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Beet Peegmic de tet raid cg oe degen Se mprsididmtertoeetparical vs} eel oto aii IT Te tt asad Sesena bn Adie athe nobibebies SI eee Aeon thd an HVE TOFD aie Gy bo how ‘ Apher per ten een ree Do ye sabe Ps@ra-DO4 the dens ya. iv agi dabei be ta9e5 ge eared esl cmenlare eset rahe me sls eee anak toll oat neb0e oe caps pede) barron el ee ete Wee on er , 4 phe he hers ee fetes Leieiey Hp tg pba: saab seta npn see: eee tohad ae > seek pear? jeied ished soe beh ada net {rete eT he ab igh niet AF We ING teeta cecesetie ie Teegasertese ts of hates" ” Sibopeyedetrhreees ysoriettl cist: iret od) opal ie O) tv Ia Hey gp adeeb wath et ehatbey Parra te ayes abetelerhtaotadehee fetret et ak GPNER EG 9% § yotrarad@ others aes ve ke ip t Peete tr '» heteiye pads cs av ausiiobiaye 4 iiiaeelat oat “Write werevery pene pire bebe be buen abe bey osha h rbot ae emee: Ss ont Bim wba se Vetere Pubs mags , 2h atte dk bein ine pets Be arbeh-dehe pra cieges Fone caeree 4aneg rts 2 aE oO sure ued rate oF whe SeedboBedeh ben Ge ORB O~ MAD Bede y: MEW tbe AEN ma wed tton. O96 NaN terete gd ott verte eet} hsb “ 5 bay peo ovanee Pinte ey ohiearent4 Lee oh. ep bois ’ 0 se reed ip tain eae rideiaieh seat |04 i oe as + eres heeea reer © aheeds date tbo, wes ve tate fhe be bathaiied Belobsa, septate eine vipeijas Shore by ah dsbheteBatheOricictets bi od Bihebs Acholes-Geacted-+ sa bedaner, Bh Pica iy in y ¥ ays 4 AGRE AEA MAU Bo A ey tT MOR ine at - ! , Pd PARA PS Omeie f be, Wien f be es iF papa ae) } Nati MR ae x Mii , RM Rhee j y & ; le = MARINE BIOLOGICAL ASSOCIATION THE UNITED KINGDOM. VOLUME, EX ENS.) 1910-13. PLY MO ULH:: PUBLISHED BY THE ASSOCIATION. Agents in London :—Messrs. DuLAv & Co. Lrp., 37 Soho Square, W. The Council of the Marine Biological Association wish it to be understood that they do not accept responsibility for the accuracy of statements published in this Journal, excepting when those statements are contained in an official report of the Council. 371159 ERRATA. P. 61, line 18. For Shpoerodoridae read Sphaerodoridae. P, 61, line 20. For p. 338 read p. 38. P. 122, line 30. After “conclusion that,” ¢nsert—“ the hybrid larvee were of all types intermediate between the paternal and the maternal. In the following year Doncaster (5) decided that,” .. . P. 187, line 12. For 40,000 read 3000. P, 295, last paragraph. or “G. Southern” read “ R. Southern.” P. 307, line 89. For “these investments” read “thin investments.” P. 331, line 34. Delete this line and read, “The species has been recorded from the Shetland Islands by Jiiderholm (Kungl. Svenska Vetenskaps-akadenviens Handlingar, Bd. 45, 1909, p. 109), and from the Clyde Sea Area and the Firth of Lorne by Ritchie (Annals of Scottish Natural History, Oct. 1911, p. 223).” P, 351, line 6. For “ Phytisca” read “ Phtisica.” P. 355, lines 23-27. Transpose “ Brachyura” to immediately above “ Leucostidae.” CONTENTS OF VOLUME IX. (NEW SERIES.) List of Governors, Founders, and Members— July, 1910 . October, 1911 Report of the Council— 1909-1910 . 1910-1911 . 1911-1912 . Balance Sheet, 1909- 1910 ditto 1910-1911 ditto 1911-1912 BULLEN, G. E. Some Notes upon the Feeding Habits of Mackerel and certain aoe in the English Channel ‘ Crawsnay, L. R. On the Fauna of the Outer Western Area of the English Channel De Moreay, W. The Echinoderms collected by the Hualey from the North Side of the Bay of Biscay in August, ‘1906 Drew, G. Haroxp. The Action of Some Denitrifying Bacteria in Tropical and Temperate Seas, and the Bacterial Precipitation of Calcium Carbonate in the Sea Drew, G. HaRoLD. A Table showing Certain Cultural Characteristics of some of the Com- “monest Bacteria found in the Laboratory Tanks at Plymouth Drew, G. Haro3p. Some Cases of New Growths in Fish Drew, G. HARo3p. On the Precipitation of Calcium Carbonate in the Sea by Marine Bacteria, and on the Action of Denitrifying Bacteria in Tropical and Temper- . ate Seas : Ewes, E. V. Notes on the littoral Polychaeta of Torquay (Part III) Focus, H. M. Note on the Early Larvee of Nephthys and Glycera GaMBLE, F. W., and Drew, G. H. Note on Abnormal Pigmentation of a Whiting infected by Trematode Larve . HaspeEr, M. On a Method of Rearing Larve of Polyzoa HEFForD, A. E. Notes on Teleostean Ova and Larvee observed at Plymouth in Spring and Summer, 1909 : Henry, HERBERT’. A List of Blood Parasites of Sea Fish taken at Plymouth MarrTHEWS, ANNIE. Notes on the Development of Mytilus edulis and Alcyonium digitatum in _ the Plymouth Laboratory . lv CONTENTS OF VOLUME IX. MarrHews, DonaLp J. A Deep-sea Bacteriological Water-bottle . Mings, G. R. The Relation of the Heart-Beat to Sac and its pee on Com- parative Physiology : : : NIcHOLLS, GEORGE E. An Experimental Investigation on the Function of Reissner’s Fibre ODHNER, NILS. Some Notes on the Genus Cumanotus Orton, J. H. An Account of the Natural History of the Shpper-Limpet (Crepidula fornicata), with some remarks on its occurrence on the Oyster Grounds on the Essex Coast Orton, J. H. The Mode of Feeding of Crepidula, with an account of the current- producing mechanism in the mantle cavity, and some remarks on the mode of feeding in Gastropods and Lamellibranchs . Sexton, HE. W. The Amphipoda collected by the Hualey from the North Side of the Bay of Biscay in August, 1906 : ‘ ‘ : Sexton, E. W. Description of a New Species of Brackish-water Gammarus (G. chevrewat, n. sp.) : : Sexton, E. W., and Marruews, ANNIE. Notes on the Life History of Gammarus chevreuxr SHEARER, CRESSWELL. The Problem of Sex Determination in Dinophilus qyrociliatus SHEARER, C., De Moraan, W., and Fucus, H. M. Preliminary Notice of the Experimental Hybridization of Echinoids StecHow, E. On the Occurrence of a Northern Hydroid Halatractus Oe eae nanus (Alder) at Plymouth Tait, JOHN. Types of Crustacean Blood Coagulation Watton, Cuas. L. Kodioides borleyi, n. sp. Watton, Cuas. L. On some Colour Variations and Adaptations in Actiniae Watton, Caas. L. Notes on Various British Anthozoa WyNHOFF, GERARDA. List of Nemerteans collected in the Neighbourhood of Plymouth from May-September, 1910 : WooptanD, W. N. F. Notes on the Structure and Mode of Action of the “ Oval” in the Pollack (Gadus pollachius) and Mullet (Mugil chelo) ; ZIMMERMANN, KATHLEEN E, Notes on the Respiratory Mechanism of Corystes cassivelaunus Abstracts of Memoirs Pecondine Work dene at the Baap eet oie Laboratory . = 944. By-laws : : Weel: Bepamont: G. H. maEcERn ee: G. H. Drew PAGE 525 « ie 566 * 82 - 437 * 444° 199 * 561 * 288 * 572 7 600 ¢ 598. Notes on Teleostean Ova and Larvae observed at Plymouth in Spring and Summer, 1909. By A. E. Hefford, B.Sc. With Plates I and II. CONTENTS. Introduction . F Table of Occurrence Pelagic Eqgs— Ctenolabrus rupestris Serranus cabrilla Caranx trachurus Capros aper Trachinus vipera Trigla Callionymus lyra Pleuronectes Solea vulgaris S. lutea 8. variegata S. lascaris Zeugopterus Gadus : F G. minutus and G. luscus G. merlangus . Onos (Motella) O. mustela ; O. ‘‘species A” (?tricirrata BI ) O. ‘‘species B” - Raniceps raninus Clupea sprattus C, pilchardus Demersal Eggs— Labrus mixtus Blennius pholis Gobius paganellus . Lepadogaster bimaculatus L, gouani Zeus faber Bibliography ~- Explanation of Plates NEW SERIES,—VOL. IX. NO, 1. OCTOBER, 1910. A PAGE A. E. HEFFORD. bh THE observations which form the subject of the following notes deal with only part of the total material on which my studies of teleostean reproduction, pursued during the last two years, have been made; but as the other and larger part of the material consists of preserved specimens of the young (chiefly post-larval) stages of fishes collected during the four years 1906 to 1909 inclusive, it is more convenient to deal with the egg collections of the past year first and to treat the whole collection of young fry separately in a further paper. Systematic examination of tow-nettings for pelagic fish-eggs began on February 11th, and in the early months before the steamer was in commission, owing to the exigencies of weather and time which attend the use of a small sailing-boat, samples were taken only inside the Sound or in its near vicinity. As will be seen from the analysis of the collections given in Table I, there was a break of more than a month's duration, beginning early in March. From April to the end of August the s.s. Oithona was available, which not only rendered collections from the open-sea areas possible, but also permitted the use of the Petersen young-fish trawl, by which large quantities of fish-eggs could be obtained. The quantities so obtaimed were generally much too large to admit of their individual examination, and therefore it was my practice to pick out a portion of the total eggs for careful examination and frequently for hatching in the laboratory, while the residue was scrutinized as carefully as possible so that the rarer specimens should not be omitted. After a certain amount of practice one can recognize many of the familiar species by size and other peculiarities even with the naked eye, so that after a confirmatory microscopic examination it is possible to obtain a fair knowledge as to the identity of the majority of the species present in a plankton sample, and then exceptional eggs can often—though not, of course, without exception—-be discerned. In the case of tow-net samples, which were taken with coarse (24 strands to the inch) or medium (50 strands to the inch) nets at various depths from the surface to about 9 or 10 fathoms, care was taken to pick out every individual egg. The hauls were in most cases of 15 minutes’ duration, so that quantitative comparison is to some degree possible. Throughout this work I have conducted my observations having in view practical fishery questions—e.g. the locating of spawning areas, the duration of the spawning period, and the relative extent of the breeding of various species of fishes in the Plymouth area—rather than details of purely biological interest, and therefore my records of the characters of eggs and larvae have had special regard to points for purpose of ready identification at the various stages of development ; hence details of embryology have little place in this paper, NOTES ON TELEOSTEAN OVA AND LARVAE, 3 The ova and larvae of the majority of species occurring here are already more. or less completely known, thanks to the labours of Cunningham, Holt, MeIntosh, Masterman, ete. in this country, and to Ehrenbaum and other continental investigators, so that the main object of the descriptive notes which follow is to fill up gaps or to amplify those previous observations which still lack completeness. It is, perhaps, unnecessary to point out that records of such essential diagnostic characters as dimensions, additional to those which have been made on an extensive scale in investigations made in various parts of the North Sea and elsewhere, are of no little importance and value, owing to the local variation which occurs in such respects. In glancing at the general constitution of my egg samples, as shown in Table I, perhaps the most striking feature is the vast preponderance of those belonging to unmarketable forms. The species which afforded the most numerous pelagic eggs was the rockling, Onos (Motella) mustela, and not far behind this in abundance come the gold-sinny or rock- wrasse (Ctenolabrus rupestris), the boar-fish (Capros aper), and the dragonet (Callionymus lyra). Doubtless more eggs of such important families as the Gadidae and Pleuronectidae would have been taken if more off-shore collections had been possible. One may assume that the relative abundance of planktonic eggs, if sufficiently numerous samples are taken, is a fairly reliable index to the proportionate numbers of mature fish occurring in the area under observation at the spawning period. It is therefore to be expected that samples of planktonic eggs from inshore areas should consist predominantly of those from the littoral species of rockling and wrasse.* The same cause, however, does not explain the predominance of dragonets, boar- fish, and Norwegian top-knot over such forms as whiting, dab, plaice, and sole, which are marketable fish of much importance to our local trawlers and line fishermen. The general aspect of the case is that species which are regular objects of the trawlers’ pursuit are poorly represented in our egg samples. How far trawling itself is responsible for this condition of things is an open question, which in any case it is not in my province to attempt to answer here. But it is a note- worthy fact that the forms mentioned above are of such small size that they would to a great extent escape through the meshes of an ordinary trawl, and therefore stand the best chance of surviving on a well-fished ground. The result cannot be entirely attributed to local distribution of the mature fish, for the Norwegian top-knot has practi- cally the same distribution here as the sole, thickback, “merry-sole,” * It should be remembered that only one species out of the four Plymouth wrasses (viz. Ctenolabrus rupestris) produces pelagic eggs. f A. E. HEFFORD. and dab, as well as approximately the same spawning period, while the same can be said regarding Capros aper as compared with the gurnards. In the course of my laboratory observations, a point which has struck me as interesting and worthy of further definite inquiry, is the relative vitality of the eggs of various species, as indicated by the extent to which they are affected by the conditions under which they are kept while under observation, in the course of their development in the laboratory. On several occasions I have kept ova of different species in the same vessel of sea-water in order to watch the process of development and examine the hatched-out larvae. In such cases it frequently happened that one species would do well and produce healthy and vigorous larvae, while another would fare badly, and, if the embyro survived so long as to hatch out, the resulting larva would be more or less moribund from the outset and frequently crooked in shape. Notable among those whose vitality in the laboratory was considerable were the eggs and larvae of Jotel/a mustela, Callionymus lyra, and Ctenolabrus rupestris, while those which most often appeared to be adversely affected were Gadus, Trigla, and especially the rare forms Raniceps raninus and Serranus cabrilla. Itis to be expected that natural selection has effected that inshore—and sometimes even estuarine—forms like Motella mustela and Ctenolabrus rupestris should produce eggs which are capable of a wider range of environmental change (e.g. of temperature, to take the most obvious factor which operated in the cases under discussion) than those species which spawn in deeper water, where the surrounding conditions are of a more uniform character. In the case of Serranus cabrilla it is not sur- prising that ova produced in this neighbourhood, which must be at the extreme limit of the natural spawning area of this species, should be of less than average health. The laboratory temperature falls below that of the sea at times in winter, and in summer is gener- ally above it. In the hot weather my vessels containing eggs were put to stand in running aquarium water for the sake of coolness. The same should be done if frost is to be feared in winter. I kept my eggs in sea-water obtained from well outside the Sound, or in aquarium water which had been treated with animal charcoal and then filtered through a “ Berkefeld” filter.* * See Allen and Nelson, ‘‘On the Artificial Culture of Marine Plankton Organisms,” Journ. M.B.A., Vol. VIII, No. 5, p. 482, an OVA AND LARVAE. NOTES ON TELEOSTEAN (1) ‘ds soug (1) snqeqound +7 (1) snzeqound (1) peytguepiay num {snpeyg (G) sngejound 7 (LT) sng] (Z ) ‘ds snaoqydosnez snaaydosnez ‘(T) snyejound ; °7 (1) ‘ds] *(T) sugeqound 4 °7 *({) sngeqound *7 (7) snqnutur ; “y ‘(T) ‘ds supey *(T) sugnurtu 4 “4 ‘(g) snosny j “4 *(T) snqeqound +7 ‘(¢) snqeqound +7 (¢)snosnqj snpey ‘(e)snqgejound +7 ‘(T) ‘ds snpesy ‘(7) snqeqound +7 supe ‘(1) snqyejgound snaeqdosuez ‘(1) ‘ds] (1) ‘ds super ‘(g) ‘ds snpeyy S55q 1ay4O SOOH-HSIA OLOV'Tad = - Silage ow loaealiaes es alee de alee “IS alts UNGE|| = || ye |} a : o4 pe Avep PURBSILIT AL = ~ eh Race lise alt: |= Al ead lee a hee (Peel AG Soft Z “*N ‘pway amry = CHIE I ae 1h Se ee te TA te i Ne Ca eam res ilGallent: : nai ee Avg puersqy Ay - - se eee ee al eet emer cee eae ocr aloes mn AG) | aap ae aot panoy euoysXppa = - See eileen th || MS cm = = = P fF ye F @) | ers Sey ST CANN ‘gurog WB J | = 5 GP cats eae Le melee eli Seale ene ie nel Sa eG IReH AB cars ia _ HMO Suey FO ee i AIS eee era eM eT cc lieneiee Wren iieee is * ole Span RSL fae : ; ‘Avg pursaey | oe ee Cee kGnimRe ne Gee a ALAS — Gi ane eg PrpAuuas | a i SH ees tel hana aed ea Pamala ces Ip Sei | GL |e er HHO §. 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Onl S B] @ *ALITVOOT 9 A ear eaeer eee eee eealeal eye laule aul tes cer alee tele ae tae lle | 6 Sis Sy elce |e lelewele lela idle |e 2 |e 1s lee ole Bee olted silo sels Wee ie lresle | olla lee ae sl 9 a a ee Fe | ees NI acl ae tate arm Ze 2 fa = +O =) = ; = ese le Se ye a) a] Ss = a Oe irene ih eee ae } fc) aw NOTES ON TELEOSTEAN OVA AND LARVAE. ih I. PELAGIC EGGS. Ctenolabrus rupestris, L. Gold-sinny or Rock-wrasse. Hott (11a), p. 465, Figs. 23, 24, 28-30. HEINCKE u. EXRENBAUM (10), p. 266, Fig.17a-d. THIS species afforded the most numerous of the pelagic eggs taken during the months of May, June, and July. Belonging to a littoral species it was taken for the most part in near-shore tow-nettings, and was practically absent from the offshore Eddystone Grounds ; but there was one notable exception to this on July 20th, when several were taken one and a half miles south-east of the Eddystone. The egg, which is easily recognized by its small size, clear homogeneous yolk devoid of an oil-globule, small perivitelline space, and embryo with slender body and uniformly distributed pigment, has been described by Holt (op. cit.) from the south-west coast of Ireland and from the Gulf of Marseilles, and by Heincke and Ehrenbaum from Heligoland. The size of the egg is liable tomuch variation. The diameters recorded for the North Sea are 0°72 to 0:94 mm., for the Mediterranean 0°70 to 0°83 mm., and for the Irish coast 0°835 mm. The diameter of my eggs ranged from 0°78 to 0:90 mm., averaging ‘878 in May, ‘839 in June, and ‘822 in July. The first specimen was taken on the 7th of May, the last on the 20th of July. The newly hatched larva has a total length of 2°16 mm., the distance from snout to anus being 1-22 mm. The post-larval stages have been taken in the young-fish trawl from late June onward, but they do not appear in that relative abundance which one might expect from the commonness of the ova. This is probably due to the fact that they early seek the environment of inshore rocks, which is the habitat of the adult. Holt (11d, p. 125) speaks of young examples being common on the zostera beds of Cawsand Bay and the Yealm estuary, but judging from his later publication of a drawing of a young wrasse taken at Fowey (Marseilles, 1899), and erroneously identified as C. rupestris, it is probable that at this time he was confusing this species with another wrasse, probably Z. maculatus, the young of which are more commonly met with off the shores of this neighbourhood in summer. The later post-larval stages, like the early larva, are characterized by their lack of black pigment, which is limited to a large spot at the base of the posterior end of the anal fin and at the base of the caudal fin, and for the rest a little in the peritoneum, on the head and on the throat. eZ) A. E. HEFFORD. Serranus cabrilla, L. The Gaper. RaFFak xt (20), p. 19, Tav. I, Fig. 5, Tav. I, Figs. 1 and 3. Hotr (11f), p. 11, PL IV, Figs. 33-40. Four eggs with homogeneous yolk and single oil-globule were taken on August 27th and six on August 30th, in each case in plankton caught by the young-fish trawl in Whitsand Bay. In both cases the eggs soon developed unhealthy symptoms, and the characters of the embryo and larva must be considered with this condition borne in mind. The diameter of the eggs ranged from 0°92 mm. to 0'97 mm. and the size of the oil-globule from 0-14 to 0:15 mm. Before the formation of the embryo the surface of the yolk presents a roughened appearance, which may be due to a slight granulation im the peri- blastic region (cf. Holt, ep. cit., p. 12). Just befere the outgrowth of the free caudal region a few round, pale yellow chromatophores have appeared along the sides of the embyro, and smaller, somewhat in- conspicuous dark stellate chromatophores are fairly uniformly dis- tributed all over the body. As development proceeds the pigment spots Increase in size, single yellow chromatophores by the anus and in the mid-post-anal region being well marked. There is a little pigment of each colour over the oil-globule, but none on the yolk-sac. The hatched-out larvae were moribund in every case. The length of the newly hatched larvae (Fig. 15) varied from 184 to 230mm. The largest and most whole specimen measured 1:00 mm. from the snout to the posterior edge of the yolk-sac, and 1:26 mm. from snout to anus. The post-anal length of 1-04 mm. should have a trifle added to it to allow for the slight shrinkage which had taken place. The pigment has a very characteristic distribution, but varies somewhat in the size and number of the chromatophores. Black pigment is confined to the dorsal (anteriorly it may be dorso-lateral) region of the head and trunk, extending as a dorsal line to the caudal extremity. The yellow, which by transmitted light shows a greenish tint, is con- fined on the body itself to a few rather widely separated chromato- phores of large size. In an average case there are one on the head; one behind the otic region; two in the pre-anal dorsal line ; two lateral spots at the level of the posterior edge of the yolk-sac: one very large one over the anus and at the mid-post-anal point in the dorsal and ventral contours. Ramifications extend into the unpaired fins in the two latter cases. All the larvae observed showed one large dorsal mid-post-anal chromatophore, but the corresponding ventral pigment may consist of as many as four chromatophores. In the dorsal fin NOTES ON TELEOSTEAN OVA AND LARVAE. 9 there is a row of four to six large chromatophores, which usually exhibit upwardly and outwardly radiating outgrowths. The most posterior of the series is a little behind the dorsal chromatophore of the caudal region. There may be corresponding pigmentation in the anal fin, but it is not so extensive, from two to four chromatophores being the usual occurrence. The pre-anal fin may have a spot of yellow pigment near its margin, and invariably there is a small patch in its antero-dorsal angle, where the contours of yolk-sac and intestine converge. There is no pigment on the yolk-sac, but over the oil- globule there are one or two chromatophores of each colour. Normally the yolk-sac is somewhat elongated and bears the oil- globule at its anterior end, but the position of the oil-globule shows some variation, and in two cases was only just within the anterior hemisphere. The general form of the larva resembles that of Holt’s Fig. 33 (op. cit.), the characteristic features apart from pigmentation being the ovoid yolk-sac with anteriorly placed oil-globule and the broad pre- anal fin membrane. My late embryos and larvae all showed a marked tuberculated epi- dermis, which was doubtless a pathological condition, and may be regarded as diagnostically unimportant. On referring to previous observations by Raffaele and Holt of Mediterranean species, one finds conspicuous characteristic features which are common to all the above and to my Channel specimens, but there are also puzzling variations which, in the case of the Gulf of Marseilles specimens, do not seem to be altogether explicable as being the result of partial observations of more than one species. Raffaele (op. cit., p. 19) deals with the eggs of three species, which I give below with the dimensions :— Diam. of egg. Diam. of mm. oil-globule. Serranus (Centropristis) hepatus. . 0-78 0.145 S. cabrilla : : ‘ . 0°90 0-15 S. seriba ; : : . 0°90 07122 He figures the larvae of S. scriba and S. cabrilla (Tav. 2, Figs. 1-4), but does not give larval dimensions nor any further indication as to the specific pigmentation than can be derived from the uncoloured drawings. In size my eggs agree most with his S. cabrilla, although slightly larger. KRafiaele’s newly hatched S. cabri/la larva has the oil-globule centrally situated, while his S. Aepatus shows it anteriorly situated. The former has large mid-post-anal dorsal and ventral chromatophores 10 A. E. HEFFORD. In the unpaired fins of the newly hatched larva there is no pigment, but a four to five days’ old larva shows two large chromatophores on the dorsal and anal fins. Holt (op. cté., p. 11) examined several Serranoid eggs from the Guif of Marseilles and found among them unusual variability. The diameter varied from 0°72 to 0°89 mm. (for the most part lying between 0°78 and 0°84 mm.), and the oil-globule from 0°14 to 0°16 mm. He found that the embryonic pigment, which first appeared before the outgrowth of the caudal rudiment (the black generally, but not always, preceding the yellow), showed considerable variation, while there appeared to be no constant relation between the variations. As to the nature of the yellow pigment (often greenish yellow by trans- mitted light, and very pale by reflected light) Holt’s records (as well as Raffaele's) agree with my observations. Both black and yellow chromatophores are shown as a rule over the oil-globule, but never over the rest of the yolk-sac. In all Holt’s observations of the newly hatched larvae the oil-globule was anterior, the pre-anal part of the body slightly longer than the post-anal, the multicolumnar notochord had its vacuoles arranged fairly regularly in two series, dorsal and ventral. Pigment was disposed along the whole length of the dorsum, and was sometimes present on the sides and intestinal region. The yellow chromatophores followed the contour of the dorsum, and post-anally there were only two spots, a dorsal and a ventral, generally well defined, marking the centze of the post-anal region. The pigment on the head varied. There was always a spot at the angle formed by the intestine and the dorso-posterior profile of the yolk-sac, and also in the region of the urocyst, and sometimes others over the intestine. It is to be noted that Holt never observed any pigment in the embryonic fins of the newly hatched larva, but after three days black pigment appears in the embryonic fin—a row of chromatophores in the dorsal and one chromatophore in the anal fin, while black pigment, which increases with the growth of the larva, appears along the dorsal contour of the intestine (op. cit., Figs. 836 and 37). My larva, on the other hand, shows soon after hatching a row of yellow chromatophores in the dorsal fin. Otherwise it agrees with the youngest larval stages which Holt represents in Figs. 33, 34, and 35, and refers to Serranus (hepatus?). It is possible that the early appearance of yellow pigment in the dorsal fin is a characteristic of S. cabril/a, which distinguishes it from S. hepatus, the species probably observel by Holt. However, a still later stage observed by him (‘‘alévin dgé de quelques jours”), of apparently the same species, does show large yellow chromatophores associated with the relatively diminutive black ones in the embryonic NOTES ON TELEOSTEAN OVA AND LARVAE. sbi | fins, and also differs from the other larvae in having the black pigment of the body predominantly ventral instead of dorsal. A still further distinct variation is shown in an older stage with the yolk almost absorbed and the mouth open (but only 2°06 mim. long), which has much yellow and no black in the embryonic fin (op. cit., Fig. 40). With the material at his disposal and the small assistance derived from Raffaele’s incomplete observations, Holt was unable to come to any definite interpretation of these unusual and irregular variations of larval pigment; and the still further difference displayed by my larvae does not add light to the problem. Holt referred them, or at least some of them, to S. hepatus as the most probable parent, on account of their smaller size as compared with Raffaele’s, and because of the greater abundance of this species in the area from which his eggs were taken. He also admits the possibility of some S. cabrilla eggs being present among his specimens. As to the specific identity of my egys I have little doubt in ascribing them to S. cabrilla, which is a constant, though not common, inhabitant of Channel waters. As far as can be judged -from Ilaffaele’s in- complete records, the egg and larva agree with his S. cabrilla from the Bay of Naples, except that his figures show no pigment in the embryonic fin of the newly hatched larva, but only at a later stage. The dimensions of the egg and oil-globule show practical agreement. The date of the occurrence of the eggs agrees with Day’s record of the spawning season of S. cabrilla. The only other member of the genus known to British waters is S. gigas, Cuv. and Val, mentioned by Day as an occasional and accidental visitor, who also states that “in warmer climates it deposits its ova in shallow water.’ Ido not know of any description of the ova of this species. It does not seem likely, however, that it would spawn successfully here. Caranz trachurus, L. Sead or Horse Mackerel. Hott (11b), p. 9; (11d), pp. 116-20 and 340. (11f), pp. 27-31, Figs. 53-63. Canu (3b), pp. 63-71. Pl. V, Figs. 1-6. HeINcKE and EHRENBAUM (10), p. 277, Figs. 28-31. EHRENBAUM (5c), p. 234. Although this species was exceedingly common off the coast in the early summer months only one egg was taken. This occurred in the catch of the young-fish trawl taken near the Eddystone on 29th June. It was not closely observed till the following day, when the larva had * See also North Sea Investigations, VI. ‘‘The Reproduction of Caranx trachurus,” Journ, M.B.A., Vol. Ill, N.S., pp. 190-4 (1893-5). 12 A. E. HEFFORD. hatched out, but in an unhealthy condition and with the tail much bent. It was recognized by the larval pigmentation and by the totally segmented yolk, which bore anteriorly an oil-globule of 0°24 mm. diameter. The black and yellow pigment had a similar distribution to that in Holt’s and Ehrenbaum’s figures, but was not so strongly marked. On account of the deformity of the tail it was impossible to ascertain the total length. It measured 0°98 mm. to the posterior contour of the yolk-sac. Post-larval stages have occasionally appeared in the catches of the young-fish trawl in July and August. It would appear, therefore, that the fish spawns out in deep water, the drift of the eggs and larvae towards the coast requiring some little time. It will be noticed that my solitary egg was at the last stage of embryonic development when taken. Its unhealthiness also is in keeping with the general rule that the more the habitat of the species les in the open sea the greater the difficulty experienced in rearing the larva in the laboratory. In spite of the abundance of scad at this time in these waters, all those I was able to examine were immature, so that I inferred that the older and spawning fishes did not approach the coasts so closely. Against this, however, it must be stated that as the result of his investigations in the North Sea, Ehrenbaum (op. cié., p. 235) finds that Caranx trachurus favours as spawning places the shallow coastal areas from 10 to 25 m. depth, while outside the 30 m. line only few eggs were found. : Previous records of the occurrence of this egg in the Plymouth neighbourhood are confined to the observations of Holt (11d, p. 116), who obtained four specimens in July, 1897. The diameter of the egg was 0°81 to 0°93, and the oil-globule 0°22 to 0°25 mm. Capros aper, Lacep. Boar-fish or Cuckoo. The pelagic eggs of this species first appeared in tow-nettings taken on 14th June, and from the end of that month to the end of August our samples from the deeper water contained a well-marked pre- ponderance of these eggs, which were especially numerous at the beginning of July in the neighbourhood of the Eddystone. The embryonic characters were first described by Cunningham (4a, p. 10), who artificially fertilized ova in August, 1897, and Holt has published descriptions and drawings of the larval stages (11f, p. 26, Pl. V, Figs. 43-8). Pelagic eggs taken by him varied from 0:93 to 101 mm. in diameter (chiefly ‘97 to ‘99), and contained an oil-globule of 0°15 to 0165 mm. diameter. The average dimensions of my ova were :— diameter of egg, 0°946 mm.; diameter of oil-globule, 6:156 mm.; and the range of size was from 0°90 to 0:98 mm. for egg, and from 0145 to NOTES ON TELEOSTEAN OVA AND LARVAE. 1s) 0:17 mm. for oil-globule, which was frequently of a yellowish tint. The species may be readily recognized towards the end of embryonic development by the characteristic yellow and black pigmentation. The yolk is homogeneous, the oil-globule of a yellowish tint, and the capsule marked with fine corrugations. At about the time of the out- growth of the caudal rudiment black chromatophores appear on the head and in a line on either side of the body. A rather large Kiipfer’s vesicle is visible at this stage. Yellow pigment appears soon after- wards. One larva soon after hatching measured 1°40 mm. from snout to anus, and about 2°46 mm. in total length. Another had a total length of 2°02 mm., the distance from snout to end of yolk-sac being 1:08 mm. Black and yellow pigment occurs in rather large, stellate or dendritic chromatophores on the head and along the sides of the body. At about the level of the anus and in the mid-post-anal region there is a tendency for it to be more concentrated. The embryonic fins and the posterior extremity of the trunk are unpigmented. In spite of the abundance of pelagic ova not a single larval or post- larval specimen occurred in our young-fish trawl collections. The young fish appear to seek early the deeper parts of the Channel. The only specimen recorded up to the present is one of 154 mm. length, taken in September, 1906, by the Danish research steamer 7’hor, to the west of the Channel Islands (21d, p. 5). Trachinus vipera, Cuv. This conspicuous and very easily identified egg occurred frequently though not abundantly in our samples from the latter part of May to the end of August. The many bright yellowish-green oil-globules and the richly pigmented embryo and yolk-sac render it a conspicuous object in the tow-nettings. The diameter lay between 1:28 mm. (in May and June) and 1:1 mm. at the end of August. Post-larval stages were frequently taken from the end of June throughout the summer. TRIGLA. The Gurnards. There are five species of Trigla occurring in the waters off Plymouth, and the specific identification of their pelagic eggs, which are similar in character and show considerable overlapping in dimensions, is a matter of great difficulty and often an impossibility. The only circum- stances which ensure certainty of determination are when the spawn- ing fish are captured in quantity at about the same time and in the same area as the eggs, so that an extensive comparison can be made between the planktonic eggs and those taken from the ripe fish, 14 A. E. HEFFORD, Such, for example, were the conditions under which Holt (11), p- 31) was enabled to identify the eggs of 7. cuewlus taken in April and May off the west coast of Ireland, and similarly Ehrenbaum (5c, p. 248) has made extensive measurements of planktonic and artificially fertilized ova of 7. gurnardus in the North Sea, the main purpose in the latter case being to distinguish between grey gurnard eggs and those of the mackerel, which may to a certain extent coincide in diameter and size of oil-globule. The following table shows the dimensions of artificially fertilized Trigla ova from four species which have previously been described by various observers :— Species. Observer and locality. Month. Diameter. | Bee of oil- globule. | l T. gurnardus | Cunningham (Plymouth) April-May 1°45 0°30 3 | Holt (west coast of Ireland) -- 1°43-1°55 | 0°28-0°33 = | Ehrenbaum (Helgoland) June 1°256-1°258 | 0°25 i ¥ A July 17163-17446 | 0-25 T. hirundo Canu (Boulogne) May-July 1:5=L7 0°27—-0°29 a Ehrenbaum (Helgoland) July 17193 0°24 as = a | August 1°1-1°352 | 0°22 T. cuculus | Cunningham (Plymouth) April-May 1°45 0-30 53 | Holt (west coast of Ireland) | May 1°47-1°61 | 0°28-0°33 T. lineata _| Holt (Plymouth) | July 1°29-1°33 | 0°24 Besides the above four species, 7vigla lyra, L., commonly known as the Piper, occurs off Plymouth in appreciable quantities, but its ova have never been described. A further difficulty is introduced by the great variation which is shown by the eggs of the same species observed in different regions and at different times. In spite of this, however, Canu (3b, p. 72) has stated that, as regards the eastern part of the Channel, the eggs of 7. hirundo exceed in diameter those of all other members of the genus, but he does not give any actual observa- tions to support this statement. Out of my own collection, during the summer of 1909, I can only refer three eggs with absolute certainty to the Trigla genus. As to their specific identity I can say nothing, except that 7. lineata appeared to be the most common species off Plymouth at the time (July) and may possibly have been the parent fish. There are few descriptions of gurnard eggs and larvae on record at present, so it may serve a useful purpose for future comparison if I briefly give here my own observa- tions. On the 5th of July an egg was taken about 2} miles north-west of the Eddystone. Its diameter was 1-4 mm., and it contained.an oil- globule of 0:22 mm. Observed on the same day, the blastodise had almost half enveloped the yolk. Next day the embryo had formed ; NOTES ON TELEOSTEAN OVA AND LARVAE. 15 the myotomes were very distinct, and faint dark pigment was just appearing. Less than twenty-four hours after this a short caudal rudiment had appeared, and the body was beset with both black and yellow pigment cells. In the caudal region yellow predominated, covering the dorsal surface almost entirely, while of black there were only a few round chromatophores. Over the yolk-sac there were many black and yellow chromatophores stellate and cruciform in shape, and the pellicle of the oil-globule was covered with large black dendritic chromatophores, each with many ramifying rays. The rudi- mentary pectoral fins occurred as relatively large flap-like outgrowths. On the fourth day, when the embryo had wholly surrounded the yolk, yellow pigment had increased in intensity, especially in the post-anal part, and had appeared on the pectoral fins. Black dendritic and pectinate pigment patches, together with yellow pigment, were visible on the embryonic fin membrane. The otocysts were relatively small, and situated so that the distance from the posterior edge of the eye to the posterior edge of the otocyst was equal to the diameter of the eye. The oil-globule had shrunk to a diameter of 0°20 mm. The egg died before hatching. On the 8th July another egg of 1:28 mm. diameter and oil-globule 0:215 mm. was taken a short distance south of the Breakwater. The embryonic character as to pigmentation, ete., was quite similar to the above-mentioned, and fortunately this proved to be more healthy. On the 12th the larva was found to have hatched out and was then prob- ably over twenty-four hours old, the yolk being partially absorbed. Its length was 4°5 mm., the distance from snout to anus being 1°7 mm. At this stage it has a general resemblance to a Zeugopterus larva, but the body is less elongate, the anus relatively nearer to the yolk-sac, and the well-developed and richly pigmented pectoral fins especially characterize it as Trigla. There is no well-defined “snout” and the frontal region is peculiarly square. The bean-shaped otocysts, with two very small otoliths, are placed immediately behind the eye. The oil-globule is situated at the posterior end of the yolk-sac, between which and the anus there is a short but deep pre-anal fin. The rectum lies at right angles to the longitudinal axis. Dorsal to the gut above the middle of the yolk-sac is a conspicuous spherical swim-bladder. The pigment is bright canary-yellow and black. The whole of the head, except the eyes, is diffusely covered with yellow. In the eye black is beginning to appear. A continuous dorsal series of rather diffuse yellow chromatophores runs back from the head to a point about 1:1 mm. from the posterior extremity, beyond which both marginal fins and body are quite unpigmented. A similar but less 16 A. E. HEFFORD. intense line runs along the ventral contour of the trunk. Over the surface of gut and yolk-sac is a diffuse covering of pale yellow and faint dendritic black markings. The pectoral fins are intensely yellow, with black etching-like markings round the margin in radial arrange- ment. On the unpaired fins there are large dendritic black and yellow chromatophores, distally distributed for the most part. In the anal fin all the pigment is marginal, while in both dorsal and anal there are fine, black, pectinate markings along the edge of the fin, similar to what is seen in Zeugopterus norvegicus. A third Trigla egg of 1°34 mm. diameter was taken on 12th July, 14 miles south of Rame Head. The embryo had formed, but the oil was still in two globules of 0°25 and 0:13 mm. diameter, which joined to make a single globule next day. On the fourth day the larva hatched out. Its total length is now 445 mm., the pre-anal length being 175 mm. In general form and pigmentation it resembles the larva above described, which was observed at a somewhat older stage. It differs slightly from that, however, in having less black pigment in the dorsal and anal fins, nor is the marginal pigment of the pectoral fin as strongly marked. Next day its length has increased to 4°74 mm. and the yolk is almost absorbed. The pectoral fins are much larger and now show the marginal fringe of black very plainly. The mouth has become distinctly enlarged and already has the characteristic gurnard form. Two days later, on the 19th, the larva, with but a very small amount of yolk left unabsorbed, has lost the brilliant yellow colouring of the younger stages. It is still fairly well marked, although diffuse, over the dorsal surface of the head and trunk, but less distinct over opercuium, along the sides of the pre-anal part of the trunk and on the pectoral fins. In the unpaired fins the marginal pigment is much reduced. Black pigment has increased, especially on the pectoral fins, which it now covers from margin to basal part, but the marginal fringe is still the most dense. Dendritic chromatophores occur at intervals along the margin of the dorsal fin, but are very sparse on the anal fin except in the hypural area, where there is a rich supply arising at the ventral edge of the tail and ramifying over the fin membrane. On the corresponding dorsal side there is a faint indication of the same thing. Fine dendritic chromatophores occur on the body, being most con- centrated along the dorsal contour of the gut and along the ventro- lateral part of the post-anal region. Large otocysts containing rela- tively small otoliths are situated immediately behind the eyes. The large fan-shaped and heavily pigmented pectoral fins, and the head which is conspicuous from its well-developed jaws and operculum, give this larva a very characteristic appearance. NOTES ON TELEOSTEAN OVA AND LARVAE. a7 Callionymus lyra. Dragonet. Although most abundant in the months of March and April and of rare occurrence during the summer, the presence of these eggs in the plankton has been recorded from 11th February to 30th August, the first and last occasion of the year on which searches for fish-ova were definitely made. The diameter of the egg varied from 0°70 to 0-91 mm., the average for February to April being 0°796, May to June 0.803, July and August, 0°74. Post-larval stages from about 3:5 to 10 mm. were exceedingly common from April to August in the young- fish trawl material. PLEURONECTES. The paucity of the eggs of this genus in our samples is greatly due to the fact that, by the time the systematic collection of samples was commenced, the spawning season of the species occurring off Plymouth was more or less over. Moreover, the regions favoured as spawning localities by the plaice (P. platessa), dab (P. limanda), “merry-sole” (P. microcephalus) are in the deep water at some distance from land, to which there is a regular off-shore migration for the colder months of the year, in which period most of the spawning of these species occurs. The flounder (P. flesus) is an exception. Seven eggs of this species were taken in four “hauls” in the Sound between 17th February and 4th March, and post-larvae were common in May. No plaice eggs were taken, the spawning being practically over by the end of January. Previously published observations of the occurrence of plaice eggs are confined to the records of the obtaining of one egg on 12th February and 7th March, 1902, by F. Balfour Browne (2, pp. 607 and 609). I myself have obtained no specimen of the pelagic post-larva, nor is there any record of such having been taken at Plymouth, due, I believe, to their off-shore spawning region. However, I hope to have more to say as to the distribution of the young Pleuronectidae in a later paper. P. microcephalus eggs have been most frequently met with, but on five out of six occasions when this egg has appeared in my plankton samples, it has been only a single specimen. The first specimen was taken on 8th April and the last on 24th May, but before that period no off-shore tow-nettings had been collected. The diameter of the egg varied from 1:30 to 138 mm, A newly hatched larva measured 484mm. Only one post-larva was taken—in July, off the Eddystone. In previous years, however, this stage has been more abundantly represented in our samples. No eggs of P. limanda were taken, and on two occasions only (in NEW SERIES.—VOL. Ix. NO. 1, ocroBERr, 1910. B 18 A, E. HEFFORp. May and June) were post-larvae obtained. There is one record only of the occurrence of the egg of the dab in the Plymouth neighbourhood, viz. on 14th April, 1902, south of the Mewstone (2, p. 613), but this, again, is probably due to the dearth of samples from the off-shore areas. Solea vulgaris, Quensel. The Common Sole. The only remark to be made concerning the occurrence of this well- known and easily recognized egg is as to its rareness in our samples, only three specimens having come into our hands through the whole season, viz. one in February at the eastern entrance to the Sound, one in May in the Sound, and one in June in Cawsand Bay. This was probably due to the fact that no samples were collected during practi- cally the whole of March. This month, according to Cunningham (4b, p. 117), represents the height of the spawning season, which extends from the middle of February to the end of April. A further cause 1s the fewness of our samples during this period from the deep off-shore waters where the spawning fishes appear to occur in greater numbers. Pelagic young fish have likewise been very few, less than ten being taken throughout the season from April to the end of August. Solea lutea, Bonaparte. Solenette. Hott (11a), pp. 460-4, Figs. 9, 10, 46-52. (11f), pp. 87-9. EARENBAUM (5a), pp. 312-17, Figs. 31-5. One specimen of the egg of this species was taken in a tow-net at a depth of about 4 fathoms 14 miles N. by W. of the Eddystone on 2nd July. It was not perfectly spherical, and measured 0°80 by 0:34 mm. The yolk contained 14 oil-globules of a pale amethyst tint. On the day of capture the embryo had developed a short caudal rudiment. Dull pale yellow pigment was present on the yolk-sac and abundant on the body, and a few black chromatophores were also visible on each. On being next examined, three days later, the larva was found to have hatched out and almost absorbed the yolk. Its total length was 3°01 mm., the distance from snout to anus being 103 mm. Dull yellow pigment in large dendritic chromatophores was present on the body, head, snout, and along the margin of the unpaired fins, with the characteristic large patch at the commencement of the posterior half of the tail. There are two large patches and a smaller anterior one on the anal fin. On each well-developed pectoral fin there is a large patch of pigment, which does not appear to have been men- tioned by previous observers. NOTES ON TELEOSTEAN OVA AND LARVAE. 19 I have not found among my young-fish trawl collections this year any post-larval stages which I can with certainty ascribe to this species. Solea variegata (Donov), Thickback. CUNNINGHAM (4a), p, 23, Fig. 15. (4b), p. 90; Pl. XVI, Fig. 6; Pl. XVII, Figs. 1 and 2. Hott (11d), p. 137. This is the species of Solea which yielded the greatest total of eggs in our samples, fifteen altogether being taken for the season. Con- sidering the relative abundance of the parent fish, previous records for Plymouth are surprisingly meagre. Cunningham first obtained a pelagic egg of 1°36 mm. diameter in July, which he identified chiefly by comparison with the ovarian egg from a ripe female taken the previous May. In his treatise on the Common Sole, 1890, he figures the newly hatched and two days’ old larva, the former being 2°42 mm. in length. He describes the eggs as measuring 1°28 to 1:36 mm. in diameter, and therefore smaller than those of the common sole. It may here be mentioned that there seems to be no evidence for Ehrenbaum’s statement (5b, p. 143) that it is larger than that of S. vulgaris, at least as regards the Channel specimens of the latter species (cf. 7, p. 23), although observations in the North Sea (where S. variegata does not occur) have given a smaller diameter for the eggs of the common sole. Holt (op. cit.) records a single egg of 1:11 mm. diameter, taken in the Hand Deeps in July, which died before being completely examined; and Balfour Browne (2, p. 615) mentions the taking of one in the West Channel on 21st April, the diameter of which was 1:44 mm. My earliest specimens (eight in number) were taken by tow-nets 7 miles south-west of the Eddystone on the 8th of April, but I was not able to examine them carefully until a day or two later, when most of them had hatched out. One of the eggs had a diameter of 136 mm. The yolk showed a superficial segmented layer and con- tained about forty oil-globules, more or less uniformly distributed below its surface. The embryo had developed a free caudal rudiment of moderate length. There were many round pale yellow or straw- coloured chromatophores and fewer small black ones on the body of the embyro and on the yolk-sac. The length of a larva, measured about one day, or possibly slightly more, after hatching, was 3°17 mm., from the snout to the posterior edge of the yolk-sac being 1°5 mm. Three other eggs taken with the young-fish trawl in the Sound on the 3rd May were able to receive more attention. Their diameters 20 A. E. HEFFORD. were 1°34, 1°56, and 1°30 mm. The first had between forty and fifty oil-globules, varying from 0:015 to as large as 0:12 mm.; the second possessed about fifty and possibly more oil-globules of diameter from 0-015 to as large as 0°12 mm.; the third had only thirty-four oil- globules of diameter 0:°024-0:'12 mm. At the commencement of the development of the caudal rudiment black and yellow pigment occurs, the former colour in rows of small chromatophores displaying stellate forms. The larva, measured during the first day after the hatching, has a total length of 2°88 mm., the distance from snout to anus being 1°38 mm. The yolk-sac is very globular in shape. Yellow is the prevailing pigment, occurring in large, stellate chromatophores over the body, embryonic fins, and yolk-sac. Large, stellate, black chromato- phores are also to be seen over the yolk-sac, but they are not very intense, and they are less numerous than the yellow. Black, stellate pigment spots are also distributed along the body, chiefly on the dorsal region, but at the posterior extremity is a short series of three along the ventral contour. The most conspicuous black pigment is in a series of twenty-five large, irregular, stellate or dendritic chromato- phores along the margin of the dorsal fin membrane, extending from the occipital region to about 0-4 mm. from the caudal extremity. There is a similar series along the margin of the anal fin membrane, but consisting of only six chromatophores. This specimen is practically identical with the somewhat older specimen figured by Cunningham in “A Treatise on the Common Sole,” Pl. XVII, Fig. 2. Subsequent specimens of this egg were taken on 29th June 1 mile S.E. by E. of the Eddystone (1 egg of 1:29 mm. diameter), and on 26th August in about the same area (3 eggs, 2 of which measured 1:26 and 1°3 mm. in diameter), in each case in the catch of the young- fish trawl. One or two post-larval stages were obtained in July and August from the deeper layers of water between Plymouth Sound and the Eddystone region. Solea lascaris, Bonaparte. Sand Sole. Hour (11a) (Species I), p. 457, Pl. XLIX, Fig. 26; Pl. L, Figs. 34, 35. (11f) (? Solea lascaris), p. 84, Pl. V, Figs. 50, 51. On 14th March, 1910, a single egg with the segmented yolk cortex typical of Solea and containing many oil-globules was taken in the tow- net between Plymouth Sound and the Eddystone. Examined on the 15th March, its diameter was about 1:36 mm., and the oil-globules, which numbered more than fifty, showed a characteristic arrangement. They were not aggregated closely together in opaque clusters as in NOTES ON TELEOSTEAN OVA AND LARVAE. Pl Solea vulgaris, nor distributed fairly regularly about the yolk as in S. variegata and S. lutea, but were arranged so that the majority of them formed a circle round the yolk outside the margin of the blasto- derm, while there were also a number closely aggregated into a group at the vegetative poles. Four days later (on the 19th) the larva was found to have hatched out (see Fig. 11) and was then probably in its first day of larval life. Its total length was 3°46 mm. and its pre-anal length 14mm. The yolk-sac at this stage is very globular, the oil-globules being mostly in the ventral part of it. The head, body, unpaired fins and pectorals (which later become rather conspicuous), and yolk-sac are liberally sprinkled with dull, pale yellow chromatophores, from the well-marked rounded centres of which ramify dendritic outgrowths. There are black pigment spots having generally the same distribution but less numerous, especially on the yolk-sac and unpaired fins. The caudal region, where embryonic fin-rays are making their appearance, is unpigmented. Black is beginning to appear in the eyes. Two days later the yolk is reduced to about the size of the head, but still shows several oil-globules also reduced. The total length is about 3°7 mm.; from the snout to the anus about 1:52 mm. The eyes are black with greenish tints. The midbrain is well developed and the frontal region prominent—of the typical Solea type. The mandi- bular region is also well developed, but the mouth is not yet open. The gut now has a ventral bend above the posterior half of the yolk-sac. The pigment is practically unchanged, except for a con- centration into marginal patches along the unpaired fins—six in the anal and nine in the dorsal fin—each patch consisting of a mass of dendritic chrome-yellow, usually with a somewhat dense black spot in the centre. The pectoral fins are large, well pigmented, and generally in active movement. The following day saw a further reduction. of yolk, but there was still some left. The gut has now a pronounced U-shaped bend. The mouth is apparently about to open. The pigmentation of the unpaired fins shows further concentration into patches along the extreme margin. Yellow (dull brown by trans- mitted light) forms the greater part of each patch. The rest of the fins and body are also well covered with dendritic and irregular- shaped chromatophores. The frontal region is further developed and certainly constitutes a remarkable feature of the larva, but is nothing like Holt’s drawing of his “Species I—Solea (7)” (11a, Figs. 34 and 35). On the 23rd (viz. four days after the larva was first observed) it showed signs of ill-health and the tip of the tail had become shrunken 22 A. E. HEFFORD. and crooked. There is still a portion of the yolk left, although the gut is now completely looped. The mouth is now open and conical teeth are visible in the lower jaw. The head is relatively enormous, the midbrain of considerable size and forwardly protruding. The dorsal fin membrane is very deep over the head and anterior part of the body. At this stage the specimen was killed. A similar egg to mine was taken in July, 1890, in Clew Bay, and has been described by Holt (11a). The diameter was 1°38 mm., and the numerous oil-globules had a characteristic arrangement, different from that of the other known British species of Solea but resembling that exhibited in my specimen. The resulting larva, however, was very different from mine, and was especially remarkable for its peculiar cephalic contour caused by a precephalic vesicular expansion of the dorsal fin. The very slight occurrence of black pigment and the limited distribution of the yellow chromatophores in the unpaired fin (op. cit., Pl. L, Fig. 34) constitute another and probably more important difference between that larva and mine. The same authority obtained a second egg in the Gulf of Marseilles (11f, /oc. cit.) of 1°36 mm. diameter and of similar character in regard to its numerous oil-globules, and referred to as “? Solea lascaris, Risso.” He points out its similarity to Raffaele’s “Sp. A” (20, p. 43, Tav. 1, Figs. 32 and 33; Tav. 3, Figs. 4-9), which, however, has a smaller egg and more black pigmentation than Holt’s, in which latter character it approaches much more nearly to mine. Holt (11f, p. 86) is inclined to minimize the importance of this difference, remarking, “que le seul alevin que j’ai vu n’¢tait pas né sous le beau soleil du midi, chose a prendre en compte lorsquon parle de coloration,” and since the dorsal precephalic prominence is probably an abnormal and certainly not a constant character (Raffaele, for example, shows it in Tay. 3, Fig. 5 only, and McIntosh and Prince (16, p. 850), referring to a similar protuberance over the brain in a single specimen of Solea vulgaris, regard it as an abnormal feature), there is much probability that Holt’s larva from the Irish egg is the same species as mine, the latter being the more normal form. Ehrenbaum (5b, p. 149) does refer it to Solea lascaris, suggesting that Raffaele’s “Sp. A” on account of the smaller diameter of the egg may probably be Solea Kleint, Bp. NOTES ON TELEOSTEAN OVA AND LARVAE. 23 THE Top-KNOTS. (ZEUGOPTERUS.) I follow Petersen* in referring all three Top-knots known in the Plymouth neighbourhood to the genus Zeugopterus. Under this heading, then, we have to consider :— Zeugopterus norvegicus, Coll. (Scophthalmus norvegicus, Gthr.) punctatus (BL) (Rhombus punctatus, Gthr.) unimaculatus (Risso), Day. (Phrynorhombus unimacu- latus, Gthr.) ” 2 I have named them in order of the abundance of adults, so far as is known, in the neighbourhood of Plymouth Sound. Zeugopterus norvegi- cus occurs fairly commonly on the Rame-Eddystone ground, and I learn from fishermen that it has been taken in increased abundance of late years. The other two species are more-littoral, rock-haunting fish, and therefore do not lend themselves to capture in a trawl to the extent that 7 norvegicus does. Z. punctatus is not infrequently taken on the rocks between tide-marks, and sometimes finds its way into the shrimp-trawl. Z. wnimaculatus has not come into our collections in my own experience at Plymouth, and I can only state that it has been known to occur here.t+ Holt (11d, p. 128) and Balfour Browne (2, p. 600) have taken pelagic Zeugopterus eggs here in the spring months of 1897 and 1902, but the observations have not yielded sufficient information to enable a definite conclusion to be made as to their identity. I have been more fortunate in obtaining a large number of eggs and in having at my disposal the solution of the identity of one of the species, viz. Z. nor- vegicus, Which Ehrenbaum’s work at Helgoland has rendered available (5b, p. 210). 3elow I give a list of all the Zeugopterus eggs taken in my collec- tions in 1909. In the majority of cases they were kept until the just- hatched larva could be observed. It will be seen that they group themselves into two classes, those in Column I having a diameter of 0°75-0:90 mm. and an oil-globule of 0°095-0'15 mm., while Column II contains the larger eggs of 0:96-1:05 mm. diameter and 0'17-0°195 mm, diameter of oil-globule. * Report of the Danish Biological Station, XII, 1902-5, p. 26. + A specimen of Z. wnimaculatus was, however, taken in the trawl off the Eddystone in May, 1910, after this paper was in manuscript. 24 A. KE. HEFFORD. I. (Z. norvegicus). II. Z. punetatus (2 all this species). Diameter Diameter Date. of egg. of oil-globule. | Date. of egg. of oil-globule. 19-IV 82 ‘13 17-I1 1:03 “19 “4 86 "132 20-IL 1:04 a | ‘; *84 13 25-11 1:02 195 35 ‘87 —_ eS 1:04 “7, a ‘87 "125 3 1:02 18 a 88 "125 4-TTT 1:05 ks) ne 84 "125 a "99 (ca) a7. 20-IV “90 15 19-IV 101 a) eS 83 13 | 20-IV 2) 175 (2) - 845 “14 28-IV coe ‘175 or 84 125 3—V 96 18 > *85 10 $3 “8 18 12-V w 3) “LI a 98 18 18-V “80 095 12-V oo 18 . 84 "12 24—V 1:01 178 3 81 13 bs 98 17 24-V 86 13 95 “94 175 14-VI 82 2 A 5 92 175 3 82 let | i “96x 9G 7 4 80 12 28-VI 82 125 | te "82 1] ss “84 12 I have not the slightest doubt in referring all the eggs of Column I to Z. norvegicus. It will be noted by reference to Table I that the localities of their capture belonged in general more to the open-sea waters than is the case for the larger eggs of Column II. Most, if not all, of the latter I ascribe to Z punctatus. Not only do the dimensions of the egg and oil-globule fall into two fairly distinct groups, but also, as far as could be observed, the hatched-out larvae from the smaller eggs exhibited one type of characteristic pigmentation (identified by Ehrenbaum, op. eit., with Z. norvegicus), and those from the larger eggs another type of pigmentation, which is identical with that shown by Holt’s “Species X” (11b, Figs. 20 and 21), referred by Ehrenbaum, with a query, to Z. punctatus. I do not doubt but that this is its species. There is only one other species possible and that is Z wni- maculatus, a much less common fish in our area. Moreover, the latter has not occurred in our collections of post-larval fishes for the years 1906, 1908, and 1909, although Z. norvegicus is common and Z. punctatus NOTES ON TELEOSTEAN OVA AND LARVAE. 25 occasionally recorded in each year for the months April, May, and June. The difficulty, if not impossibility, of distinguishing the egg of Z. unimaculatus by its dimensions may be seen by a comparison of measurements of the eggs of this species, which are definitely known from having been obtained from ripe females (‘92-93 mm. for un- fertilized, and ‘90-99 mm. with oil-globule 16-18 mm. for fertilized ova *) with Column II in the above list. There is indeed one egg (marked with a ?) in the above series which I had some ground for regarding as possibly Z wnimaculatus. This was taken on the 20th April in Cawsand Bay. Unfortunately my recorded observations of this egg and the subsequent larva are very meagre. Just before the outgrowth of the caudal rudiment there was no pigment whatever on the embryo. The larva, which hatched out on the 24th April, had a length within the first few hours of larval life of 2:4 mm., the pre-anal length being 1:07 mm. This agrees almost exactly with Holt’s newly hatched specimen from an artificially fertilized egg of Z wnimacu- latus (11¢, p. 46,and 11f, Fig. 89). My brief notes upon the larva state that yellow was the predominant pigment, occurring in moderate-sized round and stellate chromatophores over body, yolk-sac and unpaired fins, except at the posterior extremity of the latter. Along the margin of the unpaired fins the pigment was dendritic. Black pigment consisted of numerous very fine dots, scattered with the yellow all over the body, fins, and yolk-sac. However, on the 27th the pigmentation had assumed the same form, which I found at the same stage in the other larvae hatched from the larger group of Top-knot eggs and which I regard as typical of 7 punctatus. Its identity with 7. wnimacu- latus, therefore, can be based only upon the dimensions of the egg and newly hatched larva, for which comparison there is still too little material. Of course there is the possibility of a close resemblance between the pigmentation of the later vitelligerous larval forms of the two species. It will be noticed that my above lists indicate a somewhat different period for the occurrence of the two groups of eggs, Group I (Z. norvegicus) being taken from 19th April to 28th June, and Group II (Z punetatus) from 17th February to 24th May. This difference is, however, probably more apparent than real, for before April I was not able to get any samples from further seaward than the entrances to Plymouth Sound, which would leave the habitat of Z. norvegicus neglected for the commencement of the season. I may now give a more detailed account of the characters and especially the pigmentation of the two undoubtedly occurring species. * Of. Holt, 11d, p. 128. 26 A. E. HEFFORD. Zeugopterus norvegicus, Coll. Norwegian Top-knot. The oil-globule is commonly pale green and the yolk-sac slightly rugose. Before the outgrowth of the caudal rudiment the periblastic pellicle of the oil-globule becomes visible, and fine black chromato- phores appear on the embryo and yolk-sac. They are very closely set on the former but less dense and mostly stellate in form on the latter. Over the whole embryo and yolk-sac there is a pale greenish yellow tint, but no separated, coloured chromatophores are visible as yet. Yellow pigment cells soon appear on the embryo and yolk-sac, and by the end of embryonic life they become conspicuously predominant. The black chromatophores show dense, rounded centres, from which fine ramifications proceed. Some of the few yellow chromatophores on the yolk-sac are stellate, with no specially large centre. On the trunk of the embryo, just before hatching, they are large and of vague outline, and so close together as to give the effect of a diffuse tinting along the whole length of the body, except the extreme caudal end. Black pigment is fairly uniformly distributed over the body, but shows some concentration into dorsal and ventral lines. A newly hatched larva measures 2°58 mm., with a pre-anal length of 116 mm. The snout projects conspicuously forward over the yolk-sac, and the anus is situated some little distance behind the posterior end. Ehrenbaum (op. cit.) describes the position of the oil-globule as usually at the middle of the ventral side of the yolk, but I have generally found it post- median. The pectoral fins are well developed. The pigmentation is extremely rich. A diffuse yellow effect, which is evident to the naked eye, is produced by the close proximity of the large chromatophores (greenish yellow by reflected light), which are distributed over the body, fins, and yolk-sac, most densely along the dorsal and ventral contours and the upper and lower margins of the unpaired fins. Black chro- matophores are more abundant, but smaller and much less conspicuous. A very characteristic feature is the stopping short of the pigmentation of the unpaired fins some little distance from the caudal extremity, so that this part of the larval tail is quite clear. The same arrest of yellow chromatophores is seen on the body itself, but small black pigment spots occur as far as the posterior extremity. A further conspicuous cnaracter of this species is the regular, fringe-like arrange- ment of both the black and yellow pigment along the outer margin of the unpaired fin, in the early stages of the larval development. The fringe-like effect is accentuated by the finely pectinate form of the chromatophores in this region. With the absorption of the yolk the intensity of the yellow pigment increases, although undergoing NOTES ON TELEOSTEAN OVA AND LARVAE. 27 little change in distribution. At 3 mm. length, for example, with the yolk almost absorbed, the general arrangement of pigment is as above. Head and body (except posterior extremity) appear of an almost uniform bright yellow. In the anterior half of the post-anal region the notochord shines white through the yellow, but in the posterior part the yellow is so dense as to obscure it. The coloured pigment is more concentrated on the head, along the dorsum, and about the rectum. The fringe formed by the line of closely applied dendritic or pectinate chromatophores along the margin of the unpaired fins, has a some- what brownish tint, which is not seen on other parts of the larva. In the anal fin, the frige begins some distance posterior to the anus. Black pigment is most dense on the ventral surface at the level of the pectoral fins and above the anterior part of the remains of the yolk-sac, where the vestige of the oil-globule now measures about 0:03 mm. A larva in which the yolk had been fully absorbed (ca. 5 mm. long) showed less distinctness as to the marginal pigment, and a concentra- tion—more particularly of the black chromatophores—in the pigment of the unpaired fins, about half-way between the anus and caudal extremity. Reticulate markings and small papillae on the epidermis are com- monly found both in late embryonic and in larval stages. To summarize the main facts ascertained for Z. norvegicus :—The spawning season, in the Plymouth neighbourhood, extends from April to June, probably beginning somewhat earlier than the former month. The pelagic eggs have a homogeneous yolk and a single oil-globule, which is frequently of a greenish yellow tint. The average diameter of the egg is 0°854 mm. in April and 0°817 mm. in May and June, the limits lying between 0°75 and 0:90 mm. The oil-globule measures 0095-015 mm., the average diameter being 0°122 mm. Embryonic yolk-sac and larval pigments are yellow (bright greenish yellow by reflected light) and black, the former predominating and being visible to the naked eye in the larva. The body and yolk-sac of the newly hatched larva are rather elongate, the total length being about 2°58 mm. (Ehrenbaum gives 2°52-2°76 mm.). The anus is situated a short distance behind the posterior edge of the yolk-sac, and at about 3’; the body length from the snout. The characteristic feature of the vitelligerous larva is the marginal pigmentation of the unpaired fins and their total lack of pigment in the caudal region. 28 A, E. HEFFORD. Zeugopterus punctatus (Bl.). Before the outgrowth of the caudal rudiment, the body of the embryo is beset with fine, black chromatophores which are chiefly, if not. entirely, on the dorsal surface. A few black chromatophores of larger size are found over the oil-globule, and there may be a small number of black spots on the yolk-sac, which is beset with a moderate number of roundish, yellow chromatophores. The latter pigment occurs in smaller spots on the head and body of the embryo. At this stage the perivitelline space is somewhat large, and the oil-globule is contained in a very distinct periblastic pellicle. With the growth of the free caudal region (Fig. 2) a marked increase in the size and number of chromatophores takes place. The yellow spots over the yolk-sac now acquire a stellate form, but on the embryo they still occur as patches disposed over the head and body. Yellow does not extend quite to the tip of the caudal extremity as the black does. The black chromatophores over the oil-globule have increased in number, and are the largest of all. Over the yolk-sac they are small and sparingly scattered. Minute black specks occur over the whole body, from anterior to posterior extremity, being most concentrated along the dorsal and ventral contours, dendritic out- growths from which extend to the embryonic fin, The pectoral fins are relatively well developed, and rudiments of the pelvic fins are visible. Three larvae observed soon after hatching (Fig. 9) had lengths of 2°90, 2°92, and 2:95 mm., the pre-anal lengths being respectively 1°44, 140, and 145 mm, The anus is therefore appreciably nearer the median position than in the case of Zeugopterus norvegicus, which otherwise it very closely resembles both in general form and in the distribution of its black and yellow pigmentation, the former occurring in mostly small, fine specks and the latter in large, stellate chromato- phores. The yolk-sac is elongate and bears the much-reduced oil- globule at its posterior end. As in the above species, the marginal pigment of the unpaired fins is of most diagnostic importance. Im- mediately after hatching I found some resemblance to the arrangement in Z. norvegicus, but the “fringe” formation is not so well marked, and within one day a striking change has taken place which appears to be quite characteristic of 7. punctatus, at least as distinct from Z. norvegicus, I can, of course, make no comparison with Z. bimaculatus, though it is to be noted that Holt’s newly hatched larva from an arti- ficially fertilized egg of Z. bimaculatus showed no fringe-like occurrence of pigment along the upper and lower margins of the unpaired fins, which I observed in my larvae of both species. NOTES ON TELEOSTEAN OVA AND LARVAE. 29 The one-day-old larva (length 3°42 mm.) shows a complete modifica- tion of the marginal fringe of the early stage by its breaking up into a series of separate large pectinate and dendritic patches, eight of these patches occurring in the dorsal and five in the anal fin. This exactly resembles Holt’s “Species X.” (11b, Figs. 20 and 21), which Ehrenbaum (5b, p. 206) considers as probably identical with Zeugopterus punctatus. Besides the above-mentioned characteristic, large, marginal chromato- phores, in each of the unpaired fins there is a series of fairly large stellate chromatophores, midway between these and the trunk contours, a single row in the dorsal and a double row in the anal fin. The last 0-5 mm. of the tail end is free from pigment in the fin membrane, but fine black chromatophores extend to the caudal extremity of the trunk. Yellow and black chromatophores are scattered fairly uniformly over the whole body, with a certain amount of concentration along the dorsal and ventral contours. On the third day (Fig. 10) still further local concentration into patches has taken place, but the general form of the pigmentation remains the same. Black pigment now appears in the eye, and the prominent pectoral fins bear dendritic yellow and rounded black chromatophores. The yolk-sac—with the yolk somewhat more than half absorbed—is very elongate, so that its ventral contour is practically straight. To summarize:—The spawning season of 7% punctatus in this neigh- bourhood extends from the middle of February to May. The pelagic eggs have a homogeneous yolk and one oil-globule. The diameter of the egg averages 1°03 mm. in February—March and 0:98 mm. in April- May, the limiting sizes being 0°92 mm. and 1:05 mm. The diameter of the oil-globule is 0:17-0:19 mm. Embryo and yolk-sac bear black and yellow pigment, the latter becoming conspicuously predominant in the late embryo and larva. Within one day after hatching, the larva exhibits characteristic stellate or pectinate and dendritic patches of yellow pigment associated with black in its unpaired fins, usually eight in the dorsal and five in the anal fin. The newly hatched larva measures 2°90-2°93 mm. (or possibly less), and the anus is only slightly Gf at all) anterior to the median point. GADUS. Our tow-net samples of Gadus eggs have not been sufficiently numerous to enable certain conclusions as to their identity to be made in the great majority of cases. This is not due to the absence or scarcity of members of the genus from our neighbourhood, but to the fact that the open-sea water some distance from the coast was not tow- netted till April, when the spawning season of the Plymouth gadoids is 30 A. E. HEFFORD. almost over. It is a fact of common knowledge to the Plymouth fishermen that the whiting (G. merlangus), bib (G. minutus), pout (G. luscus), and pollack (G. pollachius) are to be found nearer the shore in summer and autumn than in winter and early spring, when the breeding season occurs. This habit of migrating to deeper water for the colder months of the year they have in common with the other import- ant food fishes of the Plymouth district, such as the Pleuronectidae and gurnards. Very many more observations, both physical and biological, are necessary before definite conclusions can be made as to the real causes of these phenomena. The off-shore migration in winter and the corresponding approach to shallow water in summer may, in some cases, follow the seasonal distribution of food, but I do not think this is at most more than a partial explanation. The fact that the temperature of the water in the deeper parts of the Channel is appreciably higher than that of the more inshore parts of the Channel in the coldest months of the year, may be taken as a sufficient reason for the majority of fishes preferring to seek the outer grounds at this time. That conditions directly related to the phenomenon of spawning are involved in this migration (which certainly coincides with the ripening and liberation of the sexual products of most of the species) may be con- cluded from analogy with the cases already worked out under less complicated conditions, e.g. the plaice and cod, by Johs. Schmidt (21c), who has shown that these and other species show a special sensitive- ness to external conditions, especially of temperature, in relation to spawning, and therefore make special and well-marked migrations. So far I have not been able to obtain direct proof of extensive spawning of our four common Plymouth species of Gadus on the off- shore grounds because winter samples of plankton from such regions have not been collected, but the general fact may be taken for granted. With due precautions one may accept the occurrence of pelagic post- larvae, such as were captured in the young-fish trawl in April and subsequent months, as evidence giving more or less quantitative in- formation as to the spawning times and the relative extent of the reproduction of the various Gadus species in this neighbourhood. As far as can be judged from our takings of the small fry—and evidence from the fisheries points to the same conclusion for the adults— Gadus merlangus is the most abundant, very many post-larvae of this species having been taken, especially in May and June. Next in abundance comes G. minutus, which has an almost similar period of occurrence, if anything earlier than the whiting. The early pelagic post-larval stages of the pollack have always in my experience been less common than the two foregoing, but they are very abundant, at a NOTES ON TELEOSTEAN OVA AND LARVAE. 31 size of about 5 cm., close up to the rocks on the shores of the Sound. Post-larval stages of G. /uscws, which appears to be the least abundant of our four common representatives of the genus, are not infrequent but are never numerous, and they disappear from the samples some- what earlier than the other three.* Other species, such as G. morrhua (cod) and G. virens (coal-fish), are known, but are very occasional spawners in this neighbourhood. The specific identification of the pelagic ova by the form and pigmentation of embryo and larva is difficult and sometimes impossible, owing to their great similarity. Pollack, indeed, is said to show no yellow pigment at all in the embryo and early larva (14e, p. 171), while the other three species above mentioned as common to these waters do so to a greater or less extent. We cannot feel that we are on safe ground here, however, in dealing with individual fishes on this point, since the G. minutus larva is described by Raffaele from the Mediter- ranean and by Holt from the west of Ireland as having only black pigment, and in several cases the appearance of yellow chromatophores has been noted as an accompaniment to an unhealthy condition (ef. 11d, p. 140). The impossibility of separating the species with certainty by refer- ence to dimensions of the ova is indicated by the list which I give below of measurements of eggs taken from ripe female fishes. Species. Diameter of Ege. | Month. Authority. | G. minutus 0:95-1:07 a Ehrenbaum 1-02 | April | Cunningham 0-9906-1:0287 ; = MeIntosh 1:07 i Holt | A little below 1mm. | — Raffaele G. luscus | 1-O5= 1-15 ; — Cunningham 1137 | January | Holt G. pollachius 114 _- MelIntosh (artificially fertilized egg 113 — Holt * A general idea of the seasonal occurrence of the young fry may be obtained from the following particulars taken from my records for 1909 of catches of the young-fish trawl] :— G. merlangus . . First specimens (5-10 mm.) taken 28th April. a F ast BA (9-12°5 mm.) ,, 20th July. G. minutus : . First si (ca. 5 mm.) », 28th April. 3 ‘ . Last Wa (26 mm.) », 20th July. G. pollachius . oe HIrst #5 (22 mm.) », 24th May. ‘5 : > ast A (6 mm.) ,, 6th July. G. luseus . : . First a (6 mm.) », 28th April. Last - (7 mm.) » zndJune. vs : : F + This is the diameter of the largest unfertilized ovarian egg observed. an A. E. HEFFORD. Species. Diameter of Egg. Month. Authority. G. merlangus 1125 = McIntosh and Prince (artificially fertilized) 1:226-1:352 March | Heincke and Ehrenbaum (artificially fertilized) 1-1-1:226 April | Heincke and Ehrenbaum (artificially fertilized) 1:069-1:163 May Heincke and Ehrenbaum (artificially fertilized) 1:161-1:257 April | Williamson (artificially fertilized) From this evidence we can merely state that the average size of the egg shows an increase in the order in which the species are enumerated above, but the difficulty arising from overlapping is sufficiently formid- able, seeing that the largest size for G. minutus may be larger than that of the smallest whiting. The size of the newly hatched larva shows variation, which may to some extent be taken as a guide to the species, but on this point, to an even greater degree than is the case with regard to the egg, the data hitherto available are very scanty and afford little satisfactory informa- tion. With some degree of certainty we may take it that the G. minutus larva is the smallest, but no measurement of the product of artificial fertilization is on record. Holt (11b) records as G. minutus a larva of 2°75 mm. length hatched from a pelagic egg of 1:07 mm. The G. luscus larva 1s probably bigger, and that of G. merlangus is certainly bigger still. Ehrenbaum (5d, pp. 234 and 238) gives ca. 3mm. or somewhat smaller for the former and 3:2 to 5°5 mm. for the latter species. G. pollachius is so far too imperfectly known for us to state anything as to its larval dimensions. Holt (11d, p. 141) ascribes to this species certain pelagic ova of 1°40 to 1-45 mm. diameter taken in February, and mentions that the larva was 4-2 mm. long and had black pigment only. There seems to me to be little doubt as to the correct- ness of this identification, and it seems probable that further investiga- tion of the eggs and larvae of this species will prove them to be larger in general than those of G. merlangus. The dimensions of all the Gadus eggs measured during the season are as follows :— Diameter > 7-95 (696 9 ce 89. 1:00 9 1-OWe> V6 aay No. of eggs 3 l layed 1 1 1 April and Diameter -94 1°08 1:09 1:14 1:13x1:17 mm. early May No. ofeggs 2 1 2 3 1 Diameter 1:02 1:03 .1°04 1:05 1-065 S1512) mm: No. of eggs 1 1 2 il If | February June-August NOTES ON TELEOSTEAN OVA AND LARVAE. oo Gadus minutus and Gadus luscus. On 8th April a ripe female G. minutus was taken in the trawl about 7 miles south-west of the Eddystone. Eleven ova from this measured 0°91-1:02 mm., the average diameter being 0:939 mm. Artificial fertilization was attempted, but development was stopped by death at the blastula stage. Eight eggs taken in the tow-nets on the same day and in the same locality had diameters of 0:94, 1:08, 1:09, 1:13, 1:14 (two eggs), and 113x117 mm. (the last not exactly spherical). The embryonic form of all appeared practically identical. Round and dendritic black chromatophores first appear on the embryo, and at a later stage a diffuse yellowish tint appears over both embryo and yolk-sac. Just before hatching the black chromatophores are most densely distributed in the posterior half, and there is little or none of this pigment on the head. One or two black stellate chromatophores usually appear on the yolk-sac,* to which also outgrowths from the pigment cells in the otocystic region generally extend. The head and anterior part of the embryo are covered with a diffuse greenish yellow tint; the yolk-sac occasionally shows distinct yellow chromatophores at this stage, but more usually is also diffusely tinted. Only once I noticed distinct chromatophores on the embryo before hatching. I noted no pigment on the embryonic fin at this stage. Two larvae hatched out from eggs of 1°08 and 1:09 mm. diameter measured (a) 2°65 mm., of which 1°35 mm. was the pre-anal length, and (b) ca. 2°9 mm. with a pre-anal length of ca.1°5 mm. A third larva, (c), slightly more than twenty-four hours old, measured 29 mm. total length (1-2 mm. from snout to anus). This was from the egg of 113x117 mm. diameter. A fourth larva, (d), from the egg of 1:14 mm. diameter, measured when the yolk was absorbed, had a length of 3 mm., of which 1:28 mm. was pre-anal. The pigmentation consists of black chromatophores mostly dendritic, distributed along the dorsal and ventral contours, in the peritoneum, a little on the top of the head and about the pectoral region. In specimen (a), however, the pigment extends to the tip of the caudal region, while in (b), which has heavier pigmentation, the last 0°3—-0'-4 mm. of the tail is bare. This difference, considered together with the difference of larval size, suggests a difference of species which was impossible to detect in the two eggs so similar in general character and only differing by 0:01 mm. in diameter. From which egg each larva was produced I * This was noted on eggs which gave rise to different types of larvae, e.g. (a) and (b). See above. NEW SERIES.—VOL. IX. NO, 1. OCTOBER, 1910, C aL A. E. HEFFORD. am not able to state, because I had kept them together in the same vessel, regarding them as of the same species; but the very similarity of eggs makes this point a matter of indifference. The larva (a), which is 2°65 mm. long and pigmented to the caudal extremity, I regard as Gadus minutus. Larva (b), on the other hand, which is slightly larger (ca. 2°9 mm.) and shows heavier pigmentation, stopping short so as to leave the caudal extremity bare, I consider is very prob- ably G. luscus (cf. 10, Taf. X, Figs. 20 and 21). Larva (d) (length 3 mm. with yolk absorbed) resembles (b), the posterior 0°5 mm. of the caudal extremity being unpigmented. The small size for this stage of development does not rule out the probability of its being G. luscus. Larva (c) shows black pigment of less intensity, most of it occurring along the ventral post-anal contour as far as the caudal extremity and above the gut. There are only four dorsal post-anal chromatophores, and these are not nearly so strongly marked as the ventral ones. This I regard as G. minutus, especially on account of there being a distinct resemblance to the early post-larval forms of this species, which have been carefully described by Schmidt (21a and b), and which are common in my own collections. In the same way the bare-tailed larvae (b) and (d) suggest the now well-known larval pout.* As to the occurrence of yellow pigment, all four specimens showed yellow chromatophores, with more or less distinctness on body, unpaired fins, and yolk-sae, but in the post-vitelligerous specimen (d) this colour had almost vanished except from the head (cf. McIntosh, 14e, p. 240). Three planktonic eggs of 0°95 mm. diameter, which were taken in the West Channel (entrance to Plymouth Sound) on 11th February, may be either G. minutus or G. luscus, One-day-old larvae measured 2:95 and 2:8 mm. The black pigment consists of relatively large stellate chromatophores on the head, a dorsal series which become smaller and less closely placed towards the posterior extremity, and a much weaker ventral series consisting of about nine post-anal chromatophores and a faint line dorsal to the gut. There are small specks of yellow pigment on the body and embryonic fins, most strongly marked along the body contours and along the proximal margins of the fins, but very faint in the caudal region. The larval pigment at this stage cannot be said to show any approach to either the bib or * While using this similarity of larval pigment to that of definitely known post-larval stages as evidence assisting to indicate the identity of a larva hatched from a Gadus egg taken in the tow-net, I think it necessary to point out that this similarity should not always be expected in the larval stages. As positive evidence it is helpful, but as negative evidence it is without value. Under certain conditions—chiefly of higher than normal temperature—I have noticed a precocity in development of pigment. The same phenome- non has been noted by Holt (cf. lla, p. 454). NOTES ON TELEOSTEAN OVA AND LARVAE, 35 pout type of post-larval pigment. It resembles G, merlangus, but the small size seems to preclude that species. A larva of similar character hatched from an egg of 1:00 mm, diameter, taken from the same locality on 25th February, measured ca, 2°75 mm. when more than a day old. The next noteworthy capture of Gadus eggs took place late in August, an unusual time for the spawning of any members of the genus in our area. On 26th August two eggs of 1:02 and 1:06 mm. diameter were obtained in the young-fish trawl, but they died before hatching. The following day six eggs were captured of diameter 1:03-1:05 mm, In embryonic characters these resembled what I have above described as G. luscus, An early larva from one of them had a length of ca, 2°5 mm. A second larva which had absorbed practically all its yolk was 3°32 mm. long, of which 1:22 mm. was pre-anal. At this stage the anus was still apparently imperforate ; the eyes dark blue with con- siderable black pigment, the mouth large with the relatively massive lower jaw slightly protruding. There is well-marked indentation behind the mid-brain and a typical large supra-cephalic ampullation, which extends as far back as the level of the anus. The pectoral fins are large and fan-like. The type of pigmentation strongly suggests G. luseus. Black chromatophores are distributed post-anally as very distinct dorsal and ventral lines, which stop short at a distance of ca. 0‘9 mm. from the posterior extremity, so that the last part of the tail is quite unpigmented. The dorsal line arises in the occipital region and the ventral at the base of the pectoral fin, whence it continues backwards at the level of the dorsal edge of the gut. There is also black pigment at the tip of the snout, at the end of the mandible, below the throat, and a few lateral chromatophores on the trunk. In this specimen the latter were adjacent to the dorsal series, but in another they were mainly on the ventral half of the body posterior to the anus. No yellow pigment at all was observed in the late larval stages. The early larvae were very cursorily examined and I have no notes as to the presence of this colour. In the embryonic development a diffuse yellow tint was visible on the yolk-sac and about the contours of the trunk. Gadus merlangus. Only three eggs in all were taken, which may with probability be referred to this species. The first, taken at the western entrance to the Sound on 25th February, was 1:16 mm. in diameter, but was killed by the low temperature before hatching. The second was obtained from a haul of the young-fish trawl in Cawsand Bay on 28th April. No record was made of the size of the egg, but the newly hatched larva had a length of 3°44 mm. ( pre-anal length, 1-42 mm.) 36 A. E. HEFFORD. and the yellow and black pigmentation typical of G. merlangus. A third whiting egg was taken on 12th July, about 14 miles south of Rame Head. The diameter was 1:12 mm. and the length of the larva within the first day 3°58 mm., the distance from snout to anus being 1°58 mm. The fewness and infrequency of the appearance of whiting eggs in my collections are undoubtedly due to the fact that our tow-nettings have not been taken anywhere near the off-shore breeding haunts of the species at the time of their spawning season. Onos, Risso (= MoTELLA, Cuvier). The Rocklings. It is an open question whether our knowledge of the occurrence of the members of this genus in the Plymouth neighbourhood is complete and exact, but certainly the presence of more than one species has com- plicated the task of fully identifying the rockling eggs,which have long been well known as occurring here in abundance. The most common species found here is 0. mustela, L., and O. tricirratus (Bloch) is also known, while Holt (11d, p. 143) speaks of “ the undoubted existence in the dis- trict of MM. cimbria and M. maculata, and possibly of other forms which may require specific distinction.” I have no personal knowledge of the occurrence of the two latter species, but I may mention that a form identified as Motella fusca, Moreau, by Garstang and Balfour Browne was taken in April, 1901, on the shores of Plymouth Sound.* However, the problem of denoting the species of the pelagic ova commonly occurring at Plymouth, is mainly one of deciding which other species besides the abundant O. mustela are represented. My task of identification has been aided by the recent publication by Ehrenbaum (5c, p. 237, and 5d, p. 284) of descriptions of the ova and larvae of 0. mustela, L., and O. cimbrius, L. In regard to the latter species I need only say that it does not appear to be represented in my samples of ova, nor do I know of any record of the occurrence of the adult in this district.t O. mustela, on the other hand, is the prevailing species, and what I was led to expect, from the abundance of the fish in Plymouth Sound, is confirmed by comparison with Ehrenbaum’s description of the egg and larva of the species. He gives the average diameter of the ova of O. mustela as varying (off Heligoland) from 0°878 mm. in February to 0-736 in June, and the peculiar pigmentation of the late embryo and early larva as the chief diagnostic character (at least as far as dis- tinguishes it from O. eimbrius), viz. the arrangement of the post-anal * Journ. M.B.A., N.S:, VI., p. 626. + A small immature specimen of 0, ciibrius was, however, taken in Whitsand Bay on the 3rd March, 1910, which is, I believe, the first record of the species for the Plymouth neighbourhood, NOTES ON TELEOSTEAN OVA AND LARVAE. 37 black pigment into two groups, the first a short distance behind the anus, usually confined to the ventral half of the body, the second in the form of a band from the dorsal to the ventral contour, besides which there is, in the hypural region, a small patch of pigment which has out- growths to the marginal fin. 0. cimbrius shows only one of these post-anal pigment groups. Onos mustela, L. To come to the consideration of my own specimens, they can with very little exception be designated Onos mustela. There is, of course, the bare possibility that the eggs and larvae of some other species of rockling, which are at present unknown, may so closely resemble those of O. mustela as to have been indistinguishable from them by me. But even if such were the case, the number so included would be quite inappreciable against the total, which are undoubtedly 0. mustela, These eggs occurred in my tow-net and young-fish trawl plankton samples from 11th February to 25th June, and again in August and September. The diameter varied from 0:72 to 0°83 mm., averaging 0°77 for February to March, 0°78 for April, and 0°72 for May to June. The diameter of the oil-globule varied from 0:13 to 018 mm. It sometimes happened that the size of the oil- globule was the reverse of being proportional to the size of the egg, ie. the larger eggs of a sample had the smaller oil-globules, which I thought might possibly be significant of a specific difference, but observations of the resulting larvae disproved this. The oil-globule, which may be subdivided into two or three in the early stages, com- monly has a more or less greenish and sometimes a cupreous tint. The yolk surface is somewhat corrugated. By the time the embryo has developed a caudal rudiment, small black chromatophores appear generally in a double line along the body, on the head, and in the pellicle of the oil-globule. They soon increase in size, and may become stellate, especially the anterior ones. Just before hatching the two characteristic post-anal groups (or “ zones”) of pigment are generally quite distinct. One of my smallest newly hatched larvae measured 1:88 mm. (the pre-anal length being 0°76 mm.), and the largest size I have recorded for this stage is 2°32 mm. The black pigment, which is mostly stellate and dendritic, is distributed upon the head, in the peritoneum, over oil-globule, laterally on the trunk over the anus, in the two large distinct post-anal groups above mentioned, and in the hypural region. These groups or zones are formed by the occurrence of short dorsal and ventral bars of black pigment spots, more or less fused together, from which dendritic outgrowths extend laterally. Sometimes such a bar may consist of only two or even one large 38 A. E. HEFFORD. chromatophore. Ehrenbaum states that the anterior group is usually confined to the ventral half of the body, but I have very often found it possessing a well-marked dorsal bar. The extent and intensity of these bars and groups of pigment, however, are subject to some variation. With the absorption of the yolk the post-anal pigment diminishes, especially in regard to the dorsal chromatophores, and when the yolk is entirely absorbed the latter have generally—though not always—entirely disappeared, leaving three relatively small chro- matophores along the ventral contour corresponding to the three previous groups. At this stage the pectoral fins are well developed ; but the ventrals, which soon afterwards become such a conspicuous feature of the post-larva, are rudimentary. My 0. mustela eggs were taken for the most part in Plymouth Sound and some in Cawsand Bay, while none were taken in more open water than Whitsand Bay, which is in keeping with the littoral haunts and the penchant for the vicinity of brackish water of the parent fish (cf. Holt, 11d, p. 143). Onos, Species A (2 tricirratus, Bl). Two eggs which may probably be referred to this species occur in my samples, one taken on 28th June 3 miles 8S. by W. of Rame Head, and the other taken on 30th August 5 miles W.N.W. of Rame Head. The identification is chiefly based upon the similarity of the larva to that of O. tricirratus, described by Raffaele (20, pp. 37 and 38, Tav. I, Figs. 26 and 27; Tay. III, Figs. 2 and 3) from an egg of 0°74 mm. diameter, which had an oil-globule of 0°218. The larva is characterized by the possession of less pigment than O. mustela or O. cimbrius. It is practically limited to one clearly defined zone of black pigment across the middle of the post-anal part of the body and a line of peritoneal pigment dorsal to the gut (op. cit., Tav. III, Figs. 2 and 3). The earlier of my eggs had a diameter of 0°84 mm., and the later one 0°78 mm. The oil-globules measured respectively 0°16 and 0145 mm. The embryonic pigment spots, which are small at their first appearance, become in the later stages relatively large in size, although few in number. The isolated mid-post-anal group of chromatophores is very conspicuous. In my second specimen I noted that this group consisted of six ventral chro- matophores with three dorsal and two lateral ones, which, before hatching took place, formed a dense band around the embryo by their enlargement and partial fusion. Black appears in the eyes shortly before hatching. In both specimens only one chromatophore was to be seen over the oil-globule. The newly hatched larva from my later egg (Fig. 14) measures 2°32 mm., of which 0°94 mm. is pre- anal. That from my June egg, measured when about one day old, NOTES ON TELEOSTEAN OVA AND LARVAE. 39 had a length of 2.42 mm., from the snout to the anus being 1°0 mm. The pigmentation of the former consists of a very conspicuous mid- post-anal patch, made up of a large dorsal and a large ventral chro- matophore (the latter accompanied anteriorly by a relatively small pigment spot), a small hypural patch, while pre-anally there is a group of chromatophores on the side of the body in the pectoral region, a series dorsal to the gut, one large dendritic chromatophore below the anus and extending partially over the oil-globule, and some pigment in the eye and on the head. The pigmentation of my second larva at a slightly older stage, when most of the yolk had been absorbed, differed only from the above in being somewhat more densely aggre- gated. Two large dendritic chromatophores—one dorsal and one ventral—with outgrowths extending laterally as far as the notochord, constituted the mid-post-anal group, a continuous line of dendritic pigment ran along the whole peritoneal region, one large dorso-lateral patch of pigment occupied that part of the trunk above the base of the pectorals, two small chromatophores were on the head, and the iris was now completely black. Holt (11b, Pl VI, Fig. 53) has figured a larva (“Species III (Motella?)”) somewhat similar to this, but having an anterior group of post-anal pigment spots much re- sembling that of 0. mustela, which it also approaches in having many pig- ment spots over the oil-globule. Ehrenbaum (5d, p. 278) considers this may possibly be 0. tricirratus, Bl. The egg had a diameter of 0°72 mm. (and less) and an oil-globule of 017 mm., and the larva in its first day was 2°20 mm. long. My larvae certainly have a closer resemblance to Raffaele’s 0. tricirratus, Bl., than this of Holt’s. I may mention, how- ever, that Ehrenbaum (7did.) warns one not to attach too much weight to Raffaele’s identification of the parent fish as 0. tricirratus, Bl., suggest- ing that the Naples observer did not wish to designate this species as distinguished from 0. mediterraneus (L.), but merely referred to the common tricirrate form of the Bay of Naples, which appears to be O. mediterraneus (L) (=O. tricirratus, Briinnich), and not 0. tricirratus, Bloch (=Onos vulgaris, Yarr.). I do not see, however, that there is any solid ground for doubting Raffaele’s identification in this case. Onos, Species B. A much more problematic form of larva, which I will term “Onos, sp. B,” was hatched from a rockling egg taken in the young-fish trawl 2 miles S. of Rame Head on 2nd June. The egg had a diameter of 0°68 mm., and its oil-globule 0°145 mm. When the embryo had developed a short free caudal portion, it was marked with large black pigment spots, and the stellate chromatophores in the pellicle 40 A. E. HEFFORD. of the oil-globule were noted as being especially large. On the 5th of June the larva had hatched out. Its length when about one day old was 1°84 mm. (pre-anal length = 0°84 mm.). It showed the Motella characteristics as described above, except as regards the distribution of pigment in the pre-anal part of the body, which was practically wholly dorsal (see Fig. 15). From the snout to beyond the middle of the post- anal part, there was a series of black chromatophores, most of which were large and possessed outgrowths extending over the upper part of the sides of the trunk, often as far as the level of the notochord. The most posterior group of this series contributed the dorsal bar of the typical Onos mid-post-anal zone. The corresponding ventral bar was present, as well as the usual hypural patch, but the only other pigment consisted of a single chromatophore below the anus and a group over the posterior part of the oil-globule. The absence of pigment from the peritoneal region is remarkable, and this fact especially inclines me to the view that we may possibly be dealing here with an abnormal specimen. Apart from this the pigmentation bears some resemblance to that shown by 0. cimbrius (Ehrenbaum, op. cit.). From my present knowledge of Plymouth species of rocklings, however, I will not venture to suggest a definite species. Assuming that it is normal, it is certainly not O. mustela nor O. cimbrius ; and if my previously described larva (Onos, sp. A) is indeed 0. tricirratus, Bl., that species is also excluded. It seems to me that my “Onos, sp. A” may with far more probability be referred to Onos tricirratus, Bl., than may “Onos, sp. B.” Then, by a process of exhaustion—always bearing in mind, however, that our knowledge of local Onos species cannot safely be regarded as complete —we have left Motella fusca, Moreau (which may probably be regarded with Af. maculata of the same authority as varieties of O. mediterraneus, L.). It is at least possible that “ Onos, sp. B,” the larva with a pre- anal dorsal row of chromatophores and no peritoneal pigment, belongs to this species. Raniceps raninus, L. Frog-fish. Four eggs identified with this species were taken from a young-fish trawl, mid-water haul, in Whitsand Bay on 30th August. Holt obtained eggs of the same species from tow-nettings taken at various depths off Plymouth, in June, July, and August, 1897, which he recorded as unidentified but with apparently gadoid characters (11d, p- 145). In his Irish survey the same investigator had previously met with a similar egg and had described and figured it, with the twelve- hour-old larva, as “Species VIII” (11a, p. 471, Figs. 27 and 36). He embodies his observations upon both Irish and Plymouth material in his Marseilles Museum Annals Memoir, suggesting as the possible NOTES ON TELEOSTEAN OVA AND LARVAE, 41 parent fish Phycis blennoides, a rare visitor to this coast and one which had probably arrived in the track of the shoals of mackerel and scad (Caranxz trachurus), which were present in unusual abundance in the inshore waters at the time of his Plymouth observations. Since that time, there have been no observations of planktonic fish eggs in the summer months here until the present year, so that the solution of the question as to whether the eggs belonged to a constant or intermittent visitor to these shores by the plan of noting the presence or absence of the eggs in successive years, has not been possible. It so happens that the present summer (1909) has also been characterized by a greater than usual abundance of mackerel and scad in the inshore waters of the Plymouth area, but whether this condition can be cor- related with the presence of these eggs is doubtful. Heincke and Ehrenbaum (10, p. 258) have subsequently observed the egg as regularly occurring with summer plankton off Heligoland, and since Phycis blennoides, the only other fish to which it could with any prob- ability be ascribed, is never found in Heligoland waters, they have identified it with Raniceps raninus—a quite well-founded conclusion, although the absolutely unquestionable identification by tracing back the egg to the parent still remains unaccomplished, since the ripe female is as yet unknown. Holt’s Irish specimen measured 0.775 mm. and had a colourless oil-globule of 0:14 mm. diameter. The larva about twelve hours after hatching measured 2°68 mm. Those taken by him at Plymouth at the end of June and in July measured from 0:84 to 0:91 mm. in diameter, and the diameter of the oil-globule ranged from 0:16 to 0:17 mm. In August the dimensions were 0°78 to 0°84 mm. for eggs and 0°15 to 0°17 for oil-globule, and a newly hatched larva was 2:02 mm. in length. The Heligoland eggs had a diameter of 0°755 to 0912 and an oil-globule from 0'141 to 0°189 mm. in diameter, while the length of newly hatched larvae varied from 2:26 to 2:90 mm. The dimensions of my specimens were as follows :— Diameter of egg . . 0°80, 0°78 x 0°79, 0°81 x 0°84, 0°82. oil-globule 0°145, 0°145 0-165 0:157. ”» ” Two of them were ovoidal. The yolk is homogeneous, and the oil- globule is colourless. Just before the formation of the caudal rudiment, the head and body are liberally covered with medium-sized, black chromatophores, and yellow is making its appearance along the sides of the embryo. On the yolk-sac there is pigment of both colours, which is most dense in the postero-ventral region, a feature becoming more strongly marked as development proceeds. In the two larger specimens there are black and yellow chromatophores over the oil-globule, but 42 A. B. HEFEFORD. they are absent from here in the two smaller eggs. Although pre- cautions were taken to keep the temperature low by standing the jars containing ova in circulating tank-water, the eggs became infested by infusoria, whose presence is a usual accompaniment to unhealthy con- ditions. Development proceeded apace, however, and the next day the free caudal region had grown around the yolk, so as to almost meet the head. The bright yellow pigment of the embryo is now visible to the naked eye. It occurs in large dendritic chromatophores, which ramify and intermingle so as to produce a diffuse colouration over the whole of the pre-anal part of the trunk,and appears especially dense about the anus. There is a further band-like mass of yellow about the mid-post-anal region, and an aggregation of similar chromatophores on the yolk-sac, between the oil-globule and its posterior contour. In one specimen (diameter ‘82 mm.) the oil-globule, at this stage, has a dark and smoky appearance, and is densely pigmented. In another specimen no pigment is seen over the oil-globule, and the periblastic pellicle, which is gener- ally quite apparent at this stage, showing an interspace between it and the contained oil-globule, is not distinguishable. The epidermis of embryo and yolk-sac is covered with tiny tubercles, doubtless of patho- logical origin. Next day the larva had hatched out but was distinctly moribund, and died almost immediately. The total length is 2:16 mm., ‘and it measures 1:00 mm. from snout to anus. The head projects rather considerably over the oval-shaped yolk-sac. In two of my larvae the oil-globule was almost in the centre of the yolk-sac, which was observed by Heincke and Ehrenbaum to be the case only in one instance, and may be regarded, therefore, as an abnormal and possibly patho- logical condition. The small otocysts are situated some distance behind the eyes. There is a slight swelling in the tubular gut above the pectoral region and the rectum ends blindly immediately behind the postero-dorsal edge of the yolk-sac. The pigmentation, which is on a generous scale, is very characteristic. Inter-ramifying yellow chroma- tophores form a diffuse mass of colour over the posterior part of the yolk-sac, and practically over the whole of the pre-anal part of the trunk and head, extending a little beyond the anus. Then comes a clear space followed by a band of yellow somewhat behind the mid-post- anal point. The much less conspicuous black pigment in small chromatophores which when relaxed show fine dendritic outgrowths, occurs chiefly on the dorsum in the pre-anal region and, less densely, on the head and sides. Post-anally there are about half a dozen chromato- phores along each of the dorsal and ventral contours, extending further posteriorly than the yellow pigment, although the extreme end of the tail is pigmentless for about -25 mm. Black chromatophores are NOTES ON TELEOSTEAN OVA AND LARVAE. 45 associated with yellow in the posterior hemisphere of the yolk-sac. The unpaired fins are pigmentless, except for a touch of yellow near the origin of the dorsal fin, immediately behind the otic region. Apart from its peculiar colouring the larva has the unmistakable gadoid form. The dorsal fin membrane arises over the occipital region, reaches its greatest width above the anus, and thence tapers gradually to the caudal extremity; and similarly the greatest width of the anal fin is immediately below the anus. On 2nd July, in a surface tow-netting 1} miles N. by W. of the Eddystone, an egg was taken which may possibly be identified with this species. Its diameter was ‘86 mm., and its single oil-globule measured ‘18 mm. The yolk was unsegmented. Just before the out- growth of the caudal rudiment the body was abundantly besprinkled with faint dark chromatophores. The body appeared relatively wide. Next morning the larva showed a short caudal rudiment and much increase of pigment, which is now canary-yellow as well as black. Roundish chromatophores are fairly generally distributed over the anterior part of the body, but the black appears to be mostly dorsal and the yellow ventral. Post-anally the pigment is less dense. The pellicle of the oil-globule, which has a rough, wrinkled, and rather dark appearance, bears many rounded chromatophores of both colours, rather larger than those on the embryo and constitutes the most conspicuous feature of theovum. There area few fine, chiefly yellow chromatophores in the dorsal part of the yolk-sac. The otocysts are relatively small and the rudimentary pectoral fins appear as narrow flaps. Clupea sprattus. Sprat. This egg was the commonest of those belonging to food-fishes which occurred in my samples. It was found almost continuously from the middle of February to the middle of June, after which time until 12th July it appeared with less frequency ; which may, however, be largely due to the fact that in the summer months most of my plankton was collected from the open sea, while the sprat appears to favour the close vicinity of Plymouth Sound or Cawsand Bay as a spawning locality in the warmer months, although not in winter and early spring. Very many eggs were taken in the young-fish trawl in June and July. The eggs, which are very characteristic from their segmented yolk, had . an average diameter of 1:031 mm. for February-March, 0°973 mm. for April-May, and 0°912 for June-July. The pelagic larvae, which will be treated at greater length in a subsequent paper, were most abundant in May and June. | 44 A. E. HEFFORD. Cl wped pilehard us. Pilchard. My first pilchard eggs were taken on 8th April, 7 miles 8.W. of the Eddystone, when several occurred in the tow-nettings. The diameter ranged from 1°63 mm. to 184 mm., and that of the oil- globule from 0°15 to 0°16 mm. The next specimen (of 1°6 mm. and 0-145 mm. oil-globule) was found in a young-fish trawl haul taken near the Eddystone on 26th August. More surprising was the occurrence of three eggs in a tow-netting taken inside the Sound on the 14th Sep- tember. These had diameters of 1:46, 1°52, and 1°62 mm., and oil- globules of 0°145, 0°155, and 0:14 respectively. A newly hatched larva from one of them, measured after being killed in dilute formalin, had a length of 58 mm. I may also mention that I have found numerous pilchard eggs in samples of plankton taken in the young-fish trawl in September, 1906, on the Rame-Eddystone Grounds. As Cunningham (4a, p. 44, and 4d, p. 154) has pointed out, pilchards spawn far out at sea, and it is doubtless due to the fewness of my tow- net samples from the open-sea areas that such a small number of pilchard eggs have come under my observation this season. I may mention in passing that the pilchard fishery season in 1909 has been a decided failure in the Plymouth district as off the Cornish coast, the shoals having kept out in mid-Channel 20 miles or more from the coast, and therefore out of reach of the usual fishing craft. It should be remembered that the great majority of pilchards caught by Ply- mouth drifters are not spawning fish, the usual shoreward movement of this species in summer and early autumn being apparently a feeding migration. It is hardly relevant to the present subject to discuss the possible causes of the unusual distribution in 1909, nor is there com- pletely satisfactory evidence available. We may, however, assume that the distribution of the spawners which appear to le outside the main summer shoals may show some variation in relation to the move- ments of the latter. My collections certainly sampled only the fringe of the great mass of ova spawned, or those which drifted landward with the tide and currents. B. DEMERSAL EGGS. Labrus ? mixtus, L. Eggs which in all probability belong to Labrus mixtus were found deposited among a mass of Chondrus crispus in a rock-pool on Wem- bury Reef on 17th June. The mode of occurrence is very similar to what has been described by Matthews for Zabrus maculatus (17), and my first idea was that this was the species to which the “nest” NOTES ON TELEOSTEAN OVA AND LARVAE. 45 belonged, as it is the most common wrasse in the locality, which could with any certainty be regarded as the parent fish. The smaller size of the eggs, however, and certain differences exhibited by the hatched-out larvae, led me to conclude that this was not a species identical with that described by Matthews, though certainly a closely related form. Of the other wrasses (having unknown ova and larvae) which are known to occur here—Labrus mixtus, Crenilabrus melops, and Centro- labrus exoletus—the first is the form to which every probability points as the parent of these eggs. It is the one other species known to form a nest similar to that of Z. maculatus (18, Vol. III, p. 102), and the size of the eggs is also most in agreement with this parentage. Ripe ova from Crenilabrus melops have been described by Holt (11a, p. 450) as spherical and having a diameter of 0°78 mm. Crenilabrus exoletus, of whose eggs I have no knowledge, is an exceedingly small fish, and is not at all likely to produce ova as large as my specimens. It is moreover a more deep-water form, and is not known to construct a nest between tide-marks. My specimens had a spherical or somewhat ovoid shape and a thick strong capsule. Five which were measured had the following dimen- sions—0°92, 0:94, 0°94, 0°90 x 0°94 and 1:08 x 0°86 mm. When first ob- served on the 17th June the embryonic body with well-marked myomeres had formed, but no caudal outgrowth had appeared. There was a large Kiipfer’s vesicle. No pigment was visible. The yolk was pale buff- coloured and devoid of any oil-globule. Four days later they have reached the final stage of embryonic development (Fig. 4). The yolk is much reduced, its diameter being about four-sevenths that of the egg-capsule, and the caudal extremity has grown around so that its tip in some cases overlies the auditory region. The yolk is ochreous-yellow and shows a number of small vesicles in the mid-ventral part. In the eyes there is black pigment through which shine golden tints. The body is be- strewn with black chromatophores for about two-thirds of its length, and posterior to this there are some along the dorsal and ventral con- tours, but the posterior extremity is unpigmented. Yellow chromato- phores occur on the anterior part of the body. The yolk-sac shows a few round black pigment spots and many yellow ones. The pectoral fins appear as semicircular flaps near the posterior edge of the yolk- sac. Small bean-shaped otocysts occur at a distance behind the eyes about equal to the diameter of the lens. Fig. 8 depicts a slightly earlier stage. Two larvae (Fig. 8) measured within a few hours of hatching were 3°26 and 3:28 mm. in length, the pre-anal lengths being 1°80 and 1:76 mm. respectively. The yolk-sac is relatively small, and its con- 46 A. E, HEFFORD. tents are clear and almost colourless. The head is rounded; the anus post-median. The dorsal fin membrane arises above the mid-brain and is widest above the anus. There is a well-marked pre-anal fin, The whole of the larval fin membrane has a minute vesicular structure, which is probably a mark of ill-health. Embryonic rays can be seen in the caudal region. The notochord has two layers of cells at least in its posterior part. The pectoral fins are well developed. The body is richly pigmented with black and yellow chromatophores, but the posterior third is conspicuously bare, except for a line of black chro- matophores along the ventral contour, while black pigment is lacking above the mid-brain, The sides of the body from the occipital region to a short distance beyond the anus are almost uniformly coloured with round, stellate, black chromatophores about four or five deep, These are most closely set along the dorsal contour, and are more densely distributed above and posterior to the anus than above the yolk-sac. They are not uniformly distributed in the body segments as described by Matthews for Z. maculatus, but except for the posterior continuation of the ventral line, as above-mentioned, and for the presence of black chromatophores over the sides of the gut, the distri- bution and abundance of black pigment on the body show much agree- ment with his specimen. The largest black chromatophores of all are to be seen on the yolk-sac. A group of about five to eight large, but not very intense, chromatophores occurs in the anal fin, immediately behind the anus; otherwise, except for outgrowths from pigment cells along the posterior ventral body margin, the larval fin membranes are entirely free from black pigment (see Figs. 8 and 8a). Yellow pigment is regularly interspersed with black along the sides of the trunk, but is absent over the sides of the gut. There are a few yellow chromato- phores on the head, and a single isolated one near the margin of the anal fin, about half-way between the anus and the posterior end of the notochord. Another specimen which I examined showed a less uniform distribution of black pigment on the side of the trunk in the pre-anal region, the chromatophores tending to concentrate along the dorsal contour and above the gut. On the second day the larva had a length of 3°48 mm., the increase being practically entirely post-anal. At this point I made detailed measurements, so as to compare with Matthews’ dimensions for Z. maculatus, which I give side by side below :— Matthews’ My specimen. LT. maculatus. Tip of jaw to front of eye ; ; ‘12 mm. i Pee zs =: back _,, ; : ou eee 3)// . = front of ear : : 2 ae 565 «eae 3 = centre of heart . ; Cd. aoa aye Sag oe Need end of pigmentation . 2:2 — ,, (mannig’ 2-48 ” ”? Total length ; ; : , 2 B48 3 | O00 NOTES ON TELEOSTEAN OVA AND LARVAE. 47 I also noted that my larva was distinctly more slender in dorsal view than Matthews’ L. maculatus (op. cit., Pl. XI). The pigmentation is in general the same on the second day as on the first, but the post-anal pigment in the anal fin has slightly increased. With the total absorption of the yolk, which has taken place on the fourth day, a still further increase of this pigment is seen, the group of chromatophores behind the anus now numbering fourteen or fifteen ; and there is a further extension of pigment from the ventral edge of the trunk to the proximal margin of the anal and pre-anal fin mem- branes. The embryonic fin-rays, in both the dorsal and ventral parts of the caudal region of the larval fin membrane, are now very evident. A specimen at this stage, after killing in dilute formalin, measured od mm. Blennius pholis, L. The Shanny. It is somewhat surprising that the earliest stages of this common blenny should have remained unknown for so long. McIntosh (14g) has published some observations upon eggs deposited in captivity. These were circular in outline, oblate spheroidal in lateral view, and each had a faintly pinkish attachment disc. The diameter was 1:181 to 1:219 mm., the vertical diameter being 0°763 and the height of the attachment rim 0°305 mm. He describes the yolk colour as dull pinkish or faint salmon and in certain lights having a dull brownish appearance. On June 4th some eggs of Blennius pholis accompanied by the parent fish were taken on a stone on the Breakwater rocks. In shape they were ovoid with flattened underside (Fig. 3). The length of the capsule was 16 mm., the vertical height just above 1 mm. (with the attachment disc ca. 14 mm.), When observed they were at the last stage of embryonic development, the black eyes of the embryo render- ing them very conspicuous. The yolk was of a light brown colour. The newly hatched larva (Fig. 6) is of large size—about 4:4 mm. total length and 1:8 mm. from snout to anus. Its very broad and somewhat square head gives it a tadpole-like appearance. There is a striking absence of post-anal pigment. A most conspicuous feature is the pair of large fan-like and heavily pigmented pectoral fins. These are marked with large, black and yellowish brown chromatophores, the former disposed in radial lines, the latter being most concentrated in the basal region and absent from the distal margin. Other black pigment occurs in the eyes, under the mandible where three stellate chromatophores are disposed symmetrically in triangular form, on the neck region as a single pigment spot, and in the peritoneum where there is a short double row of chromatophores. The yolk-sac, pro- 48 A, E. HEFFORD. truding on each side, has a yellowish brown tint; there are pale yellow chromatophores on the head and similar pigment, but of more intensity, in the pectoral region. The thickness and opacity of the head causes the otocysts to be hardly visible. The course of the red blood corpuscles along the circulatory system from the yolk to the body of the larva can be very easily seen. Gobius paganellus, Gm. L. Some eggs, together with a fish of this species, were taken on a stone between tide-marks on the shore of Rum Bay on 3rd June. The ova have been described by Holt and Byrne (13, p. 46) as regularly fusiform in shape, about twice as high as wide and with rather sharply pointed ends; by which characters they are distinguishable from the eggs of all other British species of goby. The above-mentioned authors give 1:84 to 19 mm. as the length. An egg which I measured at a late embryo stage was 2°3 mm. long and 0°74 mm. wide, while a second was slightly longer. The yolk was of a greyish brown colour and was darkened by the presence of many small oil-globules. When the embryo is advanced in development, the eyes become extremely conspicuous, showing abundant black pigment and a bronze-green lustre. The oval swim-bladder with strongly marked dendritic chroma- tophores and some yellow pigment over the dorsal side of it is plainly visible. This is the only really conspicuous pigment on the embryo at this stage. The large bean-shaped otocysts contain relatively small otoliths. The newly hatched larva (Fig. 7) has a total length of 48 mm., the pre-anal length being 2:2 mm., so that the anus is just anterior to the median. The head is somewhat rounded and the lower jaw slightly projecting. The large, oval swim-bladder is a conspicuous object mid- way between otocysts and anus. The gut is straight and has a slight ventral dilation below the hinder end of the swim-bladder. As in the embryo, the most conspicuous pigment is above the swim-bladder (black and yellow) and in the eyes, which are black with blue, green, and gold tints. There is a continuous row of black chromatophores from the throat to the anus along the ventral contour, the largest one with well-marked dendritic rays being below the above-mentioned bulge in the gut; the terminal one below the anus is also very pro- nounced. Except above the swim-bladder there is no peritoneal pig- ment nor any other chromatophores anterior to the anus. Post-anal ventral pigment consists of a discontinuous series of black dendritic chromatophores (about six or less in number) extending to the hypural region, the largest of which is situated in the centre of the post-anal NOTES ON TELEOSTEAN OVA AND LARVAE, 49 part and has yellow associated with it. Dorsally there is only one, relatively small, black chromatophore, accompanied by yellow, opposite the large mid-post-anal one of the ventral row. This dorsal pigment is often lacking entirely. The notochord is unicolumnar. The pectoral fins are rather large, extending to about the middle of the swim- bladder. ‘The embryonic dorsal fin arises above their base. There is a short pre-anal fin commencing below the gastric dilation. A brownish gall-bladder is visible. At the age of five or six days the ventral post-anal pigment appears to have concentrated itself more in the central part of the post-anal region and in the hypural part, but otherwise the pigment remains as in the early stages. Embryonic fin- rays have developed in the position of the second dorsal and the anal fin and a hypural lobe has formed. Lepadogaster bimaculatus, Donov. Doubly spotted Sucker. The eggs and newly hatched larva of this species have previously been described by Holt (11a, p. 447, Pl. XLVI, Figs. 1 to 7), but as all my observations have shown certain differences from the specimens he describes it will be well to give some details from my records. My observations were first undertaken chiefly with a. view to getting a knowledge of the early post-larval forms for the purpose of comparison with pelagic Lepadogaster fry taken in our young-fish trawl. The following are brief particulars as to the capture of specimens of the eggs of this species which came under my notice in the summer of 1909 :— | Date. Locality. instrament ae Other remarks, | capture. =| 8th June, 1909 Queen’s Ground. | Dredge . . | Eggs encrusting inside of a T'e7lina valve. Greatest horizontal axes of egg-capsule, 14 x 1°2 mm. 16th June, 1909 | Rame-Eddystone | Otter trawl . | Eggs in Lutraria valve with | Ground parent fish. Greatest horizontal dimensions, 1°4x1:16 mm.; height, 0°70 mm. 5th July, 1909 | Hand Deeps . | Dredge . . | Several batches of eggs with | parent fish in valves of Pecten opercularis and Lutraria, 13th July, 1909 | Hand Deeps . | Dredge . . | Eggs in Pecten opercularis valve, | | Accompanying fish a female. On 14th June a female Lepadoguster bimaculatus, with spent, flaccid and membranous ovaries, was taken in a Lutraria valve, within which a batch of eggs had been deposited, but which had disappeared, leaving traces of their former presence in the form of oval impressions. The NEW SERIES.—VOL, IX. NO. 1. OCTOBER, 1910. D 50 A. E. HEFFORD. nudibranch Calma glaucoides and a batch of its eggs were also occupy- ing the valve, and it seems probable, if not certain, that the fish-eggs had been devoured by the nudibranch, which has been recorded as commonly occurring associated with Goby and Blenny eggs, and vary- ing in colour so as to resemble the eggs which it apparently preys upon.* j Quite recently—viz. on 17th February, 1910, and after this paper was in manuscript—I have secured an early batch of Lepadogaster bimaculatus eggs. On this occasion I took particular care to examine the accompanying parent fish so as to make sure of its identity as distinct from JZ. microcephalus, a. closely similar species first dis- tinguished by Brook,t whose description, however, I have not yet been able to see. Ehrenbaum (5b, p. 121) gives as the distinctive fin-ray formula for L. microcephalus D=5, A=6, C=17-19; while Day gives for ZL. bimaculatus D =5-7, A=4-6, C=12, My specimen has clearly six dorsal fin-rays and not more than four or five anal fin-rays, which precludes LZ. microcephalus, while in its general appearance it resembled the common two-spotted sucker, Z. bimaculatus. It was not possible to count the caudal rays, as it was desired to keep the specimen alive and uninjured. Two of the eggs had the following dimensions :—Oval outline of egg-capsule, as seen from above, measured in one case 1°44 x 1:24 mm. and in the other 1°54 x 1°22 mm. The height of the capsule was respectively 0:62 and 0'70 mm. The sizes closely approximate to those noted the previous summer, and in following through the development from pre-embryonic to larval stages, the characters proved to be identical, save for very slight variation in pigmentation. The ovoidal inferiorly truncated egg-capsule and its peculiar basal attachment disc and filaments have been minutely described by Holt (op. cit.). I noted that as a rule the eggs in one batch showed several stages of development, indicating that they were depositedintermittently. In the earliest stages the finely granular yolk is quite colourless and translucent, and carries a single oil-globule of about 0:25 to 0°28 mm. diameter, which has a slightly darker appearance than the yolk and is at first the most conspicuous content of the egg. The embryo almost invariably occupies a horizontal position in the egg. Black pigment appears on the body soon after the outgrowth of the caudal rudiment, and soon forms a dense and continuous line along the ventro-lateral region from immediately behind the pectoral fins to within a short distance of the caudal tip. Anteriorly pigment is sparse, only a few * See Journ. M.B.A., N.S., Vol. VII, p. 280. + Brook, G., Proc, Roy. Phys. Soc, Edin., Vol. X, Pt. 1, p. 166, NOTES ON TELEOSTEAN OVA AND LARVAE. 51 chromatophores occurring about the neck region and shortly afterwards on the eyes. Yellow chromatophores (bright lemon-yellow by reflected light, brownish by transmitted light) next appear on the sides of the embryo above the dense line of black pigment above-mentioned (see Fig. 5). At about the same time black chromatophores frequently appear along the dorsal surface of the yolk-sac adjacent to the trunk of the embryo. The circulatory fluid is now of a red colour, which is plainly visible at the heart systole. Before the appearance of the yellow pigment the blood was colourless, but even then its circulation could be observed in vessels from the yolk-sac, and in the aorta and main arteries of the head. A day or two after the appearance of yellow, an increase takes place in the amount of black pigment, a double ventro-lateral line being formed on each side with the inferior pair, which are the more distinctly marked, coming together at the anus. A sprinkling of lateral chromatophores next appears, and the eyes become so dark as to be conspicuous to the naked eye. The only other black pigment in the anterior region is a pair of lines converging towards the occiput from the posterior lateral part of the yolk-sac. The yolk-sac and occipital region are covered with diffuse pale yellow. The newly hatched larva (see Fig. 16) has a length of 4:26 mm., of which about five-eighths is pre-anal. The remnant of yolk is relatively small, as is usual with larvae from demersal eggs, and bulges out on each side of the larva. A small oval swim-bladder is present. Rounded, stellate, black chromatophores uniformly beset the sides of the trunk in fairly regular lines, which are about four deep transversely in the pre-anal and about three deep in the post-anal region. These are larger in the anterior part of the body than posteriorly. There are similar lines of yellow chromatophores (pale lemon coloured by re- flected, brownish by transmitted light) slightly less numerous and at greater interval. These are more densely distributed posteriorly than anteriorly. The most dorsal row of chromatophores are yellow and these are of a larger size and greater denseness than the others, Over the straighv intestine -low pigment is generally sparse and sometimes quite lacking. The justerior portion of the tail, for about 1 mm., is quite unpigmented, as is also the median strip along the whole dorsum. The only pigment in the larval fins consists of a small group of about three to five black chromatophores in the anal fin immediately behind the anus. The snout is rounded. The large otocysts are situated immediately behind the eyes. The dorsal fin arises a little behind the level of the posterior edge of the yolk-sac. Tiny epidermal vesicles densely cover the embryonic fins, except along the margin of its most posterior part, 52 A. E. HEFFORD. Two post-larvae, measured soon after the yolk had been absorbed, had lengths of 48 and 49 mm. The jaws had appreciably developed, especially the mandible, so that the earlier sub-terminal position of the mouth was changed; but otherwise they resembled the newly hatched individuals. The differences between my specimens and those recorded by Holt are in size and pigmentation. His newly hatched larvae measured 2°97 to 3:15 mm., and apparently had no yellow pig- ment (op. cit., p. 448). His eggs were slightly smaller than mine, having a length of 1:37 mm., a breadth of 1:08 mm., and a height of 0°68 mm., and the oil-globule measured 0°24 mm. Guitel (9, Pl. XXV, Fig. 8) figures an early post-larval LZ. bimaculatus of uncertain age, which is about 46 mm. long and is pigmented somewhat similarly to those I have examined, except that the superior line of yellow chromatophores is not clearly shown on the side and the black chro- matophores are lacking in the anal fin. As regards Holt’s specimen, if it is the same species as those I have examined, I can only suggest that it may have been an abnormal specimen, possibly prematurely hatched under unfavourable conditions. Lepadogaster gouant, Lacep. Cornish Sucker. On the 17th June several batches of the eggs of this species were taken on the underside of flat stones between tide-marks on Wembury Reef. The parent fish was always to be found near, and generally close alongside the eggs, which cover several square inches of the stoné with a closely applied layer. Two, three, or four stages of development may be seen in one batch of eggs. In the earliest stages the yolk is bright amber coloured, which renders the mass of ova an object of much conspicuousness and beauty. Subsequently the colours become gradually darker to orange, and finally, when the embryo is advanced, they have in the mass an olive-green appearance. The egg- capsule is oval-shaped with flattened base, of length 1:90 mm. and breadth 156 mm. The yolk contains a large oil-globule of 0°34 mm. diameter. When a short caudal rudiment is developed, the embryo has a general reddish tint, and shows many stellate black chromatophores over the greater part of the body, the posterior portion, however, being unpigmented. The movement of pale reddish circulatory fluid along the vessels from the yolk to the heart is plainly visible. The newly hatched larva has a length of 571 mm., the anus is post-median, and the yellowish yolk-sac protrudes on either side of the anterior abdominal region. The straight gut shows internal con- volutions and a yellowish green gall-bladder is visible. The dorsal NOTES ON TELEOSTEAN OVA AND LARVAE. 53 embryonic fin arises in the occipital region, the caudal part of it being spatulate. The head is rounded and the mouth terminal, the otocysts situated immediately behind the eyes. Pigmentation is extremely rich. Black chromatophores are the most abundant, covering the sides of the trunk and gut in closely set and fairly regular longitudinal lines. The largest chromatophores are those on the dorsal surface of the head, and those along the dorsal contour are the largest and most closely set of those on the trunk. .Post-anally the number of chro- matophores counted transversely is four or five. About 0°8 mm. from the posterior end of the notochord, the greater part of the pigmenta- tion ceases, but there may be a few small pigment spots over the notochord behind this point. Along the anal fin there is a line of black chromatophores, extending from immediately behind the anus to the hypural area. Mid-laterally, where the black pigment is least dense, there is an irregular line of about seven large lemon-yellow chromatophores, extending from the level of the posterior edge of the yolk-sac to a little behind the anus. Between all the other chromato- phores are numerous small orange-coloured ones, with a relatively large, clear, central space, which gives them the appearance of small rings. Most of the black and yellow chromatophores also have the form of radiations from a central unpigmented spot. Small, round or stellate chromatophores of a pure red colour occur on the ventral surface of the abdomen anterior to the anus. There is no yellow or orange pigment over the sides of the abdomen, but only black, and it. is perhaps worthy of note to mention that the black chromatophores of this region appear to have a different structure from those over the rest of the body, the centre of the spot in this case being pigmented instead of clear. The proximal part of the median fin membrane shows the same minute vesicular structure as was seen in L. bima- culatus, but the vesiculation does not extend so near to the margin. A slightly older larva measured 5-7 mm., and was 3°2 mm. from snout to anus. At four or five days old the length is 6°3 mm., the pre-anal portion being 34mm. A hypural thickening is visible. The pigmenta- tion at this age is practically the same as in the newly hatched form, except for an increase of red pigment on the inferior parts of the body. A larva in which the yolk has been entirely absorbed shows small red chromatophores on the ventral surface of the lower jaw, on the ventro- lateral part of the opercular region, on the ventral and ventro-lateral surface of the abdomen and over the basal part of the large pectoral fins. The pelagic post-larval stages of L. gowani can be easily distinguished from those of Z. bimaculatus by their larger size at the same point in 54 A. E. HEFFORD. development, and by the distinctiveness of their coloured pigment. In preserved specimens in which all but the black chromatophores have disappeared, one can at once distinguish L. bimaculatus by its relatively wide unpigmented strip along the dorsum, only a very narrow line being left clear between the pair of dorsal lines of chromatophores in L. gouant. There is also a difference in the distribution of pigment spots in the anal fin. As is to be expected, however, I have never met with the young stages of Z. gowani in plankton taken away from the vicinity of the shore, while post-larval LZ. bimaculatus may be taken some miles out at sea. Zeus faber, L. John Dory. On 31st August five good-sized dories were taken in the otter-trawl 24 miles S.W. of Rame Head. One of these was an unripe male, and three were females, which had recently spawned. From the ovary of one of the latter I obtained a dead egg, which had already undergone degeneration and was opaque and pale greenish in colour. The fifth proved to be a female approaching ripeness, and from the ovary of this I obtained a few apparently ripe eggs, which occurred free in the lumen. The great majority of the ova, however, were still small and opaque, and contained firmly in the ovigerous lamellae. The ripe eggs are large and contain a relatively small greenish yellow oil-globule (Fig. 1). The rather thick egg-capsule is marked by conspicuous corrugations, which appear to be intertwined in a very irregular manner, and also by finer striations, the former of which are doubtless merely characteristic of the ovarian condition and caused by contact with vascular tissue in the ovary. The yolk is colourless and homogeneous, the ripe egg being translucent and glassy, but not of that clear transparency which is seen in all pelagic eggs, and by transmitted light it has a slightly brownish tint, which is apparently produced by the interference of the corrugated capsule with the free transmission of light rays. The dimensions taken from four eggs are as follows :— Diameter of Egg. Diameter of Oil-globule. (1) 2°04 x 2:14 mm. i? 0-44 mm. (2) 2°03 5 aes 0-44 ,, (3) 2°05 3 ie 0°28 and 0°22 mm. (4) 1:90 us i: 0-45 mm. In the third egg measured the oil was contained in two separate globules, which is commonly the case in an unfertilized egg. The fourth specimen measured was apparently not quite ripe. The eggs sink in sea-water of specific gravity 1026. Fulton (8), from. NOTES ON TELEOSTEAN OVA AND LARVAE. 55 minute observations made upon the ripening ovarian ova of a dory caught in April, concluded that the mature eggs would prove to be large, contain one or more oil-globules and be demersal; which con- clusions are confirmed by the character of my ripe eggs. Their demersal nature was indicated, even at that stage, by the comparatively dense fibrous nature of the tissue of the stroma and the follicle, by the presence of a well-defined double layer, by the character of the yolk, and by the general hardness and resistance to pressure, all these features being in contrast with ovarian pelagic eggs. One of the largest specimens examined by Fulton from the ovarian stroma measured 1°39 mm. in diameter and contained two groups of three and four oil-globules. Some other slightly smaller eggs contained a promi- nent straw-coloured oil-globule: thus in an egg of 1:02 mm. diameter the oil-globule measured 0°25 mm. All these were quite opaque and white by reflected light and still contained in the follicular investment. Holt has recorded the capture of ripe females off the west coast of Ireland in July and August, and one spent in June.* Cunningham (4g, p. 322) also has found ripe females in August at Plymouth. The demersal character of the eggs and the relatively deep-water habitat of the spawning fishes are sufficient to account for the present lack of knowledge of embryonic development. The youngest post-larval stages yet recorded are those described by Schmidt (21d) from four specimens (7? mm. to 19 mm. long) taken by the Zor in August and September, 1906 (three from various parts of the Channel and one from the Bay- of Biscay); to which must be added one specimen of 14 mm. taken by the Oithona’s young-fish trawl off Plymouth Sound on the 17th of September in the same year. BIBLIOGRAPHY. 1, Brook, G.—a. Preliminary Account of the Development of Lesser Weever (Trachinus vipera). Linn. Soc. Journ. Zool., XVIII (1884), pp. 274- 91, Pl. III-VI. b. On Some Points in the Development of Motella mustela. Ibid., pp. 298-306, Pl. VIII-X. 2. Browne, F. Balfour.—Report on the Eggs and Larvae of Teleostean Fishes observed at Plymouth in the Spring of 1902. Journ. Mar. Biol. Assoc., N.S., Vol. VI (1903), pp. 598-616. 3. Canu, E.—a. Ponte, ceufs et larves des poissons utiles observés dans la Manche. Annales de la Stat. Aquic. de Boulogne, Vol. I (1893), pp. 117-382, PI. VIII-XV. b. Ibid., Vol. II (1894), pp. 63-72, Pl. I-V. * Report of the Council Royal Dublin Society, 1892, p. 245. 56 A. E, HEFFORD. 4, Cunningham, J. T.—a. Studies on the Reproduction and Development of Tele- 5 ~J ostean Fishes occurring in the Neighbourhood of Plymouth. Journ. Mar. Biol. Assoc., N.S., Vol. I (1889-90), pp. 10-54, Pl. I-VI. b. A Treatise on the Common Sole. Plymouth (1890). c. On Some Larval Stages of Fishes. Jowrn. Mar. Biol. Assoc., N.S., Vol. II (1891-2), pp. 68-74, Pl. III-IV. d. The Reproduction and Growth of the Pilchard. Ibid., pp. 154-7, PX e. The Egg and Larva of Calhonymus lyra. Ibid., pp. 89-90, PLY: f. The Life History of the Pilchard. Ibid., N.S., Vol. III, pp. 148- 53. g. Natural History of Marketable Marine Fishes of the British Isles. London (1896). . Ehrenbaum, E.—a. Eier und Larven yon Fischen der deutschen Bucht. I. Wissensch. Meeresuntersuchungen, Abt. Helgoland, II (1897), pp. 253- 328, Taf. III-VI. b. Eier und Larven von Fischen, 1 Teil. Nordisches Plankton (1905). c. Ueber Eier und Jugendformen der Seezunge und anderer im Fruhjahr laichender Fische der Nordsee. Wassensch. Meeresunter- suchungen, Abt. Helgoland, VILI (1907), pp. 201-70. d. Eier und Larven von Fischen, 2 Teil. Nordtsches Plankton (1909). . Ehrenbaum, E., and 8. Strodtman.—Eier und Jugendformen der Ostseefische. Wissensch, Meeresuntersuchungen, Abt. Helgoland, V1 (1904), pp. 57-126. . Fabre-Domergue et E. Biétrix.—Développement de la Sole. Paris (1905). . Fulton, T. Wemyss.—The Ovaries and Ovarian Eggs of the Angler (Lophius piscatorius) and of the John Dory (Zeus fuber). Sixteenth Annual Rept. Fish. Bd. Scotl. (1898), pp. 1381-4, Pl. IIL. 9. Guitel, Fr.—Recherches sur les Lepadogasters. Arch. de Zool. experim., 2 8., 10 11 Vol. VI (1888), pp. 576-94, Pl. XXXIII-XXXV. . Heincke, Fr., and E, Ehrenbaum.—Eier und Larven von Fischen der Deutschen Bucht. II, Die Bestimmung der schwimmenden Fischeier und die Methodik der Eimessungen. JVissensch. Mecresunters. Abt. Helgoland, III (1899), pp. 131-332, Taf. IX—X. . Holt, E. W. L.—a. Survey of Fishing Grounds, West Coast of Ireland, 1890, I. On the Eggs and Larvae of Teleosteans. Sci. Trans. Roy. Dubl. Soc., Vol. IV, 8. II (1891), pp. 485-74, Pl. XLVIII-LII. b. Survey of Fishing Grounds, West Coast of Ireland, II, 1890- 1901. On the Eggs and Larval and Post-larval Stages of Teleosteans. Ibid., Vol. V., 8S. II (1893), pp. 5-121, Pl. I-XV. c. Preliminary Notes on the Reproduction of Teleostean Fishes in the South-Western District. Journ. Mar. Biol. Assoc., N.S., Vol. V (1897), pp. 41-50. d. Notes on the Reproduction of Teleostean Fishes in the South Western District. Ibzd., pp. 107-55 and 333-40 (1897). a NOTES ON TELEOSTEAN OVA AND LARVAE. ai e. On the Breeding of the Dragonet (Callionymus lyra). Proce. Zool. Soc. (1898), pp. 281-315, Pl. XXVI. f. Recherches sur la Reproduction des Poissons Osseux. -Annale du Musée @histoire naturelle de Marseille, Zoologie, Tome V, No. 2 (1899), pp. 1-128, Pl. I-IX. 12. Holt, E. W. L., and Scott, S. D.—A Record of the Teleostean Eggs and Larvae observed at Plymouth in 1897. Journ. Mar. Biol. Assoc., N.S., Vol. V (1897-9), pp. 156-71. 13, Holt, E. W. L., and Byrne, L. W.—British and Irish Gobies. Report Fisheries Trel. (1901), pp. 37-66, Pl. I and II. 14. McIntosh, W. C.—a. Further Observations on the Life History and Develop- ment of the Food and other Fishes. 9th Annual Rept. Fish. Bd. Scotl. (1891), pp. 317-34, Pl. X-XIII. b. Contributions to the Life History of the Food and other Fishes. Itid., 10th Rept. (1892), pp. 273-322, Pl. XIV, XVI. c. Ibid., 11th Report (1893), pp. 239-49, Pl. VITI-XII. d. Ibid., 12th Report (1894), pp. 218-30, Pl. II-IV. e. Ibid., 14th Report (1896), pp. 171-85, Pl. V. f. Ibid., 15th Report (1897), pp. 194-211, Pl. V-VII. g. On the Life History of the Shanny (Blennius pholis, L.). Zeitsch. f. wissensch. Zool. 82 (1905), pp. 368-78, Pl. XX. 15. McIntosh, W. C., and Masterman, A. T.—The Life Histories of the British Marine Food Fishes. London (1897). 16. McIntosh, W. C., and Prince, E. E.—On the Development and Life Histories of the Teleostean Food and other Fishes. Trans. Roy. Soc. Edin., Vol. XXXV, Pt. III (1890), pp. 665-946, Pl, I-XXVIII. 17. Matthews, J. Duncan.—Note on the Ova, Fry, and Nest of the Ballan Wrasse. 5th Ann. Rept. Fish. Bd. Scotl., p. 245, Pl. XI. 18. Moreau, Emile.—Histoire Naturelle des Poissons de la France. Paris (1881). 19. Petersen, C. G. Joh.—On the Eggs and Breeding of our Gobiidae. Report of the Danish Biological Station, II. (1891), pp. 1-9, Pl. I, a-1 b. 20. Raffaele, F.—Le uova galleggianti e le larve dei Teleostei nel golfo di Napoli. Mitteil. Zool. Sta. Neapel., VIII (1888), pp. 1-84, Tav. 1-5, 21. Schmidt, Johs.—a. The Pelagic Post-larval Stages of the Atlantic Species of Gadus, Pt. I. Meddelels. fra Komm. for Havundersog. Serie Fiskeri, Bd, I (1905), No. 4, Pl. I-1II. bi Idem. Pt. If. Ibid, Bd. If. (1906); No; 2; PL I. c. Marking Experiments on Plaice and Cod in Icelandic Waters. Ihid., Bd. II (1907), No. 6. d. On the Post-larval Stages of the John Dory (Zeus faber, L.) and some other Acanthopterygian Fishes. Jbid., Bd. II (1908), No. 9;'PI. 1. 58 A. E, HEFFORD. EXPLANATION OF PLATES I AND I, Illustrating Mr. A. E. Hefford’s “ Notes on Teleostean Ova and Larvae observed at Plymouth in Spring and Summer, 1909.” PuateE I. Fig. 1. Zeus faber, ripe ovarian egg ; diameter, ca. 2°09 mm. Fig. 2. Zeugopterus punctatus, pelagic egg ; diameter, 0°99 mm. Fig. 3. Egg of Blennius pholis, lateral view ; dimensions, 1°8 x 1:2 x 0°8 mm. Fig. 4. Egg of Labrus miztus ; diameter, 0°94 mm. Fig. 5. Egg of Lepadogaster bimaculatus ; dimensions, 1°44 x 1-24 x 0°62 mm. Fig. 6. Blennius pholis, newly hatched larva ; length, ca. 4°4 mm. Fig. 7. Gobius paganellus, newly hatched larva; length, 4°8 mm. Fig. 8. Labrus miztus, newly hatched larva; length, 3°26 mm. Fig. 8a. Labrus mixtus, sketch showing arrangement of anterior dorsal black pig- ment. Fig. 9. Zeugopterus punctatus, newly hatched larva ; length 2°90 mm. Fig. 10. Z. punctatus, larva ca. 3 days old. Puate II. Fig. 11. Solea lascaris, newly hatched larva; length, 3-46 mm. Fig. 12. S. dascaris, larva 4 or 5 days old. Fig. 13. Serranus cabrilla, early larva ; length, 2°30 mm. Fig. 14. Onos? tricirratus, Bl., newly hatched larva ; length, 2°32 mm. Fig. 15. “Onos, species B” (? or abnormal form), newly hatched larva ; length, 1°84 mm. Fig. 16. Lepadogaster bimaculatus, newly hatched larva ; length, 4°26 mm. PLATE J] Assoc. VoL. IX. JOURN. Mar. BIOL. ee mana et — Se oa \ . oe ett a @ so ASB El del: JourN. Mar. Brox. Assoc. VoL. IX, PLATE II A. E. H. del. Ae! be ad Fi . [> 7 J ne Pr hy a we es ona | i” lh 4a) , jug _—_ Prvge a se if W xy 5 a i 5 a, s @ af , wh = Fee a mu | rh i * , Vib if i a i. Y ' eh ba) : : me TNL | i +, Ti. Sw % : iy) haan i Notes on the Littoral Polycheta of Torquay (Part III). By Major E. V. Elwes. Aphroditide. A synopsis of the Aphroditide of the English Channel by Mr. T. VY. Hodgson is given in the Jowrnal Marine Biological Association, Vol. VI, No. 2, 1900. APHRODITA ACULEATA, Lin. McIntosh, Mon. Brit. Ann., Vol. II, p. 247. This species is sometimes found in some numbers, thrown up on the shore after heavy weather, especially at Anstey Cove and Tor Abbey Sands. It is recognized by the fishermen as a “ curiosity.” LEPIDONOTUS SQUAMATUS, Lin. JlcIntosh, Mon. Brit. Ann., Vol. II, p. 274. Only two or three examples found under stones on Babbacombe beach. LEPIDONOTUS CLAVA, Mont. JJclntosh, Mon. Brit. Ann., Vol. II, p. 280. Occasionally found on all the beaches. Numerous specimens were found on a large buoy in Torquay Harbour. LaGIsca FLoccosa, Sav. McIntosh, Mon. Brit. Ann., Vol. I, p. 298. Fairly common under stones. LAGISCA EXTENUATA, Gr. JleIntosh, Mon. Brit. Ann., Vol. I, p. 307. Hornell, Fauna of Liverpool Bay, 1892, p. 136, Pl. XIU, Fig. 8. Very common in roots of Laminaria and under stones. ‘The scales have the groups of papille surrounded by lines as represented by Hornell, EVARNE IMPAR, Johnst. McIntosh, Mon. Brit. Ann., Vol. I, p. 358. Rare. Recorded by Gosse from Anstey’s Cove. HARMOTHOE SPINIFERA, Ehlers. Jfclntosh, Mon. Brit. Ann., Vol II, p. 327. One example only. 60 E. V. ELWES. HALOSYDNA GELATINOSA, M. Sars. McIntosh, Mon. Brit. Ann, Vol. II, p. 384. One specimen under a stone on Babbacombe beach. POLYNOE SCOLOPENDRINA, Sav. JcIntosh, Mon. Brit. Ann., Vol. I, p. 389. Not uncommon at Corbyn’s Head. STHENELAIS BOA, Johnst. McIntosh, Mon. Brit. Ann., Vol. I, p. 408. Not uncommon in the sand at Tor Abbey Sands. SIGALION MATHILD&, Aud. and Edw. JclIntosh, Mon. Brit. Ann, Vol. II, p. 427. This is the only one of the Torquay Aphroditide which has not been also recorded from Plymouth. It is fairly common in the sand at Tor Abbey Sands and Livermead. PHOLOE MINUTA, O. Fabricius. McIntosh, Mon. Brit. Ann., Vol. II, p. 437. The most numerous of all the Torquay Aphroditidee, inhabiene especially the Laminaria roots. Glyceride. GLYCERA CONVOLUTA, Kef. De St. Joseph, Ann. Sct. Nat. Zool., Vol. XVII, 1894, p. 27. Fairly numerous in Tor Abbey Sands and at Livermead. GLYCERA LAPIDUM, Qfg. McIntosh, “On the British Glyceride,” Ann. Nat, Hist., 8. 7, Vol. XV, p. 39, 1905. One specimen in. the inner harbour of Torquay and one on the Babbacombe beach. Eunicide. This family is represented at Torquay by five littoral species. For the key to the Eunicide of the English Channel the papers by Baron de St. Joseph, entitled “Les Annélides Polychetes des Cotes de Dinard” and “Les Annélides Polychéetes des Cotes de France,” the “Notes on the British Eunicide,” by Professor McIntosh, Annals of Natural History, Vol. XI, p. 553, 1903, and the Cambridge Natural History, Vol. I, have been consulted. LYSIDICE NINETTA, Aud. and Edw. Johnst., Catalogue of Worms, p. 140. Small specimens thirty to fifty millimetres in length; extremely common amongst Laminarian roots and limestone rocks. NOTES ON THE LITTORAL POLYCHATA OF TORQUAY. 61 NEMATONEREIS UNICORNIS, Grube. De St. Joseph, Ann. Set. Nat., V, 1888; pe 207. Fairly common in the limestone rocks at Babbacombe, but as is the case with the last species it is very rarely perfect. STAUROCEPHALUS RUBROVITTATUS, Grube. De St. Joseph, Ann. Sei. Nat., V, 1888, p. 235. One specimen obtained at an unusually low spring tide at Corbyn’s Head. OPHRYOTROCHA PUERILIS, Clpd. and Meczn. Cambridge Nat, Hist., Viol Tf, p. 319, Pic. 170. This little worm is frequently seen on the sides of glass vessels containing roots and pieces of rocks. On one occasion a small aquarium in the museum of the Torquay Natural History Society was found to be swarming with this species. LUMBRICONEREIS LATREILLI, Aud. and Edw. De St, Joseph, Ann. Sct. Nat. Zool., V, 1898, p. 276. Three or four in rather coarse gravel on Babbacombe beach. Spherodoride. EPHESIA GRACILIS, Rathke. De St. Joseph, Ann. Sci. Nat. Zool., XVII, 1894, p. 33. McIntosh, Ann. Nat. Sci., 8. 8, Vol. II, 1908, p. 528 and 540. ; Two or three from Meadfoot beach. EPHESIA PERIPATUS, Clpd. nee Johnst. Claparéde, Beob. tiber Anat. und Ent. wirbellosen thiere, p. 50, de St. Joseph, Ann. Sci. Nat. Zool., XVII, 1894, p. 41. Two specimens from Corbyn’s Head. According to de St. Joseph this species differs from #. gracilis by several characters, but he only mentions two, viz. the composite bristles and the absence of the “Véventail de papilles” below the feet which exists in Z. gracilis. The bristles of #. peripatus of the Torquay examples seem, besides being compound, to be not quite so stout and not so much bulged as those of E. gracilis. Ariciide. ARICIA LATREILLI, Aud. and Edw. De St. Joseph, Ann. Sci, Nat. Zool., XVII, 1894, p. 85. Several examples were dug up from the sand at Tor Abbey Sands. In this species there are about thirty bristle-bearing segments in the anterior region, while in A, ewviert there are only twenty-one. 62 E. V. ELWES. Spionide. In preparing the accompanying key to the Spionidee of the English Channel Mesnil’s paper, entitled “Etudes de Morphologie externe chez les Anné¢lides” and Professor MeIntosh’s “ Notes on the British Spionide,” Annals of Nat Hist., 8. 8, Vol. III, have been consulted. SCOLECOLEPIS VULGARIS, Johnst. McIntosh, Annals of Nat. Hist., Suc vol, LT 909) pb: At the west end of Tor Abbey Sands; rare. SCOLECOLEPIS FULIGINOSA, Clpd. McIntosh, Annals of Nat. Hist., S, 6, Vol. Tht, 1909, p. 160. Very numerous at west end of Tor Abbey Sands and at Livermead. In December numbers were found coiled up together under stones. NERINE CIRRATULUS, Delle Chiaje. J/cIntosh, Annals of Nat. Hist., S. 8, Vol. 1111909, p, 158, Tor Abbey Sands; not numerous, AONIDES OXYCEPHALA, Sars. JMesnil, Bull. Sci. France et Belgique, X XIX, 1896, p. 242. Numerous in rather foul mud under stones at Livermead. Potypora cILIaTA, Johnst. McIntosh, Annals of Nat. Hist., 8. 8, Vol. ILI, p. 169. Very numerous in the small pools in the limestone boulders on the shore. Potypora FLAVA, Clpd. McIntosh, Annals of Nat, Hist. 8. 8, Vol-iiiy pried, Numerous on rocks and in pools. SPIOPHANES BOMBYX, Clpd. McIntosh, Annals of Nat. Hist, 8. 8, Wool buleap: 67. A few specimens at the east end of Tor Abbey Sands, Mesnil remarks that he found this species in company with Achinocardium cordatum ; this sea urchin is also common on Tor Abbey Sands. Magelonide. MAGELONA PAPILLICORNIS, Fr. Miiller, MJcZntosh, Annals of Nat. Hist., S. 8, Vol. I11, p. 174. One example at a very low spring tide on Tor Abbey Sands. Ammocharide. . - ‘ OWENIA FUSIFORMIS, Delle Chiaje. De St. Joseph, Ann. Sci. Nat. Zool., V, 1898, p. 397, NOTES ON THE LITTORAL POLYCHATA OF TORQUAY, 63 The tubes of this species are very numerous on Tor Abbey Sands; they appear to be loose in the sand, not fixed vertically, as is usual with tube-dwelling annelids in sand. They are largest in the middle, tapering towards both ends, made chiefly of small pieces of shell placed edgeways. Cirratulide. In the accompanying key to the Cirratulidee of the Channel the classification of Caullery and Mesnil in Les formes épitoques et Vevolution des Cirratuliens is adopted. AUDOUINIA TENTACULATA, Montagu, De St, Joseph, Ann, Sct, Nat. Zool., XVII, 1894, p. 49. Numerous at Meadfoot, Hope’s Nose, and Tor Abbey Sands in rather foul mud; young ones about 40 mm, in length appear to live in crevices in rocks, DODECACERIA CONCHARUM, Oersted. Caullery et Mesnil, Annales de l Université de Lyon, Fasc. XX XIX, 1898, p. 11. Very numerous in the limestone boulders at Babbacombe. HETEROCIRRUS VIRIDIS, Lang. = 7, flavoviridis, de St, Joseph, Caullery et Mesnil, Ann. de l’Université de Lyon, Fasc. XX XIX, 1898, p. 117. Found occasionally in small pools in limestone rocks at Babbacombe. HETEROCIRRUS CAPUT ESOCIS, de St. Joseph. Caullery et Mesnil, Ann, de l’ Université de Lyon, Fase. XX XIX, 1898, p. 122. Two or three examples found in the same localities as the last species. I have not seen any British records of these two species of Heterocirrus. Terebellide. The accompanying key to the Terebellidz is founded on the table given by Baron de St. Joseph in “Les Annélides Polychétes des Cotes de Dinard,” Ann, Sci, Nat. Zool., XVII, 1894, p. 180. POLYMNIA NEBULOSA, Montagu. De St, Joseph, Ann. Sci, Nat. Zool., XVII, 1894, p. 219. Occasional specimens at Corbyn’s Head and in rocks between Oddi- combe and Babbacombe beaches. ; POLYMNIA NESIDENSIS, de St. Joseph. Ann. Sci. Nat. Zool, XVII, 1894, p. 211. Very common in Laminaria roots, etc. LANICE CONCHILEGA, Pallas. De St. Joseph, Ann. Sci. Nat, Zool., XVII, 1894, p. 211. Numerous on Tor Abbey Sands, especially at the east end. 64. E, V. ELWES. Ampharetide. MELINNA ADRIATICA, Marenzeller. Sitzb. d.k. Akad. Wiss. zu Wien, LXIX, p. 472. Two at extreme low water at Livermead amongst Zostera roots. Maldanide. CLYMENE CERSTEDII (?), Clpd. De St. Joseph, Ann. Scr. Nat. Zool., XVII, 1894, p. 137. On the east side of Tor Abbey Sands; not common. LEIOCHONE CLYPEATA, de St. Joseph. Ann, Sci, Nat. Zool., XVII, 1894, p. 139. Numerous at extreme low water in the centre of Tor Abbey Sands. Capitellide. NOTOMASTUS LATERICEUS, Sars. De St. Joseph, Ann. Set. Nat., XVII, 1894, p. 117. Under stones, Corbyn’s Head and Livermead. Opheliide. ' POLYOPTHALMUS PIcTUS, Duj. De St. Joseph, Ann. Sci. Nat, Zool., V, 1898, p. 385. Common amongst Corallines, ete., in rock pools. - Arenicolide. ARENICOLA MARINA, L. Gamble, Quart. Journ. Micro. Sci., XIII, p. 419. Common on Tor Abbey Sands. ARENICOLA ECAUDATA, Johnst. Gamble, Quart. Journ. Micro. Sei., XLII, p. 419. This species seems to be very different in its habits to 4. marina ; instead of burrowing in soft mud and sand it lies under stones in gravel at Hope’s Nose and Babbacombe beach. Chlorhemide. SIPHONOSTOMA AFFINIS, M. Sars. De St. Joseph, Ann. Set. Nat. Zool., XVII, 1894, p. 96. Under stones at Corbyn’s Head. Sabellide. The accompanying key to the Sabellids of the English Channel is founded on the table given by Baron de St. Joseph in “ Les Annélides NOTES ON THE LITTORAL POLYCHETA OF TORQUAY. 65 Polychétes des Cotes de Dinard,” Ann. Sci. Nat. Zool, XVII, 1894, p. 248. SABELLA PAVONINA, Sav. De St. Joseph, Ann. Sci. Nat. Zool., XVII, 1894, p. 267. I was somewhat surprised to find several examples of this large worm in the inner harbour at Torquay only a few yards from the “Strand.” They were living in mud and gravel which could hardly be called clean. POTAMILLA RENIFORMIS, O. F. Miiller. Soulier, Revision des Annélides de la region de Cette, p. 120, Fig. 4. This species is found on the sides of the cave under the men’s bathing-place at Petit Tor. This is the cave mentioned by Gosse in the British Sea Anemones and Corals, where he found the sea anemones Halecampa microps and Hdwardsia carnea. POTAMILLA TORELLI, Mgr. De St. Joseph, Ann. Sci. Nat. Zool., XVII, 1894, p. 296. Common in the small rock pools in the limestone rocks between Oddicombe and Babbacombe beaches. FABRICIA SABELLA, Ehr. De St. Joseph, Ann. Sci. Nat. Zool., XVII, 1894, p. 319. A little Sabellid which appears to be referable to this species is very common in the little pools in the rocks at Babbacombe, in com- pany with the last species, Polydora and Dodecaceria. It lives in small holes in the rocks, with a tube of mud projecting a little from the opening. ORIA ARMANDI, Clpd. Soulier, Revision des Annélides de la region de Cette, 1902, p. 114, Fig. 2. One specimen from Babbacombe rock pools. As de St. Joseph remarks, the eyes in this species quickly disappear, while in F. sabella they are persistent even in Balsam preparations. I have also obtained this species at Newquay, Cornwall. JASMANEIRA ELEGANS, de St. Joseph. Ann. Sez. Nat. Zool., XVII, 1894, p. 316. Found occasionally crawling up the sides of glass vessels containing roots of Laminaria and pieces of limestone rock. It was first recorded as a British species by Miss Newbiggin in 1900. AMPHIGLENA MEDITERRANEA, Clpd. Soulier, Revision des Annélides de la region de Cette, p. 109, Fig. 1. Found under the same conditions as the last species. NEW SERIES.—VOL, IX. NO, 1. OCTOBER, 1909, E 66 E. V. ELWES. Serpulide. In preparing the key to the Serpulids of the English Channel the table given by Baron de St. Joseph in the Annales des Sciences naturelles Zool., XVII, 1894, p. 259, and, for the genus Spirorbis, the papers by Caullery and Mesnil, “Etudes sur la morphologie, ete., chez les Spirorbes,” have been consulted. SERPULA VERMICULARIS, Lin. De St. Joseph, Ann. Sci. Nat. Zool., XVII, 1894, p. 328. On shells thrown up on the shore at Tor Abbey Sands. POMATOCEROS TRIQUETER, Lin. De St. Joseph, Ann. Sct. Nat. Zool., XVII, 1894, p. 353. Extremely common on stones. HyYDROIDES NORVEGICA, Zunn. De St. Joseph, Ann. Sei. Nat. Zool., V, 1898, p. 440. On a stone at Petit Tor beach; numerous on buoys in Torquay Harbour. SPIRORBIS BOREALIS, Daudin. Caullery et Mesnil, Bull. Scien. de la France et de la Belgique, XXX, 1897, p. 211. Very common on Fucus. SPIRORBIS SPIRILLUM, Lin. =lucidus, Mont. Caullery et Mesnil, Bull. Scien. de la France et de la Belgique, XXX, 1897, p. 198. On Sertularia abietina thrown up on the shore. Hermellide. SABELLARIA ALVEOLATA, Linn. Cambridge Nat. Hist., Vol. U1, Figs. 131 and 135. 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: ; ‘ : : ; : ‘quar og “TT ‘squtod o\4 YT syoy[oy0.10 pesurAy “Mato peue yysiq “aeypoursq jo sated AqyueMy qnoqy “eur, ® UI sofa anoj ‘pazutod wmn1w0}s01g ‘SHCINOY snuodr) ‘ydosor 49 ap ‘suysoubuo) “NTy * : : : Y : ; : 5 : : - MIU OOT 07 OL “TT. “squtod omy YIM sjoyojoro pesuryy “wtIpoueaq 07 poutof pue sv Suoy sv eyjeuey ‘seXa anoj ‘peqyutod Alaa wNTUOyso1g "SUCINGUT NT SNUOy) (4) ysuery ‘noun pT ° 5 : : ; . 3 ; : . : : “WU OL “TT 400} YQYSto uo AT[VAQUEA QOUSWIUIOD sJayo}OID peSutAA “AMOF TA1o [vue ‘anoy sofa ‘sassoq popunor org Aq poyvUyUIIE, WUNTUIOIsSO1g ‘OldSOUDIT, SNUO+) “(j) aq ‘sewoony= ‘[TUsaTy ‘siswaurzumu “go : i ‘ ; : : ; “UI Og “TT “ANOF “TaITO TRUY «“‘quewdes yIWeEqJY 07 WE} -INY} Ye souatMIOD syutod O44 TILA sjoyoj}OIN peSura ‘sede anoj ATTVNsN “GuoIy UT popuNoI WINTWOWSOI ‘Oldg snue+) "THNNVHO AHL AO SLSVOO HSTTIONA GNV HONAYT AHL NO GNQOA WAINOIdS AO SHIONdS AHL OL ACH "cole YS a} Wol papsooe1 yak JON , “‘mseyyT ‘ovyounighjod "qy * ‘ : WU GT TT ‘Sopstaq eSarvy Jo spury yUoIEYIp OMY YIM JUOULSOS AIT ‘VIGUVOOOd Ssnuoer) ° “WUT G ST ‘apis eaeouoo ey] uO syutod yenboun omy Sur ‘Burry ‘njnuim “q+ -UL10J pooy Jo ssura ‘apis X9AUOD WO pooy Jo pUry v YIM se]}st1q [eULIONGY ‘QUOUIBES YJWOAS UO oUOUIUIOD BIYOUMTG |. 5 4 oy gaau UOA en Eee pit: : ‘UU g “TT ‘OpIs PALOTIOD UO Y4OO} T[vUIS YT | Bue gh thee Trasey (PPro) ‘d+ sepystaq TeUOUqY ‘quoulses YqUe} Wo eouEUTtMOD aeTyoUEIg cae sueyae Mae Mpa : SF oB[ng BqNOyITM soto : -JO10 pesuIa jo Weg ‘yooy ureu Sc) GET A eo. “rT ‘sorqy JO Yshaq v TILA sotmTjeu0s ‘yyQ00q Tenbo yee ogee 8 a ae hata taal d Apreou WIM diy poyeounsy v ut Surpuo soyystaq feuouqy oated cee sat ‘quowSes Surrveq-apistaq WYSto oy} wo souswMOD wioURIg [er9 qe] aie ; 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J eats ; : é : ; ; : : “UU YG 04 ; * —- MOJ IO vaIyy Aqyensn Te ie cod Hivaies 02 “TT ‘pus yoro ye porodey sapystaq ouy Aro JO so—punq — | syotpo}010 ER jo daquinN ‘quoulsas geo 1 17705 SSP lo) ied yg ang sourds OYT[-oTpeou IMOYITA syuowses aes SuLIvoq-apsluq YIYSIa oy} Wo ooweurUI0D eee : “UU QZ S wiyouriq ‘sourds 10 yY90} [B10yLT NOTIN TEE G a) elite ‘gourds oYI[-e[poou 9e1yy 10 OM} YITA syueTIses Pes ) yooy apSurs v ut Surpua so qstaq [vuIoUc Seety I desi a Sea eehoay Gea Ty \V Vi a Ait Re p : ; Wt g “TT “qUeTASes SuLIvoq-oT]SIIq YIUIAVS MO DDTOTIMIOD aIYULIg TUSO, UMN) ‘T* 1 -quouBas Isa WO Juaserd Sopystaq [RSIOC] “4S8a10 SNOAGY OYT[-GUIO B YYTM JlouFes ITY Jo sopystaq [euLoUqy “VUOGATOd Snuor) ‘pdyo ‘whquog 'g ° : MUI OG “TT ‘OMY “TAAIO [RU “W9JANOJ 07 UdAdTA ‘sJoYoJOIN pasuIM jo JoqMNN “Ao][oap oquy, ‘IX) ‘SHNVHdOIdQ snuor) ea ‘penuyuos—'VCINOIdS HHL AO SHIOUdS AHL OL AWS I~ 73 ON THE LITTORAL POLYCHATA OF TORQUAY. Ss = Us NOTI "BAI YSIJWUG OY} VIO popslooal JOA JON y f . ‘SyoUeY on ie oP ; : ; : Areyideo ev sopstug, “quesqe sea pu sjotpojorg [[t8 oa0qe “UOT (Snsuoo1ajazy) LUALOYT, » paytasttt ‘quosoid ‘ydasop 1g op ‘sepuna : ; j ; : oath * queuses paryy oy} aareadl syuaure|y (syuommeyy -oumyf = Suey ‘siprma snasvosapezy \ 104¥e spodornau ur sopystaq AreypIdeo oN “pytq syoyooIgf "6, 1enovqua y, Its ay} : a : UBIY Layout ‘yda ‘ : : ; ; ‘uu gt “TT «‘spodomou ayy [[V oe agen lt rN ah oO pe ad i a ea haat syoyoqorg | = [p18 Jo sared Las ebublly ‘49 ep ‘sr00sa yndno snamoouajazT (ut quosoad sayystaq Aaeypideg -‘xede 4v oqyvoundy szeyoj0Iy Wyse weYy ar0yy syUIWe[Y, : 5 Ie[NIRUO} o ated Peers BETA Sm TR hare De EO Fa) fll Fate career jo ard y -C[Y [IS MOTod poytosur syuoWRTY aepnorqzuay, “(sated 4ySte 07 anoj ‘Moy) syuetUeTY [TTD : * squowely aepnory -0} a} Soldavo YOU “sq U9UISas ‘phy ‘omjnovjuay DUnopny * : : : - ‘tur OOZ “TT «‘sodO ONY JUOTISIS oYy JO QUOUT UL IOIA}JUL ayy JO squewsas oy} JO a1our 10 | aUO ssodoV (SyUDTURTY auo uo avodde s]piS [eioqery pqs oy] WRT Aoyoryy AyQOUNSIp Jou) “ION ‘snyouns srynqo.wug | ; ‘ I é jin aesge et a eee ( ‘ ‘ke eet iad lead lee pue Areypidvo ‘sayystaq Jo spury omy, “quosoad saAgp sv quotes otuvs oy} UO | JO MOI ostoastta} W ‘Joy ‘semiofyy snynpoiuijy, “WU OP 04 9Z “TT ‘AavT[Idvo seystaq ey [Ty ‘seAo ON il avadde syuaueypy avpnorque y, "peaqsiyD “wnwmyrwod nr.1a9NI0p0(T "THNNVHO GFZHL AO SLSVOO HSITONA GNV HONWHYA AHL NO GANONOKX WAITOALVYNIO AHL HO SHIOUdS GNV VAHNYHO HHL OL AX ‘Role YSIJIG 9Y} Woy poprooat yok ION , BjQ “wpo9.ts9) engoouoprned 4 ; : ‘Ul OEL “TT ‘wu g ‘YIproiq ‘syueuISes Sutsvaq-apystaq 0m4-AQUeALT, "T1119 LO soda snue oy} YOIYM Jo sovjIns oy} uo ‘asepuodde oyl[-Jeay, eAvOMOD V IIA JUEUISES [UW] 420} qnoyyia ‘ydesor 4g ep “nynadhjo suoyoora'T { -o[4st1q outu-AjT9M4 07 dAY-AQUAM I, Perugeoues el ae Sra dee eee ee es ae 2 E sjuem =) a : : : 2 ° : c : : * spaeadn poqootp |-Ses 1o11eqsod > BIC) ‘saprouawfiya vruojsuyory + YOO] esIe, suo puUe {492} 27941] eo1y} YITas Sutpus syueuises Suteveq 4 UL SMO1 [BUIpNy Fae oe te 4 -a]Islaq, 901T]} ASIY JO syoyo}oIg ‘ySuey tenbs jo yje0} YPM JouUNy [RU | -Isu0T OMy UI ages a pesuriie apideg pee > \a . : ‘UU Og TT ‘WU [ ‘YYpreIg ‘saTJsTIq qnoyILA JUeUISes ran . “pd y ‘epajsuny (auaulipongy) auawhyy » peur oy) Surpaooid syusuisas OMY ay, “SNAITO [eayUeA SUMO] UO YALA SOUIT} TH 3 e ihe -oulos ‘fenboun A19A JoUUN; [vuR JO YJoay, “payUepur you pay Jo youg |- : ajueureely ee -5as 101103s0d uo |e ‘ jeuuny 31 : : : s 3 2 Ww OG “TT ‘wu g ‘yypeag ‘soyystiq | axpided qjeus oN : . °F qnouyIM jUuotIses [eur oy} Suipoosid syueurSes vey} 9 ‘yenboun sapwnuguny (auaumhjona) awawl79 ue l q *P Ee tubal REY JO Tenbo youunj [vue Jo Ye], ‘SUOT}LJUApPUT [eIAAES YIM prey Jo youd "THNNVHO HHL AO SLSVOO HSITIONA GNV HONGYA AHL NO WAINVCIVW AHL AO SAIOHdS GNV VYUANHO HHL OL AM 74 NOTES ON THE LITTORAL POLYCHATA OF TORQUAY. ‘Avg “nsnpau DUNO ‘ : ; ; q " snosetunt pur ouy AJA S][IS BY} JO SUOISTATP [vUTUIDy = omy ‘ 920} XIS 0} IMOF YITAL ‘OxtT- quoo “yq984 JO SMOI aSTOASURIY JNOTYTM TULOU YL”) 03 OOL ‘"T urs JO SoUlI]s [JIM spus oy ‘6 aitiag’s ae it acy ¢ F oO C , SQM OSEAN Ag ‘pad ianeien SpE raurouy nolo aie om qoeq peat See Jo TULOUN Jo ALor o]HOp vB YITAL SJUIUISas YJWIE}UIAAS 07 YIYSIe oI, "1[}99} poe gin * uur gg ‘Ty ‘quesoid sod ‘asuer0 FYystaq aa1Y} 10 ‘OMy pau ‘aferyg ajay ‘siswapisau nruwh; wa sepowjueT, “UMOIG 10 pod Apog ‘turun jo Mor a[surs ‘uo JO SMOI pL ge as Bug syuoUrses SULIvAK- aTslIq YJUseyUAes 07 YWYSIe oy, {| ostaasuvr] omy : a i aos - ‘wut 00% “"T "yuesod soATy Yt Turou ys), bed rm ee 4 ‘ : h { YSIPPeT 10 AyTYM sopoRpUaT, “OTA ITAA poqyods ‘umoaq TELLTC CD WOW PSONN GM MUM T vo “par Spog “Kea-J[ey Suryooyrequt turoun jo Mor arqnop Re: ata syuauides Sulrvaq-apjstlq YIUIIWOAIS 0} YAYSte op], 7 “oumpedny (maadorq) nypoqosa y 1 ; ‘uur ¢g “Ty «“quasoad sod “par sopoejueL, Y700} XIs | 0} 9a1Y} JO SMOX INOF IO ory} YIM TuIOU/) ‘€poq oy} Jnoysnoayy sapystaq Aavypidey | * ‘uur QOL “TT “quesead sodq le SMOI 9[qnOp UI WeuTOpge jo yaed yuoIy aqnay ‘syv0ns6 apuuydwp < ul turuUp § “quoay ut por Ataa Apoq ‘ozTPAs RGAE erro a seria ‘s][IS Jo saved omy, ‘squow ‘soTpoUBAg SEIT aes Oe Me iee ae \ 808 Aquasy 07 WsezUAes UY sopystIq Aaeyuder) SOIOUINU ITA ala Peds veer eh : ; a . 25 av YOM YI00} | Ve], “pua oyy geU OSG EL besa teens a aSIOASUR1] JO / AvaU 492} AY] ‘BY Uspuwnpy apupydwy < ‘asue10 soporywoy, ‘s[[ls jo sired oeayy, JO sysIsmoo i SoaOMETAR EAE ATE ae te, PES CS ‘squouIB9" wsoqtaaes UT sayyst1q Areypideg TI3 youn eee ‘hia Barn aeat . . * “urUr 00g * IS eae ae Gnaerrald “IST “2uojsuyor apupnyduy + ‘ayy sopoeyuay, ‘sq[IS jo sured seat UOISad O1ovIOY]Y Aareypidep aly UT sopystaq ted IMoj-AJUIMY UT sepystaq Aavypided Aree 5 : * MUL Og “TT ‘SyIVUL UMOIG [ILM sopovy OTTO “WO ‘99.109 ae Ua], ‘sjJUUISeas UdeqUAAeS UT SeTystaq ArvppIdeg ‘sT{IS Jo sared Leoni, ‘squUoWIRTY poouRrqun Jo 4jJn4 B JO s}stsuod [TIS yor "TH4NNVHO HHL HO SLSVOO HSITONA GNV HONDA AHL NO ANNO WdAITTHFHYAL AO SHIONdS AHL OL AX ‘seg ‘numao0ugg sapyyagasa ‘ured goreysod ut‘turug, “Apo jo qavd JUOI, UL Y[V}S SUOT B YALAd Spoor“) ‘gjounped [Sus @ WOL] SUISIIe soqye[d oxT][-quios AnoJ ‘s]atIses W9e4T[SIE ut sdiy Yyoours YI . . . . . . . . . * “Wut 09 may “Oyt[-quoo j Surystsuoo 113 aug ‘ , I ‘UU OSL “TT ‘syueuuses OMY UO STIL) urayyed Aue qnoyyia ‘asueio Apog E. V. ELWES. . . . . . . . . . . . . “ULUI 09 Agi T]ezpeq, ‘nqwpowu auorog 4 “UMOIG UYITAS peyjods samryeuos ‘per sopvjuey, “UMord yum = payjods ‘par Apog "3004 -XIS 07 eo1y} JO SMor gsdoasuely ootyy YIM TUTOUy) | 3 p . . . . . . . . . . . . . . O9L Any BYH ‘snsojas endow { ynoSas 9014} UO S[[T ‘SOULT ogTtpA YALA wuioyed ev ut poyseur Apog | : ‘wu gy “TT «“YSTpped sepowque} pue Kpog ‘soyouerq pesuvire Aqpeards / OTN (07098149 DIST 4 YA ‘OYTT-YsnIq s|Ik) “sed ay} Jo o[sue sorseysod oy} 4v uoyoofoad Suoy Ara @ pur 17390} BATIM} 0} BAIY} JO SMOT BSIOASUVIY OAY 0} e014} YALA TUTOu . . 6 . . . + 6 "I . < -(j) "qs1aQ) “wyootueyso2 = wm og “TT “quosed sad = “ja0ys “par Yavp sopowyueT, “OFM ‘ . 7 UOT “INgSNUUaA 12]002NT ya poyods ‘per Apog 9 ‘Tutoun jo Mod a[SuIs & YIIM UetUdes YIWEE;MOAS 0} WAysIe eyy, “4904 GAY 07 oI YATAS ‘SMOX OSIOASULI OMY YIIM TUTOU() ‘panuyuoo—" VW AITTAAANaL JO SHIOddS AHL OL AGM 76 sopystaq Asepptdey ‘SMOI UL pasunlse ‘squeuey apdumis jo Surystsuoo ‘s{[t) "yy004 9a1Y] 0} OMY JO SMOL BSIOASULI} OM YALA TUTOU Lf) ‘squouuses (A114 I9A0) snolotuuu ut sdiy YyOOUIS YT sopystaq Areypidep ‘s[[IS poypursq jo aved auo ¢ syUsUISas 1994XIS ul sdqy yyoous YyTM sopystaq Aaepidep ‘s[[IS poyouvaq jo saved (ao1yy Aeted) omy { SyUeUIses 1904 -UAAOS 0} UVEIJY UL sdry yoous YAM sopystaq Areppidep 17 NOTES ON THE LITTORAL POLYCHATA OF TORQUAY, ‘49 ap ‘sngnynonuap snuuohpody "BAI YS oY} WO papiooeat yok JON , ‘ydaso pr : . : : * usaqyy qnoqe ‘sapqstaq Areypideo yyIM =. “paqe[norjuap ‘syuoumdas jo gaquiny ‘“MOT[eA PTA pasuy AYYS Ys A19A 10 ‘ssaptno0pog J sapystaq Areyprdeg ser Gdenara SaAIGOINTIOn ST RaRIO : “uur §f “TT “etpraydeu jo sured vary, “juourSes Butrvoq *‘pasura So i ecb PMT ~o9sT1q qq} 04 yqusAas ou}. qe avodde TUIDUA, «“sseTanopoo ApoaUG ATVYSTS Uatyy *S[OSSOA endin ‘sopormuay snarolizo ‘mur gt “TT ‘erpraydoau jo saved xtg’ ‘syuourses jo ouI0g §=“yjoows | -pooyq Cee Ree eo SuLIwaq- apasiaq aaTouy qsty ayy Ul ruyoun ON ‘poorq pat snonordsuog J sapystaq Areppideg ou ‘urUL OOT 03 08 “Ty -‘erpraydeu jo sured sayy, ‘a nuio pur ‘aqnay ‘snanyuninn snuuohjog + ‘quautsas Surreaq- apstiq YUM 07 yYQueAds ye avodde rutoug = *A4A0F | Srewatte — a6 sys 07 IYST0- AquoMy Moqe sopystrq Aapypideo ya syuemBes yo raqumy P Re a a ON | pdr ° ‘ : jo oUON “PJOoUTS pdip “wnapuayno snamohpog + sured xig “quowtSas Burreeq-aystaq yuu qe avedde rutoug “44x18 sopsuq faerie 0} 3YySre-AzuomMy Jnoge sapystaq Arvppdvo YILA syuUoUTses Jo JoquINN ‘{UOWUBTY apSurs vB Jo Bur -4SISUOd []IS ova ‘sured aol} “ST[U) ‘s}ywauTses uaeqy wr sd yyoours WIA sapqstaq Aareypidep ; is : : : : : : ‘mU Og “TT “4a]OIA sopRyuay, ‘asue1o Apog IBY ‘eyus9n]6 enyoupsgoyonty Ee ‘penuyuoo— PY QITTHHHAAL AO SHIONdS HHL OL AX ELWES. Wee We 78 “1B[ [OO ° £410} JNoqe sjusuISes Jo 1aquin ‘apis Jove UO VAY s}UIUIe dio ‘waunssagupaus nuagbryd us i Aug T OF a gO Te opie g te @ JNOUIIM 229) He [ads letod 6 niea) “"quauSas pepstaq Jsiy Ur sys400}0 OMT, “quowBes TeuRe puR wINIUOysTI0d UT soAq pea "qUOTY “wnsopnoisaa DULUOLYyaUDA : ; ; " MMH OOT “I ‘sT[eys Jo syueusery , : each pur ‘souojs [[euts ‘puvs Jo eqny, ‘s}uoUIe[Yy [IS Jo pus ayy avau sak , o : queluses [vue : ‘Bury ‘njisows wyvunjog Ose Tino qY S19 10 xIs pure sopasiaq eqernyyeds sees "34840030 OA\J, enna jee ares I9y1OYs pue ‘jeatds IB ‘yes vyumog 4 “08 sid lee) Py aes ie racine ye eel pesuram Morieu | ‘syueUTeTYy e SULUIIOF : ; d \ -sqsko0j0 0 N — raat soko ON Ajeueu ‘spur It ey 4ou “IeT[Oo Sere cae a OM} JO XvIOY] VY} jo pua sjueMR[y == = =—s-_-& «WHF “TOTO ‘sewsofruas nyawunjog ‘e ee 1 nS d Reh saat ae JO sopqstaq [es1oq, ay} avau niga) | 04 ; ieee ahone SaaS aes saAo ON poonpoid ‘TUM OOS “TT “WLU OF ( Nepal UINIWOISLLIg jo xv10y} oyy Jo | “Avg “purzuoand mans | SOPISTIq [BS1O(, als SqUSWILTY [Ly “pnur jo eqny, | "qUOT ‘s2usoounjoa nsrdsr “WIT OST “TT “SpuTy 0M4 Jo xvI0q4 ‘aseq [aids | W* ae 12 oY, JO sop [es10q ‘jeube UMOIO TeTYoURIG O44 Jo sjavd omy OY, ®@ WLOI} “ALA ‘vzunyody svydvsbowd, "MIU 096 “YI “PUly eo Jo xeI0y} SUISTIP | HA ETL Ss ayy JO sapystaq [esiog ‘[enboun wos yeIourtq oy} Jo sjavd omy oY) syuoUIETY [ITD ‘TuIoUN pue seTystaq AreTTIdeo pesuta oyduus Ayoureu ‘SpULY JUOIO]JIp OM4 JO XVIOY} OY JO SOTASTIG [eIQUOA “WV “THNNVHO FHL AO SLSVOO HSITONGH GNV HONGYHA AHL NO GQNONOH WdAITTHAVS AHL HO SHIOddS ANV VHHNHO GHL OL ATM 79 NOTES ON THE LITTORAL POLYCHATA OF TORQUAY. ‘sdy 1104} -ydaso p-3g ap ‘sswapunuacy mjoowh py 4 . SN SO eau ie Det Te TY a cedars Ried aaa) 1) fsd1q"a1ey} Ivou pasie[Ue syueure[y [WY ‘“wmturoystied uo yueserd sokq a a curynqrpunfua poor ; ; ULUL OZS “TT “WOM OSuey ‘shOUTIeTAS 9qnq “IeT[aMp-pnut pias ay ny see Te ee eee 1 PPOKMON ) ‘spua ayy Ivau posiepuo you syuoMLTY [UD ‘wurmoystzed uo saa on waqoaTrcod symoueTy ITH) : : spud 93R[NT]} ‘ur Gy ry £4107 07 vAy-Aqatyy ‘syueurses | -eds qroys YIM s1eq0 ed He ao Saini asibueDy 2 oN ‘sJUSUISes IOIIA}Ue UT SeAd OAT, ue ‘spue Suts1edey A LR RS tla lca L ‘yySua, penbe jo 4roys seyoueiq Aepuoosg UOT YIM soTystiq ae ‘pls Yowa WO aATAy OF 4YSte syuoUETY [UH | ‘Ajoureu ‘spury omy jo qu XvIOY} JO SoTIStAQ [SLOT Ay ea . . . . wWUuI (a) ay L ‘a014y ‘MoTOpYe UL SyUEuIseS [PAI] saint “ayy “nypoqns wioLQngT Sutavaq-a[Isuq s]WsUISes oures ot) auvIq UIdUL sarastiq [eure pure somejue ul sokq Sutyovat ouy v ns "IeT[Oo [etmMoyst1ed ON ‘apts YSU] Aq poyooumos La yore wo eery} syueUIEyY TI) yenbeun spua Surodey Suoy jou ‘UU 9 “TT “WAAVS 07 XIS “UU jo S[[to Jo eye sopstaq £aeyprdeo syuouTely [[ty) -Opqe Ul Ss}UeUIseas SuLIvaq soyoursg ‘Kaueu “pury auo Jo palo Ypunusy m9 -o[JsUIg ‘syuouISes [vue puL Arepuooag xe104} JO SoTISIA [esIOG] ce 2 Joleque ut sok “quosord ‘maulOpge ; I€T[OO [BIUIOYsSLINg = “opis Youa Ur TULDU PE) UO INOF LO 9dIY} SJUSMR[Y [[UD ‘uM 9 “YT ‘syUeUTKTY ey} 07 soyouerq Arep -00e8 BUOT ON’ ‘APIs YOR UO 9AY syuoMETY IID “A[eIs Suoy v YALA Woop" UT sjoyojoIH ‘guinog ‘snuumnysa snyounsqojdy FT : . . . . : . 2 ‘mu og “ry = ‘soXa Jo ared youa avau wmody * juroun xe10qy { vain ‘y[eIS BUOT B YIIA SJoyojoIO Io TuroUN roYyy1e ‘AToUeU ‘puUTY eUO Jo xBIOY} OY} Jo SoTISTAq [BIJUOA “g ‘panuyjuoo—VCITIAGa VS FHHL AO SHIONdS CNV VYUUNAD AHL OL AU ELWES. KE. V. 80 ‘eale YSIWIG oy} Woy pepiooas yok JON , ‘pua ye aauoo ATSu014s 7 Tana ‘taupnoiedo ut yonod pooag ‘SUR au TT ‘sngnjnunab suquondy ° : : tunpnorado dy} puNol WLI YJOouIS B ‘seBpIr payet1es ON ‘pdio ‘sump srqiondyy : sespa poyedies YALA wunpnoredo jo doy, xaAuoo A]SU0.14s ‘urpneg ‘sypa.og siquondy ° OX Ipned ‘sypa1og sqao.udy uinpnoriedo uo sdUe0se10x9 10 4490} ON § BULIvAq-aTISTIq BAIT, J, jou wmnored¢ ‘tanpnotedo ut ‘TLUSOT YT 4 see : . XVIOY} OY} Ul syUsMIses SULIveq-aTISsTIq Ino U3 oy Urs} HeOq"8T9S4Q INOT | qonod poor ON pue Aroyney ‘wpunjnyy sigwomdyy ‘ydasoe 4g ap ‘snypsnd srquoudyy {° pea guowStd onedoy )- qUOUISeas SULIBA-d[ISIIq PITY eq} Ut “DIY ‘Muayoagsuabng sigsoudsy. * ¥2]OIA quowsrd onedoyy sozystaq pedeys-aTyoig 9 ‘yjoours wintnosedg * quotdses' SULIvAq-aTISIIq PATYY a} UI seTstaq podeys-a[YoIs ON ‘[olIeq we Surpquresel umpnosedyg ‘munno1ado ‘quoyy ‘snyohns109 srqsodgy 2 ur yonod pooig ‘ydasor 19 ap ‘snunor { UdTIOpYe oY} UI SpUIUISes Sur OUD = "QUOT ‘sns.Lo.1jsrurs srgsourdy | -Leeq- “OTASTLq AqueM} 0} U9E4XIG ‘sqeId pu sIeysqo, Uo onbedo aquy, ‘qUOT : * USTIOPe IY[} UI SJUeWISas SuvIq-a[JstIq We94XTs ‘snpony="T Swnypuds siqsomdy oy ua, ‘vuyagn nunniwoy Uo UoWMOD “queredsuety-TULeS aqng, ) wmyno1edo ut Td ‘sya nutoo siqwoudy wnynoiedo uo quaserd saotteosa10xe 10 ae “XBIOY} IY} UT SUIUT eee -5a fl pooiq ON ‘[eads oquy, ‘sloyjo ay} UeYy JoyUNTG pure ‘1epeRoIq ‘1aSWO] 4004 4svT oY ‘YY00} AMUN YIM TUIOUG "THNNVHO WHL 40 SLSVOO “OUT [esiop weIpsuT SO eae uo wmypno1edg ‘aul] [es1op UvIpeUt ayy JO JYSII ayy uo wnno1edg HSTIONG GNV HONGYHA AHL JO WdAITNdYHS AHL AO SHIONdS GNV VUANAD WHHL OL AWM 81 NOTES ON THE LITTORAL POLYCHATA OF TORQUAY, te * “WU OZ ‘TT “Opts Ova UO Wd9d}JUMIAVS 0} U2 Joqe s[[M) “Wards ), : c : ee as ‘uunty ‘norbamsow saprosphyy \ xe1oyy JO tu1oUN UI Y490} Jo aquINN ‘erjW89 ay} WOIF BuIstav soutds ani aIST TOG aea. foe G IY} UO suIOY} TATA ‘soutds Jo epoa1o v YALA podeys-fouuny wnNo1edg me ae eta aoe “ei aa — roy J ° “WU OG 07 OZ “TT “OAY XBIOYZ Jo TuTOUN UI Ye} JoTOquNN ‘ops |, . OP LHe EECA AEE { yous uo AqATY] Jnoqe s[[uH ‘eyeuero wuISivur ‘padeys-jouuny wmnypuosed¢g idea SG eA SEA BN USE ee * soutds dary] 07 9U0 JNOYITM IO YAM ‘Teortos 10 doq ot4 4e Wey ‘tuys ] : asnosd v aYIT YIvouropun TT Suajanbi.19 sous0jnwmog + oy4 Uo suorqoaload omy YAM tNTNoIedGQ “aodytI0 oY A9AO YAO} davys % -INO paAoT[OY Y}00 yseT eyy ut Sutqoeford aspra 01499 oy] ‘sospit sary} YIM AT[ensn yUoLay pe oquy, | *YJo0} BUTT 10 FYSIa YIM tUDUQ S lord Ba: a Ca se : ° ; . * Surumaqaoqur Agxny “wajshq¢ mupnuyny urut 9 “rT “JNO paaorfoy, puv pedevys-quyo s][L8 Jo spug “wmynosedo o x] Gapuays IEA soqny, ‘sxayI0 J F F : S ; ; : he oe a , : g poy} uvyy desiey puv “oqyuntq = 5 EON Eel aL ST { O spuy ‘satpuriq AIvpuodes YATAL siren > 5 a erie aston on CASA SUE ET 012) g alee es rome mo cael? MG IEG SL woa}INoy ynoqe yA Turug © ; : , [f.0 : a * ‘wut GT 09g “TT *xetoyy UT yuasord syTeys posura : : : : * yeards fs Sse niece erence ESE car sapistiq, pedeys-epyors eulog “queredsuvsy “aepnqops wmznoiedg|you oqny, ‘ourds qnoys v Aq z poqeurutie, ‘9009 [pews A19A . : . . : : , * ‘MU CF 0} OZ ‘STITT gnoyyIar 6 'T puv snosoumu AI9A YT yorq . ‘¢ UNE aT nnn } ‘xe10Y} 9 Url yuasord sapstaq podvys-e[yos ON ‘wnynotedo on) 4e qno poaorpoy ATdoap mmoUug paniyquoo— WOTTNddas HAL FO SHIONdS GNV VYUNHD AHL OL AUTH NEW SERIES.—VOL, IX. Some Notes on the Genus Cumanotus. By Nils Odhner. Fil. Lie., Stockholm. In 1908 Sir Charles Eliot published in the Journ. Mar, Biol. Assoc, Vol. VIII, No. 3, a paper “ On the Genus Cumanotus”; in that paper he showed that Coryphella beawmonti, discovered and named by him in 1906,* was to be referred to the above genus, which had been first established by myself in 1907 from a study of the Norwegian C’. laticeps, described at the same time as a new species.t Sir Charles Eliot also called attention to the striking resemblance of the two forms, and remarked that their identity was not improb- able, though he assumed that there might be some differences in the denticulation of the jaws and the lateral teeth of the radula. Through the kindness of Sir Charles Eliot and of Mr. De Morgan, Acting-Director of the Plymouth Laboratory, I have procured two specimens of C. beaumonti for comparison with the Norwegian C. laticeps, with a view to determine the distinguishing characters of the two forms. In exterior appearance they are quite alike, and I have found no difference of a specific value in their habitus. The proportions of the body are nearly the same, as is evident from the following measure- ments (in mm.) :— CO. beaumonti. CC. laticeps. Length of body . . 14 13 Breadth a itsuy. 2 : : : eee Dea Sea eads : 3 : Se 4 Length ,, rhinophores : . 43 4:3 * »» a few papillae in the 5th row 3°7 4 The height of the body was somewhat greater in C. beawmonti than in C. laticeps, which probably depends on their varying states of preservation. * © Notes on Some British Nudibranchs,” 1.c., Vol. VII, No. 3, 1906. + ‘‘Northern and Arctic Invertebrates, III, Opisthobranchia,” A, Sv. Vet.-Akad. Handl,, Bd, 41, No, 4, 1907, N. ODHNER: THE GENUS CUMANOTUS. 83 In both forms the soft parts fully agree in shape. On the head there are situated two small conical tentacles of the same size and position in both, connected by a low cutaneous fold. The rhinophores are close to each other, and are united at the base. The foot is extended, forming two pointed angles at the frontal sides, and is expanded laterally and posteriorly to a cutaneous border. The arrangement of the dorsal papillae is also of the same character in both. They are set in about 12 transverse rows, the 3 foremost ones being placed in front of the rhinophores. The rows are in two groups, a pre-anal and a post-anal one, the first embracing 6 rows; the anus is situated dorso-laterally, immediately in front of the 7th row. As to the number of papillae, this has been easy to determine in CO. beaumonti, for all the papillae there were intact; in C. laticeps, on the other hand, they had fallen off to a great extent, and the state- ments here given are therefore deduced from the markings. One specimen of each form was examined. The number of papillae was as follows :— CO. beaumonti. C. laticeps. In the Ist row 2 3 ee ean i, 4 6 ee Ord ay, it 5 eu sili 9 9 ae 7G ieee es 8 6 Sim SObRY TL .. 6 6 Pata Jieaee 4 4 the 4 4 The unimportant difference which was present in the two specimens examined may be quite individual, and any attempt at deducing specific characters is therefore excluded. Anatomically the papillae are of the same structure, for in both forms they are furnished, at the tips, with a saccus enidophorus, which is connected with the liver process by means of a narrow, winding canal. Of all the characters distinctive of the genus Cumanotus, the shape of the female copulatory organ is the most peculiar. To the sides of the bursa copulatrix there are attached two circular pads with a papillated margin, and these papillae are of the same number, 12, in both forms. Such difference in dimensions as was observable in homologous parts is to be explained by their different stages of maturity. Thus exteriorly the soft parts show an entire conformity, and from them consequently no specific characters are obtainable. There only 84 N. ODHNER: THE GENUS CUMANOTUS. remains the inner anatomy to be dealt with; but here I considered it unnecessary to compare the whole organization of the two forms in detail, and I have accordingly only examined the organs that are primarily of specific value, viz. the radula and the mandibulae. The rows of the radula in C. beawmonti vary in number from 16 to 24, according to Eliot; in C. laticeps I have found about 17. The teeth are furnished in both forms with long, slightly curved cusps. The form and the denticulation of the median tooth do not present any differences. The lateral teeth are denticulated only on the inner sides. In the latest formed part of the radula, I have found the number of denticles of the laterals circa 25 in C. beawmonti and 18 in C. laticeps ; in the older part above 25 and about 22 respectively. This slight difference is of no consequence, especially as the form and curvation of the lateral teeth are the same in the two specimens examined. There remains only one more character to consider, the structure of the mandibulae, but here too I have found entire agreement. Their form and colour correspond, as do those of the whole bulbus pharyngeus too. The mandibulae are lengthened, roundly quadrangular, and denti- culated in the anterior margins. In the denticulation there exists but a slight difference, the denticles seeming to be placed at somewhat greater intervals in C. laticeps than in the other. As to the shape of the denticles, I have found them in both forms to be somewhat irregular, uni-, bi- or tricuspidate, the more complicated ones being situated in the upper or anterior part of the jaw margin, a part which is most worn. The denticles are arranged in one row only at the margins. Two specimens of each were examined. In these mandibular characters also the forms agree wholly with one another. It has consequently not been possible to find out any specific dis- tinguishing points between the two forms; in all the characters they are alike. I therefore consider their identity to be proved. Nor are there any good reasons for their severance as varieties; it is hardly probable either that any would be obtainable from the characters of the living animal, though the colouring might doubtless be subject to some variation, as is usual with the Nudibranchs. As a result of the above comparison, I consider the genus Cumano- tus to consist of one species only, viz. C. beawmonti (Eliot, 1906), and regard my own species, C. laticeps, Odhner, 1907, as a synonym. C. beaumonti consequently has a wide distribution, being obtained in England as well as in Northern Norway. Further investigations will certainly show its occurrence also in the intermediate districts. [ 85 ] Kodioides borleyi, n.sp. By Chas. L. Walton. In his Report on the Actiniaria of the Norwegian North-Atlantic Expedition, D. C. Danielssen erected the genus Kodioides to receive the remarkable form which he named K. pedunculata, a single speci- men of which was dredged at Station 35, July 5th, 1876, in 1050 fathoms, Lat. 63° 17’ N., Long. 1° 27’ W., the bottom being Bilocu- lina Clay. The detailed description will be found in the section “ Actinida,” Vol. V, pp. 77-82, Pl. VI, Figs. 3-4; XXII, 8-11; and XXIII, 1-4. The genus is there characterized as follows: “The body encrusted, piriform, with a long bare stem terminating in a pedal disc. Two series, containing a few retractile tentacles. 12 pairs of septa, of which 6 pairs perfect. Suckers on the encrusted portion of the body. No gullet-groove. Mesodermal, annular muscles. Acontia.” During a visit to Lowestoft last summer, Mr. J. O. Borley handed to me for examination 2 specimens of an Actinian obtained some time previously by the ss. Huszley, Voyage XXX, Station 23, Lat. 53° 46’ N., Long. 4° 52’ E., N.N.W. of Terskelling. Depth, 20 fathoms. Bottom, mud. Conical dredge. Two specimens. A very short ex- amination convinced me that I had before me specimens referable to the genus Kodioides. The species, however, is evidently distinct, as might be expected, KX. pedunculata being obtained in the cold area, from 1050 fathoms, between the Faroé Islands and Norway; and these from 20 fathoms, and comparatively close to the Dutch coast. It is unfortunate that no description was made while the animals were living, but Sea Anemones are difficult to deal with; indeed, it is frequently impossible to do so under bad-weather conditions and during the pressure of fishery work. Danielssen mentions the diffi- culties he experienced with K. pedunculata. Further, when I examined the specimens they were by no means in the best condition, being somewhat decayed. Specimen («).—Measurements, etc., were as follows: Animal strongly contracted :— 56 Gy Le WALTON. Total length, 48 mm. Length of body, 21 mm.; breadth, 16 mm. Length of stalk, 25 mm.: breadth, just below body, 5 mm.; lower portion, 5 mm. Pedal dise irregular in outline, breadth, 15 mm. (on the average); thickness, 5 mm. On the body were remains of a slight coating of mucus and sand grains, and a thick coating of mud adhered to the pedal dise. The ectoderm was much decayed, but remains of what appear to have been suckers could be made out. The body was considerably wrinkled. Stem bare and smooth. The upper surface of the pedal disc and base of stem showed numerous fine lines. There was a considerable tumid excrescence on the lower portion of the body, much decayed, and probably due to injury received in the dredge. Neither the oral disc nor tentacles were visible. Specimen (b).—Total length, 32 mm. Body wrinkled and thickly coated with sand at the summit. Pedal dise irregular in outline, much smaller than in (@); under surface bare and much ridged and folded. This individual was only partially contracted, and showed the tentacles and portions of the oral dise. Oral dise strongly ridged, but owing to decay the number and details could not be made out ; mouth also not visible for the same reason. Tentacles (partially contracted) short, stout, and obtuse, in 3 or + series, about 90 in number; but they were difficult to enumerate, and in several places a number had been injured or destroyed. Colowrs.—Mr. Borley informed me that when alive the body was yellowish white, longitudinally striped with dull red. Tentacles, dull red(?). Nothing remained when examined but a dull uniform pinkish shade. Horizontal and vertical section with a razor disclosed a state of decay, amidst which little could be recognized; I can only say that no siphonoglyph could be found in either (@) or (2), and that the mesenteries in (a) were about 24 pairs fully developed, and a similar number incomplete. The general appearance of both specimens is very similar to Danielssen’s figures and description of A. pedunculata, as also are the measurements; the features that cause me to consider these examples as a separate species being the marked difference in the number of mesenteries and tentacles. | here repeat the generic characteristics as given by Danielssen, so altered as to include the present species :— The body encrusted, piriform, with a long bare stem, terminating in a pedal disc. Tentacles few or many, in two or more series, retractile. 12 or more pairs of mesenteries, half ‘of their number perfect. Suckers on the encrusted portion of the body. No siphonoglyph. Mesodermal, annular muscles. Acontia. KODIOIDES BORLEYI, N.SP. 87 Specific characters.—K. pedunculata : mesenteries, 12 pairs, tentacles 24. K. borleyi: mesenteries and tentacles numerous. It is to be hoped that further specimens may be obtained before long, which would enable a thorough anatomical examination to be made and the true affinities of this remarkable genus determined, as the condition of the specimens here described was not sufficiently good to permit of any observations as to acontia and many other points of interest. In view of the peculiar form of these anemones, it may be of interest to quote Danielssen’s remarks concerning A. pedunculata: “The weather was very stormy at the time, and the vessel had a constant heaving and rolling movement, which in a great degree obstructed the investigations. I was, however, fortunate enough to obtain the animal drawn in the live state, and to jot down some observations in regard to its exterior; but as it constantly kept itself pretty much shrunk together, although I had had it for several days in the glass vessel for observation, I could determine nothing in respect of the tentacles, only so much did I observe, viz. that the stem sometimes kept itself quite erect and at other times became bent, whilst the body expanded and contracted—movements which were participated in by the stem in such manner that when the body contracted the stem became attenuated, and when the body expanded the stem became tumified.” [hse T° Marine Biological Association of the Cuited Aingdow. Report of the Council, 1909-10. The Council and Officers. Four ordinary meetings and one special meeting of the Council have been held during the year at which the average attendance has been thirteen. A Committee of the Council visited and inspected the Plymouth Laboratory. The Council desire to express their thanks to the Councils of the Royal Society and of the Linnean Society, in whose rooms their meet- ings have been held. The work in connection with the International Fishery Investiga- tions, which the Council has been carrying out during the last seven years for His Majesty’s Government, has now been taken over by the Board of Agriculture and Fisheries. The Plymouth Laboratory. The Laboratory, including the pumps and engines used for circu- lating sea-water through the tanks, has been maintained in an efticient state. An air-circulation from a pump worked by the gas engine has been rearranged in such a way that an abundant supply of pure air is now available for use in small aquaria and experimental tanks. The Boats. The steam trawler Huxley, which has been used for work in connec- tion with the International Investigations, has been sold. The Oithona was again fitted out for summer work at the Plymouth Laboratory, Captain J. Tucker, who has been in charge of the Huxley, being in command. The winter collecting has been done as usual with the sailing boat Anton Dohrn. The Staff. Messrs. Borley, Todd, Wallace, Hefford, Atkinson, and Wollaston and Miss Lee have accepted service under the Board of Agriculture and Fisheries in connection with the International Investigations. REPORT OF THE COUNCIL. 89 Mr. E. W. Nelson is accompanying Captain Scott to the Antarctic as biologist. Mr. L. R. Crawshay has resigned the post of Assistant-Director owing to ill-health, and Mr. A. J. Mason-Jones has accepted an appoint- ment as lecturer in biology at the Plymouth Technical Schools. An arrangement has been made by which the services of Mr. D. J. Matthews will be partly retained by the Association. The Council are glad to say that Mr. Matthews, Mr. Crawshay, and Mr, Mason-Jones continue to work at the Laboratory. The Director, Dr. E. J. Allen, has delivered a course of twenty-four lectures on Marine Biology and Fishery Investigations at the Imperial College of Science and Technology, South Kensington. During his absence Mr. W. De Morgan acted as Deputy-Director. Occupation of Tables. The following Naturalists have occupied tables at the Plymouth Laboratory during the year :— Prof. SvANTE ARRHENNIUS, Stockholm (Experimental Embryology). W. DE Moreay, Plymouth (Hybridization of Echinus). G. H. Drew, B.a., Plymouth (Experimental Pathology). J. S. DUNKERLY, B.sc., London (Protozoa). Prof. F. W. GAMBLE, F.R.S., Birmingham (Colour Physiology), E. 8. GoopricH, F.R.S., Oxford (Fishes). G. H. Grosvenor, M.A., Oxford. Miss Harrison, Oxford (Experimental Embryology). M. D. Hitt, M.a., Eton (Alcyonium). C. Kruur1an, Freiburg (Laminaria). D. G. Linuiz, Antarctic Expedition. Prof. JacquEs Lozp, California (Experimental Embryology). W. Nicott, M.D. (The Entozoa of Marine Fishes). Miss Pootsr, Oxford (Development of Tectibranchiata). CaRR SAUNDERS, M.A., Oxford (Development of Tectibranchiata). C, SHEARER, M.A., Cambridge (Histriobdella and Dinophilus), GEOFFREY SMITH, M.A., Oxford (Bacteriology of Crabs). E. R. Server, Oxford (General Zoology). R, WHITEHOUSE, M.SC., Birmingham (Fishes). Miss GERARDA WisNHOFF, Utrecht (Nemertines). W. Wooptanp, D.sc., London (Gobius). Miss YONKER, Utrecht (General Zoology). In addition to the above, nineteen students attended the Labora- tory during the Easter vacation, when Mr. G. H. Grosvenor conducted the usual course of instruction in Marine Biology. The Library. The thanks of the Association are due for the following books and current numbers of periodicals presented to the Library during the past year :— 90 REPORT OF THE COUNCIL. Académie Imp. des Sciences de St. Pétersbourg. Bulletin. American Museum of Natural History. Bulletin. —— Memoirs. American Microscopical Society. Transactions. American Philosophical Society. Proceedings. Armstrong College. Calendar. Australian Museum. Memoirs. Records. Report. Bergens Museum. Ca). about 5°0 grammes. Sea-water . : : ; ; ; , 1000-0) ¢.c. Calcium malate is only slightly soluble in water (about .\,th per cent), and so can be added in excess. The reduction of the nitrate to a nitrite was tested for by the addition of 5 cc. of 10 per cent Sulphuric acid and 2 cc. of a 1 per cent solution of Metaphenylene diamine hydrochloride to 25 c.c. of the culture. The production of a brown coloration (due to the formation of Bismark brown) is an indication of the presence of a nitrite, and is an extremely delicate reaction. The formation of Ammonia was tested for by the addition of 5 cc. of a 10 per cent solution of Potassium hydrate, and 5 c.c. of Nessler’s re- agent: the white precipitate formed on the addition of the Potassium 146 G. HAROLD DREW. hydrate does not appreciably interfere with the test, though it renders it somewhat less delicate. The presence of nitrates or nitrites remaining in the culture fluid after the bacterial action was tested by the extremely delicate Brucine and Diphenylamine reactions. THE INVESTIGATION OF SAMPLES. OF WATER TAKEN OFF. PORT ROYAL, JAMAICA: In Jamaica, a measurement of the rate of denitrification in the modified Gran’s medium inoculated with samples of sea-water was made, but the isolation of the bacteria on solid media was not attempted, owing to lack of apparatus. Samples of sea-water were collected in sterilized stoppered bottles from the surface, and from depths of three and six fathoms, in positions about five miles from shore, where, from a consideration of the wind and tide, the water was probably under truly oceanic conditions, and unaffected by the neighbouring land. The cultures were kept in a moderately dim light, and the room temperature ranged between 25° and 31:°5° C. The average tempera- ture during the growth of each culture was noted. In a typical culture made from surface water, and for which the average temperature was 29° C., the first indication of the formation of a nitrite,as given by the Metaphenylene diamine reaction, appeared after twenty-seven hours: after thirty-eight hours the brown colour produced in this reaction was very intense, the culture became cloudy, and on testing with Nessler’s reagent, sight Ammonia formation was apparent. After forty-eight hours the culture became very cloudy and a scum of bacterial growth developed: the nitrite and Ammonia re- actions remained unaltered. After sixty-three hours the nitrite reaction was somewhat less marked, the Ammonia reaction was un- altered, and bubbles of gas began to appear. After seventy-two hours many bubbles of gas were being produced and the nitrite and Ammonia reactions were very slight. After eighty-six hours the bubbling had ceased, and no nitrite or Ammonia was present in the cultures. Test- ing the culture for nitrates by the Diphenylamine and Brucine methods, showed that no nitrates or nitrites were left in the solution. In the absence of a gas analysis apparatus the nature of the gas evolved could not be exactly determined, but considering that it was non-inflammable, did not turn lime-water milky, and that the nitrate originally present had been destroyed, it seems strongly probable that this gas was pure Nitrogen. Thus at a temperature of 29° C., 0°5 grammes of Potassium nitrate THE ACTION OF SOME DENITRIFYING BACTERIA. 147 were decomposed in eighty-six hours, and it was found that if a further 0'5 grammes of Potassium nitrate was added to the culture, it was in turn rapidly decomposed: this could be repeated indefinitely until the other constituents of the medium were used up, or the concentration of waste products became too high. The rate of denitrification varied considerably with the temperature, and in cultures kept at a temperature between 10° and 12°C., no growth or denitrification occurred. Denitrification was more rapid in cultures from water taken from a depth of three or six fathoms than from the surface. It was also especially rapid with samples taken from the thick, muddy waters of a mangrove swamp, where organic matter was plentiful. The bacteria present in the cultures were very minute, actively motile bacilli with rounded ends. THE INVESTIGATION OF SAMPLES OF WATER TAKEN ROUND THE.DRY TORTUGAS AND IN THE STRAITS OF FLORIDA. Cultures made in the modified Gran’s medium from various positions round the Tortugas gave results in close accord with those found in Jamaica, and the average rate of denitrification was the same ; hence it appears probable that much the same bacterial conditions obtain in both places. Cultures were made on various solid media, and pure cultures of the bacteria were isolated. Petri dishes with porous earthenware covers were used and were found of great advantage, as by this means the formation of drops of water on the covers is pre- vented, and the water of condensation evaporates as soon as formed: in addition a free air supply is ensured. If evaporation is progressing too rapidly, the whole Petri dish can be covered with a bell-jar lined with damp filter-paper after the first day. By plating samples of surface water from various positions as far as possible removed from the influence of the land, an average of four- teen colonies per 1 c.c. sea-water was obtained. ‘These colonies ap- peared to be of two kinds, one much more plentiful than the other. Subcultures made from these colonies in Gran’s medium showed that the bacteria forming the most common type of colony produced an active denitrification, while the others grew very slowly in this medium, and produced no denitrification. The chief characteristics of the denitrifying form are as follows :— On the Potassium malate, or Peptone Agar media, colonies are visible as minute white specks after six to eight hours, when the 148 G. HAROLD DREW. room temperature averages 29°5° C. After about eighteen hours the colonies are well developed, they are white in colour, circular, but with finely irregular outline, and have a granular appearance. Superficial colonies are much elevated at first, but as growth pro- ceeds, spread rapidly over the surface of the Agar. The deep colonies remain small, circular, and discrete. Growth is somewhat more rapid on the Peptone Agar than on the Potassium malate Agar, and the older colonies develop a brownish tinge in the centre when growing on the former medium. On Gelatin Peptone (5 per cent Peptone, kept at between 20° and 25° C. to ensure the medium remaining solid) growth was very slow; in stab cultures growth proceeded slowly from the surface downwards, forming a funnel-shaped depression of liquefied gelatin. Acid formation occurs in Glucose, Mannite, and Cane- sugar, but not in Lactose media. Growth is totally inhibited at a temperature of 10° C., but takes place slowly at 15° C. Growth is much retarded by exposure to bright sunlight, but the bacteria are not killed by a ten hours’ exposure. The bacteria are facultative anaérobes, but growth under anaérobic conditions is very slow. In Gran’s medium growth and denitrification are rapid, but no growth occurs if the Potassium nitrate be omitted, or if the Calcium malate be replaced by Calcium carbonate. Growth in a pure solution of Peptone in sea-water is very slight, but becomes abundant if Potassium nitrate be added, when denitrification quickly ensues. In its cultural reactions this bacterium is very similar to certain denitrifying forms found in the English Channel. The chief points of difference consist in its much greater denitrifying power, and the higher temperature necessary for its growth, by which it would be prevented from spreading into colder waters. The characteristics of the scarcer, non-denitrifying, form of bacterium found on the Peptone Agar plates made from surface samples are as follows :— - Growth on the Potassium malate Agar medium is very slow and indefinite. On Peptone Agar growth is somewhat slower than in the case of the denitrifying form. On the surface, circular cream- coloured colonies are formed having a brownish centre, the edges are smooth and regular, and the colony remains discrete and does not tend to spread over the surface. The deep colonies are THE ACTION OF SOME DENITRIFYING BACTERIA. 149 smaller and usually ovoid in shape, and of a somewhat darker colour than those on the surface. Growth does not occur on gelatin media, Acid formation takes place in Glucose, but not in Cane-sugar, Lactose, or Mannite media. Growth is much retarded by exposure to a strong light, and cultures on sloped Peptone Agar are completely killed by four hours’ exposure to bright sunlight. The bacterium is a strict aérobe. Free growth takes place in Gran’s medium, but develops much slower than in the case of the denitrifying form: no growth occurs if the Potassium nitrate be omitted entirely, but takes place freely if a mere trace in excess of that normally present in the sea-water be added, though no denitrification results. Attempts were made to dis- cover whether this bacterium had any nitrifying or denitrifying action in various culture media, but uniformly negative results were obtained. Nitrites were neither oxidized to nitrates, nor reduced to Ammonia or free Nitrogen, and Ammonium salts were unaffected. No growth was obtained in any culture medium that did not contain at least a trace of nitrates, so it was not practic- able to ascertain whether the bacterium had a nitrifying action without the necessary facilities for quantitative work. On one occasion samples were obtained from various depths up to 90 fathoms at a point in the Gulf Stream, 25 miles south of the Dry Tortugas. The samples were plated in the Peptone Agar medium, and counted with the following average results :— | | | | Depth | Denitrifying Non-denitrifying | Number of colonies developing fathoms, forms. forms. from 1 c.c. of sample. 0 9 2 11 10 25 4 29 40 2 2 4 60 | 5) 3 8 90 5 6 11 If any deductions can be made from one series of observations, it would seem probable that the non-denitrifying bacteria are a deep- 150 .G. HAROLD DREW. water form, and this would be upheld by the ease with which they are killed by exposure to sunlight. On the other hand, the denitrifying bacteria would appear to be a surface form, reaching their maximum a little below the surface. This would be substantiated by the results obtained in Jamaica, by which it was shown that denitrification occurred much more rapidly in cultures made from samples collected at depths of 5 and 6 fathoms, and might possibly be explained by the fact that the growth of the bacteria is inhibited by strong light. INVESTIGATION OF SAMPLES FROM A POINT 70 MILES WEST OF USHANT. The samples were plated in the Peptone Agar medium after having been kept on ice for twenty-four hours, for the reason previously explained. The colonies were well developed after forty-eight hours, and appeared to be all of one kind. A count gave the following average results :— Number of colonies developing Depth in fathoms. from 1 c.c, of sample. 10 ibs oes ae seh es 56 9 20 6 BO ea aoe ie sine ae mes 6 70 2ee ae ane aoe de ae 30 80 20 The increase in the number of colonies at 70 and 80 fathoms is somewhat remarkable, but no conclusions in this respect can be drawn from one series of observations, The appearance and cultural characteristics of the bacterium obtained from these samples were identical with those of the denitri- fying form investigated at the Tortugas, with the exception that acid formation did not take place in media containing Cane-sugar. 1000 cc. of the modified Gran’s medium inoculated on board with 10 c.c. of a surface sample immediately after collection, and kept at an average temperature of 20° C., showed the first trace of nitrite forma- tion after 70 hours. After 84 hours a very strong nitrite reaction was obtained, and a slight Ammonia reaction was given with Nessler’s reagent. The process of denitrification, even after the lapse of weeks, did not extend beyond this, and no bubbles of gas were formed. Other experiments made with subcultures from Agar and Gelatin media gave similar results, so that it appears that this bacterium cannot entirely THE ACTION OF SOME DENITRIFYING BACTERIA. 151 break down nitrates at a temperature of 20’ C. The optimum tempera- ture for denitrification produced by this bacterium appears to be about 20° C., as the process was less rapid at average temperatures of 17° C. and 25° C. At a temperature of 32° C. rapid growth took place, but no denitrification resulted. It should be noted that these temperature observations were only made with subcultures from colonies on Peptone Agar and Peptone Gelatin media, and there is reason to believe that the power of denitri- fication becomes diminished after cultivation on such media. Further and more accurate temperature experiments are required, in which the culture medium is directly inoculated with freshly collected samples of water. INVESTIGATION OF SAMPLES OF WATER FROM THE MARQUESAS KEYS, AND THE EXPERIMENTAL PRE- CIPITATION OF CALCIUM CARBONATE BY BACTERIAL AGENCY. The Marquesas Keys constitute a coral atoll which forms part of the long chain of Keys separating the Gulf of Mexico from the Straits of Florida. Within the atoll the water is very shallow, and the bottom consists of a fine chalky mud many feet deep. Samples of the water from within the atoll were sent to me at Plymouth by post, and examined fourteen days after collection. On plating on Peptone Agar, 800 colonies per 1 c.c. of the sample were obtained. These colonies appeared to be all of one species, and in appearance and all cultural characteristics were identical with the denitrifying form previously described as occurring around the Tortugas. A suspension of these bacteria from a culture on Peptone Agar was made in sterile sea-water, and a similar suspension, containing roughly the same number of bacteria, was made from a third subculture on Peptone Agar of the bacteria obtained from the station seventy mules west of Ushant. 1 ce. of each of these suspensions was then added to 1000 cc. of the modified Gran’s medium; some of these cultures were kept at an average temperature of 20°C. and others at 32°C., with the following results :— At 20° C, cultures from Marquesas showed trace of nitrite after 45 hours. 7 5 » gave strong nitrite reaction ,, 53 ,, 70 miles W. Ushant showed trace of 140 ” ” 9 ” nitrite 3 : : : 39 70 miles west of Ushant rer 162 9 = ” strong nitrite reaction . A 152 __G. HAROLD DREW. : In both cases a slight amount of Ammonia was recognizable by Nessler’s reagent when the nitrite reaction was strong, but decom- position of the nitrite did not proceed further even after fourteen days. At 32° C. cultures from the Marquesas showed trace of nitrite after 18 hours. * ‘ A gave strong nitrite reaction 22 se pa 70 miles west of Ushant never gave nitrite or Ammonia reactions. ” ”? The cultures from the Marquesas showed a slight amount of Ammonia formation, but the decomposition of the nitrite did not proceed further. . From these experiments it appears that the bacteria from sub- cultures from the Marquesas have a much greater denitrifying power than those from subcultures from a point seventy miles west of Ushant, and that as the bacteria from the Marquesas appear to be of the same species as those investigated at the Dry Tortugas, their power of causing complete denitrification in the modified Gran’s medium has been lost by cultivation on Peptone Agar, or during the fourteen days journey from the Marquesas. The presence of the thick layers of fine chalky mud within the Marquesas Keys, and elsewhere in many places near the Florida coast, led to a consideration of the possibility of its precipitation by bacterial agency. Since these bacteria grow freely in Gran’s medium, the Calcium salt of a simple organic acid is a sufficient source of organic food for them, and it seems probable that they would thrive in sea-water containing the products of decomposing vegetable matter, provided that the nitrate supply and conditions of light and temperature were suitable. Such conditions should be especially well fulfilled by the drainage into the sea of a well-wooded country with a calcareous subsoil, and the soluble organic Calcium salts carried to the sea in this way, would there be precipitated as Calcium carbonate by the action of the bacteria. In addition, the elimination of the acid radicle from the nitrate in the process of denitrification, by whatever stages it may occur, must leave the alkaline base free to destroy the normal equilibrium of the salts in sea-water, and by increasing the alkalinity, would also result in the precipitation of Calcium carbonate. To test this theory, cultures were made in a medium having the following composition,:— Calcium succinate : : : ; 2°5 grammes. Potassium nitrate . : ‘ . : 0-5 Sea-water . ‘ ‘ é : =) 200070 exc. ° 3) THE ACTION OF SOME DENITRIFYING BACTERIA, 153 Calcium succinate is soluble in these proportions, and the medium is quite clear. Free growth was manifested by the cloudiness of the medium forty-eight hours after inoculation, and nitrite formation was apparent. After ninety-six hours the medium appeared quite milky, and this milkiness was due to the presence of exceedingly fine particles of a substance which was soluble in dilute hydrochloric acid with evolution of gas, and was presumably Calcium carbonate. These particles were so minute that they remained in suspension in the liquid, and could only be satisfactorily separated from it by centre- fugalizing. The addition to this culture of very fine particles of hydrated Calcium sulphate, or of larger particles of sand, resulted in the aggregation around them of the particles of Calcium carbonate, forming a concentrically laminated concretion around a_ central nucleus. These concretions were hard, and of almost crystalline appearance under the microscope, and were soluble in dilute Hydro- chloric acid with evolution of bubbles of a gas which when the operation was performed on a microscopic slide, could be completely absorbed by running in a solution of Sodium hydrate under the cover slip. Once this process of concretion has been initiated, it appears to progress independently of the presence of particles which act as nuclei, and a large concretion may often be found having a number of smaller concretions around it, or continued into a chain of small spheres, the whole presenting somewhat the arrangement shown by freely budding yeast cells. The deposition of this form of Calcium carbonate also takes place on the sides of the flask, and more especially over any area where the glass is scratched or roughened. It would seem a reasonable suggestion that similar bacterial agency may have played a part in the formation of odlitic limestones, which are composed of concretions of Calcium carbonate around central nuclei often consisting of particles of sand or shells. In the same way bacterial action may have been of great importance in the pre- cipitation of the various chalk strata, and indeed in the formation of many rocks largely composed of Calcium carbonate. If this view as to chalk and odlitic limestone formation is correct, it would seem probable that the precipitation of these rocks must have occurred in comparatively shallow seas, which received the drain- age from a country in which extensive weathering of rocks containing Calcium salts was progressing, and where there was a considerable amount of vegetation from the decay of which the organic matter necessary for bacterial growth would be derived. NEW SERIES.—VOL. IX. NO. 2. OCTOBER, 1911. L 154 ~~ ~G. HAROLD DREW. DISCUSSION OF RESULTS. It seems that the rate of denitrification in equal volumes of the same culture medium, inoculated with equal volumes of samples of sea-water, must be a function of the number of bacteria in the sample, the tempera- ture at which the cultures are grown, and the specific power of denitrifica- tion of the individual species of bacteria. Considering the rapid multipli- cation of bacteria when the food supply is plentiful, up to a maximum determined chiefly by the accumulation of the waste products of their own metabolism, it appears that the factor of the number of bacteria in the sample may be neglected within the limits of these experiments. For example, the number of bacteria in 1000 cc. of Gran’s medium at the end of twenty-four hours would probably be much the same, whether it were inoculated from a sample containing eight or sixteen bacteria per 1 cc. Similarly, it was a matter of experience that the first trace of nitrite formation was observable at about the same time, whether 5 or 10 ce. of a given sample had been used for inoculation. Consequently it appears that, for purposes of comparison, and with- in the limits of the experiments described, if the temperature be the same for the cultures compared, the rate of denitrification is a measure of the specific denitrifying power of the particular species of bacteria. Thus it has been shown that in cultures grown at temperatures approximating that of their normal environment, the denitrifying power of bacteria in seas in the American tropics is considerably greater than that of bacteria in the water to the north of the Bay of Biscay, and that they have the power under favourable conditions of completely breaking down nitrates, which those from the Bay of Biscay do not possess. It has also been shown that these bacteria possess the power of precipitating Calcium carbonate from solutions of simple organic Calcium salts, and it is here suggested that some similar action has played an important part in geologic times in the precipitation of various rocks, wholly or in part composed of Calcium carbonate. Investigation and comparisons of denitrifying power of samples of the coastal waters of the English Channel have been purposely avoided, as previous experience had shown that the bacterial conditions in this region are so very complicated. It should, however, be men- tioned that in this coastal water denitrifying bacteria are present which are capable of entirely decomposing nitrates into free Nitrogen, e.g. the Bacillus repens (Gran), but that this complete denitrification is only possible at a temperature far higher than that of their normal environment, and that at a temperature of 15° C. even the forma- THE ACTION OF SOME DENITRIFYING BACTERIA. if3\5) tion of nitrite from the nitrate takes place with considerable slow- ness. A point that has not yet been considered is the origin of the nitrate supply in the sea. Nitrates are absorbed by diatoms and the phyto- plankton in general, and are presumably built up into complex nitro- genous compounds within the plant. If these compounds, on the death of the plant, are broken up and the Nitrogen again rendered available for use in the form of nitrates, a series of reactions must be gone through which may well be performed by bacterial agency, and this also apples to the waste nitrogenous products of animal meta- bolism. In addition, it has been shown that nitrates are actually destroyed by the denitrifying bacteria, which would thus tend to keep the nitrate concentration down to the level necessary for their own existence, and would come into competition for this essential with other forms of plant life. If the bacteria are successful in decom- posing nitrates to the extent of entirely removing the Nitrogen from all chemical combination, as seems probable from the experiments in cultures, it follows that there must be some source of nitrates in order that the concentration in the sea may remain constant. The existence of nitrifying bacteria, which are capable of oxydizing the Nitrogen of the air to form nitrates, and are symbiotic with some of the Algze found round the shores of the North Sea, has already been demon- strated, and it seems possible that forms having a similar nitrifying action remain to be discovered in the open sea. As they at present stand, the observations noted in this paper can at most be considered as offering a slight indication of the part played by certain bacteria in the metabolism of the sea. To obtain a real insight into the question, it would be necessary to make extensive bacterial and chemical investigations in tropical, tem- perate, and Arctic seas, and more especially to make a study of the bacterial flora at different depths at some spot where considerable depth, unaffected by currents, could be obtained. The probability that bacterial growth plays an important part-in causing the precipitation of certain bottom deposits in coastal waters, and conceivably also in the deep sea, is a subject that appears well worthy of further investi- gation. [> ASG 39 The Problem of Sex Determination in Dinophilus gyrocitliatus. By Cresswell Shearer, M.A. CLARE COLLEGE, CAMBRIDGE. THE group of primitive Annelids Dinophilus comprises some eight or nine species. They are remarkable for the fact that some show a well marked sexual dimorphism, in which the male is rudimentary, without any mouth or digestive tract, while in others the sexes are the same size and exhibit no signs of this dimorphism. The group as a class, therefore, is readily divisible into two subdivisions, in one of which all the species are sexually dimorphic, unpigmented, while in the other they are highly pigmented, a bright red, and are sexually monomorphic. The former may be called the Leu- codinophilidae, while the latter may be called the Erythrodino- philidae. The known species, many of which are of doubtful specific value, may be arranged under these two subdivisions, as follows :— 1. Dinophilus gyrociliatus, Schmidt, 1857. 2. Ms Conklini, Nelson, 1907. Leucodinophilidae 3. fs apatris, Korschelt, 1882. 4. 5 metameroides, Hallez, 1879. 5. “S pygmaeus, Verrill, 1892. 6. a vorticoides, Schmidt, 1848. : ‘ a 7 Gardinert, Moore, 1899. Erythrodinophilidae i NY Z : ella tae al 8. 3 teeniatus, Harmer, 1889. 9: 45 gigas, Weldon, 1886. Of the Leucodinophilidae the first three species, D. gyrociliatus, D. Conklini, and D. apatris, are closely related, and are probably one and the same. The form on which the following work has been done is one of these three species, though exactly which of the three I have been unable to decide. I have placed it under the head of D. gyrociliatus, as this is the oldest of these names. Figures of the male and female drawn to scale, are shown in Fig. 1. It will be seen that the female is very much larger than the THE PROBLEM OF SEX DETERMINATION. 157 male, and that the sexual dimorphism is well marked. It was obtained some three years ago from some sandy material collected in Plymouth Fie. 1.—Dinophilus gyrociliatus. Rudimentary male and full-grown female. The female shows the broken nature of the ciliated bands in the head region and the solenocyte bearing nephridia. Sound. I introduced it subsequently into the tanks of the Plymouth Laboratory, where it has since established itself, and breeds. Korschelt was the first to point out that a marked sexual dimorphism is present in D. apatris, where the male is small and rudimentary. He also observed that the female laid two kinds of eggs together in one capsule. Some of these were almost six times the size of the others, and were destined to give rise to the females, while the small ones gave rise to the rudimentary males. The large female eggs are laid in the proportion of three to one of the small male ones. Here is apparently a clear case in which we get sex determination in the ovary long before fertilization. Korschelt took it for granted that the eggs were only fertilized after being laid. The presence of this species of Dinophilus at Plymouth afforded me the opportunity of thoroughly investigating 158 CRESSWELL SHEARER. this question again, and of determining the manner of formation of these two kinds of eggs. As Korschelt has determined, the two varieties of eggs are laid together, but they develop immediately without the presence of the male, and are therefore fertilized inside the body of the female. In a few days the small male egg gives rise to the rudimentary male, which at the time it is ready to leave the capsule is full-grown and sexually mature (Fig. 1). The female, on the contrary, when she leaves the capsule, is very small, and still in the larval state, the ciliation and arrangement of the segments being quite different from that in the full-grown condition, which is only attained after a considerable period, and may never be assumed if she does not happen to obtain proper food. Thus in culture jars in which food is not found, the females will remain as long as six months in an immature condition, and never grow up. The young worms may be clearly watched within the capsule as segmentation and development proceed. The development is direct, and as the time for hatching approaches, the young larval females are seen to spin round within the capsule. This denotes that they are about to hatch and leave the capsule and commence their free existence. If the capsule is placed under the low power of the microscope at this stage and carefully observed, it will be seen that the little males are actively copulating with the small females at this time within the capsule. The actual transference of the sperm from the testis of the male through the penis into the body of the female can be distinctly seen. Every female as she passes out of the capsule is seen to carry a small mass of sperm, collected under the gut at the junction of stomach and intestine at the point where the ovary will subsequently appear. The rudimentary males seldom or never leave the capsule, dying shortly after the females escape. Examination of any immature free-swimming females, taken from culture dishes at random, always shows that they are all fertilized. If they are carefully fixed and sections cut from them at this stage, it will be seen that the germ cells have not been differentiated, and although a mass of sperm is collected at the point where the ova will subsequently appear, no trace of them can be detected. These only appear at a much later date, when the female has grown considerably in size. They are seen then as a few small refractive cells in the living state, and as small nuclei sur- rounded with hardly any cytoplasm in the stained condition, beneath the gut and amongst the mass of sperm. Shortly after they appear it is seen that each one is joined by a spermatozoon, the head of which THE PROBLEM OF SEX DETERMINATION. 159 has become embedded or attached to its nuclear wall, so that ultimately the nucleus of each primitive ovum is seen to be composed of one part derived from the spermatozoon, and the other part the female portion. These two elements of the nucleus never fuse, but retain their indi- viduality throughout all the dogonial divisions. The double nucleus divides amitotically, each half separately. In the majority of the divisions the male and female portions of the nucleus divide equally, so that a similar quantity of nuclear material, both male and female, gets into each daughter cell. There are probably about forty to fifty Gogonial divisions in all. In these the male and female portions of the nucleus divide and move apart simultaneously, the male portion usually dividing first. Now and again, however, the female half of the nucleus seems to divide before the male portion, so that the male portion gets left behind and is shut off entirely in one of the daughter cells. Therefore of the two resulting cells of this division, one has the whole of the male part of the original nucleus and its share of the female portion, while the other has only half the female and no male substance. This appears to be the sex determining factor ; for of these two daughter cells, the one that has received the whole of the male element, plus the female element, becomes the female, while that which has received the female portion alone be- comes the male. Both these kinds of eggs, once the sex determining division has taken place, grow rapidly. They seem to accomplish this through the power of absorbing and building up into themselves all the other immature egg cells with which they happen to come in contact, and in which the divisions of the two portions of the male and female substance has been equal. The outcome of this process is that the male egg is not fertilized, while the female egg is. It is, how- ever, impossible to speak in the strict sense of the word of the male egg as unfertilized, as it has been directly under the influence of the sperm in all the early dogonial divisions previous to the sex deter- mining one. For all the primitive germ cells are joined in the first place by a spermatozoon, irrespective of the fact that only some of these will give rise to ova later, and that the majority will be only nurse cells. It is only in the late stages, shortly before the female egg is laid, that the two portions of the nucleus, the male and female actually fuse beyond recognition. As the two kinds of eggs, male and female, are not found in a simple ratio, but in the proportion of three or two, to one male, it is probable that some other division takes place in the case of the female egg. I have been unable to decide this point so far, from my sections. I have made a careful study of the maturation divisions, and as Kor- 160 CRESSWELL SHEARER. schelt has determined in J. apatris, there are two polar bodies given off by both kinds of eggs, the first polar body in turn dividing after it has been given off. There are twenty somatic chromosomes. In both eggs there is a regular reduction in the first maturation division, ten dumb- bell shaped chromosomes going out and ten remaining in the egg. In the female egg, however, this process is somewhat different from that in the male, as there seems to be something similar to a synapsis stage in the former which is missed out in the latter. In both eggs after the extrusion of the first polar body, the ten chromosomes remaining in the egg divide, bringing back the number to twenty again. In the second polar body, in the male egg, apparently twenty or eighteen chromosomes go out and the same number remain in the egg. The second maturation division in the female egg, I have been unable to obtain satisfactorily in sections so far, and it may prove that the second polar body in this egg is simply derived from division of the first, and that only one polar body is actually given off by the female egg. In the first and second segmentation divisions in both eggs there are apparently twenty chromosomes. It is, however, very difficult to make out their number in the male egg very accurately on account of their small size, there are at least eighteen or more, and probably twenty, as in the female ege. The details of the maturation divisions I wish to reserve for my full paper on the subject, shortly appearing in the Quarterly Journal of Microscopical Science. ie dGiiea | A Table showing certain Cultural Characteristics of some of the Commonest Bacteria found in the Labora- tory Tanks at Plymouth. By G. Harold Drew. THE following table is merely intended as a rough classification of eight of the most common species of bacteria found in the Laboratory Tanks. Only the forms which occur free in the water in the greatest numbers have been considered, and the moulds and higher bacteria which can be cultivated from scrapings from the walls of the tanks have not been included. Until further and more detailed work has been done on the subject, it seemed better to designate the various species by numbers instead of names, since the majority have probably not previously been described. The culture media employed were similar to those ordinarily used for bacteriological work, with the exception that fish broth (made from dogfish) was substituted for meat broth, and that sea-water was used in place of tap-water. All media were faintly alkaline to Neutral Red. The medium designated as “ Gran’s medium,” has the following com- position :— Sodium chloride . : . 88 grammes. Potassium nitrate ; OP2 Dy = Sodium phosphate (Na HPO, 12 H. 0) G20. =, Calcium malate : ‘ . In excess of its solubility. Distilled water. ; : : » 2090°0 ¢.c; and is fully described by H. H. Gran in the Bergens Museums Aarbog, No. 3, 1901. The sample of water, from which the bacteria classified in the table were isolated, was collected from just below the surface of one of the tanks in the upper Laboratory at 5.30 p.m. on December 6th, 1909. 0:1 cc. of this sample plated on Fish-broth, Peptone Agar at 30° C., gave an average of 130 colonies of all kinds. Samples collected on other occasions show that the eight species described below may be considered as always present and abundant in the tank water. HAROLD DREW. 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CULTURAL CHARACTERISTICS uniy Aq [ItJap UL paquiosep Suadat sn])100g ou st Stud Feu UoIsNpToOUoD a4 0} yutod yor sj[nsal oars ‘ojo ‘youryg fivong auey ‘asoqov'T ‘Asoo NH SULTRY WOO eipaut auoqdag uty -B[94) SUIT YEA sjuatitiedxe daqyang “Aep Uy qe quoredde uo -BULLOJ BITOTUIY ‘sAup anof UTYGIM WOTJwWOYAZ Lap purx ‘qyaois pidey “TI MO.LS ON ab ‘skep moj ur quowwdde WOTPBIYIAWIMAp pus ‘qyjao1s piley | “YJMOIS ON -BIUIOJ Plow TFT uororjaubry prey ‘asuryo THO[OD ‘qRys ouo[E [[® YMoOAS datq | [TB YpMors “T]ZMOLS ON ‘apuByo Ino MOTJOR] -anbry padeys-jau YGMOLS 99.1] -[00 ON ‘sqjued aadeap ut woyonpat qIpST[s puoseq asueyo IMojoo ON “aoe -ivadde — ATToo Ay |*qvqgs suo[e I[® YIMOIs daly ‘aHUBITLO MOLOD ON ‘qeys suo[R daly Q “UJMOLS ON ‘gounieadde | A[[Oo\A ‘esueyo qmojoo ON *qrys Suope [fe yyparorry “TUNTpatu atyy [[B JO worjorjoubry oqo -u00 9} UT SUT -4[Nsar WoOS *LOTY -onjaubiy jo vais pedeys-[ouuny padopaaap A, prdey ‘avaddv 04 suisaq yt shep aay] Jaye juq “ysay qe uotjorjoubry ON “quqs Huoye [i@ YIMols day] “MOTIOVY -aubtt oN ‘Al uo aoRJIUs UO JAMO “SpUBMUMO Dp |“ sutpeards Apprdea worjoejanbry jo wae padeys-[au “UN “WPA. Val y] “A110 09 TMOG -Ua pos B Jo Satlooeq tan pott a[oyM oly SoaIng[no po Wy ULSIRUL T[JOOUIS WIOJIULQ “aTppiu ut Jods Jesuap SULMOYS ‘yqaous pidea L194 JO SALMOJOD azLIPA LBTUdATD “UR[UHaL pus YooUs ULSIBV “IVlNoITg "SOLUO[OD DATJORAJOL ATYSuy “yuaonsweay “TTY Ah “Oul| -quo LepNSadat APUG FTA ARMA ‘saTMO[O BUT -Oo[-Apnoypo faq tp adv] "YUIAMOLS MO[S jo pur ‘aorjins 9aoqu peyvaoja Yon sar -ofoQ ‘avpnsetat ATauy UIsivyY ‘ooursvod -de aepnuers AoA TYLA Somopoo = a4TA — Le[MIAL) | ‘spot a[tout ATaATjoR JpRMg) | ‘O[TFOUW- LOU “spod a oul | ATOATPOR aznUryL “OLIQTA ay -o ApaaTjoR [Burg Piaeeg Note on the Early Larve of WNephthys and Gl/ycera. By H. M. Fuchs, B.A. With 10 Text-figures. THE larva of MNephthys was originally described by Claparede and Meceznikow (2) in 1868, from specimens obtained from the Plankton at Naples. Since then it has been figured by Fewkes (3), Hicker (5), and Gravely (4). The larva of Glycera is, as far as I am aware, hitherto undescribed. The larvee described below were reared in the laboratory at Plymouth, at the suggestion of Dr. Cresswell Shearer, from artificially fertilized eggs. Both Nephthys Hombergi* and Glycera convoluta can be obtained from the sand in Jennycliff Bay at low water. Nephthys occurs more commonly than Glycera, but large specimens are not fre- quent. On the other hand, very young specimens are completely absent. It is at present altogether unknown what becomes of the young of these worms from the time when they sink down from the Plankton until they grow to about an inch in length. From June to August most of the larger individuals contain ripe sperm and ova, which are frequently discharged in captivity, within a day or two from the date of capture. It was found unnecessary, however, to wait until the generative products were shed naturally. The method of procedure was as follows: The worms were slit up with a pair of scissors and placed in “outside” sea-water in sterilized finger-bowls, the males being kept apart from the females. As soon as the sexual products emerged from the body cavity the worm was removed, several being treated in this manner to ensure the presence of ripe eggs. A few drops of liquid from the suspension of sperm was then added to the bowl of eggs and the latter stirred and allowed to settle. Wilson (9) has stated for Nereis and Treadwell (8) for Podarke that if the eggs are cut out of the body of the female, segmentation is irregular. This is not the case with Nephthys and Glycera. As soon as the larve swam to the surface they were pipetted off and placed in jars of filtered sea-water. All attempts to feed the larvee failed, and in consequence, although they could be kept alive for three weeks or a month, after about * It is just possible that a few small specimens of N. caeca were included with the NV. Hombergi. NOTE ON THE KARLY LARV OF NEPHTHYS AND GLYCERA. 165 fourteen days no further development took place. I have to thank Dr. Allen for providing ne with pure cultures of Diatoms and other Algee, but I was unable to find any food which the larve would eat. Nothing is easier than the rearing of Serpulids in a laboratory, and especially is this the case with regard to Pomatoceros, which is a hardy shore form, found growing even at the mouths of drains. It breeds naturally and in great profusion in the tanks at Naples, the young worms settling down and forming their tubes without having any care bestowed on them. But very few of the free-living Polychaetes have been reared to a late stage. Chaetopterus is an example of one which has been raised (Allen and Nelson 1) and Lillie (6) was successful with Arenicola. I have found it very easy to rear the larve of Nereis dumerili, which are derived from yolky eggs, as far as thirty segments or more. Quite otherwise, however, is it with those free-living Polychaetes which have a pelagic larva. It would thus seem that it is possible to rear from the egg such forms as the Serpulids, which have a typical Trochophore, and those which have yolky eggs, but no method has as yet been devised by which the majority of the errant forms can be fed and raised. The egg of Glycera is discoidal, and it does not swell up in sea- water. It is granular, but comparatively transparent, with a lighter coloured nucleus. The egg of Nephthys is also flattened at the poles, but is opaque. The cleavage is equal. Unlike the larva described by Claparéde and Mecznikow (2), the larva of Nephthys Hombergi is granular and so opaque as to render the study of the internal organs in optical sections very difficult. Neither iu shape nor in transparency can it be called a typical trochophore. An early stage is shown in Fig. 1, The upper hemisphere is more conical Fic, 1.—Larva of Nephthys. 2 days. I'rom the ventral aspect. 67. p.—Brown pigment. and of smaller diameter than the lower, and is surmounted by an apical patch of cilia. The mouth is situated mid-ventrally on the lower hemisphere, and strong cilia can be seen working within it. Brown 166 H. M. FUCHS. * pigment is scattered on the surface in small chromatophores, which are concentrated below the trochal ring and round the mouth. Fig. 2 Fie, 2.—Larva of Nephthys. 6 days. From the ventral aspect. shows a later stage, the general shape of the larva being the same as before. The alimentary canal is divided into stomach and intestine, and there is an anal patch of cilia. Fig. 3 shows a trochophore of four- Fic, 3.—Larva of Nephthys. 14 days. From the ventral aspect. Or. p.—Brown pigment. d. gr. p.—Dark green pigment. int.—Intestine. J. gr. p.—Light green pigment. sf.—Stomach. NOTE ON THE EARLY LARV OF NEPHTHYS AND GLYCERA. 167 teen days old. The upper hemisphere is larger and blunter than the lower. There is a patch of cilia extending from the lower lip of the mouth towards the anus, which is terminal. The apical cilia are usually motionless. Light green pigment has appeared in an in- complete band round the upper hemisphere, and a pair of patches on either side of the anus. These latter are characteristic of the larva of Nephthys (3). The large stomach, which fills the greater part of the upper hemisphere, has a dark green pigment in its basal walls, the remainder of the gut being yellow in colour. The intestine is sac-like. The larva described above differs very much from that figured by Claparéde and Mecznikow (2). It resembles in some points that of Fewkes (3), but is of a different shape. Hacker (5) and Gravely (4) described the Metatrochophore and later stages only. Swimming blastule of Glycera convoluta appear some ten hours after fertilization, and the trochal ring is formed in twenty-four hours. An early larva is shown in Fig. 4. It is spherical and granular with a Fic, 4.—Larva of Glycera. 26 hours. ap. ¢.—Apical cilia. ciliated trochal ring and an apical tuft of long cilia. Fig. 5 shows Fic, 5.—Larva of Glycera. 58 hours. Left side view. m.—Mouth. 168 H. M.. FUCHS. THE RELATION OF THE HEART-BEAT TO ELECTROLYTES. 181 which accompany the contraction as well as the excitation processes in muscle provide important evidence, and other facts might be adduced in favour of the idea that an essential part in the mechanism of the contractile process in muscle is played by surfaces or membranes possessed of differential permeability towards ions. It is familiar that such membranes form an integral part in the theories of excitation pro- pounded by Nernst and elaborated by Lapicque and by Lucas and Hill. The existence of membranes or surfaces in muscle is granted by the histologist to almost any desired extent. In the case of skeletal muscle indeed, it is scarcely an exaggeration to say that every histolo- gist who has undertaken the investigation of its structure has described and attached his name to some new membrane, line or band. It is clear then that we are not making any new or rash assumption in stating that certain surfaces or membranes form an integral part of the muscle mechanism, and the ionic permeability of these membranes * factor of importance in that mechanism. The possession by a membrane of differential ionic permeability is no imaginary attribute: simple experiment shows it to be a very usual property. This fact is brought out most clearly by experiments on the influence of membranes on concentration cells. Direct experiment shows also that the ionic permeability of mem- branes—and a great variety of them have been tested—is profoundly influenced by certain features of the solutions in contact with them. For a given membrane the factors likely to influence its ionic permea- - bility are— (1) The P+ of the solution with which it is treated. H (2) The presence in that solution of polyvalent ions. The simple trivalent positive ions are enormously more powerful than the divalent. sa When a membrane such as a piece of peritoneal membrane, the lining of an egg-shell or a thin sheet of gelatine, is treated with a very small concentration of acid, or of one of the simple trivalent ions of which we have spoken, its ionic permeability is modified. Direct experiment shows that this altered condition is retained, it may be for minutes or hours, while the membrane is washed with a “ neutral” solution free from the polyvalent ions—the persistence of the effect depends on the nature of the membrane, the concentration of the solution used to produce the. * Here and in the following pages we employ the word membrane with the under- standing that it need not necessarily mean anything more than the surface of separ- ation between two phases in the system. 7 7 182 GEORGE RALPH MINES. - alteration and the length of time it was allowed to act—but if the membrane is treated with a faintly alkaline solution the effect of the acid or the trivalent ion is removed at once. The parallel with the action of these same agencies on the heart is sufficiently striking. And the parallel does not end here. Seeking to test further whether this potent action on the heart is indeed a general property of trivalent ions, I employed solutions of | certain cobaltammines which yield complex trivalent ions in neutral solution. The compound used most frequently was luteo-cobalt chloride [Co(NH,),]Cl,, which yields the ion [Co(NH;),]**’. Through the generosity of Prof. Werner I was able to test also four other complex trivalent ions. All agreed in showing very much weaker action on the heart than the rare earth solutions. A concentration of the complex ion of the order of 1000 times as great as of the simple ion is needed to stop the perfused heart. A similar difference is shown in the action of these ions on the gelatine membrane. Even large concentrations have only a slight effect on the ionic permeability of a membrane of this type. Lhe influence of hydrogen ion concentration and of trivalent kations on the heart is paralleled in detail by the action of these cons on the ionie per- meability of certain artificial membranes. The ionic permeability of a membrane can be shown, both theoretic- ally and experimentally, to depend—other things being equal—on the electric charge of the membrane. Now it can be demonstrated by a variety of methods that most membranes in water, or in a solution like our “neutral saline,” possess a negative charge. This charge is reversed with great ease by acid or by the rare earth solution. This can easily be shown, for instance, by the use of dyes, which are themselves charged colloids. A piece of gelatine soaked in “neutral saline” stains very feebly with eosin, strongly with methylene blue. But after treatment with acid or rare earth it takes up eosin with great avidity and is hardly tinged by methylene blue. Losin consists of negatively charged particles, methylene blue of positively charged particles. If the membrane has . a negative charge it can hold the latter, if a positive charge, the former. This method also reveals the fact that the complex trivalent ions are much less potent than the simple trivalent ions in altering the charge of the gelatine membrane. In these membranes we_are, of course, dealing with substances in the colloidal state. If instead of using extended sheets of material we turn our attention to colloidal solutions, where one phase of the colloidal system consists of discrete particles floating freely in the * THE RELATION OF THE HEART-BEAT TO ELECTROLYTES. 183 other, it is possible to study the action of electrolytes more simply still. For in such systems the alteration of electric charge on the particles is often followed by an alteration in their state of aggregation, which may be manifested as an alteration in the colour of the solution, or as is very usual by actual precipitation. For instance, when the particles are negatively charged, as in colloidal gold or in a solution of egg-white, the addition in suitable concentrations of electrolytes capable of conferring positive charges on the particles causes agglutination of these particles. Testing a large number of colloidal solutions by this method, it becomes apparent that there are great differences in the relative efficiencies of different ions in affecting the charge of different surfaces. Thus considering the simple and the complex trivalent ions one finds that many colloidal solutions are quite as readily precipitated by them as by the simple trivalent ions (e.g. colloidal gold, arsenious sulphide, boiled diluted egg-white, and many more), while others which are readily precipitated by the simple trivalent ions are unaffected by even large concentrations of the complex ions (e.g. unboiled diluted egg- white, haemoglobin, etc.). Of a large number of colloidal solutions, those which are very sensitive to complex as well as to simple trivalent ions are found to belong to that class of colloids known as lyophobe or -suspensoid; those sensitive to the simple but insensitive to the complex trivalent ions, to the class called lyophil or emulsoid. Clearly, what has been stated would indicate that the membranes in the heart muscle are of an emulsoid colloidal material. This indeed confirms in one special instance a conclusion drawn from entirely different considerations by Martin Fischer as to the nature of the body proteins in general. The study of the action of these electrolytes on the heart will give more precise information than this. When the ratios of the activities of the simple and the complex ions on surfaces of various compositions have been worked out in detail, we shall be enabled to define precisely some physico-chemical constants for each surface. As has been remarked, the investigations of colloid chemistry have not as yet been carried far enough for this to be done, but ~ the methods are available, and the research would present no insuper- _ able difficulty. Just as the classification of liquids of two phases into suspensions, colloidal solutions and true solutions, is now recognized to be an arbitrary though convenient subdivision, there being no real boundaries between the classes; so it will probably be found among the colloidal solutions themselves. We shall learn in course of time to describe each colloidal solution in terms of certain essential factors which will include such terms as the viscosity of the phases, the 184 GEORGE RALPH MINES. amount of surface presented by them, and the electric charge they carry. At present this cannot be done in any quantitative sense, yet we may illustrate from material already available the nature of the biological interest which will accrue from knowledge of this kind. In the first place, new light is thrown on the relations between the living heart muscle and its normal environment. We have considered the importance of the P+ of the solution. lel Take now the case of magnesium. Mg*: is the only simple divalent ion which can be expected to act on the heart simply qua divalent ion. As we have shown, Ca‘:, Sr‘, and Ba‘: enter probably into special chemical relation with some constituent of the tissue, while the other metals which give divalent ions (such as beryllium, zinc, cadmium, ete.) form solutions which are hydrolyzed.* It is generally stated that an emulsoid colloid is very insensitive to salts. This is not true, at any rate in the case of some typical emul- soids, when simple trivalent ions are considered. Quite small con- eentrations of the rare earth solutions can produce large effects on colloidal materials which present all the other characters of emulsoids. Many experiments on a variety of colloidal solutions lead me to the conclusion that a fairer statement of the characteristic relations between emulsoid colloids and electrolytes would be this: that as compared with the suspensoids there is a very great difference in the. - relative concentrations of simple di- and tri-valent ions needed to produce the same effect. While for suspensoids the ratio of the activities of di- to tri-valent ions is taken ordinarily as of the order of 3 to 100, for emulsoids it is rather of the order of 1 to 10,000. It is certain that these ratios vary widely with different colloids, and that from a more precise study of these ratios than has at present been undertaken much more will be learnt about the colloids themselves, ‘but the broad distinction appears to hold. - On the artificial scheme of membranes, then, magnesium produces effects tending in the same direction as those produced by the simple trivalent ions and by the hydrogen ion, but is enormously less potent than these. Precisely the same is true of the action of Mg-: on the heart. ~ * Incidentally it should be noted that the effect of hydrolysis, as regards the activity ‘of the solution on the heart and on colloids such as gelatine, is precisely opposite in the case of these solutions of divalent ions from that described for trivalent ions. For the H ions, though less powerful than the equivalent concentration of simple trivalent ions, are much more powerful than the equivalent concentration of divalent ions in these rela- tions. Consequently a solution of beryllium, which is hydrolyzed, affects the heart, and colloid systems of the class sensitive to H*, much more powerfully than the non-hydrolyzed solutions of magnesium. THE RELATION OF THE HEART-BEAT TO ELECTROLYTES. 185 In the ray heart, for instance, a concentration of -000001 M to ‘00001 M of Ce*‘': or Nd‘: is sufficient to cause prompt arrest in diastole. The same effect can be produced by Mg°’, but the concentration needed is about ‘02 M. If our explanation of the action of magnesium on the heart is true, it should of course be found that the dose of Mg needed to arrest the heart depends greatly on the P+ of the solution in which it is applied. An increase in the H numerical value of the P+ should tend to counterbalance the effect H of magnesium. This is abundantly confirmed by experiment. For example: a ray heart was brought to rest by raising the concentration of Mg-: in the solution, perfusing it to (025 M. The diastolic arrest occurred after two or three contractions. When the heart had remained completely at rest for a minute with this solution running through, without changing the concentration of Mg:: the P+ of H the solution was raised from about 6°5 to 9. Within a few seconds the heart started beating, and continued to beat in this solution as strongly and regularly as in the original solution which contained no excess of Mg°-. — Consideration of the relations of colloidal materials to simple and com- plex trivalent kations enables us to make certain inferences as to the nature of these colloidal substances from which we can predict their relations to certain other tons. Applying this principle to the heart, we arrive at conclusions as to the nature of the colloidal membranes in its structwre Jrom which we can predict the action of magnesium upon the heart-beat. Finally, I wish to indicate certain directions in which experiments of this kind may assist in the development of a scientific comparative physiology. Two instances which I have encountered in the course of work at the Plymouth Laboratory this summer may serve as illustrations.* The facts already stated about the general similarity in the re- lations of the heart muscle of widely different species to Ca, Sr, and Ba encourage the belief that the general character of the mechanism is the same in these varieties of heart muscle. Can we find differences in the relations of the membranes to “charging” electrolytes? What kind of differences are to be looked for ? A study of the properties of colloids by a variety of methods leads to the following conclusions with respect to the part played by the * I am much indebted to Miss Dale, of Newnham College, for her skilful assistance in this work. NEW SERIES.—VOL. IX. NO, 2. OCTOBER, 1911. N 186 GEORGE RALPH MINES. nature of the colloid itself in determining its relations to electro- lytes. (1) Different surfaces in the same solution may take on different electrical charges. (2) Different surfaces possessing the same charge in some par- ticular solution may require different concentrations of the same electrolyte to produce equal changes in their charges. The physiological expression of a difference in the heart membranes of the kind indicated in the first of these conclusions would be of this type: that one kind of heart should behave in a neutral solution as another would behave in an acid or in an alkaline solution, other conditions being equal. The heart of the mollusc Pecten maximus differs from all the vertebrate hearts studied in this way. On perfusing it with “neutral saline” it stops instantly in systole. It will beat, however, if we alter the P+ of dul the solution in the direction of slightly increased acidity ; if the change is carried a little further, diastolic arrest—the characteristic acid effect . —is then produced. Now the P+ of the solution which perfuses the H heart of the living Pecten—the blood of the animal—is about 6°5 to 7, that is to say, very slightly on the alkaline side of our “ neutral saline.” How then can the heart beat in such a solution? The answer is simple. The blood of Pecten contains a large concentration of mag- nesium. The divalent Mg‘* is present in sufficient concentration to bring the charge of the membranes to the value required for their proper differential permeability. The assertion that the blood of Pecten is practically the same as sea- water overlooks one important fact about it. Though its osmotic 4 pressure and concentrations of Na, Mg, K, Ca, etc. are closely the same | as in sea-water, there is a distinct difference in the P+ of the two H fluids. The P+ of sea-water varies from about 7°9 to 8:3, while that H of the Pecten blood is rather below 7. The difference may be demon- strated by adding a drop of a solution of rosolic acid to a little sea- water and to a similar quantity of the colourless blood of Pecten. The colour given by the former is bright pink, that of the latter yellow or yellowish pink. Thus we find that sea-water, despite its high concentration of magnesium, on perfusion instantly stops the Pecten heart in systole. 5 & THE RELATION OF THE HEART-BEAT TO ELECTROLYTES. 187 But if by the addition of a little dilute hydrochloric acid we change the P+ of the sea-water to the same value as that found for the blood H of Pecten (about 1°5 ce. of -1 M HCl in 100 cc. of sea-water), we have at once a solution which sustains the beat of the heart admirably. A solu- tion of the same composition as this but without magnesium causes the heart to stop in systole. Further, as is predicted theoretically, it is possible to replace the Mg by a very small concentration of a simple trivalent ion. Thus the Pecten heart which has stopped beating in “neutral saline” can be made to beat again and be kept beating for a considerable time by the addition to the solution of a concentration of about 00003 M of Ce’*': or Nd‘. Here we have one tri- valent ion doing the work of some 40,000 divalent ions, and it is therefore not surprising that the solution with rare earth does not, as a rule, keep the heart going so regularly as the solution with mag- nesium. Higher concentrations of the simple trivalent ions (e.g. ‘(0001 M) stop the Pecten heart in diastole exactly in the same way as a too acid solution. , We may express the difference between the heart of Pecten and the hearts of the vertebrates which have been studied, by saying that the membranes have different iso-electric points. This kind of difference is most probably related to a difference in the chemical composition of the membranes; it being a well-established fact that different protein substances found in the animal kingdom do show differences of this kind, according to the predominance of the “ acidic” or “ basic” amino- acids included in their composition. It is by no means a new suggestion that the differences between different species are at root differences in the chemical composition of their tissues; but as far as I am aware it has not before been pointed out by what mechanism such differences can affect the physiological behaviour of the living heart. The second conclusion formulated above as to the relations between different surfaces and the same electrolytes finds biological application in the differentiation of more closely allied species. The hearts of the elasmobranchs Raia and Scyllium are kept beat- ing satisfactorily for many hours by perfusion with the same “neutral” solution containing sodium, potassium, calcium, and magnesium ~ chlorides and urea, the solution being thoroughly aerated and of P+ H about 6:5. The concentration of Mg in the solution generally used was (005 M. Removal of the Mg from the solution only causes the beats to get rather faster. Increase of the concentration of magnesium 188 GEORGE RALPH MINES. causes slowing of the heart-beats, and if carried far enough, stoppage. Qualitatively these phenomena are alike in the hearts of the ray and of the dogfish. But studied quantitatively the ray heart is found to be much more sensitive to Mg than is the dogfish heart. Thus taking the mean of a number of observations on ray* hearts, the con- centration needed to give a 50 per cent reduction in the rate of beat (calling the rate in the solution with 005 M Mg=100) was only (009 M. For Scyllium it was (05 M. To stop the ray heart the concentration of Mg:: needed was about ‘02 M, while to stop the heart of Scyllium a concentration usually greater than ‘1 M was required. A corresponding difference is found in the sensitiveness of these hearts to the simple trivalent ions. Thus ‘00001 M Ce°:: readily stops the heart of Raia, while a concentration approaching ten times this magnitude is needed to produce the same effect on the heart of Seyllium. In these respects the heart of the angel fish, Riana squatina, was found to resemble that of the dogfish rather than that of the ray. The curve representing its behaviour in the presence of various con- centrations of Mg lies between that plotted for Raia and that for Scyllium, but much nearer the latter than the former. This is in- teresting, as in the ordinary morphological classification Rhina is placed between the Scyllidae and the Raiidae. It is said that Rhina resembles rays rather than dogfish in its general habits, yet it is possible that the character indicated by these experiments represents something deeper seated than the details of the responses given by the central nervous system. It is at any rate conceivable that by the extension of work along these lines we may learn to express genetic affinity in physico-chemical terms. One further point. It will be recollected that the behaviour of a membrane depends not only on its original composition and on the’ nature of the solution immediately surrounding it, but also to some extent on its previous history. I have found that the blood plasma of Scyllium and of Rhina is of P+ numerically slightly higher than H the plasma of Raia (clavata or blanda). It is possible that the difference in sensitiveness of the hearts of these animals to the “ Two species, &. clavata and R. blanda, were used in these experiments. No difference could be detected in the behaviour of the hearts of these two species towards electrolytes. Using rays of very different sizes no constant differences were found between the large and the small specimens. THE RELATION OF THE HEART-BEAT TO ELECTROLYTES. 189 agencies we have named is to be explained in part by the fact that their membranes have been bathed for years before the experiment in solutions of slightly different P+. . H There are striking differences in the relations of the hearts of different species of animals to hydrogen ion concentration and to polyvalent ions: these correspond exactly to differences found in the effects of these ions on the ionic permeability of different colloidal materials. A mechanism is thus sugyested which explains how differences in the chemical constitution of a tissue component may cause differences in the physiological behaviour of the living tissue. I venture to hope that these fragmentary remarks will make it apparent that a field of research hes open here, containing treasure not alone for the physiologist who desires to elucidate the mechanisms by which vital processes are wrought in the individual, but also for the zoologist, who in looking for clues of genetic affinities has learnt the limitations of pure morphology and the truth of the axiom of physiology that by studying what a cell can do we shall best learn what it is. For the furtherance of the science of Comparative Physiology the co-operation of morphologists and physiologists with physical, organic, analytical, and colloid chemists is necessary. This country is not lack- ing in any of these. The study of the science can be carried on best where the greatest variety of animals, living and in good condition, can be provided. This condition is most perfectly fulfilled at a marine station. _. At Plymouth the fauna is rich, and the facilities provided in the b Laboratory for keeping animals in condition for physiological experi- ment are admirable. It is earnestly to be wished that more workers in this country would interest themselves in a branch of zoology -and of physiology pregnant with interest for both sciences, and that their efforts might secure for the Plymouth Laboratory per- manent accommodation and equipment for physiological work, on a scale approaching that provided by marine stations in other countries. : In conclusion, | would express my warmest thanks to the Director and Staff of the Laboratory of the Marine Biological Association at Plymouth for much valuable assistance and advice.* September, 1911. * Iam indebted to Professor Stanley Gardiner for the use of a University Table at the Plymouth Laboratory during July, August, and part of September. The expenses of the researches were defrayed in part by a grant from the Government Grant Committee of the Royal Society. 190 GEORGE RALPH MINES. BIBLIOGRAPHY. References to work on the subjects named below will be found in the following :— Measurement of hydrogen ion concentration, ete. : Sorensen, C. R. Lab., Carlsberg, VIII, p. 1. 1909. Palitzsch, Publ. de Circonstance, No. 60. Cons. perm. pour Pexplor. de la mer. TOU: Action of chemical substances on the heart : Langendorff, Ergebnisse d. Physiol., IV. Luciani, Human Physiology, trans. by Welby, p. 337. 1911. Electric charge of membranes : Perrin, Journ. de Chimie Physique, II, p. 601. 1904. Action of trivalent ions on heart and on colloids : Mines, Journ. Physiol., XL, p. 327. 1910. Ibid., XLII, p. 309. 1911. png Types of Crustacean Blood Coagulation. By John Tait, M.D., D.Sc. (From the Marine Laboratory, Plymouth, and the Physiological Laboratory of Edinburgh University. ) HAVING been engaged at intervals during the last few years in studying from a physiological point of view the coagulation of crustacean blood —Tait (08), (10, A and B), (11)—and finding more variation in this regard in different crustacea than has hitherto been recognized, it suggested itself to me to inquire if the observed differences are correlated with any special physiological peculiarities, and, further, if they depend in marked degree on phylogenetic relationship. It is not a simple matter to settle either of these questions, and I make no pretence to have done so. At the same time, I have thought it worth while to put my observations on record, in the hope that the matter may thus sooner arrive at a satisfactory settlement. The literature of the subject and details regarding the technique of examination of the blood and other particulars I hope shortly to publish elsewhere. Suffice it here to state that at least three distinct modes of blood coagulation may be recognized in crustacea :— A. Simple agglutination of the blood corpuscles without any subsequent jellying of the blood plasma. (This is probably the most primitive and essential device both in invertebrates and vertebrates for procuring arrest of hemorrhage from a wound.) B. Agglutination of the blood corpuscles with subsequent general jellying of the plasma. C. Jellying of the plasma in two successive stages, the pre- liminary cell-agglutination being relatively insignificant. The first plasma coagulation consists of localized (primitively globular) clots, which occur around or in immediate relation to special blood corpuscles, originally discovered by Hardy (92), and by him named “explosive corpuscles.” At a later stage a second jellying process occurs, which this time involves the whole of the remaining plasma. 192 JOHN TAIT. Although it is not my intention to discuss in detail the basis on which crustacean coagulation is separated into these three types, it would be a mistake to convey the impression that a hard and fast line of separation can be drawn between them. The classification is’ arrived at by examining the blood always under the same conditions— that is to say, entirely removed from the vessels and placed cither in a dish or on a glass slide. These are, in fact, the conditions under which observations on coagulation are generally made. ‘The classifica- tion here given represents a more extended knowledge than that hitherto in vogue—originally due to Heim (92)—which recognizes two groups, viz. A and B above. COAGULATION C. It will be convenient to begin with coagulation C, which is associated with the presence of explosive cells, and which from its complexity must be regarded as a somewhat specialized form of coagulation. This form seems to be specially frequent in Isopoda. I have found it in Conilera cylindracea, in Idotea baltrca and emarginata, in Ligia oceanica, in Oniscus, and in one or more species of Porcellio, i.e. in individual members of three sub-orders out of six.* On the other hand, I have - failed to find it in Gnathia maxillaris, in Dynamene rubra, in Sphaeroma serratum,+ and in Jaera marina, though possibly owing to the fact that the amount of blood obtained from these relatively small specimens was too meagre to allow of a satisfactory examination. Among Amphipoda the occurrence of coagulation C seems to be much less frequent. It is not present in Gammarus marinus, Gammarus pulez, in Orchestia littorea, nor in certain specimens of Caprella examined by me. On the other hand, I have recorded its presence in Gammarus locusta—Tait (10, B). This was not, however, under the conditions above specified—that is to say, in blood wholly removed from the animal and examined separately on glass, nor have I as yet observed it under these conditions. As regards the Mysidacea coagulation C is absent in at least one species of Mysis (the only member of this order examined by me). Crustacean blood coagulation has been chiefly studied in the * The zoological classification referred to in the present paper is that given by Calman in Part VII of Lankester’s Treatise on Zoology. + In the course of this work I found that Sphaeroma serratum (with black chromato- phores) undergoes colour change in response to its background, like that undergone by Idotea—V. Bauer (05)—and by Ligia—Tait (10, C)—whereas Oniscus and Conilera, the latter possessed of orange and lemon-yellow chromatophores, do not. CRUSTACEAN BLOOD COAGULATION. 193 Decapoda, and it is of especial interest to know if coagulation C occurs among them. Of this order I have examined Pandalus montagui and P. brevirostris, Hippolyte varians and H. viridis, Palaemon serratus, Crangon vulgaris, Palinurus vulgaris, Homarus vulgaris,