m l 5? O m o THE RAY SOCIETY. INSTITUTED MDCCCXLIV. This volume is issued to the Subscribers to the BAY SOCIETY for the Tear 1879. LONDON: MDCCCLXXX. BfJ A MONOGBAPH FREE AND SEMI-PARASITIC COPEPODA OF THE BRITISH ISLANDS. BY G. STEWABJDSON BEADY, M.D., F.L.S., PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF DURHAM COLLEGE OF PHYSICAL SCIENCE, NEWCASTLE-ON-TYNE ; CORRESPONDING MEMBER OF THE ZOOLOGICAL SOCIETY OF LONDON, OP THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA, ETC. VOL. III. LONDON: FEINTED FOR THE RAY SOCIETY. MDCCCLXXX. PBINTED BY J. E. ADLARD, BABTHOLOMEW CLOSE. A MONOGRAPH BRITISH FREE AND SEMI-PARASITIC COPEPODA. VOL. III. GENERAL ANATOMY AND DEVELOPMENT OF THE COPEPODA. IN preparing this monograph it was not part of my plan to enter at all into the consideration of the physiology or internal anatomy of the Copepoda, but, in compliance with the wishes of some of my friends, expressed while the first volume was passing through the press, I have put together, in the form of a preface to this third volume, a condensed account of some of the more important observations which have hitherto been made on this part of the subject. It will be at once apparent that what I have attempted is nothing more than a general outline, in the production of which I have been greatly indebted to the works of Glaus, Gegenbaur, and Huxley. Had I included the truly parasitic species, in whose anatomy, physiology, and general habits of life so many points of the highest interest occur, I should have been travelling beyond VOL. III. A 45279 BRITISH COPEPODA. the professed limits of my work, and have extended this preface beyond reasonable bounds, — on which account those species are noticed in the most cursory manner, and only in elucidation of the proper subject of the memoir. The non-parasitic COPEPODA may be described as ENTOMOSTRACA, having an elongated body of variable form, but generally cylindrical, without a bivalved shell, and showing more or less completely a division of the body into numerous rings, or somites. There are two pairs of antenna, three pairs of prehensile and masticatory, or suctional, mouth organs, and five pairs of feet, adapted chiefly for swimming. The females are mostly fertilized by means of spermato- phores ; the ova are usually carried in external ovisacs ; when first hatched the larvae have only three pairs of limbs, and go through several metamorphoses before attaining the mature form. The parasitic species, at one end of the series approach very nearly in structure and general appear- ance to the non-parasitic ; at the other end they are extremely different, exhibiting, especially in the males, many most remarkable examples of retrograde development, so that without the study of their metamorphoses it would be quite impossible to re- cognise them as Copepoda, or even as Crustacea of any kind. Yet even in these degraded forms — at any rate in the females — natatory limbs in a very much atrophied condition are almost constantly found. GENERAL FORM. — The animal is usually somewhat pear-shaped, rounded in front and tapering towards the GENEEAL ANATOMY. 3 hinder extremity, convex on the dorsal and flattened on the ventral surface. The degree of dorsal convexity, however, is variable, the most common form of the body being sub-cylindrical, but in many cases — notably amongst the Poecilostoma, Siphonostoma, and in some genera of the Harpacticidse (Zaus, Peltidium, For- cellidium, Idya, Scutellidmni) — the animal is markedly flattened, constituting a type of structure almost as widely distinct from the normal Copepoda as are the Isopoda from the Amphipoda, amongst the sessile-eyed Crustacea. But in the genera of Harpacticidae here referred to, the flattened form does not coincide with any deep-seated differences of structure, for, indeed, in some normal genera (Thalestris, Laophonte) we find certain species assuming this form, but retaining all the other characteristics of the genus to which they belong. The directly opposite type of structure, in which the animal is compressed laterally, occurs only in the curious genus Amymone, a group of rare occur- rence so far as our present knowledge extends, not having been noticed outside of the European area. The front of the body is covered by a mem- branous shield or carapace, which falls rather loosely over the bases of the limbs below, but behind is continuous with the coverings of the thoracic somites ; in front it is usually projected from the forehead in the middle line, forming a rostrum, short or entirely absent in some few cases, but often moderately long and curved. In the Calanidse it is usually long and slender, much curved, and furcate nearly to its base. Behind the rostrum is commonly BRITISH COPEPODA. placed the median eye, often single to all appearance, but really composed of two closely approximated lateral eyes, embedded in a mass of black or crimson pigment. The head and the first thoracic segment are usu- ally fused together, a fact which may be recognised by the position of the first pair of swimming feet (the first thoracic appendages), these limbs being generally fixed to the hinder part of the first body- segment, which is thus seen to be composed of all the cephalic, and the first of the thoracic, somites. In some cases a transverse indentation maybe noticed, probably a trace of the " cervical suture " which is so conspicuous a feature of the carapace in Crayfishes and Lobsters. Instances are frequent, however, in which the head is quite distinct from the thorax. Theoretically, the Copepoda, like other Crustacea, are composed of twenty or twenty-one somites (twenty-one according to most authors, but twenty if we follow Huxley, who does not look upon the telson or last abdominal segment as a true somite)! the entire series not being developed, how- ever, in any one animal. Sometimes one or many of the somites are suppressed, at other times several are united into one segment, the real nature of which is rendered evident by the attachment to it of several appendages, each pair indicating the position of an anchylosed somite. Probably in no case amongst the Copepoda can more than sixteen or seventeen somites be recognised by enumeration either of appendages or segments. The somitic appen- GENERAL ANATOMY. 5 dages or limbs, though very various in form and function, may all be reduced to three component parts — a basal portion or protopodite, which gives support to two branches, termed respectively endo- podite and exopodite. These parts are most clearly developed in the swimming-feet, which are distinctly made up of a peduncle and two branches, but in the mouth organs the same structure may be traced, though often modified to such an extent as to be obscure and difficult of recognition. The cephalon in the Copepoda is composed of six somites, its appendages being one pair of eyes, two pairs of antennge, one pair of mandibles, one pair of maxillas, and two pairs of foot-jaws ; the thorax consists of five somites, and has five pairs of appen- dages in the form of swimming-feet ; the abdomen has no appendages, but consists of five somites and terminates in a forked tail, which ought probably to be considered as a sixth somite ; in the female the first two abdominal somites are generally united, forming one large genital segment with a pair of vulvar apertures. In parasitic species the abdo- men is often very much, reduced in size, both as respects the number and bulk of its somites, and this is the case also in some genera which are only partially parasitic, as Corycaus, Acontiophorus, Arto- trogus and others. As regards the cephalic appendages, it must be noticed that the so-called two pairs of foot- jaws are in reality portions of one and the same somitic appendage, but having the appearance of perfectly distinct organs they have come to be con- 6 BRITISH COPEPODA. sidered as such, and their characters so taken note of by all systematists. The process of development, however, has been traced by Glaus, so as to leave no doubt as to the true nature of the organs. APPENDAGES OF THE HEAD. — The eyes in their simplest form — in Cyclops, for instance— appear as a red or black spot in the middle of the frontal region, directly over the brain, with which they are connected by a large nerve, the spot when closely examined being found to consist of two lateral eyes, closely approximated and embedded in a mass of pigment ; the visual part of the apparatus is composed of two refracting bodies, or crystalline cones, and when more highly developed may possess numerous lenses, so as to form something like a facetted cornea. In some cases the eyes are widely separated, and have between them, in the median line, a simple, globular, pedunculated eye (PontelUnce) ; in other cases, as in some Corycceidce, the median eye is very small, while the lateral eyes are large, destitute of pigment, and consist of simple highly refracting lenses. In some parasitic species only are the eyes entirely wanting. In Pleuromma there is a supplementary eye, consisting of lenses with black pigment matter, on one of the thoracic segments. The anterior antennce are usually large and conspicu- ous organs, rising from hollows in the front of the head on each side of the rostrum. They act in many species (Calanidce, Cyclopidce) as very powerful swim- ming organs, it being by their agency chiefly that these animals propel themselves through the water; the GENERAL ANATOMY. 7 motion is thus a succession of rapidly recurring jerks corresponding with the separate sweeps of the antennas. These organs vary greatly in length and in the cha- racter of their setose armature. In the Harpactinidce, as well as in most parasitic and semi-parasitic species, the length of the anterior antenna falls short of, or at any rate does not usually much exceed, that of the first division of the body (carapace), while in the Cyclopidce it often equals, or even exceeds, that of the cephalothorax. In many Galanidce the length of the antennae is still greater, not unfrequently exceeding — sometimes very much exceeding — that of the entire animal. There is, however, no instance of this kind among British species, nor, so far as I know, among any but distinctly pelagic forms. And it is remarkable that, with this extreme length of the antennae, there is usually asssociated a greatly in- creased development of the apical lash of swimming- set^ with which the organ is armed. Sometimes also, as, notably, in the pelagic genus Euchceta, the mar- ginal hairs, though few, become wonderfully long. The antennas have, however, other important functions besides those of locomotion ; some of the variously formed setae with which they are in most cases largely provided act, no doubt, as organs of special sensation— the more simple hairs, perhaps, as tactile, the flattened and club-shaped setae as olfactory organs. The flat- tened, ensiform kind of appendage is seen on the antennae of most of the Harpacticidce (e. g. Idyafurcata, Plate LXVII, fig. 2 a), and the club-shaped form is seen in great force in Isias clavipes (Plate VII, figs. 4, 5) as 8 BRITISH COPEPODA. well as in other Calanidce ; these last-named organs are always most largely developed in the males, and it is not unlikely that they are subsidiary sexual organs, pos- sibly endowed with a highly-developed sensuous faculty. Besides the forms of antennal appendages here noted other modifications exist, of which the special uses are at present unknown. The anterior antennas serve yet another essential purpose, being adapted in the males as clasping organs ; they, together with the fifth pair of feet, are in very many species specially fashioned so as to insure a firm grasp of the female. The number of joints found in the anterior antennas varies from five or six in some semi- parasitic species (Lichomolgus) and eight or nine in the Harpacticidce, to twenty-four or twenty- five in the Calanidce. In the females the antennas are always alike on both sides of the body, but in the males of many of the Calanidce the right antenna is modified for the purpose of clasping ; while in all other non- parasitic species the male antennas of both sides are specialized for that purpose. In the semi-parasitic Corycce-idce, Saphirinidce, &c., there is little or no sexual difference in the anterior antennas, the clasping function devolving on the posterior pair. The par- ticular structural adaptations differ in different cases ; in the Calanidce, as before stated, the right antenna only is differentiated, the alteration consisting in one of the joints not far from the apex being so articulated as to form a hinge, by means of which the distal portion of the limb can be flexed upon the basal portion. Above and below the hinge the inner margins of the GENERAL AXATOMY. 9 antennae are frequently armed with denticulated plates, giving a firmer grasp. A very remarkable instance of this structure exists in Gandace pectinata (Plate X, fig. 2), also in the genera Centropages, Pontella, Para- pontella, &c. ; other genera, such as Temora and Diaptomus (Plate VI, fig. 7), are provided with one or more strong spines in lieu of, or in addition to, the denticulated plates. These spines are situated at various points of the internal surface of the antenna above the hinge-joint, and the limb itself is more or less swollen in the same situation to give room for a powerful flexor muscle. In the Pontellince this enlarge- ment of the limb is excessive (Plate X A, fig. 2 ; Plate XI, fig. 2). In the males of many Calanidce, however, there is little or no difference of form between the antennae of the two sides ; the difference is very slight also in the Misophriidce. In most Cyclopidce there are found differences of a kind similar to those already described, but affecting equally both right and left antenna (Plate XVII, fig. 5 ; Plate XXII, fig. 16) ; there are here no denticulated plates and few spines, but the limb is distinctly geniculated near the base as well as near the apex, and the articulations of the terminal segments are likewise very mobile. Amongst the Notodelphyidce the structure of the male antennas is somewhat similar to that of the Cyclopidce, but by no means so well marked. In the Harpadicidce, where the anterior antenna are very much smaller in com- parison with the size of the animal, the hingement of the male organ is not quite so obvious, though still sometimes quite of Cyclopid type (Cantlwcamptus 10 BRITISH COPEPODA. minutus, Plate XLIV, fig. 3 ; Eobertsonia tennis, Plate XLI, fig. 3) ; the apex is, however, often strongly clawed (Longipedia, Plate XXXIV, fig. 2 ; Euterpe, Plate XL, fig. 2 ; Tachidius, Plate XXXVII, fig. 3, &c.), and very frequently the two or three joints at the proximal side of the claw are coalescent and greatly enlarged, forming a pyriform or subglobose swelling for the reception of strong muscular bands ; examples of this structure are seen in the genera Tachidius (Plate XXXVII, fig. 3), Harpacticus (Plate LXIV, figs. 2, 13), Jonesiella (Plate XL VIII, fig. 3), and in many others. The spinous and setose armature of the antennae is, as a general rule, more profuse in the males of all families ; a good illustration of this cha- racter is seen in the case of Longipedia (Plate XXXIV, figs. 2, 3). In Pcecilostoma and Siphonostoma the sexual distinctions of the anterior antennas are not strongly marked, consisting chiefly of imperfect hinge- ments or contractions of the articulating surfaces between various joints of the male organ. The posterior or second pair of antennce are generally much smaller than the anterior pair, and consist in most cases of two branches ; they seem to be used both as swimming and prehensile organs, and in para- sitic and semi-parasitic species are specialized for prehension much more decidedly than the anterior pair. The main branch consists usually of three or four joints, to the basal or second joint of which is attached a " secondary" or " inner" branch of smaller size, and composed of one or several joints. This branch is in some cases altogether absent ; in others GENERAL ANATOMY. 11 it is equal in size to the primary branch, and may be numerously jointed (Longipedia, Calanidce, Misoph- riidce). In Cyclopida} the secondary branch is want- ing, and in most of the Harpacticidce it is reduced to small dimensions or, in some few cases, is quite absent. The limb is generally curvate, dilated at the apex, and provided with numerous curved or geniculated setse at the extremities and over the margins of both branches. In the PcBcilostoma and Siphonostoma it consists of only one branch, which in the male of the Corycceidce (Plate LXXXIII, figs. 13, 14) is powerfully clawed, and is used chiefly as a prehensile and clasping organ. The mandible consists, in its fully developed form, of a masticating portion and a " palp ;" the first-named division is in the form of an elongated, more or less triangular plate, dilated at the distal extremity and cut up into a variable number of tooth-like processes, these being sometimes only slight serrations, some- times large and powerful. The palp is variously formed ; in the Calanidce, most of the Harpacticidce, and in many Cyclopidce, it is composed usually of a large basal joint, from which spring one or two small setif erous branches ; these branches are generally 1- or 2-jointed, but may be absent altogether. In the sub-families Porcellidiince and Idyince the setiferous portions of the organ are very largely developed, form- ing large laminated appendages, which are fringed with densely ciliated filaments (Plate LXVIII, fig. 4 ; Plate LXIX, fig. 11). In the genus Cyclops, while the biting part of the mandible is well developed, 12 BRITISH COPEPODA. the palp is atrophied, and is represented only by two or three ciliated setee. In the Corycceidcv the mandible is small and weak, its palp obsolete or reduced to very small dimensions, while in some Siphonostoma* the mandible itself is converted into a long and slender piercing style, which is enclosed in a tube resembling considerably the antlia of Lepi- dopterous insects, and composed of prolongations of the upper and lower lip (Plate XCIII, fig. 3) ; in Cyclopicera, however (a genus here included amongst Siphonostoma), there is an intermediate condition of things ; the mandible is very much elon- gated, slender, and finely toothed at the apex, being, in fact, almost stylet-shaped, but not enclosed in any sheath; the palp, also, is quite rudimentary (Plate LXXXIX, fig. 4). The maxillce are small appendages, composed of a chewing portion, which is divided at the apex into numerous rather long and slender curved teeth, and of a complex, lobed, and setiferous palp, which frequently has filamentous appendages, possibly of a branchial character. In Cyclops the maxillae are very small in all their parts ; in the Calanidce, on the contrary, they are largely developed and possess numerous plumose filaments (Plate I, fig. 5) ; in the Notodelphyidce, also, the filamentous appendages are large (e.g. Plate XXIX, fig. 5). The Harpacticidce * The Pomologies of the mouth-organs in this group have been dis- cussed by Glaus, Thorell, Buchholz, and others. The subject is not without difficulty, and is treated under the description of the Siphono- stoma (p. 26) at greater length than would be suitable in this prefatory notice. GENERAL ANATOMY. 13 have the maxillas very small, but numerously divided ; there is usually a chewing portion pretty strongly toothed, and a palp which is divided into two or three setiferous digits, and has likewise two lateral (? branchial) offshoots (Plate LYII, fig. 2, Plate LX, fig. 6, Plate LXII, fig. 5, &c.), but this arrangement is subject to endless variation as to the number and development of the various parts. In parasitic species the maxillae, though usually present, are often atro- phied, and in the semi-parasitic forms belonging to the groups Pcecilostcma and Siphonostoma, are ex- tremely minute, and sometimes attached to the base of the mandibles. The anterior and posterior foot-jaws (which are in reality only the exo- and endo-podites of a single somitic appendage) do not present features requiring any lengthened description. They are in all cases adapted for prehension. In the Calanidce, Cyclopidce, and Notodelphyidce, the two pairs are not very dissimilar in structure, consisting generally of from four to six joints, which are in most cases marginally produced into digitiform or wart-like eminences on their inner side, and are more or less densely setiferous. In the Harpacticidce, while the first foot-jaw is like those of the preceding families, the second is usually in the form of a strongly-clawed hand, and in some sub- families (Tdyince, &c.) both pairs assume this cha- racter; such, also, is the case in the semi-parasitic Pcecilostoma and Siphonostoma. The subfamily Longi- pediince, unlike the rest of the Harpacticidce, has the posterior foot-jaw destitute of a clawed hand, those 14 BRITISH COPEPODA. organs in Ectinosoma being excessively slender, and almost linear in form. THE APPENDAGES OF THE THOEAX are, in their simplest form, five pairs of swimming-feet, each foot consisting of a 2-jointed base and two 3-jointed branches, but the number of joints, though never exceeding three, is within that limit, subject to a good deal of variation. The second, third, and fourth pairs are in almost all cases constructed as simple swimming-feet, without any other function, but the first pair, not unfrequently (as in most Harpacticidce) has one of its branches con- verted into a clawed prehensile limb, and the fifth pair shows very extensive modification in almost every case, often differing very considerably in the two sexes, and sometimes being very much reduced in size, or even altogether wanting. Though, as has been already said, the second, third, and fourth pairs of feet are constructed simply as swimming organs, and present no peculiarities of form, there are some exceptions to that statement. In Metridia armata the inner branch of the second foot in the male has a very remarkably excavated notch with spinous margin (PI. LVI, fig. 20 &), and some foreign species, belong- ing, probably, to two distinct genera, Metridia and Undina, exhibit a similar structure,* sometimes in both sexes. Again, in the genus Harpacticus the inner branch of the second foot of the male has the middle joint produced downwards into a strong spine, which varies in character according to species, but, in some * These species will be described in the ' Report on the Copepoda ' taken during the voyage of H. M.S. " Challenger." GENERAL ANATOMY. 15 shape or other, is always present (PI. LXIV, figs. 7, 16; PI. LXV, fig. 11). The third foot of the male in the same genus is sometimes (perhaps not always) con- verted into a clasping organ, the outer branch being bent across the inner and having its last joint armed with several strong spines (PI. LXIV, fig. 20). In the Calanidce and Misophriidce the fifth pair of the male is usually specially adapted as a clasping organ, the limb of one or both sides being reduced to a single branch, and provided with an armature of spines or hooks, which either entirely supersedes the swimming function, as in Temora, or is superadded, as in Centropages; but in other genera belonging to these families the sexual alteration of the limb is not very great (Calanus). Some pelagic genera, which are not represented in the British seas (Euchceta, Undina}, though possessing in the male a strongly developed prehensile fifth foot, have in the female only four pairs of simple swimming- feet. In Undina the male fifth foot is remarkably long and very fantastic in shape, reaching sometimes even beyond the extremity of the caudal segments. The organ may not unfrequently be seen with sperma- tophores adherent to its apex,* and is possibly used as the means of conveying these bodies to the vulva of the female. f It is remarkable, too, that in species so constituted (especially in Undina Darwinii, Lubbock) the spermatophores are very commonly found affixed in a futile manner to the back of the thoracic rings of * A similar condition is figured by Dr. Claus in the case of Euchata prestandrece (' Die frei lebenden Copepoden,' pi. xxx, fig. 9). f A similar function is performed by tbe maxillary palps (chela?) of male spiders and by the hectocotylised arms of some cuttle-fishes. 16 BRITISH COPEPODA. the female. In the Cyclopidce and Notodelphyidce the fifth feet are usually rudimentary and alike in both sexes, and in the HarpacticidcR they take the form of small, marginally setose, foliated expansions, slightly different in the two sexes, but generally larger in the female, in which sex they serve sometimes as a covering and support for the external ovisacs. In the semi-parasitic groups these organs are generally small, 1- or 2-jointed, and alike in both sexes. THE NEEVOUS SYSTEM of the Copepoda is described as consisting of a brain, which gives off various sensory nerves, a sub-oesophageal ganglion, and a ventral nerve cord, on which are situated ganglionic enlarge- ments ; the antennary nerves are also thickened, forming ganglionic rings. From an investigation of the nervous system of Cyclops, Mr. Marcus M. Hartog, F.L.S., of Owen's College, Manchester, has recently made out some other points which he kindly allows me to insert here. The most important of these are, that " gangli- onic swellings are found near the terminations of all sensory nerve fibres ; that the ventral nerve cord gives off at the end of the third segment of the body a pair of superficial cutaneous nerves, and at the fourth segment two pairs, one to the rudimentary fifth legs and another to two ventral muscles which rise from the sternal portion of the fifth segment. In the first abdominal segment is the fork described by Glaus and Leydig, but this takes its origin from the superficial (ventral) aspect of the cord which is continued onwards under the colleterial gland. After GENERAL ANATOMY. 17 running obliquely outwards each branch of the fork subdivides into two, an anterior and a posterior branch, both sensory. At the commencement of the third abdominal segment the ventral cord forks, its branches diverge slightly in this segment, but more in the next, rising to the sides of the intestine, and having the ventral muscles of this segment superficial to them. In the last segment they have left the intestine and run about the horizontal median plane straight into the axis of either branch of the f urea. " The ventral cord in Cyclops is not differentiated into distinct ganglia up to the second free (third) thoracic segment : beyond this is an enlargement con- taining ganglion cells at the posterior end of the fourth, and another (very small) in the last thoracic segment." THE ORGANS OF SENSE, so far as they exist in the tactile and olfactory rods of the antenna?, have already received brief notice. Mr. Hartog has recently de- scribed certain vesicles in the frontal region of Cyclops, and others attached to the bases of the fifth pair of feet and seated on a ganglionic enlargement of the nerve supplying the feet, which vesicles he believes to be auditory organs. In the male they contain one or more highly refracting bodies floating freely in the interior. Claus found a pair of these vesicles in the brain of Calanella, and has figured them in * Die frei lebenden Copepoden ' (plate vii, fig. 9). THE DIGESTIVE CANAL has a short, straight gullet, a large stomach, often with two ca3cal tubes, and an intestine opening on the dorsal aspect of the last (or VOL. in. B 18 BE1TTSH COI'EPODA. last but one ?) abdominal segment. The hinder por- tion of the alimentary canal is perhaps also uriniferous, but there exist near the bases of the foot-jaws other glandular organs, which may have a renal function. In the males of parasitic Copepoda the digestive canal disappears entirely. RESPIRATION. — If we are right in assigning to certain appendages of the mouth-apparatus (to which refer- ence has been already made; see pp. 10, 11 *) a branchial function, then we cannot altogether assent to the commonly - received belief that respiration in the Copepoda is carried on entirely by the dermic and intestinal surfaces of the body, without the interven- tion of any specialized respiratory apparatus. A sub- rhythmic contraction of the hinder extremity of the gut has, however, been noticed by Mr. Hartog in several Copepoda (Cyclops, Diaptomus, Canthocamp- tus), and by several observers in other Crustacea (e. g. Astacus, Limnadia, Daphma) : this is, no doubt, a respiratory movement. CIRCULATION. — In many Copepoda no special circu- latory organs have been found ; but in some there is a tubular heart, situated in the last thoracic segment, which drives forward the blood by a short vessel to the brain and anterior parts of the body, the blood * There can be no doubt that these setiferous plates are homologous with the structures called in the Ostracoda branchial lamince by G. O. Sars. But they do not appear to have any special internal circulation of the vital fluids, and if their function be branchial, they must act only by propelling waves of aerated water over the neighbouring surfaces. It is, pei'haps, on the whole, most probable that the currents produced by these ciliated appendages are subsidiary to nutrition rather than to respiration. GENERAL ANATOMY. 19 then passing through lacunae scattered throughout the tissues of the animal, and finding its way back to the heart, which it enters by slits in the walls of that organ. REPRODUCTION. — The sexes in the Copepoda are always separate, sexual differences showing themselves even externally in the form and structure of the body ; in some, especially in parasitic species, the dimorphism is most remarkable, the male becoming little more than a motionless sperm-sac attached to the body of the female ; but in the species which come within the scope of this memoir the males are usually smaller, more active, and less numerous than the females, the chief external distinctions being found in the almost constant conversion of the anterior antennae — less constantly of the fifth pair of feet, and occasionally also of the posterior antennas and foot-jaws — into clasping organs. The ovaries and testes are placed in the middle or in the sides of the cephalothorax, com- municate with accessory glands, and have efferent canals, which open by distinct apertures on the sides of the first (or conjoined first and second) abdominal somites. The efferent canal of the ovary may be simple, or may give off laterally a number of pouches, which hold the eggs (Corycceidce), while in some para- sitic species it forms several terminal coils, in which the eggs are detained; in the Notodelphyidce the duct is converted into a large dorsal pouch or pseudo-uterus, covered only by the integument, in which organs the ova undergo partial development. In the free Cope- poda, however, the ova pass at once into two (often coa- 20 BRITISH COPEPODA. lescent) external ovisacs attached to the first abdominal segment the coating of the ovisac being formed by the secretion of a special gland, situated near the termina- tion of the efferent duct, an enlargement of which forms in many cases a " receptaculum seminis." In the males the free-living forms have a simple testis ; many of the parasitic and semi-parasitic (Corycceidce, Sapphirinidce) a double testis, with two distinct vasa deferentia, the right duct being sometimes atrophied. In the coiled portion of the duct are formed the spermatophores — masses of spermatozoids enclosed in a capsule of hardened mucus, and usually fusi- form or club-shaped. During copulation the male affixes one or more of these bodies near the vulvar aperture of the female, the contents passing into the receptaculum seminis, and fertilizing the ova either in the interior of the body or during their passage into the ovisacs. In some cases the seminal fluid appears to be inserted directly into the vulva without the intervention of a spermatophore. THE DEVELOPMENT of the free Copepoda from the moment of rupture of the ovum to the attainment of matured form presents a complex series of metamor- phoses. The parasitic species present some of the best marked examples of " retrograde development" to be found in the whole animal kingdom, but these do not come within the limits of our present subject. The form of the young Copepod on its escape from the egg is that known as NaupUus, having been de- scribed by Muller under that name before its relation to the Copepoda was known. The larva in this GENERAL ANATOMY. 21 stage is oval, has a single frontal eye, three pairs of limbs arranged round the mouth, and no frontal appendages ; the mouth-organs proper are entirely absent, and the posterior part of the body has no appendages except a couple of setae in the neighbour- hood of the anus. The anterior portion of the body is equivalent to the three anterior cephalic somites, its three pairs of limbs becoming eventually antennae and mandibles. At the first moult the body becomes elongated and new limbs appear in the following order : — a fourth and fifth pair representing respec- tively the maxillae and foot-jaws ; a sixth and seventh which become the two anterior pairs of swimming- feet. At this stage the larva still resembles a Nauplius, and does not take on a Cyclopoid appear- ance until after the next moult.* It then resembles more closely the adult Cyclops as to the antennae and mouth-organs, but the number of limbs and somites is smaller ; the body in this condition is composed of an oval cephalothorax, three thoracic and one long terminal segment, which in succeeding moults be- comes forked. In the Cyclopidce the posterior antennae and the mandibles lose their accessory branches, but in other families these parts are usually retained. All the free, and many of the parasitic species pass through a further series of moults, in the course of which the still-wanting limbs and body- segments appear, the limbs attaining, by successive * In some Nauplii, if not in all, the terminal part of the intestine is subglobular, and contracts periodically like the " contractile vesicle " of a Rotifer. 22 BRITISH COPEPODA. steps, their full number of joints and perfect develop- ment.* Those parasites which miss the Nauplius stage are hatched in Cyclops-form ; many of the retrogressive species become fixed to some animal, segmentation is lost, limbs and eyes disappear or become atrophied ; the males, often dwarfed, being permanently fixed near the sexual apertures of the female. * For details of this process in the genus Cyclops, see vol. i, p. 100. NOTE. CHANGE OF GENERIC NAMES. The generic name LopJwphorus (vol. i, p. 121), having been previously used to designate a genus of Phasianidse, must be withdrawn. I therefore propose to substitute the word Pterinopsyllus Cylindrosoma (in Table of Genera, vol. i, p. 31) is for a like reason discarded for Cylindropsyllus, and Solenostoma (loc. cit.} for Acontiophorus. BEITISH COPEPODA. Section II. — PCECILOSTOMA, Thorell.* THORELL'S division of the Copepoda into three groups, G-nathostoma, Poecilostoma, and Siphonostoma, the distinctions between which are found in the characters of the mouth-organs, is disapproved by Glaus and some other authors, chiefly, as I understand them, on the ground of the gradual lapse of one series into the other rendering it impossible to draw perfect lines of demarcation, but partly, also, on the ground of a difference of interpretation of the homologies of some of the appendages. While differing from M. Thorell as to the nature of some of these organs, I myself think that his proposed division is a very natural one, the three groups presenting characters which, though differing in degree in various species, do point, on the whole, to habits of life very remarkably different, and deserving of expression in any natural classification. The three groups are denned by M. Thorell as follows : Series 1. GNATHOSTOMA. Os mandibulis duabus libens tribusque paribus maxillarum instructum, siphone nullo. * See p. 31, vol. i. 26 BRITISH COFEPODA. Series 2. PGEOTLOSTOMA. Os mandibulis et siphone carens, maxillarum paribus 3 — 1 ( — 0) instructum. Series 3. SIPHONOSTOMA. Os in siphonem, mandibulas 2 plerumque inclu- dentem, productum, et maxillarum paribus 3 — 0 in- structum. As regards the debateable anatomical points, it may be useful if I quote, in the first place, some remarks of M. Thorell, taken from a letter which he was good enough to address to me some few years ago — before the publication of Dr. Claus's 'Neue Beitrage zur Kenntniss parasitischer Copepoden.' M. Thorell writes as follows : — " You know, of course, that I have pro- posed to divide the Copepoda into three parallel series, Grnathostoma, Poecilostoma, and Siphonostoma, and that I consider the Poecilostoma to be characterized by having the parts of the mouth free, and formed for stinging or licking, as also by the mandible being absent. This view of the oral apparatus of the Psecil- ostoma has been accepted by Claparede and a few others, but it is not admitted by, for instance, Glaus, who considers that the Poecilostoma have true man- dibles, and who rejects the subdivision proposed by me. The reasons which induced me to believe that the Paacilostoma were destitute of mandibles, and that what Glaus calls mandibles are the true maxillge, and that his maxillas are maxillar-palpi, were — first, that the " mandibles " are always placed far more back- wards than the true mandibles of the Gnathostoma; P(ECILOSTOMA. 27 secondly, that I sometimes, as, for instance, in Licho- molgus albens, found a small longitudinal, semi-pipe- formed depression or groove exactly at the place where the sipho of, for instance, Dyspontius and Ascomyzon is inserted, and which I therefore considered to indicate the place where the sipho and mandibles ought to be found if any mandibles existed ; and thirdly and chiefly, that sometimes, as in the genus Lichomolgus, the so- called ' maxillae ' are fixed on the ' mandibles ' (quite as an ordinary palpus is fixed on a mandible or a maxilla), and directed from the oral aperture, a cir- cumstance with which I could find nothing analogous in the class Crustacea, supposing Glaus' ' mandibles ' really to be mandibles." The greatest difficulty which besets the discussion of this question is the minuteness of the mouth-organs in these animals, and the liability to displacement or mutilation of the various parts in conducting a dissec- tion, so that the organs of one and the same species will often present very different appearances in differ- ent preparations of the animal. There can be no doubt, however, that the fact so strongly insisted on by M. Thorell, — that of the coalescence, in Lichomolgus, of the maxilla and mandible (or maxilla and palp) — does really exist : the question remains, What is this palp- like organ ? In appearance it is not unlike the poorly- developed mandible- or maxilla-palp of many Grnathos- toma, but it is also much like a single branch of such a maxilla as we find in the genus Cyclopicera or Arto- trogus, so that not much can be learned by comparison of structure only. The point next to be considered is 28 BRITISH COPEPODA. the nature of the appendages, — by Thorell called the first pair of maxillae, by Glaus the mandibles, — to which the palp-like organ is attached. In considering this question we shall do well to take a somewhat wider survey than merely of the order Copepoda. Among the nearly related order Ostracoda, for the most part consisting of true Gnathostomous Crustacea, we find a group, — including chiefly the genus Paradoxostoma, — in which the mouth is modified for suctorial purposes in a manner at once reminding one of the siphonostomous Copepoda. In Paradoxostoma the tubular mouth is formed by the coalescence of the labrum and labium, and the man- dible assumes the form of a stilet, having a very slender filiform palp, the almost exact counterpart of the same organs in A.contioplwrus , Cydopicera, &c. (see PI. LXXXIX, fig. 4, and PL XC, fig. 4). There can be no doubt, I think, that in the well-marked siphonostomous Copepoda, such as Acontiophorus, Dys- pontius, and Artotrogus (Ascomyzon), the tubular mouth is formed, as in Paradoxostoma, by the union of the upper and lower lips, and that the filiform organs lying immediately by the side of the siphon (see PI. XC, fig. 1 c, and PI. XCI, fig. 6 c c) are modified man- dibles and palps; in Artotrogus, indeed (PL XCIII, fig. 3 b &'), we find this stilet-shaped mandible dis- tinctly toothed at its apex. In the genus Cydopicera, of which I have fortunately collected and examined many specimens with great care, all the mouth- organs are largely developed; there is an unmistak- able mandible with a well-developed palp, a distinct PCECILOSTOMA. 29 2-branched maxilla, and a- stout proboscidiform suc- torial mouth (PI. LXXXIX, figs. 4, 5, 6). And it can scarcely be doubted that the two-branched organ shown in Plate LXXXVII, fig. 10 (Acontiophoi*us armatus), is homologous with the mandible and palp of Cyclopicera nigripes represented in* Plate LXXXIX, fig. 4, and of G. gracilicauda (Plate LXXXIII, fig. 3). To recur to the genus Lichomolgus. If we examine again the disputed organ (PI. LXXXV, figs. 4, 12, PL LXXXVII, fig. 3, PL LXXXVIII, fig. 11) we shall find that it bears a very strong structural resemblance to those appendages of siphonostomous genera which have just occupied our attention, and the mandibular nature of which is, I think, pretty conclusively shown. I have therefore little doubt that this organ in Licho- molgus ought to pass for a mandible, and inasmuch as the mandibular palp amongst the Siphonostoma is sometimes nearly or quite suppressed, but the maxilla never, and, moreover, as when the mandible-palp does exist it assumes a form totally different from the palp- like appendage of the mandible of Lichomolgus; for these reasons I am. disposed to regard this appendage as a rudimentary maxilla. The opinion is confirmed by a comparison of the maxilla of Gorycceus, which is connected with the mandible in a manner very like that of the supposed maxilla of Lichomolgus to its mandible (see Plate LXXXIV, fig. 10). 30 BRITISH COPEPODA. [Genus CYLINDROPSYLLUS, nov. gen. Animal cylindrical, much elongated ; head united with the thorax and having a sharp rostrum ; abdomen 4-jointed, as wide as the thorax, and not distinctly separated from it. Anterior antennas short, 5-jointed ; posterior 2-jointed, destitute of a secondary branch. Posterior (?) foot-jaw small, provided with an apical curved spine and several marginal setas (rest of the mouth-organs unknown). First four pairs of swim- ming-feet having the outer branch B-, the inner 2- jointed; branches of the first pair very short and nearly equal ; inner branch of the second, third, and fourth pairs very short ; fifth pair rudimentary, 1 -jointed, foliaceous. In the Table of Classification (vol. i, p. 31) this genus is given as Cylindrosoma ; which name, however, being already in use, is here altered to Cylindropsyllus. Though given in the table amongst Harpacticidse, it more probably belongs to the Pcscilostoma. For the present I content myself simply with a description of the species, without attempting to assign it to any recognised family. 1. CYLINDROPSYLLUS LMVIS, nov. sp. PI. LXXXIV, figs. 1—8. Animal of equal width throughout, the abdomen being as wide as the cephalothorax and not separated from it by any constriction; first body-segments as 31 long as the two next following ; rostrum as long as the first joint of the anterior antenna ; abdomen equal in length to the cephalothorax, and composed of four nearly equal joints. Anterior antenna (fig. 2) about as long as the first segment of the body, 5- jointed, joints nearly equal in length, except the fourth, which is much shorter than the rest; posterior antenna 2- jointed, the last joint bearing about eight apical seise. First pair of feet short; second and third (fig. 5) longer ; fourth (fig. 6) rather longer than the third ; the external branches of the second and third pairs bear long and slender marginal spines, but those of the fourth are very weak and small. The fifth pair (fig. 7) consists of one small subquadrate joint, which is fringed distally with five or six rather long seta?. The caudal segments are very small, about twice as long as broad, but scarcely more than one fourth the length of the last abdominal somite; the tail-setas (fig. 8) are three, two very short and one of consider- able length, the latter being sharply geniculated above the middle, and equalling in length the last three abdo- minal segments. Length -rgth of an inch (1'4 mm.). One specimen only of this remarkable Copepod has come under my notice. It was dredged off Hartlepool, in a depth of five fathoms, amongst muddy sand. Without more knowledge of the mouth-apparatus, it is impossible to assign to the species more than a provisional place, but it seems to me not unlikely that it may be found to be of parasitic or semi-parasitic habits. It is very similar, in general character, to the genus Ophthalmopachus, Hesse.] 32 BRITISH COPEPODA. Family 7. COEYC^ID^;, Thorell. Body composed of eleven or twelve segments, elongated or subpyriform ; abdomen elongated, much narrower than the cephalothorax ; head usually anchy- losed with the first thoracic segment. Anterior antennae alike in both sexes, 5 — 7-jointed ; posterior simple, 3- or 4-jointed, forming a prehensile hand, which is armed at the apex with a claw. Mouth- organs (except the second pair of foot-jaws) minute, and destitute, or nearly so, of palps. Posterior foot- jaw forming a prehensile organ, and, in the male, powerfully clawed. First four pairs of feet alike or nearly so, and adapted for swimming, 2-branched. Fifth pair of feet rudimentary, alike in both sexes, seldom altogether absent. Heart wanting. In addi- tion to the small median eye, there are usually two large lateral eyes, each composed of a single lens. The sexual organs, in both male and female, are double and symmetrical. Ovisacs usually two. The British genera belonging to this family are Corycceus, and Monstrilla. Genus 1. CORYG^EUS, Dana (1845). (Dana, Proc. Acad. Nat. Sc. Pkiladelph.) Body elongated, subcylindrical; abdomen 2-jointed, penultimate segment of the cephalothorax produced ventrally into two hook-like processes (PL LXXXIV, CORYOEUS. 33 fig. 14 a, and PI. LXXXIII, fig. 11 a a). Anterior antennae 6-jointed, short; posterior uncinate, power- fully prehensile, terminal claw much longer in the male than in the female. Mandibles (PI. LXXXIY, fig. 10) slender, divided into two apical teeth (a), and bearing a minute setiferous palp (6) ; maxilla (fig. lOc) composed of a single lamina, which bears several marginal Iacinia3, and near the middle a crescentic row of small setse (see also PI. LXXXIV, fig. 9). First pair of foot-jaws short and stout, alike in both sexes, apex produced into a stout hook-like spine, inner margin bearing several setas (fig. 11). Posterior foot-jaws elongated, 3-jointed, forming a strongly uncinate prehensile hand (PL LXXXI, figs, 16, 17), the claw very much larger in the male. First, second, and third pairs of feet having both branches 3-jointed, (PI. LXXXIY, fig. 12) ; inner branch of the fourth pair (fig. 13) rudimentary, 1 -jointed; fifth pair (PI. LXXXI, fig. 18), rudimentary, composed of a single small setiferous joint. Last joint of the cephalothorax (PL LXXXIV, fig. 14 b) very small and partially over- lapped by the penultimate joint. Frontal eyes two, each composed of a single large, colourless, highly refracting lens, situated near the bases of the anterior antenna3. o s oraei loe vis . 9JCopyccens furcifer. 10-14- ,, sm. g lie -as. ' PLATE LXXXV. Lichomolgus fucicolus. Fig. 1. Adult male. 2. Anterior antenna. 3. Posterior antenna. 4. &, mandible ; b, maxilla. 5. Anterior foot-jaw. 6. Posterior foot-jaw of female. 7. Posterior foot-jaw of male. 8. Foot of first pair. 9. Foot of fourth pair. 10. Abdomen of female ; a, fifth foot. 11 . Abdomen of male. Lichomolgus forficula. * 12. f Mandible. 13. Posterior foot-jaw of male. 14. Inner branch, fourth foot. 15. Foot of fifth pair. 16. Abdomen of male. * In the Plate, for forcicula read forficula. f In the Plate, for 12—18 read 12—16. Plate 85. Tff.Wtst*C? imp- . omolg-us facie olus. forci:f ulsu PLATE LXXXYI. Lichomolgus liber. Fig. 1. Adult male. 2. Anterior antenna. 3. Posterior antenna. 4. Mandible. 5. Anterior foot-jaw. 6. Posterior foot-jaw of female. 7. Posterior foot-jaw of male. 8. Foot of first pair. 9. Foot of second pair. 10. Foot of fourth pair. 11. Foot of fifth pair. 12. Abdomen of female. 13. Abdomen of male. Lichomolgus forficula. 14. Anterior antenna. 15. Posterior antenna. 16. Anterior foot-jaw. 17. Posterior foot- jaw of female. 18. Abdomen of female. * u? imp. 1 — 13 LiclioTnolgixs liter. 141"! 8 forficula,. PLATE LXXXVII. Lichomolgus arenicolus. Fig. 1. Adult male. 2. Anterior antenna, female. 3. Mandible; a, maxilla (?). 4. Anterior foot-jaw, 5. Posterior foot-jaw of male. 6. Foot of first pair. 7. Foot of fifth pair. Acontiophorus armatus.* 8. Anterior antenna. 9. Posterior antenna. 10. Mandible. 11. Maxilla. 12. First foot- jaw. 13. Second foot- jaw. 14. Siphon. 15. Abdomen and fifth foot. * In the inscription of the Plate, for Solenostoma armatum read Acontiophorus armatus. 13 1 — 1 Lac]ao:m.olg-u.s arenicolus. 8— 15 SolenostoxnaL a,zixaa,'tum. WWestkCe.ivy PLATE LXXXYIII. Lichomolgus Thorellii. Fig. 1. Anterior antenna. 2. Posterior antenna. 3. Maxilla. 4. Anterior foot- jaw. 5. Posterior foot-jaw of female. 6. Posterior foot-jaw of male. 7. Foot of fourth pair. 8. Abdomen and tail of female. 9. Abdomen and tail of male. Lichomolgus furcillatus. 10.* Anterior antenna. . 11. Mandible. 12. Anterior foot-jaw. 13. Foot of first pair. 14. Abdomen and tail ; a fifth foot. * In the inscription of the Plate, for figs. 1 — 14 read 10—14. 1 — 5 Tlaorellii. fun?eillatas . Wffat&Ct.tmp PLATE LXXXIX. Cyclopicera nigripes. Fig. 1. Female seen from below. a. Anterior antenna. 6. Posterior antenna. c. Mandible. d. Maxilla. e. Anterior foot- jaw. /. Posterior foot- jaw. g-k. 1st — 5th pairs of feet. I. Labrum. 2. Anterior antenna of female. 3. Posterior antenna. 4. Mandible. 5. Maxilla. 6. Labrum. 7. Anterior foot-jaw. 8. Posterior foot-jaw. 9. Foot of second pair. 10. Foot of fifth pair. 11. Abdomen and tail of male. Cyclopicera lata. 12. Abdomen of female, Plate 89. '•2 GSBraJy Ad. ,A tith-. 1— H,Cy clop ic era TII gripes latsL. PLATE XC. Acontiophorus scutatus* (female). Fig. 1. Female seen from below, a. Anterior antenna. b. Posterior antenna. c. Mandible. d. Maxilla. e. Anterior foot- jaw. /. Posterior foot-jaw. s. Siphon. 2. Anterior antenna. 3. Posterior antenna. 4. Mandible. 5. Maxilla. 6. Siphon. 7. Anterior foot- jaw. 8. Posterior foot-jaw. 9. Foot of third pair. 10. Foot of fifth pair. Gyclopicera lata. 11. Anterior antenna of female. 12. Anterior antenna of male. 13. Posterior antenna. 14. Maxilla. * In the inscription of the Plate, for Solenostoma scutatum read Acontiophorus scutatus. Plate 90 1" 10, SolerLOstoTna. scru-tatimi? 1H.41, Cyclopic era, . lata, . PLATE XCI. Artotrogus Boeckii* (female). Fig. 1. Anterior antenna. 2. Posterior antenna. 3. Mandible. 4. Maxilla. 5. Posterior foot-jaw. 6. First segment of body. a. Anterior antenna. b. Posterior antenna. cc. Mandibles. dd. Maxillse. ee,ff. First and second foot- jaws. g. Siphon. 7. Foot of third pair. 8. Abdomen ; a, fifth foot. 9. Fifth foot, var. Gyclopina gracilis (male). 10. Abdomen and fifth foot. 11. Anterior antenna. Artotrogus Normani (male). 12. Anterior antenna. 13. Posterior antenna. 14. Second foot-jaw. 15. Foot of third pair. * In the inscription of the Plate, for Artotrogus lAlljcborgii read Artotrogus BoecJcii. Plate 91. 1—& Artotrog-us Lillj eJoorgii 10,11 Cyclopina graciliso. 12-15 Art otrogiis PLATE XCII. Dyspontius striatus. Fig. 1. Adult male. 2. Anterior antenna of female. 3. Anterior antenna of male. 4. Posterior antenna. 5. Maxilla. 6. Anterior foot- jaw. 7. Posterior foot-jaw. 8. Foot of third pair. 9. Foot of fourth pair. 10. Appendage of first abdominal segment, male. 11. Terminal spines of swimming-foot. 12. Abdomen of female. 13. Shell-structure. Artotrogus Nbrmani. 14. Shell- structure. Pleute 9 2 1 — ISDysp orituxs striatuis. ll' Artotrogizs Normani. PLATE XCIII. Artotrogus magniceps (female). Fig. 1. Adult female. 2. Anterior antenna. 3. Siphon. a a. Extremities of siphonal limbs. a. Same more highly magnified. b. Mandible. b'. Extremity of same more highly magni- fied. 4. Maxilla. 5. First foot-jaw. 6. Second foot-jaw. 7. Foot of fourth pair. 8. Foot of fifth pair. 9. Shell-structure. Artotrogus Normani. 10. Adult male. I 10 INSTITUTED 1844. FOR THE PUBLICATION OF WORKS ON NATURAL HISTORY. ANNUAL SUBSCRIPTION ONE GUINEA. 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Colman, Jeremiah J., Esq., M.P., Carrow House, Norwich. Cooke, Benjamin, Esq., 103, Windsor Road, Southport. Cooper, Colonel E. H., 42, Portman square, W. Cooper, Sir Daniel, Bart., 6, De Vere gardens, Kensington Palace, W. Coppin, John, Esq., Kingfield House, by Corbridge-on-Tyne, R.S.O. Cork, Queen's College, Cork. Cornwall, Royal Institution of, Truro. Craven, Alfred E., Esq., 36, Princes Gate, S.W. 8 Cresswell, Rev. R., Teignmouth, Devon. Croft, R. Benyon, Esq., R.N., F.L.S.,, Farnham Hall, Ware, Herts. Crowley, Philip, Esq., Waddeu House, Croydon, S. Cruickshank, Alexander, Esq., 12, Rose street, Aberdeen. Cunningham, Professor R. 0., Local Secretary, Queen's College, Belfast. Currey, Fred., Esq., F.R.S., Sec. L.S., 3, New square, Lincoln's inn, W.C. Curtis, William, Esq., Alton, Hants. Darwin, C., Esq., LL.D., F.R.S., Down, Kent. Dawson, Professor J. W., F.R.S., F.G.S., M'Gill College, Montreal. Deane, Jas., Esq., F.L.S., 17, The Pavement, Clapham, S.W. Devon and Exeter Institution, Exeter. Devonshire, Duke of, 78, Piccadilly, W. Dickie, Prof. G., M.D., F.L.S., 16, Albyn terrace, Aberdeen. Dickinson, Wm., Esq., Jun., Warham road, Croydon. Dickson, Professor Alexander, Glasgow. Dohrn, Dr. Anton, Naples. Douglas, J. "W., Esq., 8, Beaufort gardens, Lewisham, S.E. Douglas, Rev. R. C., Manaton Rectory^Moreton Hampstead, Exeter. Douglas, W. D. R., Esq., Orchardton, Castle Douglas, N.B. Drewitt, Drewitt 0., Esq., Jarrow Hall, Newcastle-on-Tyne. Drosier, Dr. W. H., Cambridge. Dublin, National Library. Dublin, Royal Irish Academy. Dublin, Trinity College. Dublin, College of Surgeons. Dublin, Hon. Society of King's Inn. Ducie, Earl of, F.R.S., F.G.S., 16, Portman square, W. Dunning, J. W., Esq., M.A., F.L.S., 24, Old buildings, Lincoln's inn, W.C. East Kent Natural History Society, Canterbury. Edge-worth, M. P., Esq., F.L.S., 6, Notham gardens, Oxford. Edinburgh College of Physicians. Edinburgh, Library of University of. Edinburgh Museum of Science and Art. Edinburgh, Royal Society of. Edinburgh Royal Physical Society. Edinburgh Royal Medical Society. Egerton, Sir P. de M. Grey, Bart., M.P.,F.R.S., G.S., President, 28s, Albemarle street, W., and Oulton park, Tarporley, Cheshire. Elliot, Sir W., F.L.S., Hawick, Roxburgshire. Elphinstone, H. W., Esq., F.L.S., 2, Stone Buildings, Lincoln's Inn, W.C. England, Royal College of Surgeons of, Lincoln's-inn-fields, W.C. England, Bank of, Library, London, E.G. Enniskillen, the Earl of, D.C.L., F.R.S., F.G.S., 65, Eaton place, S.W. Eyton, T. C., Esq., F.G.S., F.L.S., Eyton, Wellington, Salop. Ferguson, W., Esq., F.L.S., F.G.S., Kinmundy House, near Mintlaw, Aberdeenshire. Ffarington, Miss M. H., Worden HaH, near Preston. Fitch, Fred., Esq., F.R.G.S., Hadleigh House, Highbury New Park, N. Flower, J., Esq., M.A , F.Z.S., Fairfield road, Croydon, S. Flower, W. H., Esq., F.R.S., Royal College of Surgeons, W.C. Ford, J., Esq., 1, Market street, Wolverhampton. Foster, C., Esq., Thorpe, Norwich. Fowler, Rev. W. W., Repton. Fox, Rev. W. D., Broadlands, Sandown, Isle of Wight. Friedlander & Son, Messrs., Berlin. Fuller, Rev. A., Itchenor, near Chichester. Galton, Capt. Douglas, F.R.S., F.L.S., 12, Chester street, Grosvenor place, S.W. Garner, Robert, Esq., F.L.S., Stoke-upon-Trent. Garneys, W., Repton, Burton-on-Trent. Gatty, C. H., Esq., F.L.S., F.G.S., Felbridge Park, East Grinstead, Sussex. Geological Society, London, W.C. Geological Survey of India, Calcutta. George, Frederick, Esq., 10, Finchley road, St. John's wood, N.W. Gerold and Sons, Messrs., Vienna. Gibson, G. S., Esq., F.L.S., Saffron Walden, Essex. Glasgow Philosophical Society, Glasgow. Glasgow University, Glasgow. 10 Godman, F. D., Esq., F.L.S., 10, Chandos street, Cavendish square. Goode, J. F., Esq., 3, Regent place, Birmingham. Gordon, Rev. George, LL.D., Manse of Birnie, by Elgin, N.B. Gore, R. T., Esq., 6, Queen square, Bath. Gottingen, University of, Gottingen. Gould, John, Esq., F.R.S., L.S., 26, Charlotte street, Bedford square, W.C. Gray, Wm., Esq., 75, Petergate, York. Graham, W., Esq., F.R.M.S., Ludgate hill, Birmingham. Green, R. Y., Esq., Newcastle-on-Tyne. Grut, Ferdinand, Esq., 9, King street, Southwark, S.E. Giinther, Dr., F.R.S., British Museum, W.C. Hackney Microscopical and Natural History Society, 1 94, Mare street, Hackney, E. Haeckel, Professor, Jena, Prussia. Hailstone, Edward, Esq., F.S.A., Walton Hall, Wakefield. Hamburgh Town Library, Hamburgh. Hamilton, Dr. E., F.L.S., F.G.S., 9, Portugal street, Grosvenor square, W. Hancock, John, Esq., Newcastle-on-Tyne. Harford, F., Esq., Ocean Marine Insurance Company, 2, Old Broad street, B.C. Harmer, F. W., Esq., Local Secretary, Oakland House, Cringleford, Norwich. Harper, P. H., Esq., 30, Cambridge street, Hyde Park,.W. Harris, Edw., Esq., F.G.S., Rydal Villa, Longton Grove, Upper Sydenham. Harris, Dr. F., F.L.S., 24, Cavendish square, W. Harvey, Dr. J. R., 7, Upper Merrion street, Dublin. Harvard College, Cambridge, U.S. Hawkins, Dr. B. L., "Woburn, Beds. Hayek, Herr Gustav Edler von, Vienna. Hepburn, Sir T. B., Bart., Smeaton, Preston Kirk, N.B. Hey, Samuel, Esq., 36, Albion street, Leeds. Hicks, Dr. John B., F.R.S., 24, George street, Hanover square, W. Hicks, Dr. J. Sibley, 2, Erskiue Street, Liverpool. Hillier, J. T., Esq., 4, Chapel place, Eamsgate. Hilton, James, Esq., 60, Montagu square, W. 11 Hinde, R., Esq., Lancaster. Hoest, Dr., Copenhagen. Holdsworth, E. W. H., Esq., F.L.S., 84, Clifton hill, Abbey road, N.W. Hooker, Sir J., C.B., M.D., F.R.S., Kew, W. Hope, A. J. B., Esq., M.P., 1, Connaught place, W. Hopkinson, J., Esq., F.Z.S., F.G.S., Wandsford House, Watford. Houghton, Rev. W., F.L.S., Preston Rectory, Wellington, Salop. Hovenden, F., Esq., Glenlea, Thurlow Park, Dulwich, S.E. Howden, Dr. J. C., Sunnyside, Montrose. Howitt, Thomas, Esq., F.R.C.S.E., Local Secretary, Lancaster. Huddersfield Literary and Scientific Society. Huddersfield Naturalists' Society. Hudson, R., Esq., F.R.S., F.G.S., Clapham common, S.W. Hughes, W. R., Esq., F.L.S., Local Secretary, Thome Villa, Hands- worth, Birmingham. Hull Subscription Library. Humphry, Professor, F.R.S., Cambridge. Hunt, John, Esq., Milton of Campsie, Glasgow. Hutchinson, R., Esq., 29, Chester street, Edinburgh. Huxley, Professor T. H., F.E.S., Museum Practical Geology, Jermyn street, S.W. Indian Museum, Calcutta. Jeffreys, J. Gwyn, Dr. F.R.S., F.G.S., Ware Priory, Herts. Jenner, Charles, Esq., Easter Duddingsten Lodge, Portobello, Edin- burgh. Jordon, Dr. R. C. R., 35, Harborne road, Edgbaston, Birmingham. Kenderdine, F., Esq., Morningside, Old Trafford, Manchester. Kibbler, Dr. R. C., 61, King Edward road, Hackney, E. Kilraarnock Library, Kilmarnock. Kitson, J., Esq., Elmete Hall, Leeds. Knapp, A. J., Esq., Llanfoist House, Clifton, near Bristol. 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Swain, E., Esq., 34, Elsham road, Kensington, N. Swanston, W., Esq., F.G.S., 50, King street, Belfast. Tebbs, H. V., Esq., 15, Knight Rider street, Doctors' Commons, B.C. 16 Tennant, Professor James, F.G.S., 149, Strand, W.C. Thomson, Professor Allen, Glasgow. Thomson, Dr. Thomas, F.R.S., 6, Bower terrace, Maidstone. Thomson, Prof. Sir Wyville, F.R.S., Edinburgh. Toronto, University of, Canada. Torquay Natural History Society, Torquay. Townsend, F., Esq., M.A., Honington Hall, Shipston-on-Stour. Trail, Prof. W. H., M.B., Local Secretary, King's College, Old Aber- deen. Tristram, Rev. Canon H. B., LL.D., F.R.S., The College, Durham. Trubner & Co., Messrs., London. Tudor, Richard A., Esq., 21, Church View, Bootle, Liverpool. Turner, Professor W., F.R.S.E., Anatomical Museum, University of Edinburgh. Tyler, Captain Charles, F.L.S., F.G.S., 317, Holloway road, Hollo- way, N. Upsala, University of, Sweden. Varenne, E. G., Esq., Kelvedon, Essex. Vass, M., Leipzig. 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T., M.A., F.L.S., G.S., Secretary, 25, Granville park, Lewisham, London, S.E. Wollaston, G. H., Esq., 117, Pembroke road, Clifton, near Bristol. Wood, E., Esq., Richmond, Yorkshire. Woodd, B. T., Esq., Conyngham Hall, Knaresborough, Yorkshire. Wright, Professor E. P., Trinity College, Dubliu. Yale College, New Haven, U.S. Yeoman, T. P., Esq., 4, St. Hildas terrace, Whitby. York Philosophical Society, York. Young, Dr. J., College, Glasgow. Zoological Society, 11, Hanover square, W. LIST OF THE ANNUAL VOLUMES RAY SOCIETY. FKOM THEIE COMMENCEMENT, IN 1844, TO AUGUST, 1880. LIST OF THE ANNUAL VOLUMES ISSUED BY THE RAY SOCIETY. FOR THE FIRST YEAR, 1844. I. Reports on the Progress of Zoology and Botany. Trans- lated by H. E. Strickland, Jun., M.A., F.R.S., E. Lan- kester, M.D., F.R.S., and W. B. Macdonald, B.A. 8vo. II. Memorials of John Bay : consisting of the Life of John Ray, by Derham ; the Biographical Notice of Ray, by Baron Cuvier and M. Dupetit Thouars, in the ' Biographic Universelle ; * Life of Ray, by Sir J. E. Smith : the Itine- raries of Ray, with Notes, by Messrs. Babington and Yarrell. Edited by E. Lankester, M.D., F.R.S. 8vo. III. A Monograph of the British Nudibranchiate Mollusca. By Messrs. Alder and Hancock. Part I. Ten Plates. Imp. 4to. FOR THE SECOND YEAR, 1845. I. Steenstrup on the Alternation of Generations. Translated from the German, by George Busk, F.R.S. Three Plates. 8vo. II. A Monograph of the British Nudibranchiate Mollusca. By Messrs. Alder and Hancock. Part II. Thirteen Plates. Imp. 4to. 22 LIST OF ANNUAL VOLUMES III. Reports and Papers on Botany, consisting of Translations from the German. Translated by W. B. Macdonald, B.A. ; G. Busk, F.R.S.; A. Henfrey, F.R.S.; and J. Hudson, B.M. Seven Plates. 8vo. FOR THE THIRD YEAR, 1846. I. Meyen's Geography of Plants. Translated from the German by Miss Margaret Johnston. 8vo. II. Burmeister on the Organization of Trilobites. Translated from the German, and edited by Professors T. Bell and E. Forbes. Six Plates. Imp. 4to. III. A Monograph of the British Nudibranchiate Mollusca. By Messrs. Alder and Hancock. Part III. Eleven Plates. Imp. 4to. FOR THE FOURTH YEAR, 1847. I. Oken's Elements of Physio-philosophy. Translated from the German by Alfred Tulk. 8vo. II. Reports on the Progress of Zoology. Translated from the German by Messrs. Geo. Busk, A. H. Haliday, and A. Tulk. 8vo. III. A Synopsis of the British Naked-eyed Pulmograde Medusse. By Professor E. Forbes, F.R.S. Thirteen Plates. Imp. 4to. FOR THE FIFTH YEAR, 1848. I. Bibliographia Zoologise et Geologiae. By Professor Agassiz. Vol. I. 8vo. ISSUED BY THE RAY SOCIETY. 23 II. Letters of John Ray. Edited by E. Lankester, M.D., F.R.S. Two Plates. 8vo. III. A Monograph of the British Nudibranchiate Mollusca. By Messrs. Alder and Hancock. Part IV. Twelve Plates. Imp. 4to. FOR THE SIXTH YEAR, 1849. I. 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FOR THE TWENTY-SECOND YEAR, 1865. I. The British Hemiptera Heteroptera. By Messrs. J. W. Douglas and John Scott. Twenty-one Plates. 8vo. II. A Monograph of the British Spongiadse. By J. S. Bower- bank, LL.D., F.R.S. Vol. II. 8vo. FOR THB TWENTY-THIRD YEAR, 1866. I. The Miscellaneous Botanical Works of Robert Brown, D.C.L., F.R.S. Vol. I, containing Geographico-botanical, and Structural, and Physiological Memoirs. Edited by J. J. Bennett, F.R.S. 8vo. ISSUED BY THE EAT SOCIETY. 27 II. Recent Memoirs on the Cetacea. By Professors Eschricht, Reinhardt, and Lilljeborg. Edited by W. H. Flower, F.R.S. Six Plates. Imp. 4to. III. Nitzch's Pterylography, translated from the German. Edited by P. L. Sclater, M.A., Ph.D., F.R.S. Ten Plates. Imp. 4to. FOR THE TWENTY-FOURTH YEAR, 1867. I. A Monograph on the Structure and Development of the Shoulder-girdle. By W. K. Parker, F.R.S. Thirty Plates. Imp. 4to. II. The Miscellaneous Botanical Works of Robert Brown, D.C.L., F.R.S. Vol. II. 8vo. FOR THE TWENTY-FIFTH YEAR, 1868. I. Vegetable Teratology. By M. T. Masters, M.D., F.L.S, 8vo. II. The Miscellaneous Botanical Works of Robert Brown, D.C.L., F.R.S. Vol. III. Thirty-eight Plates. Imp. 4to. FOR THE TWENTY-SIXTH YEAR, 1869. A Monograph of the Gymnoblastic or Tubularian Hydroids. By J. Allman, M.D., F.R.S. Part I. Twelve Plates. Imp. 4to. FOR THE TWENTY-SEVENTH YEAR, 1870. A Monograph of the Gymnoblastic or Tubularian Hydroids. By J. Allman, M.D., F.R.S. Part II. Eleven Plates. Imp. 4to. 28 LIST OF ANNUAL VOLUMES FOR THE TWENTY-EIGHTH YEAR, 1871. A Monograph of the Collembola and Thysanura. By Sir J. Lubbock, Bart., M.P., F.R.S. Seventy-eight Plates. 8vo. FOR THE TWENTY-NINTH YEAR, 1872. A Monograph of the British Annelids. By W. C. M'clntosh, M.D., F.R.S.E. Part I. Ten Plates. Imp. 4to. FOR THE THIRTIETH YEAR, 1873. A Monograph of the British Annelids. By "W. C. M'clntosh, M.D., F.R.S.E. Part I. continued. Thirteen Plates. Imp. 4to. FOR THE THIRTY-FIRST YEAR, 1874. A Monograph of the British Spongiadse. By J. S. Bowerbank, LL.D., F.R.S. Vol. III. Ninety-two Plates, 8vo. FOR THE THIRTY-SECOND YEAR, 1875. A Monograph of the British Aphides. By G. B. Buckton, F.R.S. Vol. I. Forty-two Plates. 8vo. FOR THE THIRTY-THIRD YEAR, 1876. A Monograph of the British Copepoda. By G. S. Brady, M.D., F.L.S. Vol. I. Thirty-six Plates. 8vo. ISSUED BY THE RAY SOCIETY. 29 FOR THE THIRTY-FOURTH YEAR, 1877. A Monograph of the British Aphides. By G. B. Buckton, F.R.S, Vol. II. Fifty Plates. 8vo. FOR THE THIRTY-FIFTH YEAR, 1878. A Monograph of the British Copepoda. By G. S. Brady, M.D., F.L.S. Vol. II. Forty- nine Plates. 8vo. FOR THE THIRTY-SIXTH YEAR, 1879. I. A Monograph of the British Copepoda. By G. S. Brady, M.D., F.L.S. Vol. III. Eleven Plates. 8vo. (Com- pleting the work.) FEINTED BY J. E. ADLAED, BARTHOLOMEW CLOSE.