S. CHALLENGER.
and rightly gave his figure four eyes ; and Dr. Baird ^ likewise observed the true condition in several species.
When closed, the mouth presents a simple puckered orifice with two prominent fleshy lobes running to the anterior border of the head, and which are probably the homologues of the same parts in Euphrosyne. They are not merged into the gape during the extrusion of the proboscis, but form two sides of a special triangular fold superiorly. The
posterior border of the mouth occurs at the margin of the fourth segment. The
extruded proboscis probably agrees with that of the Mediterranean form in being in three tiers, viz., a membranous basal portion, a denser median, and a firm cartilaginous distal part in the form of two lateral lips.
Each of the typical segments of the body bears a branchial tuft, a well-marked dorsal papilla with a fascicle of bristles, and immediately behind a long madder-brown cirrus upon a pale basal portion, while the ventral division has a very long tuft of bristles, and a shorter pale cirrus placed inferiorly and posteriorly. The first three body-segments possess no branchial tuft,' but, as mentioned by Pallas, have a second small pale cirrus at the base of the dorsal one.
The dorsal bristles spring’ in a radiating fan-like manner from the papilla, and consist of long tapering organs with yellow tips and a secondary spur at the base (PI. Ia. fig. 7, which represents one from the middle of the dorsum). The distal end is pointed — smooth on the side with the basal spur (absent in Dr. Baird’s Cliloeia tumida from India), but on the opposite edge furnished with from twelve to twenty large recurved fangs, besides having the tip beyond the latter slightly serrated. Kinberg’s artist does not clearly define the spur.^ In the preparations the yellow pigment is most intense just above the latter, and this colour is very characteristic in glancing at the bristles en masse in spirit-preparations, especially when contrasted with Notopygos and allied forms. These bristles have a slight curve, and terminate inferiorly in a slender and almost pointed extremity, so that they are fusiform. The anterior dorsal bristles differ in having in some cases smooth tips and a longer basal spur (PI. Ia. fig. 8), while in others there are very few recurved fangs. The mere number of the latter is therefore immaterial. The bristle-papilla to which the dorsal fascicle is attached is peculiar in rising out of a kind of pit in the adjoining skin, which forms a free margin all round, with the exception of the posterior third. The whole tuft can thus be readily moved.
The ventral bristles consist of long yellowish structures having tips like that repre- sented in PI. Ia. fig. 9. They terminate inferiorly in long tapering roots, which appear to be solid for a considerable distance. There is little difference between the anterior and posterior forms, except that the former are more slender. It will thus be apparent that all the bristles of this form are modifications of a single type.
The dorsal cirrus is a simple subulate process springing from the bristle-paj^illa
^ Journ. Linn. Soc. Land., vol. x. p. 230. ^ Qp p]_ xi. fig. 1, G.
EEPORT ON THE ANNELIDA.
11
towards its posterior border and having a basal segment and an elongated terminal region, which is purplish-brown throughout, wdiereas the former region is less deeply coloured, with a pale band in front. From the base of the process a fan-shaped patch of pigment extends downward and inward on the bristle-papilla. The ventral cirrus is quite pale, and lies below and rather behind the ventral tuft of bristles. A slight enlargement of the base occurs anteriorly. The cirrus is filiform, tapering, and considerably shorter than the dorsal.
About thirty-two pairs of arborescent branchise are ranged along the dorsum, the number var5dng according to the length of the specimen. Each is situated at the posterior border of the segment, a short distance from the cirrus, and slightly overlapped by the radiating dorsal bristles. Their shape is dendritic ; a central stem rising from the dorsum and giving off, first, a large external branch, which generally divides into two, and these give off others, which bear the ultimate pinnules. The other branches of the main stem pass off transversely (nearly opposite in arrangement) and subdivide into pinnules. Those at the tip are simple, i.e., have no subdivisions. In the ordinary condition the branchiae fold backward, and since the pinnules are all grouped posteriorly they are thus in the most protected position. In minute structure the stem' consists externally of a firm cuticular investment, a considerable layer of hypoderm, a -central area of muscular and connective tissue, and the two main vessels (afferent and efferent). Each of the ultimate pinnules also carries its two blood-vessels in the midst of a considerable amount of hypoderm. The muscular fibres in the centre of the main stem are complex- longitudinal and radiating as well as transverse. If a longitudinal section of any of the larger branches be made, the tranverse fibres form a series of septa.
The branchise continue to the penultimate segment, but diminish in size posteriorly. They are marked by a deposit of pigment, best seen in small specimens, along the main branches.
The dorsum of the animal has a very conspicuous row of brownish-purple spots, which occupy the posterior two-thirds of each segment, in the middle line. A brownish band also occurs at the anterior part of each segment, from the branchise outward. The anus is dorsal, with two elongated cirri behind it.
When withdrawn the proboscidian region presents three layers — an outer membranous, a denser middle portion, and an inner division of cartilaginous consistence. The latter forms two longitudinal folds, which in extrusion become transverse. In the quiescent state of the parts {i.e., when withdmwn) in the Mediterranean Chloeia procured during the “Porcupine” Expedition, the lining tissue of the buccal cavity forms some simple folds superiorly, above the two muscular external lips. These folds gradually increase in complexity, and when the lips disappear they form a series of vertical and horizontal papillae. The latter are covered by the usual dense cuticle, with h}q)oderm beneath, and followed by a complex muscular layer varying according to the situation examined
12
THE YOYAGE OF H.M.S. CHALLENGEE.
Immediately behind the mouth-slit a continuous ring of the following structure is formed by the buccal wall : — Superiorly are various leaf-like glandular papillae, succeeded on each side by an inferior dense margin composed of the two muscular processes which become terminal in extrusion. A variously folded papillose region occurs beneath and completes the ring. The chief muscles are massed at the sides of the dense superior folds. The great ridges increase in size and muscularity as we proceed backward, and the wall beneath becomes more compact. The muscular fibres moving the dense lips are evidently important, and are probably the chief agents in the biting movements which the part performs in alimentation. The fibres form a complex transverse and longitudinal meshwork, the free surface of the part having a glandular structure. Passing inwards, another fold then occurs externally on each side, and the superior papillae occupy a larger space, while the region next them has its surface covered with pointed papillae, and the dense portion is confined to the ventral part of the massive plait. The wall of the buccal organ presents the following minute structure ; — Below the dorsal thin glandular portion the massive lateral wall is formed by an interwoven series of transverse and longitudinal fibres, the conical papillae being on the inner surface. Beneath is a dense fold of the hard pad, which in section has the inner part of its area filled with longitudinal fibres. The secondary fold below is now made up of interwoven longitudinal and oblique fibres, and this and the former plait are bound to the dense outer wall of the organ by strong muscular bands. Instead, therefore, of forming the upper part of the lateral wall, the dense portion now constitutes the lower, the two accessory folds (enveloped in a firm layer) meeting in the middle line beneath, while the papillae have disappeared from the wall outside them. Immediately behind, the fold on each side unites with its fellow, the larger superior mass being formed chiefly of longitudinal fibres within the glandular layer, and the inferior of a complex interlacement of longitudinal and transverse fibres, the latter forming spaces for the former. The upper lateral wall is boldly papillose, while the dorsal margin has a row of blunt papillse. The lower part of the organ gradually merges into the upper, with similar radiating and circular fibres, the whole (with the exception of the dorsal arc) forming a dense muscular tube. Beneath is a great retractor of longi- tudinal and vertical fibres. Internally the long papillse of the lateral surface are continued over the ventral curve. The outer wall of the rounded tube is composed of a well-marked series of longitudinal fibres, which gradually increase in thickness from the dorsal to the ventral median line. The papillse on the inner surface then have a tendency to form ridges, and the thin glandular wall is greatly extended dorsally; while the canal diminishes in calibre, assuming a uniform outline, the inner surface being covered with the glandular rugse and papillse resting on a coat of circular muscular fibres — externally bounded by the longitudinal layer.
The whole muscular mechanism of the organ is suited for performing the functions of a pair of flesh}^ biting pads, ’for the extrusion and retraction of these in varying degrees.
REPORT ON THE ANNELIDA.
13
and probably also for their employment as crushing surfaces in the passage of the food inwards in the ordinary state of the parts. The arrangement of the fibres in the thickened retractor would further assist in the expulsion of the organ, as in Magelona.
In extrusion the basal or membranous portion of the organ is formed by a plait continuous with the lips, which passes inward to the posterior retractor muscles, so as to join the denser inner fold of the proboscis ; the middle region is formed by the latter, and the distal by the denser yellowish part, the two divisions of which, by the nature of their connections, are directed vertically in the extruded condition.
The food of this species probably consists of the same nature as that of Euphrosyne. The cardiac opening of the stomach, in the retracted condition of the proboscis, projects into the organ on a similar long rugose eminence. The inner surface of the stomach is remarkably ridged and glandular, while in transverse section the intestine presents a closely arranged series of glandular lamellae.
In vertical section the caruncle is found to be composed chiefly of hypodermic tissue enveloped in cuticle. In the centre a strong vertical muscular band proceeds from the body-wall to the crest, and numerous longitudinal fibres appear at the base (in the peduncle). In front a deep indentation exists on each side a little above the base, surmounted by numerous narrow folds. The latter occur from base to summit posteriorly, and have the aspect of a single series of zig-zag folds on each side. The madder-brown pigment is deeply imbedded, and close to the central vertical muscular septum. In front a series of fibres proceed from the latter along the median tentacle, and similar bands occur in the other cirri. The anterior part of the caruncle lies over the cephalic ganglion, but the function of the organ is doubtful.
In regard to the structure of the body-wall, the cuticle is comparatively dense, the hypoderm somewhat less developed, while the circular, longitudinal, and other muscles are strong. The nerve-cords are somewhat small and flattened, being placed in an area bounded internally by a transverse band of fibres, and externally by the circular muscular layer and hypoderm. The oblique muscles are attached at the outer border of each trunk.
Dr. Baird’s Chloeia tumida seems to be rather like this form, but devoid of spots on the dorsum. One collected by H.M.S. “Alert” off Torres Straits Islands had taken a hook, which is now firmly grasped by the proboscis of the blackened preparation.
A small specimen measuring about an inch in length is marked with doubt as a native of the Arafura Sea. The segments amount to about thirty, exclusive of head and tail. The bristles agree in structure with the foregoing, allowing for the difference in size. The branchiae are less branched, a condition probably due to the latter. The coloration and other features correspond.
14
THE VOYAGE OE H.M.S. CHALLEHGEK.
Chloeia fasca, n. sp. (PL II. figs. 1 and 2 ; PI. Ia. figs. 14, 15 ; PI. IIa. figs. 1, 2).
Habitat. — Dredged near Banda (a Station ofi* the Moluccas), 1st October 1874.
The entire body of this species is iridescent dusky brown, with a well-marked pale median dorsal strijDe from the caruncle to the tail. This pale band is rendered very distinct by a darker brown belt on each side, so that there are really three stripes on the dorsum. The body is about 18 mm. long, and consists of twenty-three segments. The latter are broader from before backwards than in Chloeia Jlava, and on the whole are more evidently marked.
The head is well-defined, and bears posteriorly a caruncle, which extends to the fifth body-segment. The folds of the organ are more lax than in Chloeia Jlava ; indeed they form a series of distinct vertical lamellae, which are easily separated externally down to the ventral fold. The four eyes are distinct, two being at the anterior border and two toward the posterior border of the head in front of the caruncle. At the anterior end of the latter is a very long tentacle, which exceeds the caruncle in length. Like the caruncle it is pale buff in the preparation, without any special development of pigment. Two shorter tentacular cirri spring from the anterior border of the head ; and the inferior cirri at the sides of the labial folds are pale, short, and slender. The labial folds are large and prominent, and the mouth opens immediately behind, i.e., at the anterior border of the third body-segment, which has two curved rugm in the middle line.
The branchise commence on the fifth body-segment, and continue to the tail, about seventeen being visible; while the first four feet, on the other hand, have a second (smaller) dorsal cirrus, placed to the inner side of the bristle-tuft. In the preparation the organs are directed backward, with the pinnse placed outward and backward, so that the main stem is internal, a feature less prominent in Chloeia Jlava. There are generally five branches on each side of the main stem, each furnished with secondary pinnae, the basal branch on the outer side being larger than the rest. Their colour throughout is slightly ferruginous.
Most of the bristles had been swept from the dorsal tufts, only the anterior and posterior segments having escaped. The bristles on the whole are more opalescent than in Chloeia Jlava, though a tinge of the same characteristic greenish-yellow occurs in all. As, however, many of the best marked and most typical bristles are absent, the following remarks will probably require qualification when a complete example is obtained. Three types are apparent in the dorsal tuft ; first, a very slender elongate kind, which diminishes from the lower third to a little below the fork (PI. Ia. fig. 14), the slender tip extending far beyond the point figured; and one tinted of a deep yellow throughout this region, the long limb of the fork being extremely produced, and, like all the other tips of the dorsal bristles, minutely granular from microscopic projections. The next kind (PI. IIa. fig. 1) has either a deep yellow tip and pale shaft, or is more elongated, with a pale attenuated
REPOET ON THE ANNELIDA.
15
tip beyond tbe fork, and very conspicuous grains toward the end of the longer limb. The third type (PL IIa. fig. 2) has the usual serrations on the edge of the long limb, and each is tinted deep yellow from a little below the dilatation of the fork to the tip. The figure represents an intermediate form, some being longer, and with more distinctly- marked serrations.
The ventral bristles in structure resemble PI. IIa. fig. 1, some having the tips deep yeUow, others being pale or very slightly tinted. As a rule the long limb of the fork is somewhat more slender than in the case of the dorsal, and posteriorly this feature is much more developed. The granular (shagreen) aspect of the tips is also present in the ventral bristles, an average example of wliich is shown in PI. Ia. fig. 15, from one of the anterior feet.
The dorsal cirri spring from the usual position at the posterior border of the dorsal bristle-papilla, and are slender, the tip being very 'finely and gradually attenuated. The base and tip are somewhat pale, the rest deep purplish-brown. The ventral cirrus tapers from base to apex, is pale and much shorter than the dorsal. The caudal styles are absent. A little granular matter and sand only were present in the stomach, and it is curious that both this and the previous species should have the intestinal tract so empty.
Kinberg ^ mentions two species from the east besides Chloeia jlava, viz., Chloeia heiigalensis and Chloeia malaica, but as no description or figure is given, it is impossible to identify them. Accordingly it has been deemed prudent to give the present form a new name. It will readily be distinguished when descriptions of the two first-mentioned are published. The Chloeia loarva of Dr. Baird resembles this form in the outline of the body, but differs in other respects. Its locality is unknown. The Chloeia niacleayi of Mr. Haswell, from Australia, differs in the structure of the bristles (so far as description goes) and in coloration. Prof. Grube’s Chloeia ceylonica, likewise appears to have a dark body.
Chloenea, Kinberg.
Chloenea atlantica, n. sp. (PI. I. fig. 4; PL Ia. figs. 10-13).
Haloitat. — Dredged at Station 3 (south of the Canaries) 18th February 1873, lat. 25° 45' N., long. 20° 14' W., in a depth of 1525 fathoms; bottom temperature 37°'0, surface 68° '0 ; hard ground. Two specimens were found adhering to the remarkable sponge Poliopogon amadou, Wyv. Thoms., which grew upon the calcareous axis of a dead Corallium, coated with peroxide of manganese.
The body (measuring 1 2 mm. in length and 5 mm. in breadth) is short and somewhat ovoid, wide at its anterior third but tapering posteriorly, and composed of about
^ Op. cit, p. 86.
16
THE VOYAGE OF H.M.S. CHALLENGER.
nineteen segments. The opalescent bristles (milk-white in life) project nearly as much on each side as the breadth of the body. Posteriorly it terminates in two elongated styles. The head bears a large and somewhat spongy, rugose caruncle, which extends backward at least to the fourth segment. The folds almost separate from each other on manipulation, and their looseness affords a distinctive feature when contrasted with those of Chloeia jiava and others. No eyes are visible in the specimens. In front of the caruncle is the median tentacle, which has about the same proportional length as in Chloeia. It is tinted of a dark greenish hue superiorly. After a longer interval than in Chloeia a pair of tentacular cirri, also slightly tinted superiorly, occur at the anterior border of the snout. The other pair (similarly coloured) are situated at the sides of the anterior labial processes. The latter are well developed, and homologous with those in Chloeia. The posterior border of the mouth is formed by the second body-segment instead of the^ fourth as in Chloeia. The segments of the body have a comparatively large antero- posterior diameter, the median five or six being conspicuous in this respect. A tyjDical segment bears on each side of the middle line a dorsal branchia, a tuft of long opalescent bristles, two dorsal cirri, a ventral tuft of long bristles, and a ventral cirrus.
The branchiae seem to commence on the fifth segment, and when fuUy developed consist of a short trunk bearing three or four simple pinnae, and an external basal division of a few processes. These organs are softer than in Chloeia jiava, and their hypodermic tissue is less compact. A series of muscular fibres runs from base to apex along the main stem, and in the pinnae fibres occur within the hypoderm at each side in ordinary views.
The dorsal bristles are opalescent and elongated, especially posteriorly, so that the body of the animal is enveloped in a kind of chevaux defrise by the crossing of the bristles anteriorly and posteriorly. Each bristle may be described as a long fusiform structure, the base equally with the tip tapering to a fine point. The shaft gradually dilates upward, then diminishes somewhat below the fork, where a little enlargement occurs. The elongated limb beyond the bifurcation has a slight bend opposite the tip of the shorter, and in the best developed a series of serrations occurs on the outer border of the long limb (PI. Ia. fig. 10), the points being directed downward in the ordinary way. Others show a slighter array of serratures along the edge next the fork (PL Ia. fig. 11), while a third and somewhat shorter set (PL Ia. fig. 12) appear to be quite smooth. These bristles have a very thin external crust, and are somewhat less brittle than in Chloeia Jiava. The air alluded to by Claparede probably gained entrance into the bristles of the latter through cracks. The anterior dorsal bristles have stouter though shorter tips, with all the markings boldly developed, especially the serrations on the side next the fork. The point of insertion is much more slender than the distal end. Those towards the extremity of the tail, again, are characterised by very elongated tips and distinctly serrated outer margins, as in PL Ia. fig. 10.
The ventral bristles form long asbestos-like tufts, which make a remarkable fringe.
REPORT ON THE ANNELIDA.
17
especially in the posterior half of the animal. About the middle of the body the structure of the tip of each bristle resembles that in PI. Ia. fig. 13. The long limb beyond the fork has serratures like those seen in the corresponding region of the dorsal bristles, and the shaft is extremely long, nearly cylindrical for the greater part of its length, and terminates interiorly in a pointed extremity, as formerly indicated in the dorsal bristles. Moreover, the same shortening of the tips anteriorly and the elonga- tion posteriorly characterise the ventral bristles. In the posterior bristles the longer limb of the fork is in many shghtly curved inward at the tip.
At the inner margin of the dorsal bristle-tuft, and a little in front and to the exterior of the branchia is situated a long fihform cirrus, which is tinted brownish-green superiorly, such forming a distinction, therefore, between this form and Chloeia. Another cirrus, the homologne of that in Chloeia, springs from the posterior part of the dorsal bristle-papilla, and like the former is extremely long and attenuated, with the pigment placed distally. The ventral cirrus occurs in its usual position, viz., rather below and behind the ventral bristle-tuft. It is also furnished with pigment distally.
"When dredged up, it was referred {vide Atlantic, vol. i. p. 176) by Dr. v. Willemoes Suhm to the family Amphinomidae, sub-family Euphrosyninae, with many of the char- acters of the genus Euphrosyne. It approaches Grube’s genus Notopygos in general configuration, and in the presence of the branchial cirrus, but differs materially in regard to the structure of the branchiae and the nature of the bristles, both of which show a nearer approach to Chloeia. In the present unsatisfactory condition of the Chloeia group, and though Kinberg’s description of the genus can hardly be followed, I have thought it best not to make a new genus, but to place it under his Chloenea} In the structure of the body-wall and the arrangement of the nerves it agrees with Chloeia. The perivisceral corpuscles are largely developed, and the wall of the stomach is loaded with refracting cells and granules, the contrast between this region and the more rigid intestinal canal with its lobose and more translucent glandular papillae being well marked.
Notopygos, Grube.
Notopygos megalops, n. sp. (? crinitus, Grube, var.) (PI. I. fig. 1 ; PI. IIa. figs. 3, 4).
Habitat. — Dredged at Station 36 (off the Bermudas) April 22, 1873, in 30 fathoms, amongst coral.
The body is about 9 mm. in length and 3 mm. in breadth, is fringed by a dense series of pellucid bristles, and consists of about nineteen segments besides head and tail. The segments are distinctly marked, those in the middle of the body having the greatest antero-posterior diameter, as in the previous form.
1 Op. cit., p. 86.
(ZOOL. CHALL. EXP. — PART XXXIV. — 1885.)
LI 3
18
THE VOYAGE OF H.M.S. CHALLENGER.
The head bears a caruncle which extends backward to the fifth body-segment, and has the usual folded structure, a little more lax than in Chloeia. At its anterior border is a short, slender, filiform tentacle, and the two pairs of tentacular cirri (one in front of the head and the other beneath) are also very small and short. All these organs are quite pale in the preparation. A pair of very large black eyes lie immediately behind the tentacular cirri at the front of the head, and a smaller jDair behind them, a little anterior to the base of the tentacle. In front of the mouth are the fleshy lips, which extend to the anterior border of the third body-segment.
The branchiae commence on the first segment clear of the caruncle (sixth), and continue apparently to the last. When fully developed each consists of a small tuft of about four branches, viz., a lateral on each side, and a median, which becomes bifid shortly after its commencement ; and occasionally a third small process proceeds from the larger division of the latter. These organs are highly vascular, a large afiferent and efferent vessel passing along each process.
The dorsal fascicle, projecting from the usual papilla, consists of a series of stiff glassy bristles, which at the anterior third of the body present a boldly forked tip, with a smooth outer margin, and having about three serrations on the inner surface of the longer process, near the tip (PL IIa. fig. 3), thus differing from the Notopygos fidvus of Haswell.^ The central canal is very distinctly marked, both in the shaft and the pro- cesses beyond the fork. After remaining for some hours on the slide in distilled water, a peculiar exudation (of an oleaginous appearance) took place from the fractured bristles, the majority of the isolated drops having a pyriform aspect with a pointed end ; and sometimes they formed a concentrically arranged group like certain fatty crystals.
The ventral bristles form a similar stiff glassy fringe. An average example is shown in PL IIa. fig. 4, the form being more slender than in the dorsal series, but the curves at the fork similar. The serrations along the inner margin of the long limb are, however, much more distinctly marked than in the dorsal bristles, about four being present in each. The type of bristle, therefore, is identical in both dorsal and ventral groups.
A slender filiform dorsal cirrus exists in front of the branchial tuft, and anteriorly it occupies a similar position though the latter is absent. A cirrus with a stout buff- coloured basal division arises from the usual situation behind the dorsal papilla. The tip is pale, filiform, and long. The ventral cirrus is pale, subulate, and comparatively short.
In this form the nerve-cords lie within the circular coat, the hypodermic insertions of the oblique being at their external border. The proboscis has internally numerous ridges composed of hypoderm with a chitinous covering, while a well-marked circular muscular layer encircles their bases in section.
This species has certain resemblances to Grube’s Notopygos crinitus,^ from the neigh- l)Ourhood of St. Helena, but the structure of the bristles as shown by Kinberg differs,
1 Proc. Linn. Soc. N~. S. Wales, 1878, p. 343. ^ Arthiv f. Naturgesch., Jahrg. xlL, 1853, p. 93.
EEPORT ON THE ANNELIDA.
19
since the ventral bristles in the form procured by the Challenger are serrated like the dorsal. The size of the eyes also diverges. Further investigation is therefore necessary
Notopygos labiatus, n. sp. (PL II. fig. 6 ; PL IV, fig. 2 ; PL IIa. figs. 5, 6).
Habitat. — Trawled at Station 201 (in Basilan Strait, to the south of the Philippine Islands), in lat. 7° 3' N., and long. 121° 48' E., in a depth varying from 84 to 102 fathoms; surface temperature, 83°‘0; bottom — stones and gravel.
It was accompanied by Polynoe, Eunice, Serpula, and a small Holothurian with long ambulacral processes. As further evidence of the hard nature of the ground is the occurrence amongst the bristles of fragments of tubes formed solely of sponge-spicules, ranged round the wall in a transverse manner or reticulated together to form the branched end of the tube {Terebellaf). There were also in the same shelter a few fragments of gravel.
This is a comparatively large form, measuring about 30 mm. in length by 9 mm. in breadth, and having the sides and dorsum protected by a dense series of stiff bristles, which are pale green throughout. The body has the usual shape, and consists of from twenty-five to twenty-eight well-marked segments. On the dorsum a triangular brownish area indicates the middle line at each segment-junction. The ventral surface is marked by a median line. The head is furnished with a caruncle which extends to the posterior part of the fifth body-segment. It is of a deep blackish hue below the inferior frills. The lamellae are somewhat lax and spongy, but symmetrically arranged ; the lower being very regularly folded and more rigid. The front of the organ is bounded by a median and two lateral folds, while posteriorly the pointed tip is formed by a coalescence of the transverse plaits. Close to the front of the caruncle is the comparatively short median tentacle, while after an interval in front of the head are the two superior tentacular cirri, which resemble the former in shape. The inferior are placed on each side of the prominent lips. The rounded head lies in front of the caruncle, and has a large eye on each side anteriorly, and a smaller, separated by a considerable interval, behind. The great oral folds or lips project in front of the head when viewed from the dorsum. The hinder part of the mouth is formed by the third body-segment. Posteriorly the body terminates in two thick club-shaped processes.
The branchiae in contraction form small tufts composed of two divisions, viz., a small outer and a larger inner. The outer bears a series of pinnae, with occasionally secondary pinnae, while the inner, in the form of a miniature bush, likewise shows minor branches. All have specks of pigment, so that they are slightly greenish in spirit. In their ordinary position in the preparation the main stem is external, the pinnae being directed inward toward the middle line.
The pale greenish dorsal bristles are very prominent, and are large, stiff, and fragle.
20
THE yOYAGE OF H.M.S. CHALLENGER.
As usual they increase in length from before backward. Their chief peculiarities are the large size, smoothness, and the great length of the median peninsula before the canals from the processes join (PL II a. fig. 5).
A slightly opaque granular dej)osit exists outside the canal in the longer process, and often also in the smaller, as well as in other parts of the tij). The canals show very distinct granules below the fork. The whole of the tip, to a point below the fork, is covered with a microscopic shagreen. The arrangement of these dorsal bristles is always more lax than that of the ventral.
The ventral bristles form a dense fascicle with the broad axis of the fan directed vertically. The larger bristles are generally dorsal, indeed, when the tuft is viewed from the ventral aspect a somewhat regular gradation of the tips is observed, so that the slope from below upward is bristled with a continuous series, which the few shorter dorsal bristles do not affect. In a lateral view the outline narrows from below upward to the long bristles. These are paler than the dorsal, and much more slender (PL IIa. fig. 6). The inner border of the long process of the fork has two (or in some three) serrations, and the isthmus at the base of the fork is proj)ortionally shorter than in the dorsal. The type of both corresponds very closely except that the surface of the tip in the ventral does not show the microscopic shagreen. When a bristle is broken and left in water the shaft exhibits in its centre a series of obliquely curved lines, and in many a number of regular transverse lines occur below the isthmus at the fork, and throughout a considerable extent of the shaft beneath. The yellowish oleaginous contents of the bristles are well seen after fracture, both externally and in the hollow of the shaft, where they sometimes assume a lenticular shape. Mr. W. A. Haswell, B.Sc., who has so carefully and skilfully investigated many of the Annelids of New South Wales, describes two species of Notopygos ^ in which both dorsal and ventral bristles are quite smooth. The dorsal bristles of the present species, Notopygos labiatus, have no serrations, so that too much weight need not be attached to this character of Kinberg’s.
The branchial cirrus arises at the inner border of the bristle-papilla, a little in front of the branchia, is pale at the base, but tinted madder-brown throughout the rest of its extent. It is covered with rows of long cilia. The dorsal cirrus proper (in the usual j^osition behind the papilla) has a madder-brown large basal division and a filiform pale distal region, which is constricted just below the soihewhat cylindrical tip.
In transverse section the body-wall shows the features of the group, besides certain definite characters of its own. Instead of the little bifid papillse of the hypoderm of .the central dorsal region of the Chloeia from the Mediterranean, this form shows a central and two median longitudinal ridges, by the great increase of the central oblique muscles of the part. The circular muscular layer (beneath the hypoderm) is also largely developed. The nerve-cords are especially large and distinct, and are much less flattened out than in
1 Proc. Linn. Soc. N. S. Wales, November 25, 1878.
REPORT ON THE ANNELIDA.
21
Chloeia ; but the attachment of the oblique muscles at the sides, and the relation to the other parts are the same. The circular muscular coat of the intestine is also strongly developed. It is noteworthy that this and the previous kinds so often have empty intestinal tracts. As in many fishes, the digestive process is probably rajDid. In vertical longitudinal section the posterior part of the great muscular cushion or tongue presents an easily separable series of the usual vertical muscular lamellse, transversely arranged. They possess a somewhat fan-shaped appearance.
This form cbfiers from Grube’s Notopygos crinitus in regard to the branchiae and the structure of the bristles as shown by Kinberg.^ There is considerable doubt in regard to its connection with Notopygos macidata, Kinberg,^ from the island of Panama, which Semper also found at Bohol in the Philippines;^ but in the characters above-mentioned it also deviates from this species. A re-examination of the bristles of Notopygos maculata would help to remove ambiguity.
Amphinome, Bruguiere.
Amphinome rostrata (Pallas) (PL I. fig. 7 ; PI. IV. fig. 1 ; PI. Ia. fig. 16 ; PL IIa. figs. 8-12).
Aplirodita rostrata, Pallas, Misc. Zool., p. 106, Tab. viii. f. 14-18, 1766.
Amphinome rostrata, Auctorum.
Habitat. — Several fine examples were procured at the surface of the sea near the Bermudas, some being captured along with a large Hermodice on a log. Amphinome vagans is occasionally found under the same circumstances on floating timber.
All the specimens are distinguishable by very evident external characters in spirit, viz., the peculiar slate-blue of the entire body, and the ferruginous hue of the tentacles, cirri, and branchiae. The smaller specimens have the bluish colour of the body less marked, especially ventrally. A small example of Amphinome vagans, Leach (Sav.), procured from the Godefiroy Collection, and named by Grube, shows this colour on the dorsum, while the ventral surface is pale buff ; and the latter colour characterises another all over.
Since the description by Pallas, no author has entered minutely into the characters of the animal, and therefore it is necessary to do so on the present occasion, as at least one species [Amphinome vagans) is closely allied.
The body is elongated and somewhat tetragonal, the following numbers of body-seg- ments occurring respectively in the examples : — 56, 56, 50 (imperfect), 52 (imperfect), 46 (small), 48 (imperfect), 50 (imperfect), 49, 51, 54, 52, 58, 59, 57, 51. Pallas gives 66 as the total number in his small specimen, 55 in the larger. Segments are so readily throvm
1 Freg. Eugen. Resa, Tab. xi. fig. 3, G.
^ Ofversigt k. Vetensk.-Akad. Fdrhandl., 1857, p. 12 ; and Freg. Eugen. Resa, Taf. xi. fig. 5 (as Lirione 'niaculata).
® Anuel. Fauna d. Philippinen., p. 8, Taf. i. fig. 3.
22
THE VOYAGE OF H.M.S. CHALLENGEE.
off in such an animal, and the anal region accommodates itself with such ease to the altered circumstances, that little weight is to be put on this feature. Both dorsal and ventral surfaces are somewhat convex, the former being marked transversely with the segment-lines and slightly corrugated throughout ; the latter showing the segment- junctions even more distinctly, each being coarsely rugose, and having a median longitudinal furrow. The three segments behind the mouth are somewhat smoother.
For so large a body the head seems disproportionately small. In front the snout, which consists of the anterior border of the buccal segment and corres]3onds to the prominent anterior labial folds in Euphrosyne and Chloeia, bears two tentacular cirri (palpi) which are larger than the succeeding processes of the kind. They are conical and of a ferruginous hue from base to apex. The next organs are the two cirri situated at the anterior border of the slightly differentiated cephalic region, which indeed appears at this part to be continuous with their buccal division. Their bases are separated by a slight interval, and each has a basal bluish segment, and a terminal conical ferruginous division. Immediately behind is the small tongue-shaped caruncle, which springs nearly in a line with the anterior border of the first bristle-papilla, and terminates in a groove in front of the posterior border of the first body-segment. The organ is attached along the middle throughout its entire length, but is free at the edges. Its upper surface is quite smooth, or in some slightly wrinkled, and the whole is somewhat darker (bluish- black) than the surrounding parts. In some the edges are of the latter shade, while the central region is of the ordinary bluish colour of the dorsum. It is deeply cordate in front, and at the termination of the sinus bears a short ferruginous conical process — the median tentacle. The mouth opens inferiorly at the anterior border of the third body- segment, and from the oral aperture two well-marked folds (corresponding with sjDecial thickenings in Chloeia) run forward to the anterior prominence of the snout which bears the cirri. These folds are wide in front, narrow behind. Two additional plaits pertain to the second segment, and another to the first.
The branchiae commence on the third body-segment, and have the form of dense arbuscles, the first being nearly as large as any of the succeeding. So dense are they, indeed, that Pallas likened them to the cotyledons of the ruminant chorion. Each arbuscle consists of six or seven main stems springing from a common base, and dividing somewhat dichotomously into a dense bush of filaments. The outer stems have shorter terminal processes, the inner possess both longer stems and longer terminal processes. The latter are ferruginous in colour throughout, while the stems and branches are of the usual bluish tint. The tuft is situated close to the base of the dorsal bristle-papilla, and somewhat to the inner or dorsal side. The larger (dorsal) branches of the arbuscle envelop the bristle-papilla in the preparation, so that only a small portion of its anterior and inferior region is bare. The branchiae are continued to the tip of the tail. Towards the posterior fourth of the body, as Pallas says, they meet on the dorsum, again separat-
REPORT ON THE ANNELIDA.
23
ing toward the anal region. Pallas also correctly observed that the branchiae do not proportionately decrease posteriorly.
The dorsal cirrns arises from the posterior part of the dorsal bristle-papilla, and has a basal and a distal division. The former is bluish in colour, the latter ferruginous and tapered towards the extremity, which extends as far outward as the tips of the bristles. Occasionally the dorsal cirrus of the second segment shows a small inner filament. The ventral cirrus is very short, but it possesses a similar structure. It is attached to the posterior border of the ventral curve of the foot.
The dorsal bristles form a curved row, with the convexity directed forward, and consist of two kinds, viz., a stout series with grooved and serrated tips (PI. IIa. fig. 8), and a longer series with finely tapered, curved tips, the latter being minutely roughened on the convex side with small points (PI. IIa. fig. 9). The shafts of the longer kind are much more granular and opaque than in the stouter shorter series. The tips of the spinose bristles are peculiarly curved, and the surface from which the rough points spring would seem to be somewhat flattened. The points resemble the bases of a brush-like series of spikes (PI. Ia. fig. 16). The ventral bristles, again, are few in number and comparatively short, but powerfully developed (PI. IIa. fig. 10). The tip of each is strongly curved (hook -like), and marked by a series of fine striae in the opaque or whitish portion, these being continued downward on the posterior part of the bristle.
The spines of both dorsal and ventral divisions of the foot are peculiarly expanded at the tip, and each differs from the other in the form of the dilatation. Thus the dorsal (PL IIa. fig. 11) has a more elongated tip; while the ventral (PL IIa. fig. 12) has a larger and shorter tip, and the entire organ is stronger. Besides the bristles and spines projecting externally, each bristle-sheath has various spines and bristles in course of development, and each from its earliest condition bears its characteristic features. The solid tips are the first parts to appear, and in the case of the ventral series the bristles are in pairs with a spine at intervals.
The anus in most of the specimens is a great circular opening, embracing several segments at the posterior end of the body ; indeed the size of the aperture is a remarkable feature. Internally the mucous membrane forms a prominent mass, occasionally with a somewhat radiate arrangement of rugse. Pallas states that the anus is bilabiate, and in some conditions the description is very appropriate. Most of the examples in the present case may have lost the tip of the tail. The large size of the anus would indicate a peculiarity in the digestive function.
In the intestinal canal fragments of sessile-eyed Crustacea were found in considerable abundance, generall}?" mixed with brownish-purple debris. The cirri of barnacles were also common, and occasionally a valve of the same animal, so that they would appear to feed on them ; they would readily be procured on the floating timber.
The general structure of the proboscis conforms to that in Chloeia. The alimentary
24
THE VOYAGE OF H.M.S. CHALLENGER.
canal is capacious, its first division being largely dilated and covered by transverse rugae. A powerful layer of longitudinal muscular fibres is developed externally at the posterior part of the proboscis.
The central organ of the nervous system occurs in the caruncular region, a little in front of the median tentacle, in the form of a bilobed mass. It is interesting that the separate nerve-cords in front are smaller in proportion than the trunks after they form the double cord. The nervous system on the whole is feebly develoj^ed.
On each side, in a line with the median tentacle, is a large ocular (?) organ, which appears to be single. Section, however, shows that there are two deep centres of the pigment, and that there is a tendency to facets on the surface. Each mass forms a pro- minent oval projection, and appears minutely dotted under a lens.
On the dorsum of the second body-segment are a number of prominent warty rugae of the hypoderm.
Two small specimens from the Atlantic, named by Prof. Grube Amphinome vagans, differ in no respect from the foregoing. They came from the Godeffroy Museum. This species is probably the Amphinome pallasii of De Quatrefages,^ from the Azores and the Antilles, and is certainly the Pleione tetraeda of M. -Edwards.^
Ilermodice, Kinberg.
Hermodice carunculata, Pallas (PL V. ; PI. IIIa. figs. 1-4).
He7'modice carunculata, Auctorum. It is doubtful -whether Seba’s Millepod amarina amhoinensis, Seba, Tbes. Ser. Nat., i., tab. Ixxxviii. p. 131, be the same.
Habitat. — A large specimen measuring upw'ards of a foot in length was procured at the surface of the bea near the Bermudas, while a bleached example about a fourth the length comes from the littoral region of the same islands. Two others were found at St. Vincent, Cape Verde Islands, in July 1873 ; and two were collected at St. Thomas, West Indies, in March of the same year.
In those best preserved the characteristic features are the dull greenish or slate-bluish finely corrugated dorsum, the small size of the branchial tufts as compared with Amphi- nome rostrata, the pale buff of the ventral surface, and the alternation of the dorsal processes.
The body is even more distinctly tetragonal than in Amphinome rostrata, this character being heightened by the issue of the bristles from the prominent angles. The diminution of the body posteriorly is well marked. The segments are clearly defined, and range from sixty-seven to seventy, and in the large example to one hundred and fifty-five. On the ventral surface a large vessel runs along the median line and is visible externally. The anus opens' as a somewhat symmetrically corrugated orifice on the dorsum, and there is a peculiar papilla, projecting posteriorly at the lower border.
1 Anneles, i. p. 394. Efegne Anini. ill., pi. viii. his, fig. 1.
EEPOET ON THE ANNELIDA.
25
The head is at once differentiated from that in Amphinome by the greater caruncle and the modification of the anterior border of the snout. Instead of the broad median region anteriorly there is in this form only the small rounded upper surface of the pre- labial folds. The caruncle is much larger than in Amphinome, and extends to tlm posterior border of the fourth body-segment. It is composed of a series of folds (eight in number), each of which is again deejDly notched in the preparation, so that it resembles a rope with its strands. The primary folds run from behind forward and outward, and are alternate with regard to the central axis. The median tentacle arises immediately in front of the latter, from the prominent top of the head. It is much larger than the two tentacular cirri which lie outside and in front of it, or than the inferior cirri which spring from the sides of the labial folds beneath. The eyes are two on each side, and by no means large. One is situated in a line with the anterior margin of the median tentacle ; the other has nearly the same relation to the posterior border of the organ. In one a third pigment-spot occurred on the left, near the base of the tentacle ]30steriorly. In the large example the anterior pair of eyes are brownish, the posterior black. The top of the head and the caruncle are minutely flecked with whitish grains.
The mouth opens interiorly in the middle line between the third and fifth segments. The thick pre-oral folds, so well marked in some of the genera, are here comparatively small, and chiefly occupy the middle of the first segment. There is a decided interval between their posterior border and the oral aperture. The pouting masses of buccal tissue are softer and more rugose than in Amphinome.
The relative difference in the size of the branchise forms a characteristic feature when contrasted with Amphinome ; moreover, they commence on the first bristled or body- segment, the first three or four being less than the others. Each tuft (when fully formed) consists of two main divisions, of a pale greenish colour, an outer smaller, in the form of a separate process immediately behind the bristle-tnft, and a larger inner arl^uscle which divides into several main Ijranches with short terminal processes. Three seems to be a common numl^er in the terminal groups. The whole organ is minutely dotted with white grains, which are densest on the posterior aspect of the process, and is somewhat dichotomously branched. The branchise are continued to the posterior extremity of the body, and only slightly diminished in that region.
The dorsal bristle-tufts are distinguished from those of Amp)hinome and the others yet examined by their evidently alternate arrangement. So well-marked is this feature that at first sight the dorsal series seems to be double. The bristles are very fine and slender, and in this respect in strong contrast to those of Chloeia, Amiohinome, and Notopijgos. Their beautiful asbestos-like whiteness is also striking, as well as their slightly twisted appearance as they emerge from the socket, like a tuft of camel’s hair from the cjuill of a hair-pencil. They do not spring from a raised papdla as in Chloeia and Notopijgos, but emerge from a pit. Each tuft consists of a dense series of
(ZOOL. CHALL. EXP. PART XXXIV. 1885.) LI 4
26
THE VOYAGE OE H.M.S. CHALLENGER.
simple, long, slender, and finely tapered bristles wliicli have tlie tips (PL IIIa, fig. l) minutely roughened, from the point a considerable way downward. Such microscopic processes on the surface are homologous with the bolder condition in Amphinome. The other kind of bristle is distinctly serrated at the tip (PI. IIIa. fig. 2), upwards of twenty teeth being visible, and a fold of the chitine passes from the apex of each serrature down- ward like a hood. These bristles show in a remarkable degree the albuminous contents (the brownish masses of Grube) after drying, for each bristle collapses, and the contents form nodular masses all over the exterior. Glacial acetic acid causes many bubbles of gas to escape from the central cavity of the bristles, but has no effect on the globules. Sulphuric ether and absolute alcohol show as little action on the latter as the acid. They would therefore appear to be albuminous, though their appearance is decidedly fatty.
The ventral bristles are all of one kind (PL IIIa. fig. 4, which represents one of the inferior bristles), presenting a slightly striated shaft and a curved, slightly bifid tip, while a series of very distinct serrations occur on the edge of the latter, from the tip to the spur. Like the dorsal bristles, each of the points or serrations does not indicate merely a spike, but also a slight wing. The upper bristles possess much longer and more slender tips, and the spike is more distinct (PL IIIa. fig. 3). In the bleached example from Bermuda the serrations and tips of the ventral bristles are much abraded, so that only those sheltered by position show the ordinary structure. The animal, therefore, makes some use of them, for progression or otherwise, and probably under stones. In the specimens from St. Vincent, Cape Verde Islands, these bristles form a somewhat regular vertical row, the longest bristles (with the longest tips) being superior. In the very large example many of the ventral bristles are more distinctly bifid than usual.
The dorsal cirrus arises behind the bristle-23apilla, and has a very attenuated tip. It is composed of a basal and a distal division. The ventral cirrus is smaller and occupies the ordinary position behind the ventral bristle-tuft. Both increase in length posteriorly.
In the stomach of the large sj)ecimen were muddy masses containing sponge-spicules, Diatoms, Eadiolarians, sand-grains, fragments of chitine, and cellular and granular debris.
The nerve-cords are quite within the body-wall, for the oblique muscles meet in the middle line beneath them, the fibres apparently decussating vdth the circular muscular coat. Connective tissue and granular matter, moreover, separate them from the muscular fibres, and in the sections the perivisceral corpuscles also often intervene. A considerable hy|3odermic canal occurs immediately below the decussation just mentioned. This seems to be filled with opaque material. When a section is made in the line of the ganglia, a large nerve-cord on each side is seen to arise from the cells in their interior. The upper and lower walls are strengthened by a dense capsule. Superiorly also a peculiar conical
EEPOET ON THE ANNELIDA.
27
band rests on the cords, the lower part (next tlie ganglion) being more translucent and granular, the outer more opaque.
The perivisceral lining is bordered with large nucleated granular cells — the perivisceral corpuscles.
The proboscis has the usual structure.
This species, subsequently to the remarks of Pallas and Savigny, formed the subject of the late Prof. Grube’s inaugural dissertation,^ chiefly with regard to the digestive, vas- cular, and nervous systems. It is a characteristic western form. The Ampliinome gigas in the collection of the British Museum is this species from the Atlantic Ocean.
Prof. Grube mentioned it as one of the Annelids of Madeira in 1857, and it has since l3een found there by Prof. Langerhans, whose figures ^ of the bristles, however, deviate from those procured by the Challenger, since the serrations of both dorsal and ventral bristles are much fewer. This author notes the calcareous nature of the bristles, which in this respect agree with those of others of the group, e.g., Chloeia fucata.^ The form described by Dr. Baird under the name of Amphinonie didymoh7xmchiata,‘^ from the Island of Ascension, seems to approach this genus {Hermodice) in the size and structure of the caruncle and in other particulars.
Eurijthoe, Kinberg.
Eurythoe pacifca, Kinberg? (PI. II. figs. 3, 4; PI. III. fig. 3; PI. IIa. fig. 13; PI.
IIIa. figs. 5-9).
Habitat. — Several specimens occurred at Bermuda between tide-marks. It seems to extend to both great oceans, the coral reefs of the Pacific being a favourite haunt. Grube ® describes a form from Tahiti and the Nicobar Islands, which he doubtfully refers to the same species. Kinberg’s specimens came from the shores of the islands of Eimeo and Foua.
The body is somewhat flattened and rectangular in section, slightly pointed in front, and gradually diminished posteriorly. The specimens are comparatively small, the longest measuring about 65 mm., with a breadth of 7 mm. One had one hundred and five body-segments.
The snout differs from that in Hermodice in having a broad lobe in front, shaped somewhat like a hoof, and bearing the two tentacular cirri, one pair superiorl}^ in front of the eyes, and another interiorly toward the posterior border. The head carries the rather elongated and linear caruncle, which impinges] on the fourth body-segment, and has anteriorly the median tentacle ending in a filiform tip, the organ being
1 De Pleione carunculata, Befiiomontii Prussonm, 1837.
^ Die Wurmfauna Madeiras, Zeitschr. f. wiss. Zool., xxxiii. p. 277, Taf. xiv. fig. 8.^
2 Trans. Zool. Soc. Loncl, ix. p. 396. ^ Trans. Linn. Soc. Lond., xxiv. p. 449, pi. xlv.
^ Annel. Novara-Exped., p. 8.
28
THE VOYAGE OF H.M.S. CHALLENGEE.
shorter than the foregoing cirri. The caruncle shows a dorsal ridge, indistinctly separated from the horizontal basal portion, and thus decidedly different from that in Hermodice. The larger pair of eyes is situated in a line with the anterior margin of the tentacle, and the smaller a short distance behind. The four mark out an oblong, probably nearly a square in life.
The mouth opens on the ventral surface at the anterior border of the fourth body- segment, the central region in front being occupied by the wedge-like lobes which pass forward to form the snout, and the sides by the convergent lines of the three anterior body-rings. The anus occupies the dorsum of about three of the small terminal segments, and has posteriorly a prominent papilla. In the intestinal canal of specimens from the Seychelles are masses of algse and considerable pieces of coral and nuUipore. In the present case pieces of olive-brown sea- weed are most conspicuous.
The branchiae commence on the second body-segment each as a tnft of two processes. They are continued along the dorsum as somewhat less conspicuous organs than in Hermodice. When fully formed each tuft consists of a series of slightly divided processes, chiefly after a dichotomous type. They are continued to the last or the penultimate segment. A peculiar infusorial parasite occurs in numbers on the branchiae.
The dorsal bristles consist (1) of a few simple distinctly curved bristles (PI. IIIa. fig. 5), which are frequently alisent in the preparations ; (2) of a large group of charac- teristic bristles with slightly bifid tips (PI. IIIa. fig. 6), one of the divisions being a mere spur, while the other is extremely elongated and tapering, with serrations along the inner edge ; and (3) of a well-marked serrated series (PI. IIIa. fig. 7), vdth the terminal portion striated in a peculiar manner.
The ventral bristles are bifid, one or two superiorly (PI. IIIa. fig. 8) having slender shafts and very long tips, with serrations on the inner edge ; while the majority have stout tips (PI. IIIa. fig. 9) with three or four serrations on the inner margin. Anteriorly the tips of these bristles are somewhat longer. Amongst the inferior bristles are some short simple spines with a dilatation in the centre (PI. IIa. fig. 13). Similar forms are also shown by Kinberg in Eurythoe corallina} These hastate spines have the tips projecting only a short distance beyond the cuticle, the stoutest part of the bristle being internal.
The dorsal cirrus has the usual basal segment, and though fairly developed does not project quite as far as the bristles. It becomes more elongated jDosteriorly. The ventral is a short process in the ordinary position.
The arrangement of the nerve-trunks differs from that in CJdoeia, since the oblique muscles decussate beneath them. In some sections (anterior) a regularly arranged series of longitudinal fibres occurs within the circular muscular layer, so that the nerve- cords are carried further inward at these points.
1 Freg. Eugen. Resa, Tab. xii. fig. 12.
REPOET ON THE ANNELIDA.
29
This species agrees with the Eurythoe pacijica, Kinberg, as named by Prof. Grube in the Godeffroy Collection, except in slight particulars due to indifferent preservation and the wear of the bristles in the larger specimens. The same author, moreover, had under examination other examples from Tahiti and the Nicobar Islands.^ Some from Samoa measure about a foot in spirit. The same form was found by Prof. Percival Wright in the Seychelles, and it appears to be identical with the Amphinome hruguieresi of De Qnatrefages.^ The Eurythoe alcyonia of Savigny from the Eed Sea (Dr. Eiippell) is also closely allied, to judge from an example in the British Museum. The ventral bristles are smooth. A near form is the Amjohinome alba, Baird, the ventral bristles of which are mostly smooth, only one showing a serration. In the British Museum the Eurythoe [Amphinome) complanata from the West Indies seems to have the tip of the ventral bristles quite smooth.
In the present state of our knowledge it is almost impossible to diagnose the forms referred to by the various authors, and it is probable that the same species has received several names. The laxity in this respect is shown by the fact that Kinberg in his description states that the ventral bristles are smooth, while his artist figures them with serrations.
One of the Ceylonese individuals of Eurythoe pacijica in the British Museum shows a few notches in the ventral bristles towards the tip, but nO' distinct serrations.
Eurythoe pacijica, var. levuhaensis (PL XVI. fig. 5; PL IIa. fig. 14; PI. IIIa. figs, 10-12).
Habitat. — Levnka, Fiji.
The examples are small, the longest measuring about 18 mm., and the diameter of the fragmentary larger sj^ecimen being 3 mm. One has fifty segments.
It is distinguished from the foregoing by the much more distinct segmentation, the rings being both larger and more deeply cut. The snout is more evidently bi-lobed, and the caruncle is proportionally broader, but it is doubtful how much reliance can be placed on spirit-preparations in this respect. The median tentacle is smaller than in either of the others. The eyes occupy the ordinary position, and are of a dull brown colour The anterior pair (as usual) are considerably larger. The caruncle impinges posteriorly on the fourth segment. Interiorly the mouth opens at the anterior border of the fourth body-segment, the three anterior somites sloj)ing oblicjuely from the sides inward. The median folds (prsestomium) show a distinct central hollow in front, though in this respect it agrees with the previous form.
The only indication of branchiae on the first body-segment is an eminence at the base of the cirrus, but on the next is an inner tuft of two or three erect processes, 1 Annel. Novara-Exped., Zool. Th., Bd. ii. p. 8. (sep. Abd.). ^ Anneles, i. p. 398.
30
THE VOYAGE OE H.M.S. CHALLENGES.
and an enter of two filaments. When fully formed the organs differ from those of the previous form by their greater size and more evident divisions, features borne out on microscopic examination. The processes are larger and longer, as well as more distinctly lanceolate at the tip, whereas in the former case they are blunt. The branchiae are continued to the posterior end.
In examining the dorsal bristles, a feature at once strikes the observer, viz., their greater translucency and much more flexible nature. The bifid kind (PL IIIa. fig. 10) have somewdiat larger shafts than in the former examj^les, and the serrations on the inner side of the tip are less distinct. The serrated kind (PL IIIa. fig. 1 1 represents an average specimen) is not marked by distinct lines or grooves toward the tip as in the previous form, and the teeth are less prominent. These bristles seem to be more numerous in the tuft than in the typical Eurytlioe 'pacijica. Another feature is the great prominence of the smooth curved form (PL IIIa. fig. 5) in the foregoing specimens, and their comparative paucity and slenderness in this.
The ventral bristles are on the whole more translucent, and have the tips more dilated than in Eurytlioe pacijica proper. One of the average bristles from the anterior third of the body is shown in PL IIIa. fig. 12, though it has to be mentioned that the number of serrations on the inner edge of the tip is sometimes three or four in the anterior feet. If the bristle is slightly turned round the tip appears narrower, so that there is lateral flattening. The hastate spine (PL IIa. fig. 14) is somewhat larger than in the previous form. The elongated series, corresponding to PL IIIa. fig. 8, is also present, and closely approaches the latter. The resemblance in the characters of the bristles is striking, and it has been thought unnecessary to make specific separation.
The nerve-cords have the same relations as in the preceding form, and the alternation of the ventral longitudinal muscles is even more apparent. This alternate passage invurd and outward of the ventral longitudinal muscle is noteworthy. A small canal exists in the central line of the hypoderm. It is interesting to find the canal independent of the nerve-oords.
Hipponoe, Audonin and Milne-Edwards.
Hipp>onoe gaudichaudi, Audouin and Milne-Edwards (PL I. fig. 5; PL IV. fig. 3; PL IIIa. figs. 13-17).
Elabitat. — Several examples occurred on a log at the surface of the sea, 100 miles north of Bermuda, 28th May 1873. Another was found adherent to Lepas fascicularis on the surface of the North Pacific, 6th July 1875.
This form certainly presents a very considerable divergence from the Amphinomidse in external appearance. The body is much more flattened, somewhat fusiform in outline, and composed of about thirty-two boldly-marked segments, including head and tail. The length of the largest specimen is 24 mm., and its breadth fully 5 mm.
EEPOET ON THE ANNELLDA.
31
Instead of the preoral folds the minute head of the animal forms the anterior border of the snout. In shape it is somewhat ovoid or occasionally lozenge-shaped, from the pointed nature of the anterior edge. It bears on the dorsal surface four eyes, all nearly equal in size, and forming a rectangle, those of each pair being close together, thus diverging from the ordinary type in the Amphinomidse. Somewhat behind the posterior pair of eyes is the long median tentacle, a simple subulate process which projects backward as far as the middle of the second body-segment. A little in front of the eyes at each side is a tentacular cirrus, considerably shorter than the foregoing ; while on either side of the tip of the snout is another of similar length. The mouth opens on the under surface at the anterior border of the second body-segment, and a flat surface, marked by two narrow longitudinal ridges, leads forward to the tip of the snout. The posterior end terminates in a bulbous anus, which in some appears pediculated, from the constriction immediately in front.
The branchiae commence on the fourth body-segment as a considerable tuft in each case, and when fully developed consist of a slightly divided arbuscle richly coated with cilia, and thus diverging from the condition in the Amphinomidae. Moreover, the hypo- derm of the organ presents a lax and cellular appearance, while the cuticle is very thin on the distal processes. In transverse section the translucent areolar hypoderm is especially marked at the base of the organ. The central area is occupied by large granular globules, apparently coagulated blood, and the latter also occurs in the two channels in the distal branches, v/hich are irregularly dichotomous. The cilia appear to form several rows on the sides of the processes. The branchiae in the Hippoyioe cranchii of Baird are decidedly larger.
The dorsal bristles form a broad fan-like tuft at the anterior border of each segment, extending from the dorsal to the ventral aspect of the foot, and with the slight concavity of the curve directed backward. The bristles are translucent, simple, and tapering (PI. IIIa. fig. 13). The outline of the lower part of the shaft (PI. IIIa. fig. 16) is slightly irregular, from indications of the points or spikes, which become more distinct distally, and there is a single median streak probably due to a ridge externally, the rest of the surface being marked by fine longitudinal lines. In the figure the shaft is comj)ressed and cracked, but the spikes are thus rendered more evident. In its normal condition the lower part of the shaft appears as a clear tubular bristle (PL IIIa. fig. 15), only a slight roughness along one of the margins indicating the spikes. The mere evaporation of a small portion of the water between the cover and the slide suffices to destroy the outline by breaking the tube and causing it to resemble fig. 16. The tips of some are peculiarly bifid (PI. IIIa. fig. 14). Acetic acid has very little effect on these bristles except in rendering them more translucent. No bubbles of gas escape, and they are less brittle than in the Amphinomidse ; so that on the whole they do not seem to be of the same characteristic calcareous nature.
82
THE VOYAGE OE H.M.S. CHALLENGER,
The dorsal surface of the body is slightly convex, whereas the ventral is marked hy a deep median groove from the month to the posterior extremity. On the prominent portion of the second body-segment, bounding the lateral and j^osterior part of the mouth, on each side is a slight pit containing the characteristic hooks of the species. The succeeding segments show a similar fossa on each side, about midway between the median groove and the external border. The hooks (PL I II a. fig. 17) are strictly bifid, with a long curved sharp terminal claw, and a shorter blunt one beneath ; the former being marked by coarse, and the latter by fine striae. The deep pits with their prominent fleshy margins are probably of importance in keeping the hooks always sharj) and uninjured. The latter are perhaps capable of a certain amount of extrusion when the animal wishes to anchor itself to floating timber or other structures. In Hipponoe cranchii, Baird, the hooks quite differ in structure.
A single cirrus of considerable size, in the form of a simple subulate process, is situated at the inferior margin and slightly behind the bristle-tuft, and therefore is best seen from the ventral surface.
Posteriorly the body terminates in a simple rounded papilla bearing the anus on the dorsum. The last pair of branchise occur at each side of the papilla.
The specimens are all so distended with ova that the structure of the body-wall cannot be seen with clearness. The chief points noticed are the great thickness and translucency of the hypoderm, which is almost Nemertean in its areolar and cellular condition, along the ventral and lateral surfaces ; but on the dorsum between the bristle- bundles it is much thinner and more compact. Moreover, the cuticle along the ventral region is not readily distinguished as a separate layer (the surface resembling that in the Nemertean which has a ciliated coat); while on the dorsum the cuticle is of considerable thickness. There is thus some evident difference in the physiology of the surfaces. Beneath the hypoderm is the circular muscular coat. The longitudinal and oblique muscles were so interfered with by the ova in the somewhat softened preparation, that nothing definite could be determined. So far as could be observed, the nerves seemed to lie above the oblique muscles.
In the example from the North Pacific there are twenty-four segments, including head and tail. The branchiae commence on the left side on the second, as a simple process, and on the right on the fourth segment as an arbuscle. The ventral hooks and dorsal bristles correspond in minute structure with those from Bermuda.
Dr. Baird speaks of certain small parasitic animals being attached to the under surface of the specimens in the British Musuem. None occurred in the present forms. The species aj)pears to have a very wide range, stretching from Port Jackson, Australia, to Madeira and the Bermudas. Dr. Baird,^ has the credit of correcting the erroneous descrip- tions of his predecessors in regard to the eyes, hooks, and other parts. It is remarkable
^ Proc. Linn. Soc. Land., toI. x. p. 239.
REPORT ON THE ANNELIDA.
33
that the two former organs escaped the notice of MM. Audouin and Milne-Edwards, in their original description of the species from Port Jackson, where it was obtained by Capt. Freycinet in his celebrated voyage round the world. In two instances in the British Museum the specimens occurred amongst barnacles.
Family Aphroditidjs.
The descriptions of the family given by Kinberg and other authors render it unnecessary to define it on the present occasion. All the genera procured in the Expedition of the Challenger fall under those already described, yet there are some forms which, from their intermediate structure, help to clear up the relationshi]3s between Aphrodita, Lcetmonice, and Hermione, and especially indicate the steps between the two first mentioned. Something of the same kind, however, is brought to light without traversing the great oceans^ viz., in watching the development of the bristles in the ventral branch of Hermione hystrix.
Those authors who, like Savigny, Milne-Edwards, De Quatrefages, Grube, and Ehlers place the Aphroditidse, Polynoidse, Acoetidse, and Sigalionidse in one family have certain grounds for this step. Thus all have biramous segments, all have ventral cirri on every foot; the scales are borne by similar segments ; the head has two eyes on each side, a median tentacle, a pair of palpi, and four tentacular cirri. Moreover, there is a certain agreement in regard to the arrangement and relation of the great nerve-cords, which in Aphrodita occur in a transversely elongated space between the ventral attachments of the oblique muscles, bounded externally by the hypodermic basement-tissue of the cuticle ; in Polynoe they occupy a hypodermic area between the ventral longitudinal muscles, the oblique muscles piercing the vertical at the upper and outer angle of the space, and being attached externally and superiorly to the cords. In the Acoetidse they are situated in the hypodermic region between the ventral longitudinal muscles (which are closer than in the Polynoidse), a thin layer of the former occurring between them and the cuticle. The great oblique muscles pass down do their upper and outer border. Lastly, in the Sigalionidse the space between the ventral longitudinal muscles is still more narrowed than in the previous group, and the hypodermic area for the nerves is thus increased in depth. Superiorly the arch is completely covered by the insertions of the vertical and oblique muscles; and the latter do not pierce the former (which occupy the middle line), but are attached to the basement-tissue below them on each side of the nerve-area. It will be observed that there is a gradational narrowing of the ventral longitudinal muscles between the first and last mentioned groups.
On the other hand there are fair reasons why several authors adhere to the anmigement of these groups in separate families. Thus, for instance, the Polynoidse diverge
(zOOL. CHALL. EXP. PART XXXIV. 1885.) LI 5
84
THE VOYAGE OF H.M.S. CHALLENGER.
considerably from the Aphroditidse in the structure of the head, the arrangement of the eyes, the armature of the proboscis, the form of the bristles, the presence of the antennse (lateral tentacles of Grube), and other points, including the relationships of the great nerve-cords (which in the Aphroditidse have the ganglia of the three first feet sepa- rate, i.e., not united by transverse commissures), while such features are coaimon to the rest. Taking the known facts into consideration, it wonld appear to be prudent at present to leave the various groups above mentioned in separate families as Malmgren has done.
The family of the Aphroditidse is well represented in the collection, as may be inferred when it is stated that Kinberg in his Annelids of the Voyage of the Swedish frigate “ Eugenie ” mentions seven ; that only two are noticed by Grube in Semper s great Philippine collection, and the same number (two) amongst the Annelids dredged in the German exploring ship “ Gazelle” ; further, that none occur in the same author’s Annulata CErstediana, and none in Schmarda’s series ; while the present collection affords no less than thirteen.
All the Aphroditidse are deep-water forms, a feature at once dividing them from the other groups formerly mentioned. They are evidently active feeders, though their opportunities for such may sometimes be limited.
Mr. HaswelG has recently added to our knowledge of the segmental organs of Aphrodita. Their external apertures, he observes, are situated on the ventral surface close to the base of the parapodia, but there is no perforated papilla as in Polynoe. The segmental organs consist of small, flattened, and somewhat sigmoid sacs.
Aphrodita, Linnaeus.
Aphrodita acideata, Linnaeus.
Habitat. — Dredged by H.M.S. “Knight Errant,” in the Faeroe Channel at Station 7, August 12, 1880, in 530 fathoms; bottom temperature 46° '5, surface temperature 5 7°; ooze. A small specimen.
AjAirodita australis, Baird (PL VII. figs. 6, 7 ; PI. VIa. figs. 4-7).
Aphrodita australis, Baird, Journ. Linn. Soc. Lond., voL viii. p. 176, 1865.
Habitat. — Dredged at Station 163b, off Port Jackson, Australia, 3rd June 1874; lat. 33° 51' 15" S., long. 151° 22' 15" W. ; depth, 35 fathoms; bottom temperature 63°, surface temperature 69° ; hard ground.
The single young specimen is about 26 mm. in length and 18 mm. in breadth. Contrasted with an example of Aphrodita acideata of the same length, it is very much broader, wdiile the more abundant lateral hairs give it a woolly aspect. The dorsal felt
1 Proc. Linn. Soc. N. S. Wcdes, vol. yii. p. 608.
REPOET ON THE ANNELIDA.
35
is coated with greyish mud, amongst which a Sabdlaria was found, so that there must have been soft parts amongst the rocks. The colour of the beautiful lateral hairs differs from that of the common species, being of a more delicate green, the whole indeed inclining to a mixed pale bluish appearance. These lateral hairs are also much longer. The segments are forty-two, the posterior region, which as usual in the group is distinctly narrowed, bearing a proportionally larger number. The dorsal felt is extremely tough and dense, more so than in Aphrodita actdeata. The head is a rounder and comparatively larger organ than in the latter, having in front a short blunt conical tentacle or papilla, immediately behind which (in a transverse line) are two eyes on each side, the anterior pair being wider apart than' the posterior. They are small but distinct black points, entirely sessile.
The body is more depressed than in Aphrodita actdeata, and instead of the rows of stiff, sharp, dark brown spines which flank the sides in the latter, a series of light bronze- coloured spines project outwards amongst the hairs. The tips of these do not taper much, but end in somewhat broad points (PI. VIa. fig. 4), which are covered with minute chitinous spikes, so that the surface is rasp-like. In consequence of this structure the tips are generally coated with extraneous organisms of various kinds. The inner tufts of bristles, which curve round and backward amongst the felt of the dorsum, are broad at the base but taper to a slender tip, also marked by slight roughnesses or points. A translucent filiform prolongation of the tip is common (PI. YIa. fig. 5).
The ventral bristles anteriorly are for the most part dart-shaped (PL VIa. fig. 7), but the tip is seldom entire. It is easy to pass from this form to the slightly hairy kinds as we proceed backward, and then to the form typical of the group (PI. VIa. fig. 6), the figure rej^resenting one of the smaller bristles from the inferior division of the ventral branch of the foot, these being somewhat less liable to injury than the strong upper ones. In the latter the tips are almost all broken, a mere trace of the hairy part remaining. Posteriorly the ends of the stout superior ventral bristles are rather more tapered.
The dense tufts of fine iridescent bristles which spring from the middle of each foot are longer than in Aphrodita aculeata, and on the whole the basal regions of the shafts are more slender. They taper to an extremely fine tip.
The dorsal cirri occur on alternate feet, and are long and finely tapered toward the tip, which is slightly clavate. The surface is smooth, while internally a very distinct muscular band occupies the middle. A peculiar network of fibres, apparently sub-hypo- dermic, is also present. The ventral cirri have the same position as in Aphrodita aculeata, but the tips are more finely tapered and more definitely bulbous. Though the cirri of the dorsum are alternate in Aphrodita, the alternate arrangement of the bristles present in Lcetmonice does not occur.
Dr. Baird does not mention eyes (though they are present) in his examples, which were considerably larger than the foregoing. They came from Port Lincoln and Van
36
THE VOYAGE OF H.M.S. CHALLENGER.
Dieman’s Land. The larger example, in the British Museum, measures upwards of 4 inches in length, and an inch and a half in transverse diameter at the widest part. It is much tapered posteriorly. The coarse nature of the dorsal felt alluded to above is conspicuous in this large specimen. The large flattened yellowish-brown bristles with a metallic lustre, curve backward over the felt. The sides are furnished with the long iridescent hairs, which on the ventral surface have a rusty brownish hue. The dorsal and ventral cirri agree in having a dilatation below the slender clavate tip, a feature not seen in the small example from the Challenger. The dorsal bristles on the w^hole agree with the latter in structure, but the ventral show no trace of the woolly region, probably from friction. A curious arenaceous Foraminifer is parasitic on the middle of the ventral region of the body. The other large specimen (from Van Dieman’s Land) unfortunately had been sent in a dried condition, so that it is doubtful if it quite agrees wdth the former. All the dorsal bristles appear to be much worn. Mr. Haswell, in his excellent account of the Australian Aphroditacese,^ states that this species is not so common as the European form ; indeed, he had only seen a spirit-preparation. He mentions Port Stej)hens in addition to the above localities.
Aphrodita echidna, De Quatrefages ? (PI. VII. figs. 1,2; PL VIa. figs. 2, 3).
Aplirodita echidna, De Quatrefages, Anneks, i. p. 197.
Habitat. — Trawled at Station 307 (in the Strait of Magellan), January 4, 1876 ; lat. 49° 24' 30" S., long. 74° 23' 30" W.; depth, 140 fathoms ; bottom temperature 7°’6 C., surface temperature 53'°0; blue mud. Also at Station 309 in the same Strait, January 8, 1876 ; lat. 50° 56' S., long. 74° 15' W.; depth, 40 fathoms; blue mud.
The body of the large example measures about 53 mm., and has a diameter of 25 mm. at its widest part, viz., the middle.
The dorsum has a dull muddy green hue, marked at the sides by the points of the spines which barely project through the investment. The lateral regions are of a dusky ferruginous colour, and the same tint occurs on the anterior third of the ventral surface. The odour of the specimens is peculiar. The segments are about thirty-five. Numerous long-stalked Pedicellinse occurred on the large specimen.
The head is smooth, prominent, and rounded, with the tentacle in the median line at the anterior border ; the tentacle is barely the length of the head, and the swelling below the tip of the long distal division seems to be slight. It is covered with a blackish deposit. No eyes are visible. The palpi have the usual length.
The dorsal felt is formed of very distinct and nearly equal fibres, which have the ordinary hooked tip. The dorsal bristles (PL VIa. fig. 2) are short, and for the most
1 Proc. Linn. Sue. N. S. Wales, vol. Yii., June 1882.
REPOET ON THE ANNELIDA.
37
part enveloped by tbe felt, and they constitute a kind of double row along the dorsum. The shafts of these are somewhat ovoid in transverse section in the middle, and more Hattened towards the tip. Their texture is brittle, and they have the usual lustrous brown appearance. Below the dorsal spines is a tuft of very long, simple, flexible bristles, generally coated from base to apex with fine mud. They are most conspicuous posteriorly, forming a dense mass on each side superiorly.
The envelopment of the dorsal spines by the felt causes the upper ventral bristles to appear very prominent. They resemble stiff brown spines with a slightly pilose tij?, which is frequently broken. The inferior group (a short distance outside the cirrus) are much more slender, but conform to the same type, their distal regions remaining more perfect from their sheltered position (PL YIa. fig. 3 — one of tlm inferior or smaller group, it may be wdth the tip in process of reproduction). Anteriorly, as in allied forms, the tips of the bristles are in many cases quite smooth. The internal portion of the bristle is also less brittle than the exterior, the former remaining as a long appendage after fracture of the latter.
The dorsal cirrus is long and tapering, and ends in a slightly enlarged tip. Its surface, with the exception of a microscopic q»^^lpocil or minute papilla or two, appears to be smooth. The ventral cirrus is subnlate, tapering from base to apex, and with more numerous papillae. The tip appears also to be slightly bulbous, but it must be recollected that spirit greatly alters these delicate structures.
The scales as usual are fifteen pairs, and both dorsal and ventral surfaces are coated with a ferruginous deposit. A few microscopic papillae occur on the upper surface. Some fragments of long hair-like greenish confervae were attached to the felt on the dorsum of one specimen.
The proboscis {'pharynx exsertilis of Kinberg) and alimentary canal conform to the typical structure. The former is much compressed laterally. The body-wall appears to agree in structure with Aphi'odita aculeata, except that the dorsal cuticular papillae are somewhat longer and have simple pointed tips, whereas in Ayylirodita aculeata they have a slight terminal button or dilatation. The nerve-cords occupy the middle of the same wide area between the oblique muscles inferiorly.
In one specimen the intestinal canal contained much granular debris, forming a pulpy mass, in which shreds of cuticle and numerous bristles apparently of one of the Spionidae occurred.
This difiers both from the Aphrodita alta and the Aphrodita longicornis of Kinberg, the former coming from the South Atlantic near Rio Janeiro, and the latter from the same ocean off the River Plate. From the first it is distinguished by the prominent spines of the dorsum (they are hidden under the felt in Kinberg’s form), the absence of the elevated ocular region on each side, and the shape of the tentacle, which has a shorter and wider distal region in Aphrodita cdta ; whilst from Aphrodita longicornis it diverges
88
THE VOYAGE OF H.M.S. CHALLENGER.
botli in regard to the shape and length of the tentacle (the organ being much shorter in Aphrodita echidna), the form of the head and the absence of eyes. There is nothing in the description of the Aphrodita echidna of M. de Quatrefages to give certainty in contrasting it with others, but it agrees with the present form in having the felt in a single layer, transfixed by the dark brown spines, and in all probability the forms are the same. Nothing further is known of the French specimen than that it was procured off South America by A. d’Orbigny.
Aphrodita intermedia, n, sp. (PI. I. fig. 6 ; PI. VIa. fig. l).
Dredged at Station 23a, off Sombrero Island, West Indies; lat. 18° 26' N., long. 63° 31' 15" W. ; depth, 460 fathoms; Pteropod ooze: also at Station 2a, off Culebra Island, West Indies, lat. 18° 38' 30" N., long. 65° 5' 30" W. ; depth, 390 fathoms; Pteropod ooze.
A minute ovoid form, measuring about 5 mm. in length and about half as much in breadth.
The whole dorsum is enveloped in a whitish coating of Foraminiferous mud, which obscures all the parts of the animal except a few of the ventral bristles along the anterior edge. In the natural condition it is, indeed, difficult to distinguish the anterior from the posterior end. The dorsal surface is convex, the ventral flattened. On the latter surface the spaces between the feet are occupied by masses of the same granular whitish material, which also in some parts projects beyond the tips. The whitish coating over the dorsum is composed of minute round spicular bodies, white sand-grains of various shapes, and debris. The layer is friable, and has only a few fine hairs in its composition. The latter have nearly the delicacy of those of Aphrodita, and pass from the upper division of the foot in beautiful pale iridescent tufts. They are so mixed with the little white spiked bodies and other debris, that it is difficult to detach them, especially as there are traces of minute serrations at the tip. No dorsal spines are visible in the example, a feature probably associated with the presence of the dense coating of the whitish granules.
The specimen, unfortunately, is much softened, so that a minute description of the cephalic region is difficult. It is at once seen, however, that the shape of the head corresponds to that in Aphrodita, not Lcetmonice — being pear-shaped with the broad region posteriorly. Onl}" the base of the tentacle remains in front. No eyes are visible, and no ocular peduncles. The palpi are of moderate length, and furnished with elongate papillse, which from their pointed tips resemble spines.
The ventral bristles occur in groups of three or four, supported by a spine in the slender foot. The tip of each passes off from the shaft at a characteristic angle (PL VIa. fig. 1). There is a very well-marked spur as in Lcetmonice, above which is a region densely joilose almost to the terminal hook, and placed on one side like the setose tip in the
REPORT ON THE ANNELIDA.
39
ordinary examples of tlie genus just lueutioued. lu tins remarkable form, however, the region is still further continued in the shape of a translucent filiform villose process which projects considerably beyond the hooked tip. The morphology therefore of this bristle is most interesting, since it shows an intermediate condition between the simply villose Aphroditacean bristle, and that with hook, spur, and pinnae characteristic of Lcetmonice.
The scales are very thin and quite smooth.
Hermione, Blainville.
Hermione hystrix (Savigny) (PI. YIII. fig. 3).
Halitliea hystrix, Savigny, Systeme des Ann61ides, p. 20.
Habitat. — Two small specimens were procured at St. Vincent, Cape Verde Islands, July 1873.
They quite agree with those from the Channel Islands, the South of England, and the Mediterranean. It seems to me that the two closely allied forms, viz., Hermione hystrix and Hermione hystricella, De Quatrefages, require further investigation.^ The prickly dorsal bristles which pass from tlie great spines inward over the back occur in many of the family, and a ventral bristle with a blunt tip from Hermione hystrix would quite agree with Kinberg’s Hermione hystricella, De Quatrefages. The definitions given by M. de Quatrefages as to the distinctions of the species are also open to doubt.
Lcetmonice, Kinberg.
Lcetinonice jilicornis, Kinberg (PI. Va. fig. 7).
Lcetmonice filicornis, Kiuterg, Ofversigt k. Vetensk.-Akad. Eorliandk, 1855, p. 382; andFreg.
Eugen. Resa, p. 7, Tab. iii. f. 7, &c.
Habitat. — A single small specimen was dredged in the Fseroe Channel by H.M.S. “Knight Errant,” August 12, 1880, at Station 7; lat. 50° 37' N., long. 7° 19' W.; depth, 530 fathoms ; bottom temperature 46°'5 F., surface temperature 57° ; ooze.
This form ranges over both sides of the Atlantic, being found on the shores of the United States and Canada, and off various parts of Northern Europe.
Lcetmonice producta, Grube (PI. IVa. figs. 1-8).
Lcetmonice j)roducta, Grube, Monatsber. k. Akad. zu Berlin, August 1877, p. 512.
Habitat. — K large number of all sizes were dredged off Kerguelen. One specimen at Station 149, on January 9, 1874 ; lat. 49° 8' S., long. 70° 12' E., from Accessible Bay ;
1 The Hermione chrysocoma of Baird is one of the Palmyracere, with long spinous dorsal bristles.
40
THE VOYAGE OF H.M.S. CHALLENGER.
depth, 20 fathoms; volcanic mud. Two examples at Station 149c, Balfour Bay, Eoyal Sound, Kerguelen, January 19, 1874; lat. 49° 32' S., long. 70° 0' W.; depth, 60 fathoms; volcanic mud. A considerable number from Balfour Bay in 20 to 60 fathoms. A large number at Station 149k, on January 29, 1874, off Christmas Harbour, in 120 fathoms. Further, two small specimens were dredged at Station 151, February 7, 1874, off Heard Island; lat. 52° 59' 30" S., long. 73° 33' 30" W..; depth, 75 fathoms; volcanic mud.
Many of the sjDecimens are large, some measuring about 100 mm. in length, and including the bristles about 50 mm. in breadth. The outline of the body is broadly fusiform. Segments from forty-four to forty-seven.
The somewhat triangular head has a slender elongated median tentacle, with a tip to which is attached a pear-shaped process, the bulbous end being suj^erior. On each side is a prominent rounded peduncle, bearing on the dorsal surface the rather small eye. The palpi spring from the anterior border of the snout above the oral aperture, and are separated at their bases by a peculiar fimbriated lobe (facial tubercle of Kinberg), which may well subserve a tactile function. They appear smooth and glistening to the naked eye, but microscopically their whole surface is covered with pointed cuticular papillae. The palpi are gradually tapered from base to apex, but do not appear to be quite round, a ridge occurring along their dorsal aspect. The aperture of the mouth is directed forward on the ventral surface, the post-oral cuticle being boldy ridged longi- tudinally. Behind the ocular peduncles a multilobate process- occurs, extending forward as a boundary along the outer border, almost to the eyes. It is apparently better developed in some specimens than in others.
The first foot is directed forward and bears a double tuft of simple bristles and two cirri (tentacular cirri of Grube), the dorsal longer than the tentacle or the inferior cirrus. They are situated on the posterior surface of the foot, but as the latter is directed forward they become external. Both have a stout basal segment, and the peculiar pear-shaped process at the tip. The bristles of this foot form a sort of tufted fringe all round the anterior region — dorsally, anteriorly and ventrally, and they consist of simple tapering bristles as in Lcetmonice jilicornis, coated with mud and parasitic growths of various kinds (PL IVa. fig. 1, representing one from the ventral group of the first foot). It is easily noticeable that the internal structure of the bristle differs from that in the Amphinomidse in being chitinous or fibro-chitinous throughout. The dorsal tufts are more elongated than the ventral, but they have a similar structure.
In the second foot the upper branch consists of a lower division of simple smooth bristles with finely tapered acute tips, and an upper group of stouter bristles (PI. IVa. fig. 2) having their shafts covered with chitinous nodules, while the tip is finely pointed, though curved, and in some slightly serrated. The lower branch of the foot, again, has two groups of bristles. (1) A dense brownish series, increasing in strength
REPOKT ON THE ANNELIDA.
41
from below upward, and furnisbed with elongated spinous tips. The stronger upper series are much more boldly spinous than the lower, each, however, having the same armature, viz., a double row of spines from the point where the shaft begins to diminish nearly to the tip, which is bare (PL IVa. fig. 4). (2) The upper division shows in some
only a single powerful brownish bristle (PL IVa. fig. 3) with a strong hook at the top of the shaft, and a series of about twenty-five spines beyond it ; while the tip for a considerable distance is quite smooth and gently hooked. This kind of bristle has much more numerous processes than the homologous forms in Lcetmonice Jilicornis from Shetland, yet the cirri of the latter are throughout of a more elongated character, even to the terminal pear-shaped processes.
The third foot has all the foregoing nharacters more decidedly developed, and the number of strong bristles in the upper group of the ventral series is increased. As in the second foot the ventral cirrus has undergone a change, being now a short process with a filiform tip, thus diverging from the elongated dorsal with its pear-shaped extremity.
The fourth foot, perhaps, presents a maximum amount of complexity in regard to its bristles. From below upward it shows— above the greatly diminished ventral cirrus — a dense series of the doubly serrated kind, the upper groups presenting a strong tooth at the commencement of the spinous row on each side, and since they are not opposite, an alternate character is impressed on the rows. From the papilla above spring one or
two very powerful bristles, with a basal spur at the tip, and apparently only a single
row of spines. Then from the upper papilla arises a still stronger bristle with the hooks and spines better developed. In the superior division of this foot there are no less than four groups of bristles : (1) An inferior dense series composed of bristles with compara- tively smooth slender shafts and simple tips. The shafts are finely striated longitudinally (by transmitted light), and the tips are shaiq), smooth, and finely tapered. (2) A vertical series of strong brown bristles (few in number), distinctly curved, with granular or
minutely nodulated shafts and delicately tapered tips. (3) Immediately in front of
the former is a group of simple bristles with long sharp tips like the inferior division of the dorsal series. (4) Superiorly is a set of bristles, more slender than the second group, but stronger than the third, and showing granulations, especially towards the tip.
The eighth foot still presents the four groups superiorly, the more robust being in front and somewhat below the dorsal cirrus. Ventrally there are four or five very strong though very brittle bristles which show an increase in the gap between the spine and the more numerous spikes beyond. The seventh foot bears dorsally a series of very long- spines, and a scale but no dorsal cirrus, the eighth having neither of the former. The spines of the seventh are not barbed at the tip, but they are fully so in the ninth group.
In an average specimen the following is the arrangement of the feet in regard to cirri and bristles : — the first presents a dorsal and ventral cirrus ; the third, a dorsa^. cirrus ; in the fifth the bristles have undergone great elongation, but are without traces of barbs ;
(ZOOL. CHALL. EXP. — PART XXXIV. 1885.) LI 6
42
THE VOYAGE OF H.M.S. CHALLENGES.
tlie sixth, a dorsal cirrus; the seventh, long dorsal bristles; the eighth, a cirrus; the ninth, abundantly barbed long dorsal bristles, with a few simple ones ; and so on to the twenty- eighth, which has a cirrus ; the twenty -ninth, a cirrus ; the thirtieth, dorsal bristles ; the thirty -first and thirty-second, dorsal cirri ; the thirty-third, bristles ; the thirty-fourth and thirty-fifth, cirri ; the thirty-sixth, bristles ; the thirty-seventh and thirty-eighth, cirri ; the thirty-ninth, modified small bristles ; the fortieth and forty-first, cirri ; and the forty-second, bristles.
Thus the general arrangement in regard to the structure of the feet throughout the chief jDart of the body, viz., from the fifth to the twenty-eighth foot, is that the segments bear cirri and long dorsal bristles or spines (with scales) alternately, the former having the chief bristle-tufts directed laterally, the latter dorsally. A typical spine-bearing segment, e.g., the nineteenth, bears dorsally a series of boldly curved dull golden bristles which sweep backward and inward over its own and the succeeding scale. These bristles are rather broad and flattened inferiorly, but taper to attenuate simple extremities, the greater part of the latter region being minutely nodulated. They seem to attain their greatest development in those species devoid of a dorsal felt, and thus probably subserve a protective function.. The next inferior group consists of a tuft of lustrous brown spines {setce glocliidece), about three-quarters of an inch in length, which slant backward in a conspicuous manner (PI. IVa. fig. 5- — representing an example in which the sheath at the tip has been removed). A characteristic feature when contrasted with LcBtmonice JUicornis is the opposite or nearly opposite position of the fangs, some showing them very slightly alternate at the. base. There are usually five pairs of these recurved fangs, though a few show six, the first and last being smaller than the others. This formidable defensive apparatus is sheathed in two chitinous flaps (PI. IVa. fig. 6 — from the ninth foot, and somewhat broader than usually seen). As mentioned, a few in the ordinary feet show alternate fangs (PL IVa. fig. 7); and one or two equally powerful bristles of the same length, but with smooth tips, also^occur. Below the foregoing is a tuft of simple bristles having striated shafts and very attenuate tips. With the latter exception the whole of the bristles are advanced upward, so that they alternate with neighbouring feet. In young specimens the strong spines on the dorsum are often simple at the tip, and the dorsal dull golden simple bristles stand nearly erect instead of curving over the scales as in the adult. The number and variety of parasitic growths and adherent debris on these and other bristles are remarkable, and indicate the rich nature of the ground on which they feed.
The ventral bristles (PI. IVa. fig. 8) have lustrous brown shafts, and project far beyond the feet on each side. The shaft has a long and curved hook at its distal end, then after an interval this is followed by a series of chitinous processes (arranged in a pectinate manner), which gradually diminish in size to the slightly hooked tip. Posteriorly, the bare part at the tip again lengthens as observed anteriorly (PI. IVa. fig. 3), and the spines diminish in length. There is thus a transition anteriorly and posteriorly. The central
REPORT ON THE ANNELIDA.
43
portion (tliat wliicli surrounds the cavity, and is streaked by transmitted light) is tougher than the exterior glistening part, and is often torn out in the form of a long appendage to the tip.
The scales amount to twenty pairs, and entirely cover the dorsum, the first and last being small. They are smooth, parchment-like, slightly iridescent organs, showing under a moderate power granular rows radiating from the attached portion. The segments to which they are fixed in an example having twenty pairs are : — -second foot, fourth, fifth, seventh, ninth, eleventh, thirteenth, fifteenth, seventeenth, nineteenth, twenty-first, twenty-third, twenty-fifth, twenty-eighth, thirty-first, thirty-fourth, thirty-seventh, fortieth, forty-third, and forty -fifth. They are symmetrically arranged, as usual, on both sides. No dorsal felt exists.
The dorsal cirri are long smooth organs, tapering from base to tip, and each is furnished with the distal pear-shaped process. As usual they occur on the feet devoid of scales. The ventral cirrus is a short subulate process, with a basal division. The tip scarcely reaches the point of exit of the inferior bristles.
The anus is dorsal and well-defined. The last pair of small scales arch over it superiorly.
The ventral surface of the body has many brownish cuticular warts, which also cover the sides of the feet. Some of the younger specimens dredged off Christmas Harbour show a much more villous condition of the ventral surface, and the villi are pale. This papillose state is very conspicuous over the post- oral area.
The play of colours even in the spirit-preparations is varied. The dorsum has a pale iridescent hue from the scales, upon which the lustrous dark golden inner dorsal bristles rest. These are flanked by the boldly developed long spines, which are deep brown, while the pale golden lateral tufts form a conspicuous fringe from head to tail along their outer border. Thus from the first the long spines form a median row flanked by two golden rows of varying shades. Lastly, the long lustrous ventral bristles constitute an interrupted series of palisades below. A large number of parasitic growths — sponges, Foraminifera, Diatoms, hydroid zoophytes, Polyzoa, Loxosomse, Ascidians, entangled worms, and others in tubes of sponge-spicules — occur amongst the bristles.
One half of the alimentary system of the animal is formed by the powerful oesophageal apparatus or proboscis, the comparatively short intestinal canal succeeding the latter- having its surface augmented by lateral diverticula, which in these subserves the purpose of the spiral valve in higher forms. The intestine generally contains a cjuantity of greyish pulp, showing a vast number of sponge-spicula, 'fragments of Diatoms, bristles of Polynoidae and other Annelids, fragments of sessile-eyed Crustacea, ova, a few small Mollusca, and all the organic debris usually found in rich mud. The proboscis is much more flattened from side to side than in Aphrodita acideata, but it seems to be composed of the same densely arranged series of fibres. It differs in its relation to the alimentary
44
THE VOYAGE OF H.M.S. CHALLENGER.
canal posteriorly, for its conical termination projects freely into the dilated digestive tube, which throughout is larger proportionally than in Aphrodita aculeata, and does not exhibit the siphonal bend at the termination of the proboscis. The jDapillee of the cuticle are less globular than in Aphrodita acideata, having a somewhat ovoid shape.
This species evidently occurs, as many of its congeners do elsewhere, in great pro- fusion at Kerguelen. It is briefly described by Prof. Grube^ from specimens procured by the German exj)loring ship “ Gazelle.” It w"as the only species of the genus got in the Expedition. Although perhaps not a typical representative of the group, it shows how insecure the notion of Kinberg and De Quatrefages was, that Lcetmonice diverges from Hermione in having the dorsal covering of felt over the scales.. This has been dwelt on by Baird, Grube, and Claparede.
Lcetmonice produGta, var. ivyvillei (PI. YIL fig. 3 ; Ph IVa. figs. 9-11).,
Habitat. — Dredged along with many other Annelids on very rich ground at Station 144a off Marion Island, December 26, 1873 ; lab 46° 48' S., long. 37° 49' AV. ; in 50 to 70 fathoms; surface temperature, 41°’0; volcanic sand. Another Station is off Prince Edward Island, in 150 fathoms ; and a third is Station 157 (about midway between the Antarctic regions and Australia), on March 3, 1874; lat. 53° 55' S., long. 108° 35' E ; depth, 1950 fathoms ; surface temperature, 37°'2;. Diatom ooze. The latter Station was also prolific in novelties..
This form on careful examination differs from the foregoing, even in external apjiear- ance. Thus the colour of the long dorsal' bristles is much darker and duller than in the typical Lcetmonice producta, and they also have less ‘‘sheen.” The ventral surface is more villous or papillose, especially over the- oral region. The head has somewhat shorter and more globular ocular peduncles, which, however, show no trace of eyes. The median tentacle is considerably longer, though it follows the same type. The process behind the ocular peduncle on each side is simple and much smaller. The length of the largest example is about 68 mm., and its breadth 25 mm., exelusive of the bristles. The number of segments is forty-three.
The great dark brown spines of the dorsum, traces of v/hich occur on the fourth foot, quite differ from those of Lcetmonice producta. They are rather stouter, especially at the tip (PI. ITa. fig. 9), and as a rule there are only three recurved fangs on each side, rarely four. The differences in general structure are shown by contrasting the figure just mentioned and fig. 5 of the same plate. The recurved teeth are directed more to one face than another, so that both are not well seen at the same time. Another divergence is the length of the inner dorsal bristles, which in the previous species form the con- spicuous series over the scales. In Lcetmonice producta, var. loyvillei, they are longer, and
1 Monatsher. d. h. preuss. Akad. d. Wiss. Berlin, August 1877.
REPORT ON THE ANNELIDA.
45
tteir curve at the tip less marked. The minute nodes toward their distal region are also larger and more evident (PL IVa. fig. 11), while the tip is smooth.
The semi pen n ate ventral bristles (PI. IVa. fig. 10), are distinguished from those of the previous species in being furnished with much more slender and elongated tips, but the terminal part or hook is shorter. The secondary processes or pinnae are more numerous, more slender, and more elongated. Moreover, throughout the lower two thirds it is observed that these show a shght enlargement toward the centre, so that the process is somewhat fusiform. Instead of the comparatively small number of these pinnae in the typical LcBtmonice producta, there are upwards of seventy in the present form. The basal hook is pro]3ortionally smaller, and it is often broken ; indeed the entire bristle is very easily injured, so that it is rare to have the pinnae perfect. Parasitic on the bristles were many beautiful thecate Infusoria with Diatoms in their interior, and stalked Acineta-ioxms, externally; and to one bristle a small Terehratida adhered. Foraminifera and Diatoms occurred in swarms on these organs.
Only eighteen scales appear to be jDresent ; otherwise there is complete similarity between this form and the foregoing, in regard to both scales and cirri. The* number of the scales is, however, of comparatively little moment ; the main distinction lies in the structure of the dorsal spines and ventral bristles. It is an interesting feature also that the long lateral bristles of the cirriferous feet have a nodular surface towards the tip, and that the head dilfers in form as well as in the- absence of eyes. The papillae of the cuticle are ovoid. This appears to be a well marked variety of the foregoing.
The example from 1950 fathoms had evidently fed on one of the Polynoidse allied to Evarne, and its intestinal canal contained shreds of skin, hosts of bristles, pieces of scales, numerous Radiolaria, and much