THE LIBRARY OF THE UNIVERSITY OF CALIFORNIA DAVIS BE1TISH SESSILE-EYED CRUSTACEA. LONDON : PRINTED BY WOODFALL AND KINDER, AKGEL COURT, SKINNER STRKET. A HISTORY OF THE BRITISH SESSILE-EYED CRUSTACEA. C. SPENCE BATE, F.R.S., F.L.S., ETC., AND J. O. WESTWOOD, M.A., F.L.S., HOPE PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF OXFORD, ETC. IN TWO VOLUMES. VOL. I. LONDON: JOHN VAN VOORST, PATERNOSTER ROW. M.DCCC.LXIII. LIBRARY UNIVERSITY OF CALIFORNIA DAVIS INTRODUCTION. THE term "Sessile-eyed" has been applied to the order of crustaceous animals forming the subject of this work in contradistinction to that of the " Stalk-eyed " order, of which Professor Bell has given an account in a pre- ceding volume of this series. The name, with its Greek equivalent, Edriophthalma, was first given by Dr. Leach, and has been recognized by all subsequent naturalists. It must not, however, be understood to characterize every genus that should be classed in the order. Among the Isopoda, the genera of Tanais, Paratanais, Apseudes, and Munna, have their eyes fixed on pedicles. In the first of these genera, the structure differs so much in character from that of the normal Isopoda, that it has been classed with the Stalk-eyed Crustacea by Fritz Miiller and Anton Dohrn. In this work we have placed it in an interme- diate position between the Amphipoda and the Isopoda its most important structural characters holding a position nearer, but intermediate in relation between, these two orders than they bear to the Stalk-eyed Crustacea. While, therefore, the eyes may be considered (as they have been since the days of the Swedish naturalist, Linneeus) as a ready and convenient means of classifi- cation, separating one great division from another, this character must be received as only an approximation to a a 2 iv INTRODUCTION. general law. So common, however, is it, and so ready of discernment, that it will probably be retained, even after a more perfect, but less readily detective, system of natural arrangement be discovered. The term was at first applied so as to embrace all Crustacea that were not contained in the Stalk-eyed order, with the exception of the Cirripedia. It is still so retained in Mr. Dana's " Classification of Crustacea," and consequently embraces a large number of forms, exclu- sive of those described in this work, which vary so con- siderably from each other, that we believe it is neither natural nor desirable to group them under one definition. In the present volumes, we speak of the Sessile-eyed Crustacea as constituting a legion between the Stalk-eyed (Podophthalma) and the Entomostracous Crustacea. But the great difference of character in some animals of this legion from the others induced Latreille to divide it into O two orders, naming them respectively after the structure of their locomotive appendages, Amphipoda and Isopoda. Another division was proposed by the same author, and very generally adopted, namely, the Lcemipoda, orLamodi- poda. The animals that constituted this supposed order differ from the normal species of the Amphipoda only in the absence and deficiency of parts ; consequently, in this work, they are viewed as an aberrant group of the order; whereas Latreille first placed the animals of this group in the order Isopoda,* and Lamarck united them with the Amphipoda and Isopoda as members of one family only, under the name of Arthrocephales, or Capites. Dumeril, in his " Zoologie Analytique," united the Amphipoda with the Stomapoda, the point of similarity being the sepa- ration of the head from the body. The term Tetradecapoda has been proposed for the * Dictionnaire d'Hist. Nat. INTRODUCTION. v Sessile-eyed Crustacea by M. Blainville, in contradistinc- tion to that of Decapoda : the one being defined by having fourteen legs, the other having only ten. But this, upon the most superficial examination, will be found to be the most imperfect character, not only in usefulness, but also in appearance. Not only all the Sto?napoda, but even the Macrura, below the family of Palcemonida, possess fourteen fully developed pediform limbs ; and even in the Brachyura and Anomura, the anterior appendages that protect and supply the mouth are legs altered for a necessary purpose, and not really oral appendages; conse- quently, the distinction in structure that the two separate names would lead a student to expect, does not exist. The only true Decapoda are Caprella and Anceus, and these belong, in the present system of classification, to the Tetradecapoda. The term Choristopoda, or separate-footed, has been applied by Mr. Dana, who uses it as synonymous with Tetradecapoda of Blainville and our term of Sessile-eyed, over which it appears to possess no advantage, without which it is unwise to add to the already too numerous list of synonyms. Thus it will be perceived that, in our con- sideration of the orders treated of in this work, we consider that the Sessile-eyed Crustacea bear a nearer structural affinity with the Stalk-eyed Crustacea than with the Trilobita, Entomostraca, and Rotatoria, which Mr. Dana unites into the one division under the term of Sessile-eyed Crustacea. The classification that we have adopted nearly resembles the system of arrangement adopted by Milne Edwards in his " Histoire des Crustaces ;" but, in his classification, the aberrant Amphipoda are admitted to a rank of equal im- portance to that of the Amphipoda, whereas certain very exceptional forms of Isopoda are only distinguished as a separate family of Isopoda. vi INTRODUCTION. The aberrant group of Isopoda, although containing, and perhaps based upon, the most characteristic genus of Dana's supposed order of Anisopoda. yet must not be con- sidered synonymous with it, since all the parasitic forms that possess such extremely aberrant characters in the adult females, possess the true character of the normal Isopod, both in the young and adult male. Thus the genera Arcturus, Bopyrus, and the rest of the parasitic Amphipoda, we have classified with the normal Isopoda. Nor can we think that the only feature that assimilates Arcturus to the Amphipoda (the forward direction of the second pair of pereiopoda), can be considered of suf- ficient importance to narrow the distinction between it and the Amphipoda, whereas other characters of greater importance induce a natural separation that is strongly marked. The consideration of the structure of the Sessile-eyed Crustacea has, until recently, but little attracted the attention of zoologists. The observations of Loven, Lilljeborg, Goes, De la Valette, Grube, Fritz Miiller, Anton Dohrn, Schobl, Schiodte, and others, have done much to show the large amount of novel and interesting subjects of biological knowledge that have been, and still are to be, developed by the study of this hitherto much neglected class of animals. The structure of these animals, though offering a very palpable distinction from the higher forms, is indubitably formed upon the same common type. So clearly can this be demonstrated, that we are somewhat surprised to find that Mr. Dana ("United States Exploring Expe- dition," vol. i. p. 1404) should say that " they have not a macrural characteristic, but have a body divided into as many segments as they have legs (hence our name Choristopoda) ; the antennae, legs, and whole structure are distinct in type." INTRODUCTION. vii That every segment has its appendage is a law common to all Crustacea. In the Stalk-eyed order, the develop- ment of the cephalou is carried to a monstrous extent as a shield or carapace, covering and protecting, in some cases, all the segments of the pereion. When the cara- pace is so developed, the necessity for perfect segments in the latter does not exist, consequently the dorsal sur- face is wanting ; but the lateral portion is always present. In the Sessile-eyed Crustacea this enlargement of the cephalon does not exist, and the absence of a carapace permits the development of the dorsal surface of the segments of the pereion. A careful examination of the appendages of the head will clearly show the same number of segments associated together as is found to exist in the macrural forms, consequently the head or cephalon in the Sessile-eyed Crustacea homologizes with the carapace in those Crustacea that have their eyes supported on foot-stalks. Gradually, from the Brachyura, it decreases through each succeeding order, and this, apparently, in relative degree with the separation of the nervous system into separate ganglia, obedient to a common law of depreciation, which in the Sessile-eyed Crustacea appears to reach a lower limit in the Isopoda than in the Amphipoda. The appendages that are supported by the cephalon are various in form, and generally associated with the senses. The first, or most anterior pair, are the eyes, which, from the circumstance of being closely impacted within the dermal skeleton, give the name of Sessile-eyed to the legion, as above mentioned. This position is not invari- ably the case, since in the genera Tanais, Paratanais, Apseudes, &c., the eyes are carried on elevated stalks. In the Isopoda these organs appear to be more perfectly developed than in the Amphipoda, except, perhaps, in Hyperina, where their monstrous development deprives viii INTRODUCTION. the head of its normal form. In the Isopoda generally, the lenses of the eyes are well developed, and lodged in the texture of the skeleton of the animal, which is fre- quently thinned out to an extreme tenuity, and marked with numerous facets, corresponding with the many lenses belonging to the organ. In the Amphipoda, the lenses either are not so numerous or are less apparent, and the dermal tissue that covers the organ is thick and un- changed in character. This condition is carried to the greatest limit in the Phoxides, Ampeliscides, and those Gammarides that are inhabitants of deep and dark wells, where no rudiments of eyes are apparent, except in the presence of some coloured and ill-defined pigment cells, which in the Phoxides coalesce into a single organ. In the genus Ampelisca this pigment of colouring is associated on each side with two solitary lenses, that appear to be built into, and form part of, the dermal covering. It appears to be a law in the decreasing structural importance of Crustacea, that the segment supporting the appendages shall disappear before the appendage that it supports. In the Sessile-eyed legion, the eyes alone remain, the segment and the articulating portion of the appendage not being developed ; the eyes are developed in most families so deeply within the head, that they generally appear to be behind the antennae, and some- times, as in Phoxus, at the extremity of the frontal rostrum ; in others, as Ericthonius, on a projecting lobe of the head, situated between the two pairs of antennae, in which position, owing, probably, to the insufficient depth of structure, the eye is borne on the internal surface, where it is lodged as a protuberance. But what- ever may be the position of these organs, the variable- ness of situation can only be consistent with certain advantages under peculiar conditions. In the young animal the number of facets is fewer in INTRODUCTION. IX the eye than in the adult state. In the genus Gam- marus, the number of lenses in the young is first eight or ten, whilst in the adult they number from forty to fifty. The superior or first pair of antennae we consider, con- trary to the opinion of Mr. Dana, to be formed on the same type as those of the Macrura. Each of them con- sists of three distinctly formed joints and a flagellum, with sometimes a more or less important secondary appendage. We have long since expressed our opinion that in these organs lies the seat of auditory consciousness, and we are still inclined to retain that opinion. We are aware of the elaborate experiments of Dr. Von Hensen, which tend to demonstrate the existence of auditory cilia on several parts of the animal, as the superior antennae, (in which Professor Huxley was the first to demonstrate, in some exotic Macrura, the presence of highly refracting otolithes,) on the inferior antennae, as well on the caudal appendages as in the external branch of the posterior pleopoda, on which Van Beneden has discovered, and we have seen, what appear to be well-formed otolithes, of the same type as those found in the first joint of the anterior pair of antennae in Mysis, &c. But we have always attributed to certain very delicate membranous cilia of various forms, found on the primary flagellum only of the superior antennas, and present, under normal conditions, in nearly every family of Crustacea, the power of convey- ing impressions of sound. But these membranous cilia are very distinct from the auditory hairs of Dr. Von Hensen.* That the superior antennae are, in their most normal development, purely aquatic organs, we see in the depre- ciation of their character in the partly marine genera * An elaborate memoir on the auditory organs of the Crustacea, by Dr. V. Hensen, was published in Zeitschr. f. Wissensch. Zoologie, xiii. Bd. 3. Hft. 1863, an abstract of which may be seen in the Zoological Record for 1864. X INTRODUCTION. Orchestia and Talitrus, and their rudimentary condition in the terrestrial Isopoda. The inferior or second pair of antennae are formed on the simplest character of the Macrural type, and consist of a peduncle with five joints, of which the first two, (the hornotypes of the coxa and basis joints of the true leg.) are very closely associated, and carry the olfactory denticle. In the higher groups, the two basal joints are fused together, and often with the nearest part of the segment to which they belong. Sometimes, so perfect is the union, that not the slightest trace of the relation of one part to another is capable of being detected. This complete association of the appendage with the body of the animal lessens with the degradation of the creature, until we find the five separate joints distinguishable from each other and from the body of the animal. The denticle at the base of the second pair of antennae in the Amphipoda (Fig. 1), homologizes with a perforated FIG. 1. tubercle situated on the ventral surface of the cephalon in the Brachyura, laterally anterior to the oral apparatus, and indeed covered by some of the appendages, in the higher groups of the class. The denticle in the Amphipoda, upon close examination, appears to have an open extremity, through which a cylindrical tube, retained in its place by membranous ligatures, protrudes. This tube closes at the INTRODUCTION. xi internal extremity rather suddenly, and encloses the elongated bulbous extremity of a nerve-thread, that pro- ceeds from a second bulb or nerve-ganglion implanted at the base of the denticle. This denticle, though frequent, is not invariably present. In the genera Orchestia and Talitrm, the two basal joints of the antennae are built into the anterior wall of the cephalon, so as to be generally mistaken for it ; while in others, as also in the Isopoda, every trace of the denticle is lost (Fig. 2). FIG. 2. There is no secondary appendage to the inferior antennae, and, with the exception of the squamiform plate in the Macrura, it is never found in Crustacea ; nor is it invariably a macrurous condition, since in some genera it is entirely absent ; and even in Palinurus, a most typical form, it is lost as an appendage, being distinguishable only in the outline impressed in the walls of the fourth joint of the antennae. The flagellum in all Crustacea originates, in the upper antennae, after the third perfect joint ; in the lower, after xii INTRODUCTION. the fifth ; and in every case the secondary appendage, whether in the form of a scale attached to the lower, or a filamentary appendage, or several, invariably in upper and lower alike arises from the distal extremity of the third. This appears to be a very constant condition with all the appendages of the cephalon, pereion, and pleon. The most freqnent exception exists in the first joint or coxa, as exemplified in the branchial appendages and the ovigerous plates of the female Amphipoda and Iso- poda. According to our experience, whenever any secondary appendage is developed from the second joint or basis, it exists more as a rudimentary effort than as a true organ. After the third joint, we are not aware that any secondary appendage is ever produced, though in some genera, as in Palamon, the primary flagellum of the anterior antennae occasionally divides or sends off a smaller one. The flagellum in the Sessile-eyed Crustacea is generally multi-articulate. It attains its most filamentary character in the sub-family Gammarides ; but in some genera many, and sometimes all, the numerous articuli coalesce into one or more joints, as in Podocerus, Corophium, Chelura, the terrestrial Isopods, &c., in all which cases they become organs assisting in climbing and grasping. Unlike the superior antennae, the inferior pair appears to be always present, and we only know of their being reduced to an immature condition in those Crustacea that pass their lives as parasites upon others, as the Bopyrida, Hyperiida, and Cyamus, a circumstance that induces us to believe that the second pair of antennae is the seat of a sense which undergoes but slight modifications to enable it to be equally distinguishable whether in air or water, since the Isopoda and Orchestia, in which the antennae are well developed, are terrestrial. The oral apparatus in most Crustacea is a somewhat INTRODUCTION. xiii complicated series of organs. It is built up of many separate pairs of appendages, those belonging to the higher groups of Crustacea being the most numerous. In the Sessile-eyed orders, the mandibles are separated from the second or posterior pair of antennae by the ven- tral surface of the fourth or mandibular segment, and a protuberance that, from its position, is called the labrum, or anterior lip. In the Amphipoda, the epistome is generally placed vertically, and occasionally produced anteriorly into a sharp spear-like process. In many, however, as also in the Isopoda, it exists as a plate that gives strength and solidity to the fulcrum on which the mandibles rest. The labrum is divided into two parts, the lower of which moves on the upper by a slight hinge, and assists in perfecting the shutting of the mouth. The free margin is generally clothed with short hairs, often of club-shaped and deformed appearance. The mandibles are powerful organs, impinging against each other at their extremities, the biting edge being in the median line. In the Sessile-eyed Crustacea, they bear a near resemblance to the same appendages in the larval condition of the highest order of Crustacea. The anterior or biting margin of the mandible is generally divided into several short and strong denticles, though in some genera it is smooth and even. Within the denti- cular margin a second process generally exists, a smaller repetition of the first, and which commonly, when present, is attached by a movable joint. Near the centre of the mandible is a large internally projecting process, that corresponds with and meets a similar process in the opposite mandible, and is evidently adapted for masti- cation, and may with propriety be named the molar tubercle. It forms, generally, with the anterior or XIV INTRODUCTION. incisive margin, the two extremities or horns of a cres- cent. The second, or articulated process, is situated between the two, but somewhat nearer the anterior margin. It appears to be able to assist in carrying the food from the one point to the other, from the biting to the grinding surfaces, between which and the molar tubercle are frequently a row of strong and curved spines that facilitate the process. The mandibles are moved by powerful muscles attached to the inner surface of the dorsal part of the cephalon, corresponding with the homological parts that are attached to the inner dorsal surface of the carapace of the higher Crustacea. The surface of the molar tubercle is granulated with rows of minute denticles that are only visible under a strong magnifying power. In some species, a long and slender ciliated filament is appended to the margin of the tubercle that may be associated with the sense of taste. The mandibles are no exception to the fact that all appendages are but modified legs. In all Crustacea, we think that it can readily be demonstrated that the man- dible consists of the first three joints being closely anchy- losed. The small appendage, that generally consists of three freely articulated joints, represents the fourth, fifth, and sixth joints; the seventh, or dactylos, being seldom present. An homological examination of the genera Nebalia and Pontia, with Homarus, together with the homotypical parts in other appendages in the same ani- mals, we think will readily confirm this opinion. The small three-jointed appendage to the mandible is wanting in but few genera, excepting in the terrestrial Isopoda and Amphipoda. In aquatic species it is, with few exceptions, always present, and appears to be of efficient use in directing floating material towards the mouth. INTRODUCTION. XV In some parasitic families these organs undergo an extreme amount of modification. This is much more exaggerated in the Isopoda than in the Amphipoda. Among the Cyami, the oral appendages are all reduced and somewhat modified, but in the Cymothoida, Bopyridce, and Anceida, among the Isopoda, they appear to lose much of their normal character, and fulfil the office of a sucking apparatus. In the formation of this organ one or more pairs of the appendages may be implicated, as is shown in an elaborate memoir by Schiodte* on the subject. The manner in which the organ is developed in lone from the mandibles, we have described at page 253, vol. ii. of this Work. In the Anceida, the appendages of the mouth in the young stage are sharp and lanceolate, the sucking organ being apparently modified from the labrum, where, as in the adult animals, the oral aperture, with the sup- plying appendages, are lost, or converted into members useful for other purposes. In the genus Brachyscelus, and others of the family Platyscelida, the appendages of the oral apparatus are reduced to a single pair of membranous leaf-like organs; nor have we been enabled to trace any different character of organ to take the place of the lost ones. Both in the adult and young animal, the mouth appears to be reduced to a rudimentary and simple character : an aperture with the probable power of opening and closing at will being the most that we have been enabled to determine. The first or anterior maxillae (Siagnopoda) are separated from the mandibles by a posterior lip, which differs in the Amphipoda or at least in some genera in being cleft lon- gitudinally in the median line, and is termed the labium ; * Natur. Hist. Tidssk. 1866, p. 168206. xvi INTRODUCTION. it appears to be capable of being slightly moved, and probably assists the mandibles in the process of manduca- tion. There are three pairs of Siagnopoda, the two ante- rior of which are extremely delicate foliaceous appendages, whilst the third is much more robust, yet still possessing a foliaceous character, particularly as regards the three or four basal joints. In some genera, as in Sulcator, some of the plates, particularly of the two anterior pairs, are folded so as to become two or three parallel leaves, one of which, on the first pair in Sulcator, is developed into a prominent lobe, containing large nucleated cells. Of the office or use of this gland-like organ we can offer no sug- gestion, not having met with any analogue in the order. The two anterior pairs, the maxillae of authors, vary somewhat in their form in genera, and very much between the Isopoda and Amphipoda. In the parasitic species of both orders, they are defective, and sometimes wholly wanting. The third Siagnopod, or first maxillipede of authors in these orders, is a true cephalic appendage, and covers the organs of the mouth as a protecting operculum. These last three pairs of appendages are concentrated about the mouth, the segments to which they belong being represented by the ventral portions only, and these are closely fused together, from the sides of which, in the genus Talitrus, originate two bony processes, that meet, without uniting, near the internal centre of the head, there spreading out into flattened plates, from each of which a thin and somewhat delicate process is directed anteriorly and slightly upwards ; the stomach is supported by them in its position. This osseous internal arch, that we described in the British Association Report " On the British Edriophthalma," 1855, Professor Huxley has, in his lec- tures at the Royal College of Surgeons, published in the INTRODUCTION. XVll Medical Times and Gazette, vol. xxxvi. p. 467, 7th Novem- ber, 1857, named the Endophragmal arch (Fig. 3, En.). FIG. 3. The seven segments which succeed the cephalon, or head, are, in the higher orders, protected by the carapace. This becomes gradually smaller in the descending series, until, in the Sessile-eyed Crustacea, each segment is exposed and developed into a perfect ring, analogous in appearance to the segments of the pleon in the Macrura. The several appendages that belong to the segments of the pereion are locomotive in their charac- ter, some being perfectly natatorial or ambulatory, others adapted for climbing and grasping. In this respect the two anterior pairs in the Amphipoda are most constant in their adaptation. The probability is, that these last are never in the Amphipoda used, except for carrying food to the mouth, or more rarely for climbing, or occasionally grasping the female. In this they are found to possess b XVlll INTRODUCTION. a feature that, with the exception of the Isopoda, is common to most Crustacea, even including the aberrant Isopods. We have thought it convenient to describe them iinder a name distinguishing them from the true ambulatory legs, although by doing so we must include some genera of Isopoda, where they assimilate to and fulfil the conditions of true walking-legs. In the Brachyura, the gnathopoda are developed, so as to serve chiefly as protecting the oral apparatus. In the Macrura, they assume a pediform appearance, and are used in seizing and holding food. In the Stomapoda, the Squillidce have them developed into formidable pre- hensile organs. This change takes place gradually from the highest Crustaceans to the Amphipoda. The cha- racter is still increased in some of the aberrant genera, until it becomes a perfectly didactyle chela. In the Isopoda, the prehensile character may be said to be lost, presenting itself only occasionally in the anterior pair, in the male animals. The five remaining pairs of walking-legs (the pere- iopoda) homologize with the five pairs of legs in the Stalk-eyed Crustacea, that give the name of Decapoda to the order. These are produced on a somewhat different plan from the walking-legs of the Stalk-eyed Crustacea, the modification, as it appears to us, taking place in accordance with certain necessities that have arisen from the depreciation of their general develop- ment. The two anterior pairs of legs, or gnathopoda, are developed upon one type; the two succeeding pairs, or first and second pairs of pereiopoda, on a second; and the last three on a third. The normally developed appendage of every kind in Crustacea consists of seven joints. In the Brachyura, the first, or coxa, is anchylosed with, and forms part of, INTRODUCTION. xix the sternum. In the Macrura, it also forms part of the sternum, but the separation is distinguishable by a free and movable articulation. In the Sessile-eyed Crustacea, the coxa is more laterally situated, and very firmly attached, without being fused to the segment of the body. With few exceptions, it is developed into a broad and scale-like joint, and is so large in the Stegocephalidte that it covers the greater part of the animal. The object of this development is evidently to cover and protect the branchial appendages, when situated beneath the pereion. These scale-like coxae have been considered as parts of the segments of the body of the animal to which the legs belong, and are described under the name of epimera, or side-pieces, by Professor Milne Edwards. There is a peculiar tendency in the Amphipoda for the joints of the legs to be produced in a scale-like form. Besides the coxa3, the basis, or second joint of the three posterior pairs of pereiopoda, are almost always so de- veloped. In Orchestia, the males in some species have the carpus and posterior pair of pereiopoda enlarged ; in Podo- cerus and Cerapus, the two anterior pairs have the basis so produced ; but in Sulcator this predisposition appears to reach the culminating point, where it is apparent in almost every joint of the appendages of the head and body. The next division of the animal is that which we deno- minate the pleon. It consists of seven segments, as in each of the former divisions, and carries three kinds of appendages. The segments generally resemble those of the pereion, and, like them, carry on each side squamiform coxa3, which Professor Milne Edwards has again mistaken for epimera, or side-pieces, belonging to each respective segment. These are, both in the Amphipoda and Isopoda, XX INTRODUCTION. generally fused closely with the dorsal surface of the seg- ment ; but in the genus Apseudes, as we have shown in fig. p, page 148, vol. ii., they are free. Here we have a distinct exposition of the relation which the squamiform side-piece holds both to the segment and the movable bifurcate appendage. The segment is distinctly separated from the squamiform side-piece, which, articulating with it, forms the first joint of the pleopoda or swimming-leg, and is developed into a large scale-like process, to the base of which the second joint is articulated, from whence is suspended freely a third, which in its turn supports the two free plates which form the terminal appendage of the anterior pleopoda. In the Isopoda, as well as the Amphi- poda, this interpretation illustrates the relation of the parts of the pleopoda to the segments of the pleon. The forms of the pleopoda may and do change, according to the law of modification of parts, to suit their require- ments ; but under whatever condition they may exist, they consist of three normal joints, more or less fused together, and with the segments of the pleon and a depreciation of the four terminal joints into one or a pair of movable plates, as in the Isopoda, or articulated flagella, as in the Amphipoda. The three anterior pairs in the Amphipoda are deve- loped upon this type ; the two succeeding have the double appendages stiff and unyielding, and the posterior is generally variable in the different genera. In the Isopoda, the four anterior pleopoda are developed upon one type, while the fifth is converted into an operculum. Some variation of the anterior pairs also takes place in relation to the sex of the animal. The last, or twenty-first segment, differs from the rest in most Crustacea by not carrying any appendage. To this we know of but one exception among the Crustacea, and that is in a genus in the family INTRODUCTION. xxi My sides, discovered by Mr. Norman. The telson in the Sessile-eyed Crustacea is generally an abortive., and fre- quently a rudimentary, part. In the Isopoda, except in the genera Apseudes and Anthura, it is always fused with the preceding segment. The composition of the dermal skeleton is, in all Crus- tacea, the same. In the Sessile-eyed order the texture is very thin, and seldom consolidated into a firm structure, except in certain parts of some few genera where strength is required, as in the chelse of large-handed species. This circumstance offers the advantages of enabling the observer to examine the internal structure of the animal without the necessity of dissection. During the life of the animal, we are enabled to trace the currents of circula- tion of the blood, the motion of the cardiac vessel, and the position of the internal organs in relation to each other. This delicacy of the structure also enables us to dis- cover the very diverse and varied arrangement of the material of which it is built up. and demonstrates (con- trary to our anticipations) that in species often closely allied, there is a very distinct appearance in the micro- scopic structure. It may prove to be of some importance in determining species, but care should be taken that the several specimens examined should be taken from the same part of the skin of each animal. We have illus- trated many of these varieties of structure throughout the work, in connection with the animals to which each belongs. Frequently, besides the markings that illustrate the manner in which the skin is built up, there is another that is not always constant, consisting of a series of small perforations through the tissue, which in some species assume a waved appearance, as may be observed in the genus Ampelisca. xxii INTRODUCTION. Although we believe that the microscopic examination of the skeleton in these animals would frequently facili- tate the determination of doubtful species, yet it is a con- dition that is not to be trusted to alone, inasmuch as it is not uufrequently found that similar appearances are repeated in very distinct genera. Examples of this may be found on comparing the structure of Megamosra Otho- nis with that of Chelura terebans. The form and structure of the hairs that are found on these animals, when microscopically examined, are of a very distinct and different character. They not only vary in separate species, but differ in several parts of the same animal. In Sulcator there are no less than twelve varieties. Some are plain, stiff, bristle-like spines of various lengths, which are generally attached to the margins of the limbs. A second variety, longer in general form, fringed on one side with a series of fine, straight, teeth-like processes, possessing a rake-like cha- racter, is attached to the third siagnopod ; as is also A third, that differs from the preceding in having the teeth bent in a curve directed to the base. A fourth is found on the carpus of the second pair of gnathopoda. In this position are also two varieties, which originate from closely approximating bases. One is long, slender, and clean to the tip, where a few exquisitely fine cilia appear, which give to the extremity a bulbous appearance, that can be resolved only with a high (700) magnifying power. The other, or The fifth, is short, broad, flat, terminating in a point that is sharply bent upon itself; the lateral margins are like- wise furnished with a series of sharp denticles, ranged on each side, pointing to the base for about two-thirds of its length. A sixth is found on the propodos of the same appen- INTRODUCTION. xxiil dage; comprising two forms moulded on the type of the two preceding ; the shorter changing the hooked extremity for a bulbous termination, and the shaft being armed with teeth on one side only. A seventh exists on the mandibular appendage : it is straight, enlarged and rounded at the apex, and serrated on one side; while An eighth differs from the preceding in being more robust, slightly turned at the extremity, and smooth along the margins, excepting a single short, straight, distally directed cilium. A ninth resembles the sixth, but wants the serrated margin, and carries on the convex side a fine cilium. This variety is found 011 the first pair of gnathopoda. The tenth, eleventh, and twelfth varieties are plumose, and found mostly on the second pair of antennae, though a few are present on several other parts of the animal. One is short and obtuse, being crowned with numerous radiating cilia. It is to this variety that we understand Professor Hensen attributes the power of hearing. This great variety of form in the hairs of a single species is not constant. In the genus Talitrus, there is but a single form of hair, which is but little modified in the various parts of the animal. It is short, stiff, and blunt, and exhibits under the microscope a tendency to a spiral condition for about one-fourth from the extremity, at which distance a second but smaller process exists, so that the hair might be characterized as being forked, but for the unequal proportion of the two branches. This kind of hair is by no means rare in the Amphipoda. Those found in Orchestia, Talorchestia, Nicea, Gammarus, &c., are but modifications of the same form. This great variation in the form of the hairs is more or less common to all Crustacea. Those in Carcinus mcenas have been xxiv INTRODUCTION, described and figured by Dr. Mclntosh in the " Linnsean Transactions" for 1862, p. 79. The hairs are not only various in form, but sometimes they will be found con- stant in number. Thus, in the genus Phoxus, we have found the number of hairs on the coxse of the three or four anterior pairs of legs to be constant in the respective species. EXUVIATION AND REPRODUCTION OF LIMBS. The power of Crustacea to throw off their skin and replace it by a new one, has long been a recognized fact in all the higher orders. It is, however, on the authority of Mr. Couch, stated by Mr. Bell, in a note to his intro- duction to the " Stalk-eyed Crustacea," p. Ixi., " that the families in which the eyes are sessile in their adult growth .... do not exuviate, or voluntarily throw off their limbs." These Crustacea, however, like their higher congeners, renew their integumentary tissues periodically. This is equally true with regard to the alimentary canal, which is cast in connection with the skeleton. The animal shows no appreciable difference in its habits at the time imme- diately anterior to its throwing off its exuviae. It swims about very actively until the hour of moulting arrives, when it seeks a place of comparative security, where it may remain uninterrupted the necessary length of time for the completion of the process. In this position it grasps with the anterior pair of gnathopoda some fixed and conveniently secure material for an anchorage. Here the labour is commenced, and, judging by the quietness and rapidity of the process, appears to be one of no great discomfort. During the INTRODUCTION. XXV operation, at almost any stage, the animal, if disturbed, is capable of removing itself to a more quiet and secure place. The process appears to be the result of an internal growth of the animal, which becoming too large, the skin splits at the margin of the dorsal and sternal arches of the three anterior segments of the pereion, the inferior arch carrying the legs, inclusive of the coxae. The anterior segment of the pereion extends over the posterior margin of the cephalon. At this point the attachment is broken anteriorly, and the lateral disunion of the three anterior segments allows their upper surfaces to be raised as a movable lid, through the opening of which the animal escapes from the old integuments. With some exertion, the posterior portion of the body, together with the limbs, are withdrawn, after which the head and the anterior members are removed, and the entire animal is free from the old exuvia3, which, resem- bling a dead individual, is left, attached to its old position. Unless disturbed, the animal, which is now extremely soft, generally rests for some time, as if exhausted, near the cast-off skeleton. Upon being disturbed, it is capable of swimming away immediately. Mr. Harry Goodsir, in the Edinburgh Philosophical Journal for 1842, has described the process of exuviation, as observed by him in the genus Caprella. He says that the animal, previously to the commence- ment of the process, " lies for a considerable time languid, and to all appearance dead. At length a slight quivering takes place all over the body, attended in a short time with more violent exertions. The skin then bursts behind the head in a transverse direction, and also down the mesial line of the abdominal surface ; a few more violent exertions then free the body of its old covering. After xxvi INTRODUCTION. this the animal remains for a considerable time in a languid state, and is quite transparent and colourless." The new creature is a perfect representation of the old one, slightly enlarged. According to our observations, every hair is produced complete. We have often seen them, convoluted and bent up within the old case, from which they only wanted to be freed to assume the erect position of the perfect hair. It has, however, contrary to our anticipation, appeared that all the hairs are not de- veloped within each corresponding one. We have fre- quently observed them as a second armature, independent of the old one. This remark is particularly distinguish- able in the teeth that fringe the first two siagnopoda. These have generally a dentated arid forked character, which might be injured in their removal from the old and hard tissue of the rejected skin, an accident that not unfrequently befalls the branchial sacs, which are occa- sionally torn off and retained behind in the old case. The power of Crustacea to throw off any of the limbs upon receiving an injury, and sometimes in consequence of fright, is well known in relation to the higher orders. The manner in which this is done has been described by Dalyell, Goodsir, and ourselves. It certainly is a remark- able power and law of reproduction, and which always takes place at the same homotypical position in every limb that is, between the coxa and the next succeeding joint. The wound that is caused by this sudden rupture is simultaneously glazed over by a thin membrane, which must be very suddenly formed, and probably is the ampu- tating power. Observers have very generally added as an appendage to- the above interesting fact, that it is exceed- ingly fortunate that there is this power of voluntary amputation of the limbs, for otherwise, in consequence of the non-contractile character of the dermal covering, the INTRODUCTION. XXVH animal, upon being wounded in either of the limbs, would of necessity bleed to death. That such would be the case would appear to be extremely probable, but, like all nega- tive evidence, is only of value in the absence of direct testimony. In the Sessile-eyed orders the animal appears to want the power of voluntarily throwing off any of its appendages, no matter how severely it may be wounded. If a leg be cut off, or in any way injured, the wound very soon after becomes cicatrized with a black scar, which remains until the next exuviation of the animal, when the entire limb is thrown off with it, and a new one commences growing. TASTE AND DIGESTION. The sense of the enjoyment of food, even in the highest types of the animal kingdom, exists rather in the power of parts to receive impressions than in the presence of any especial organ for the purpose. Arguing, therefore, from analogy, we should suppose that the sensation of taste in the lower animals (such as the Crustacea, and other groups in which mastication is of an imperfect character), must necessarily be rather a faculty peculiar to the mouth in general, than the result of any especial organ adapted for the purpose. From the mouth the oesophagus leads directly to the stomach. The passage is very short, being directed up- wards and forwards ; it enters the stomach at the infero- anterior margin, and, as in all Crustacea, is within the limit of the cephalic region. Just within the anterior opening of the stomach are situated two rake-like organs, the teeth being placed in a row on an arched base ; they are slightly curved and dentated on the margins. They are so placed as to have XXV111 INTRODUCTION. the points directed inwards, so that food can readily pass into, but cannot return again from, the stomach. The teeth on each side appear to correspond, so that they probably play an important point in tearing and lacerating the food as it passes into the stomach. Posterior to this FIG. 4. triturating apparatus there exists four leaf-like plates, fringed with long and powerful cilia. These are attached to the lateral walls in pairs, one anterior to the other; immediately above the second or posterior pair, appa- rently in a chamber of its own, is a gizzard-like apparatus. We observed this most distinctly developed in Sulcator and Talitrus, and we believe it to be present in all the Amphipoda, and we take it to be the same appendage which Bruzelius and Loven figure and describe as the " mellanbalkan," which is situated within the "blind- sacklikt organ," and not, as their figures * would lead one to believe, on the floor of the stomach. * 6'fversigt af K. Vetensk. Akad. Forhandl. 1859, pi. i., figs. 1, 3, 8. INTRODUCTION. XXIX This apparatus, under a high magnifying power, is seen to consist of several closely packed rows of fine strong short hairs, very commonly arranged together in the form of a heart, the apex of which, directed anteriorly, is truncated. This appears to be the most general appear- ance, though in various genera it is different in form. Its appearance suggests its capability for triturating and grinding food, though it is curious that two such kinds of apparatus should exist at each end of the stomach, the one at the cesophageal entrance, the other near the pyloric outlet. The cavity in which the latter is placed has the walls thickly covered with very short hairs. In the genus Talitrus, posteriorly to this apparatus are placed two long ca3ca, one on either side of the posterior opening of the stomach. These caeca are not universally present in other genera. They are delicate prolongations of the walls of the stomach, and gradually narrow towards their free extremity. They probably supply the stomach with a gastric juice. Still more posteriorly, at the point where the stomach terminates and the alimentary canal commences, are situated from four to six long cseca- like lobes, filled with hepatic cells. These are attached to the inferior surface, forming the liver, and are carried] parallel with the alimentary canal. In the Amphipoda, as illustrated in Gammarus, the liver con- sists of four sub-equal lobes ; in the Isopoda, as exempli- fied in Ligia, it is formed of six lobes, two of which are much longer than the other four, and have a slightly waved or tremulous-looking appearance towards the free extremity. From the pyloric orifice of the stomach the alimentary canal in all Crustacea passes, without curve or inflection^ straight to the anal termination under the telson. To this we know of but a single exception, and that on the authority of Professor Alman, who says : " In Chelura, the alimentary canal is so arranged as to XXX INTRODUCTION. shut one part within another, so as to admit of the head being projected forward, that the animal might eat its way into the wood that it penetrates." This we have not been able to verify, nor can we see the necessity for the disarrangement of the stomach with all its attachments, when a prolongation of the oesophageal canal would enable the animal to accomplish the work on far easier conditions. The structure of the alimentary canal is longitudinally fibrous. In the genus Ligia, a little anterior to the anal termination, a series of transverse muscular bands sur- round it without uniting on the under surface, and probably fulfil the office of sphincter muscles. About two-thirds of the distance between the stomach and the telson, one or two appendages are attached to the alimentary canal in the Amphipoda. We say one or two, because we have distinctly dissected out two in Sulcator (Fig. 5), but have failed to determine more than one in FIG. 5. Gammarus (Fig. 6), Mcera, and other genera. The organ is free at one extremity, and is borne in a forward posi- tion, resting on the dorsal surface of the primavia. It is more important in appearance in some Amphipoda than in others; in Sulcator it is very long. We have never seen it in any of the Isopoda that we have examined, but, as far as our experience supports us, it is present both in the male and female Amphipoda, in the adult as well as in the INTRODUCTION. XXXI larval stage. In the younger form (Fig. 7) it is rudi- mentary, but scarcely more so than in Mcera (Fig. 8). FIG. 6. Immediately posterior to the point of attachment of this organ with the alimentary canal are a series of muscular bands lying transversely across the latter, which probably fulfil the office of sphincter muscles by compressing the passage just posteriorly to the efferent orifice of this sup- posed urinary organ. Muscles very similar in appearance are situated near the terminal exit of the alimentary tube, and probably fulfil the office of sphincter muscles to the anal outlet. The contents of the appendage that we call the urinary organ are, under an object-glass of one-fifth focus, resolved into small round cells, containing a granular nucleus (Fig. 9). These cells are closely packed together, FIG. 7. FIG. 8. FIG. 9. but riot so as to lose their rounded character, and the whole are confined within stout walls. Xxxii INTRODUCTION. CIRCULATION. The circulatory system in the Amphipoda differs very importantly from that of the Isopoda. According to some researches of Professor Wagner * on the genus Porcellio, there exists a well-developed arterial system in the Isopoda. To establish this he adopted the method that was first shown to be practicable by M. Emile Blanchard, and which has since been successfully pursued by M. Kowalewsky on Idotea. A mixture of glycerine and water coloured with carmine injected through the heart into the circulatory system, demonstrates the existence of distinct vessels for the passage of the nutritive fluid. The greatest amount of arterial development, as might have been anticipated, is found to exist in the cephalic, branchial, and generative regions, which the author illustrates by diagrammatical figures. In the Amphipoda, the heart is situated in the dorsal region of the pereion, reaching from the posterior extremity of the first segment to the posterior of the fifth. It is a long, simple, sack-like vessel, consisting of elastic fibrous walls, possessing more the features of a great arterial vessel than that of a true heart. The blood corpuscles pass posteriorly from the pulsating heart through the entire length of the animal immediately above the alimentary canal, and the great venous course returns along the dorsal surface, probably on each side, until it reaches the last segment of the pereion, where it dips to the ventral surface and enters into the branchial sacs, where it passes down the anterior margin and up the posterior, then direct to the heart, which it enters by three lateral pulsating oblique aper- tures. The heart of the Isopoda is situated within the dorsal surface of the pleon, except in Tanais> and probably * Ann. des Sc. Nat. p. 37, vol. iv., 1865. INTRODUCTION. XXXlll other general of the aberrant type, where it is situated, according to the observations of Fritz Miiller, in the dorsal surface of the pereion, corresponding in position with that of the respiratory systems of the various orders. In the Amphipoda, the branchiae are by no means the simple sacs that they have been described. They are situated upon the inner surface of the coxse, and assume the form of leaf-like hollow plates, ranged in parallel lines on each side of the sternum (Fig. 10), and are attached to every pair of legs except the first in the females, and generally the last in males ; though, in Gammarus, we have seen the seventh pair fur- nished with branchiaB as well as the preceding. In the Aberrantia, the number of sacs is reduced to two or three pairs. In this order they homologize with the branchiae of the decapod type, each branchial appendage being viewed in the light of a single plate of the compound organs of the higher type; or rather, perhaps, they bear best comparison with the same organ as it appears in the larval condition in the Brachyura. The great distinction in their character is derived mostly from the appearance which these organs assume in the higher forms, FIG. 10. being that of an internal position. But this is one of appearance only. The branchire are overcapped by the monstrous production of the cephalic shield in the Stalk-eyed orders of Crustacea, a circumstance that gives to the portion of the dermal skeleton that it covers the c XXXIV INTRODUCTION. character and appearance of an internal skeleton. The branchial organs are covered and protected, but they are, nevertheless, essentially external appendages. In the Amphipoda this condition does not exist ; consequently the branchiae are pendant in the water, and placed on the inside of the pereiopoda, the first joints of which are developed into large squaminiform plates for their more efficient protection. The internal structure of these organs appears to consist of thick fibrous tissue attached to the inner surface of the wall of each sac (Fig. 11). The fibrous tissue is arranged FIG. 11. in patches of irregular form, but which correspond in their arrangement with one another. These patches are largest near their centre, and thin out towards their mar- gins: the result is that a channel is left between each. All the channels so formed are connected together throughout the entire organ, and exhibit a continuous labyrinth, through which the blood circulates in many small streams. Should the animal become feeble, a gradual accumula- tion of corpuscles takes place in different parts of the gills, INTRODUCTION. XXXV mostly at first out of the reach of the stronger currents. As the vitality of the animal diminishes, the arterial current is observed to lessen in force, until it is propelled only by jerks, coexistent with every pulsation of the heart. RESPIRATION. The organs of respiration in the Isopoda are homo- logically distinct from those of the Amphipoda. We have already stated that Professor Wagner has shown, in the genus Porcellio, and M. Kowalewsky in Idotea, that the blood in the Isopoda runs in arterial channels. We are FIG. 12. not aware that any of the Amphipoda have been put to the same test as the two genera named in the Isopoda ; and certainly, to microscopic observation, the structure of the xxxvi INTRODUCTION. branchial appendages and other parts of the system that from their transparency and tenuity may be conveniently examined, afford presumptive evidence against the circula- tion of the blood being confined to walled channels. In the Isopoda, the branchial organs are variously diffe- rentiated. In some, as Ligia, for example, the passage of the circulating fluid through the branchial plates is clearly and distinctly defined (Fig. 12). The main artery, com- mencing at the base, gives off numerous lateral branches, that divide and sub-divide into a rich plexus with abundant capillary vessels. In the genus Spharoma, the branchial organs consist of a series of plates attached to the posterior wall of the fourth and fifth pairs of pleopoda (Fig. 13). In the degraded family of the Bopyridce, the bran- chial organs are depauperated to the lowest degree, being in some genera little more FIG. is. than excrescences on the ventro-lateral margins of the pleon. In Tanais, the true branchiae have not been clearly determined. It is the opinion of Dr. Fritz Miiller, Van Beneden, and Doctor Anton Dohrn, that an appendage attached to the first pair of gnathopoda is not a branchial organ, but a flabelliform ap- pendage, that by its constant and unvarying motion induces the surrounding medium to flow over the branchial appendages that as yet have not been discerned. At page 122 of the second volume of this work we have described and figured one of the pereiopoda with a sac-like appendage attached, that we considered as the homologue of the branchial sac in the normal Amphipoda. INTRODUCTION. XXXV11 This appendage appears not to be constant in all species, nor in all specimens of the same species. If, therefore, it be the homologue of a branchial sac, it can only be an organ of repetition. Fritz Muller is quite positive in the assertion that no corpuscles of the circulating fluid pass into the caudal appendages, which are the seat of the branchia3 in the normal Isopoda. The terrestrial Isopoda have the respiratory organs some- what modified from those of the aquatic species. These have been described and figured by MM. Duvernoy, Sa- vigny, Lereboullet, and Professor Wagner. M. Savigny, however, was the first to show that in the genus Tylos the system of respiration was carried on by two separate means ; the one by branchia?, as in aquatic Crustacea, the other by the spiracular air-tubes. This has been recently confirmed by Professor Wagner, who shows the relation of the opercular valves to the respiratory system, and contends that, besides their power of protecting the branchial plates from injury, and precluding the too rapid escape of moisture, they fulfil, by means of a plexus of minute vessels, situated at the base of the operculum, a pulmonary function. This organ, which he figures, has, he says, a kind of tracheal division into numerous rami- fications. Seen by transmitted light it is opaque, but viewed under a direct light it is silvery white; and he contends that it is a pulmonary or tracheal chamber, which serves as a supplementary organ to the true branchiaB. This view is supported by M. Milne Edwards, as may be seen by the reference to the " Atlas du Regne Animal," (Pl.lxx.fig.l.m.), and "L^ons sur la Physiologic et 1'Anatomie comparee," t. ii. p. 141. Our own opinion relative to these organs on the branchial operculum is that they are glands for the secretion of a fluid that xxxviii INTRODUCTION. assists in lubricating the branchial plates in warm and strongly evaporating atmospheres. We have been led to this conclusion from finding that they diminish in size in those specimens that have been long detained in dry places. GENERATION. The organs of generation in the male of the Sessile- eyed Crustacea are not to be determined without great nicety in dissection and care in manipulation. We have, however, in Sulcator among the Amphipoda, and Ligia among the Isopoda, been able to examine them clearly, besides less perfectly so in the animals of other genera in both orders. Bruzelius and Loven have given their atten- tion to the former order, and demonstrated the arrange- ment in the genera Gammarus and Podocerus. The male organs internally consist of a more or less oblong pair of testes, which are liable to vary somewhat in form in different genera. These testes are fitted with numerous small seminal cells. A narrow passage, or vas deferens, connects this organ with a second oval chamber, or vesicula seminalis, which is filled with long fine hair-like spermatozoa, lying thickly coiled one upon another. From the vesicula seminalis a narrow passage leads to the inner surface of the first joint of the seventh pair of legs, where it penetrates in each into a soft membranous external penis. We have kept species of Amphipoda long under observation, and paid close attention to their habits, but have hitherto failed to detect any communication between the sexes which would admit of a direct passage of the penis into the vulva of the female. The male Amphidod grasps the female by one of its strong subcheliform gnathopoda, inserting its claw beneath the anterior edge of the first segment of the INTRODUCTION. xxxix pereion, whilst another is inserted beneath the posterior margin of the fourth or fifth segment. Grasping the female in this way, the male draws it into immediate contact with itself, so that the dorsal surface of the female presses against the ventral surface of the male. In this attitude, more or less firmly compressed, they swim about or rest on any convenient surface for many days. If the two be driven asunder through fear of any danger, the female seeks a place of shelter, while the male swims more actively about. Should the male swim within some little distance of its late companion, it becomes imme- diately aware of the circumstance ; and we have seen it, after having passed the spot, abruptly turn back, seek her out, and seize her with avidity from amidst a numerous mass of others. Immediately after securing, he strikes her with two or three strong lashes of his tail. The female, rolling herself closely up, is carried off by her more powerful mate. This contact between the two sexes is either occa- sionally repeated, or it may last throughout the entire period of incu- bation. We have frequently taken them so coupled, even when the young have been so far developed as to be enabled to leave the care of the parent. We are induced, from this fact, to believe that a series of broods may take place successively through the year, and that the erotic state of the female may exist during the period of incubation. The penis (Fig. 15) is a soft membranous tube, that terminates in a small orifice. It probably has, under certain conditions, the power of becoming harder, but FIG. 15. xl INTRODUCTION. it generally lies pendant from the inner side of the coxa, and is longer in some species than in others. In the genera Proto and Caprella, the penis seems to be formed out of the anterior pairs of pleopoda, just as is the case in the Brachyura, among the Stalk-eyed Crustacea. These observations are further confirmed by those of M. Rousel de Vauzeme on the genus Cyamus. In the Isopoda, these organs have been carefully worked out by Siebold, Lereboullet, and Schobl. In the genus Ligia (Fig. 16), we have observed on each side three testes, consisting of long narrow vesicles, thinning away to exquisitely fine filamen- tary prolongations. These vesicles increase in dia- meter as they approach towards the efferent duct, where they rapidly be- come constricted before uniting with the vas defe- rens. These vesicles are filled with seminal cells, and are, we believe, the true testes. M. Lere- boullet, however, in his researches on the Onis- cida* states that he has observed that each of FIG. i6. these fusiform sacs has attached to its extremity other irregular sacs, which he regards as the principal secreting organs, and con- sequently the spermogenic glands or testicles. These * Mem. sur les Crustacea de la Famile des Cloportides, par A. Lereboullet. Strasburgh, 1852. INTRODUCTION. xli organs, which have previously escaped the observation of anatomists, the author says, "are very irregular sacs, variable in form, simple or compound ; they are generally about three-quarters of a millimetre in length, but some- times less. They are situated deeply on each side the stomach, and are retained in their position by delicate but strong ligaments, which are covered with black pig- ment, which lose themselves between the muscular fasciae, of the segments of the body. These organs are full of cells, that M. Lereboullet considers as the spermatic cellules. The second vesicles, or those which we thought to be the true testes, M. Lereboullet calls testicules acces- soires. They are, he says, three in number on each, enlarged towards the middle ; they thin out insensibly towards the extremities : at one end they unite with the organs that M. Lereboullet calls the testes, and at the other they open into the spermatic reservoir the vesicula seminalis. These accessory testes contain cells which are of two kinds, the larger being less numerous than the others. From these vesicles an efferent duct leads to the vesicula seminalis, which in Ligia is a long and narrow vessel, increasing in breadth gradually as it approaches its extremity, where it is suddenly constricted to a narrow outlet, which, covered with black pigment cells, leads direct to the external penis, which is situated near the centre of the ventral arch of the seventh segment of the pereion. In the males, processes of the branchial appen- dages are developed into stylets, (vide fig. 12), that we suppose must have some secondary influence in the pro- cess of fertilization. The anatomy of the reproductive organs in the females has been carefully worked out by MM. Loven and Bru- zelius in the Amphipoda, and by Lereboullet and Schobl in the Isopoda. xlii INTRODUCTION. According to the former authors, corroborated in part by Mr. H. Goodsir on the genus Caprella, by Roussel de Vauzeme on Cyamus, and from our own direct observa- tion on Gammarus, &c., the internal organs consist of two sets of ovaries. These are long cylindrical bodies, having a duct near the middle, on the inner side, that opens into the vulva, which is situated on the inner side of the coxa of the third pair of pereiopoda, or fifth pair of legs. According to the latter authors, the structure of the same organs in the Isopoda is very similar ; but M. Lereboullet has failed to trace the connection of the ovaries with the vulva. Herr Schb'bl has been more suc- cessful in his researches on the genus Typhloniscus, and has figured them attached to the inner surface of the fifth pair of legs. He has also described and figured a pair of receptaculce seminales, in which the male animal deposits the spermatozoa that fructifies the ova3. Accord- ing to this statement, in the Isopoda, if not in the Amphi- poda also, the male impregnates the female by direct intromission a circumstance of which we have entertained some doubt, partly arising from the formation of the animals themselves, particularly of the Amphipoda, in which the development of the coxa3 and the narrowness of the animal would almost, it would seem, preclude the possibility of the sternal portions of the animals being brought into immediate contiguity, and also from the circumstance of having watched the animals, particularly Asellus, from previous to impregnation to the birth of the young, we have never seen the male in any position relative to the female except in that previously described. The incubatory pouch, in which the ova are deposited, from the period of their fertilization until the young are developed sufficiently for independent existence, is the result of the folding over of several lamelliform plates, INTRODUCTION. xliii generally fringed with hairs. One of these plates is developed on the inner side of each of the two pairs of gnathopoda (Fig. 17), and the two an terior pairs of pereiopoda. These plates overlie each other in a compact form, securely protecting the ova, or the immature young, from external accidents, as shown in fig. 10, p. xxxiii. It is the opinion of Von Siebold that these appendages are periodically developed at the " epoque du rut." FIG. 17. This we have not, from our own obser- vation, been able to verify, having taken females during all periods of the year with these appendages fully de- veloped. They are absent on the young females. We believe, however, that, when they are once developed, they continue permanent organs, only disappearing as the result of accident. In the Anceida, the incubatory pouch appears to belong to the three posterior segments of the pereion. By the continued growth of the ova, the pereion is reduced to a most impoverished state. The alimentary canal being in a collapsed condition, and always empty, the animal can only be viewed in the light of a great egg-producer, after the' development of which an empty sac only is left, the poor remains of a worn-out animal. The^ history of the development of the ovum from its impregnation to the development of the perfect larva has been best worked out by Valette St. George in the Amphipoda, and Anton Dohrn in the Isopoda. We must refer the student to the memoirs of these two authors for a detailed account of the germination and growth of the ovum in all its stages. It will suffice for us to say, that it appears to be clearly established by xliv INTRODUCTION. all observers, that in the progressive growth of the ovum, the embryo of the Amphipoda is rolled within the egg in an opposite manner from that of the Isopoda. The latter is folded backwards, so that the ventral appendages are developed on the external surface, whereas the Amphipoda is bent on itself, the ventral appendages being developed on the inner surface. Dr. Fritz Miiller states that, in Tanais, one of our aberrant genera, the development of the larva is after the manner of the Amphipoda, and not of the Isopoda, among which it is classified. The length of time between the epoch of the deposi- tion of the ovum in the incubatory pouch, and the period of the emancipation of the young animal from the care of the parent, is probably about six weeks. We have ob- served that to be the time required in the genus Asellus. At first the egg is perfectly round. It shortly after- wards increases in one direction, becoming also somewhat larger in Amphipoda at one extremity. Indistinct seg- ments are now observable. The wall of the ovum is of an elastic character, and yields to the movement of the internal embryo. Probably about the middle of the period of incubation the embryo quits the egg, for we have constantly taken it from the pouch in a very immature condition, without being enclosed in the egg-case. The larva at this period is very immature, and enclosed within a general tunic, which, without having any apparent vital connection with the animal more than the original egg-case had, adapts itself in general form to the whole creature, and fulfils the duty of a protective tissue. As the embryo increases in dimensions and completeness of form, so the tunic cor- responds in size and form. At length, freeing itself from this case, the larva strengthens in its own development, but does not immediately quit the care of the parent. INTRODUCTION. xlv We have frequently observed the young Talitrus escape from the mother, upon the capture of the latter; and from the active state of their existence at this time, they appear as if they had long been capable of so acting, if they had required it. The observation of Dr. Salter on the common Gammarus, detailed at page 380 of the first volume of this work, fully confirm this fact as does the circumstance that the young of Ardurus are protected by the mother, who supports and carries them about on the antennae. Also we have been able to corroborate the observation of Mr. H. Goodsir, that the Caprella carries about its young attached to its body. These, together with the fact that many genera, particularly of the Podoceridce, protect and nurse their young for some time within nests, which they build apparently for no other purpose, afford abundant proof that in these animals there is a conscious love of offspring that appears to be less marked in animals far higher in the scale of scientific classification. When the young of Gammarus first swims about as a free animal, it only resembles the parent in a modified degree. The antennae show no distinction between the peduncle and the flagellum. The latter is shorter, and consists of but five articuli, while thirty to forty may be present in the parent. This relative proportion is visible also in the lower antenna, and in the secondary appen- dage of the upper, which increases with advancing age, until the adult stage is acquired. In the structure of the eye we see the same gradual increase going on after the animal has become free. The lenses in the young are from ten to twelve in number, whereas, in the adult, from sixty to eighty may be counted. In many genera it also changes its colour, as does also that of the animal itself. xlvi INTRODUCTION. The young are generally white, or of a deep orange colour ; in the adult, the colours vary apparently in rela- tion to the presence of light and other surrounding cir- cumstances. Occasionally the males vary in colour from the females. We see in Orchestia a rosy tint frequently ornamenting the great claw, and some other parts. We have also observed in Amphithoe littorea the well- matured males assume a yellowish appearance. This may also be the case in other genera of which we have not had the opportunity of exact observation. In Orchestia, the second hand in the larva bears a near resemblance in form to the same appendage in the female a fact that is, we believe, consistent throughout the entire class. The warty development of one of the pos- terior legs also increases with age. In Hyperia, the larva bears but little resemblance to the parent. This was first pointed out by M. Milne Edwards, and next by Mr. Gosse. But more extended observations of the forms of these young animals were detailed by us in a memoir published in the " Annals of Natural History for 1861," on some exotic species. Our observations on the larvse of the parasitic Isopods show a wonderful similarity between the larvae of families in distantly separated orders. NERVOUS SYSTEM. The nervous system was first made out in a general memoir on the subject by MM. Milne Edwards and An- douin. The observations of these authors have since been generally verified by HH. Loven and Bruzelius in the Amphipoda, and Lereboullet in the terrestrial Isopoda. We have also carefully dissected out most of the system in both the genera Talitrus among the Amphipoda, and Ligia among the Isopoda. The plan of the nervous INTRODUCTION. xlvii system in these two orders is that of a typical crustacean. A ganglion corresponds to every segment of the animal ; those belonging to the organs purely of sensation being amalgamated together into a cephalic lobe. This is very beautifully shown by HH. Loven and Bruzelius (Bidrag till Kannedomen om Amphipodernas inre byggnad*). Every ganglion of the several segments after the head is united to the others by two parallel cords in the Amphipoda, and one in the Isopoda, although in the genus Ligia we distinctly made out two, as in the Amphi- poda : from each ganglion, on the right and left, is given off two main branches, and in Ligia we observed two other less important threads. These supply the legs and internal viscera. From the cords, about midway between each ganglion, branches off, on the external side of each, a single branch, which in the Oniscida M. Lereboullet places nearer to the preceding ganglion. In the Amphi- poda, we found it rather nearer to the succeeding ganglion. In Ligia, it appears to be just midway between the two, from the base of which, both before and behind, spring other thread-like branches. The diagrams of the arrangement of the caudal supply of nerves, given in the memoir of Lereboullet, differ from that given by M. Milne Edwards in his " Histoire des Crustaces." The latter author figures a distinct ganglion to each of the caudal segments, illustrating his view from observations on Cymothoe, in which the six segments are separate, while Lereboullet illustrates the caudal ganglia as being consolidated into a single mass, from which numerous threads are sent back to the extremity of the animal. Moreover, this author only figures six separate ganglia after the cephalic mass, which would make (even allowing the oral appendages to be supplied with small filaments of nerves, instead of branches springing from * Ofversigt af Kongl. Yetenskaps-Akademiens Forhandlingar, Jan. 1859. xlviii INTRODUCTION. a well-developed ganglion), the seventh segment of the pereion to have its ganglion consolidated with those that supply the caudal region a view that our own observations lead us to believe has been founded on a misconception. GEOGRAPHICAL DISTRIBUTION. The Geographical Distribution of the two orders of the Sessile-eyed Crustacea, if made under careful and extensive observations, would (from the great amount of the modifi- cation of parts, while a close assimilation of general form is very persistent throughout great numbers of genera) afford one of the most interesting and, we believe, instructive chapters in the distribution of life over the globe. The subject has not yet sufficiently been worked out so as to approximate to correct information ; for so much of the earth's surface has yet to be searched, that it is by no means improbable that new arid intermediate forms may frequently be found in places that are yet" unknown, so that forms that as yet are described as species or genera may be only modified forms of one species, or, as has been demonstrated by M. Hesse with respect to Anceus and Pra- niza, that animals placed by authors in separate genera and in distinct families may be only sexually distinct. Such imperfect information as is at our command, while it does not enable us to grasp the subject so as to do justice to it as a whole, has yet enabled us to observe some points of interest that our British species possess in relation to exotic forms. With the exception of a single specimen, brought from Algiers by M. Lucas, the genus Talitrus is only known as an inhabitant of the northern and western coasts of Europe, while its closely allied form, Orchestia, and its congeners, excepting Nicea, of which we know but one or two species (which tend to corroborate the assertion), appears to be INTRODUCTION. xlix very abundantly scattered over the whole world. Like Talitrus, Orchestia lives out of the sea, choosing moist places, but not burrowing a habitat for itself as Talitrus does. With us, Orchestia lives within the reach of the spray of the sea ; but some species in the Southern Hemi- sphere live many miles inland, choosing terrestrial plants for their abode, sometimes at an elevation of fifteen hundred feet above the level of the sea. In these specimens the males, and, we believe, the males only, have some one or other of the joints of one of the posterior pairs of legs developed into a large internally concave scale, which, we believe, assists in retaining moisture, so that the branchial sacs may not suffer from desiccation. The genus Montagua appears to be wholly confined to the northern temperate latitudes, the species gradually diminishing in size as they approach the warmer seas. The close assimilation of this with Stegocephalus and Pleustes of the colder latitudes, is shown by the inter- change of certain parts in their structure. In Montagua, the superior antenna? have no secondary appendage, neither have the mandibles a palpiform one, and the posterior pair of pleopoda terminate in a single ramus. Pleustes resembles Montagua in the former characteristics, but has the posterior pair of pleopoda terminating in two rami. Stegocephalus resembles Pleustes in its charac- teristics, but it has a rudimentary appendage on the superior antenna?. We have little doubt but that the others have also such an appendage in the larval con- dition, since it is a common feature in young Amphi- poda. Stenothoe, in the Southern Hemisphere, represents the StegocephalidcR in the Northern, and agrees with Montagua in all important characters ; it differs in having a very large hand to the second pair of gnathopoda, a doubtful generic character, in our estimation. d 1 INTRODUCTION. The genera of the sub-family LYSTANASSINA appear to be very generally diffused over the entire globe, increasing in dimensions in those species that approach nearer to the Arctic and Antarctic latitudes, in some instances reaching to the largest known of the order, equalling three inches in length, as may be seen in Lysianassa Magellanica, from the Straits of Magellan, and L. gryllus, from Spitzbergen. These two so closely resemble one another, that they can- not be characteristically distinguished. The genus Ampelisca, and its near ally Haploops, we only know as belonging to the Northern Hemisphere, but in that region extending from Japan to Europe, from Greenland to North Carolina on the coast of America, and in Europe to the Mediterranean Sea. In the sub-family PHOXINA all the genera but one are only known in the north temperate region, but with a widely diffused area, extending from Japan to Europe. One species of the genus (Ediceros has been taken in New Zealand, and one of Iphimedia in Terra del Fuego. Of the former we have our doubts in its relation to the genus ; the latter has a very near resemblance to /. Eblance of Europe. Most of the genera of this sub-family are burrowers in mud or sand. Isaea dwells, without being parasitic, on the back of hairy crabs, and the only specimens of Darwinia, that have been taken alive,' were found adhering to the throat of a cod-fish. The genus Sulcator lives on sandy shores, making tracts along the margin of the sea, somewhat similar to those found in older slate and sandy rocks; and it may be interesting to remember that we have attributed to this sub-family the only Amphipod that has been hitherto discovered as fossil, the Prosoponiscus problematicus of the magnesian limestone of Durham, and Zechstein- dolomite of Gliicks- brun. INTRODUCTION. li The family of GAMMARID/E belongs to the Arctic and north temperate zones. With but few exceptions of the closely allied congeners Dexamine and Atylus, which consist together of twenty-one species, we know of only one taken, near Valparaiso : all the rest are northern species. Of the genus Aora but two species are known; one from the British seas, the other from the western coast of South America (Valparaiso). Judging from the figures in Gay's " Hist, de Chile," the resemblance of the two species is remarkably close, an apparently useless tooth on the anterior margin of the first pair of legs of the southern form alone distinguishing it from the northern. The subterranean fresh-water genus Niphargus, which lives generally in closed pump-wells in England and many parts of Europe, has its nearest congener in Eriopus, from the deep sea off Bohusia. Judging by the figure given by Bruzelius, there is little that distinguishes one genus from the other ; and it is highly probable that Gammarus pungens, from the warm springs of Italy, is also a species of Niphargus. Of the two species of Cran- gonyx, another fresh-water subterranean genus, one is found in England, the other in Kamschatka, and these bear a very close resemblance to the female form of the marine Gammarella, a genus, though only having three species, found in the European seas, as well as on the South American coast and at Pitt's Island. Species of the genus Melita have been taken in European, Brazilian, and Indian seas, and Moera extends all over the temperate zones of both Northern and Southern Hemispheres. The genus Amathia is essentially an Arctic form, the species losing their size and spinose character as they approach the temperate seas. No species has been recorded south of the English Channel, while a species found on the Crimean Hi INTRODUCTION. shores of the Black Sea is as large and well developed as the Arctic specimens. From Pondicherry, also, a specimen is recorded that closely resembles the large specimens of the northern type. The genus Gammarus, even as we have restricted it, contains between forty and fifty species, all of which are Arctic and north temperate, and extends round the globe, except one taken at Jamaica, another at New Holland. Fresh-water species of the genus inhabit the rivers and streams of Europe and North America, Megamcera, a near congener of Gammarus, has the largest and most spinose species in the northern regions, while others are found at Peru, Borneo, and the Zooloo seas. The genus Amphitoe contains between thirty and forty species, and is very universally spread over the globe, species having been taken in the Arctic seas and all round the coast of Europe, in the Black Sea, and the Medi- terranean ; they have been found at the Cape of Good Hope, and on the eastern and western coasts of South America, on the Australian shores, as well as in Zooloo and Japanese seas, in the islands of the Pacific and Atlantic Oceans, also on the weed in the Saragossa Sea, of the Atlantic, and on floating plants in the Pacific ; and one species is recorded from the fresh-water marshes of South Carolina. Podocerus is mostly northern, extending, however, down the coasts of Europe and America. One species is recorded by Dana from the Bay of Sunda, and another from the shores of Brazil. Cerapus, including its female, Leucothoe, has a wide range, species having been taken on the European and North American shores, on the eastern coast of South America, and in the Indian and Zooloo seas, while its near ally, Siphonacetus, has only been found on the north- INTRODUCTION. liii western shores of Europe. The genus Ncenia, all the species of which are closely allied in form, has only been recorded from the British shores. Four species of Cyr- tophium have been discovered, one of which is from the East Indies, one from Rio Janeiro, and two from the north- west of Europe. Corophium, so abundant when found, has been taken on the western shores of Europe, the Mediterranean, on the coasts of Japan and Brazil. It burrows in mud; but there is reason to doubt either that it preys on the Annelids or migrates at particular seasons. That terrible wood-destroyer, Chelura, so devastating to the piles and submarine timber all round the shores of Europe, has not been recorded from other lands. We have generally looked upon the Hyperina as pelagic species; but recently it has been pointed out by Mr. Edward, that some of our British genera burrow into and hide themselves in sand on the shores of the Moray Firth. The two British species of Hyperia (which we have great reason to believe to be but one, being male and female), have an extensive geographical range, from Greenland to Cape Horn, from Rio to the Zooloo seas. Vibilia has apparently an equally extensive range, though fewer species have been determined. Themisto, also, has been recorded from Greenland to sixty-three degrees south latitude in the Atlantic Ocean, while species of Phronima have been taken as far north as the Shet- lands, as well as in the Atlantic, at Naples, and at Borneo. The Capreltidce appear to be very universally and abundantly diffused. The very close resemblance of the species from very distant and opposite localities is suggestive of a close affinity in the respective forms. Specimens from Japan, and the eastern coast of North America, are not appreciably distinct from others found on the eastern coast of South America, as well as on our English coast ; and when we take into consideration the liv INTRODUCTION. changes in the forms that the animals of this genus undergo in their growth to an adult state, it is not im- probable that immature specimens may be misinterpreted for adult varieties. Cyamus lives parasitically on the whale, and probably thrives on no other animal. The one or two solitary specimens that have been found attached to the dolphin are probably young creatures that have strayed from their natural habitat. The genera of Isopoda appear to be more generally diffused throughout the various regions of the sea ; and from the various distant localities in which that species have been found, some may be inclined to think that they are universally distributed. The genus Tanais has been found on the coasts of North-Western Europe, Brazil, in the Zooloo and Feejee seas, as well as on the western coast of North America ; and equally varied have been the recorded habitats of the nearly allied genera, Paratanais and Leptochelia, which latter Fritz Miiller believes to be the male of Tanais. The near ally, Apseudes, is only known in Europe and Egypt, where but few specimens of two closely resem- bling species have been found. The genera Anthura and Paranthura are also sparsely represented, both in the species and specimens. They have been taken on the southern and western coasts of Europe, at New Zealand, the Mauritius, and the Cape of Good Hope, as well as on the eastern coast of North America. Of the genus Anceus, of which eleven species have been determined on the north-west coast of France, by M. Hesse, three at most are known to the rest of Europe, and but a single species to the eastern coast of North America. The Bopyrida are tolerably abundant in the temperate regions, but few in the more tropical or Arctic latitudes, the genera confining themselves with considerable exacti- tude to peculiar species of Crustacea. Thus we have failed INTRODUCTION. Iv to detect Cryptothiria, which we have found to be tolerably abundant in the genus Balanus, in Cthamalus, whose habits and general appearance are so closely allied to it. The several genera of the family ^Egida are animals peculiarly belonging to the temperate seas, and adequately represent the Cymothoidae of the torrid zone. It is remarkable that, being parasitic upon fishes, no species of the latter family has been hitherto detected on our own coasts. The Asellidce nourish chiefly in the temperate regions of the seas, being scarcely represented in the frigid zones, and not at all in the torrid. Arcturus is peculiarly an Isopod of the colder zones, where its species grow to the greatest dimensions in both the northern and southern seas ; but a single specimen has been taken in the torrid zone, in thirty-one fathoms of water, north of Borneo. The Idoteidce flourish every- where, the largest specimens being in the Baltic Sea and near Cape Horn. They live amongst the weed, either fixed or floating, and species have been often taken swimming free in mid-ocean, where they assume, as Crustacea under the same condition frequently do, a deep indigo-blue colour. The Sphceromida are a family that are very littoral in their habits; they range from the equatorial latitudes to the colder regions of the temperate zones, but die out before reaching the Arctic and Antarctic isothermal lines. In hotter latitudes, some species, in their depredations on submarine timber, take the place of Limnoria, a genus of the Asellidce, and surpass it in the extent of their capability of injuring submerged wood. Liffia, and the other terrestrial genera, appear to find their home best in the temperate latitudes, but live from the equator to within a short distance of the frigid climate. These few observations, imperfect as they naturally Ivi INTRODUCTION. must be, demonstrate, we think, the great amount of interesting information that a more complete study of the subject must elucidate. (C. S. B.) As the information conveyed in the following letter reached us too late to appear in the Appendix, we think it but just to the author to publish it entire ; the more so since, during the progress of his researches, we repeated them and know their accuracy. MY DBA* SPENCE BATE, You are kind enough to ask me for a short abstract of my investiga- tions in the anatomy of Anceidae which I tried to make when staying with you in Plymouth. I am the more glad to follow your request, since it is especially your Memoir upon these animals that made me desirous to work on them. You were quite right in directing the attention of observers to the internal structure of these little Crustacea, for there are some points in their organization which were not followed up by Mr. Hesse in his elaborate Memoir, and some points in which, your opinion differing from that of the French naturalist, we had no certainty about their real nature. I do not think that you are right in speaking of an early distinction between the male and female Anceus. There is no doubt that the outward aspect of some of the little Pranizae, just having left the parent, makes more the impression that they are to become Anceus, whilst others resemble more the female, or Praniza form. But in giving special attention to that point, I found that this impression was only due to the expansion of the segments of the pereion being greater or smaller than to any real difference. Besides that, I kept some animals, which had rather the aspect of females than males, during some time in a glass, and had the opportunity of watching their moult. Two of them enabled me to see the large projecting mandibles of the males within the head of what I thought was a female. I examined immediately the sexual parts of the t-pecimen, and found a well-developed penis on the last exceedingly small segment of the pereion. There c^n be no doubt, therefore, that Praniza changes into Anceus.* This is what Mr. Hesse contended. But though I must agree with him in this, I cannot but have another interpretation regarding the so-called larval or Praniza state. Mr. Hesse says, that only the Anceus state is the adult state, and that, "quelques jours avant la transformation des Pranizes fumelles en Ancles les ceufs qui preexistent s'apercoivent a travers la peau," fifth joint of peduncle ; 6, flagellum. d Mandible; d" mandibular ap- pendage. (II e First maxilla, or first siago-) e First nopodos* j f Second maxilla, or second) / Secon siagonopodos . . \ Maxillipede, or third siago- > g Foot- nopodos $ h 1st Gnathopodos, represent-") ing the 2nd maxillipede, > or fourth siagonopodos. J h 1st ' i 2nd Gnathopodos, represent- } ing the 3rd maxillipede, > or fifth siagonopodos . j 1st Pereiopodos t 2nd k 3rd I 2nd do. ... I 4th m 3rd do. ... m 5th n 4th do. ... n 6th a 5th do. o 7th. P 1st Pleopodos. p 1st 9 2nd do q 2nd r 3rd do r 3rd s 4th do. or 1st Uropodos * 1st t 5th do. 2nd do. t 2nd V 6th do. 3rd do. v 3rd z Telson ... z Term Foot-jaw or pedipalp. legs. natatory legs. caudal appendages. (P to Z) p 2nd do. .... 3rd do 4th do. or 1st Uropodos 5th do. 2nd do. 6th do. 3rd do. z Terminal joint or middle tail-piece. The corresponding joints of all the limbs are marked with the same number, and bear the same names throughout. We take one of the Gnathopoda as the type. 1 Coxa Coxa. I// Branchia 1"' Plate of incubatory pouch 2 Basos 3 Ischium 4 Meros; 4", inferior angle . 5 Carpus; 5", inferior angle . Thigh. Knee. Metacarpus ; 4", inferior angle. Wrist ; 5", inferior angle. 6 Propodos ; 6',palma ; 6", inferior angle ; Hand; 6', palm ; 6", inferior angle ; '" superior 6'", inferior margin ; 6"", supe- 6'", inferior margin; 6 margin. 7 Dactylos; 7", Unguis . nor margin. Finger ; 7", Nail. * This name is here suggested as the Greek equivalent for the Latin name of the five pairs of appendages succeeding the mandibles, which were collectively termed pates-mdchoires by Cuvier, Savigny, &c. The 16vres superieure and infe"rieure of these authors are omitted for the reasons assigned in the following page. (I. 0. W.) B 2 4 AMPHIPODA. The animal is naturally divided into three parts : the head (or cephalon, c), formed of a single segment ; * the body (or pereion), consisting of seven segments (H to o) ; and the tail (or pleon), formed of six segments (p to v), exclusive of the terminal scale (or telson, z). These divisions are distinctly visible, and never encroach upon each other ; while the appendages assume characteristic forms in each division. Those which belong to the head are more or less connected with the organs of sense. The eyes (a) are sessile and compound. Their normal position is between the bases of the superior and inferior antennae. In the Orchestiidae and near allies they are on the top of the head, to which position they are thrust by the great increase of the size of the two basal articula- tions of the antennae and their absorption into the ante- rior portion of the head. The outer integument of the eyes is never divided into facets, except in some genera of the Hyperina. In many of the Phoxides the eyes appear to be wanting ; but this is probably caused by the absence of any colouring pigment, or its dispersion after death, rather than from the absence of the organ of vision. In Ampelisca they appear like four simple organs, resembling the ocelli of true insects. The anterior or superior pair of antennae (U] are formed * Adopting the theory of Oken, that each pair of limbs or organs indicates a separate segment (often, indeed, coalescing with the adjacent one), the head would consist of nine segments, namely : 1st, that supporting the eye ; 2nd, the upper antennae ; 3rd, the lower antennae ; 4th, the upper lip, formed of two lateral halves united ; 5th, the mandibles, or jaws ; 6th, the lower lip, formed like the upper lip ; 7th, the first pair of maxillae ; 8th, the second pair of maxillae ; and 9th, the foot-jaws. If to these are added the seven segments of the body, the six segments of the tail, and the segment represented by the terminal scale, we have twenty-three segments as the normal number in the Ampldpoda. As, however, Mr. Spence Bate regards the two lips merely as the calcined extremities of the alimentary canal, the number of head -segments would be reduced to seven, and the entire number to twenty-one. (I. 0. W.) GENERAL CHARACTERS. 5 of a peduncle consisting of three joints, and a terminal multiarticulate filament, supplied with auditory cilia. Oc- casionally there is a second appendage, generally rudimen- tary, but in some of the Phoxides it is of almost equal importance to the primary filament ; the secondary ap- pendage is, however, never furnished with auditory cilia. The posterior or inferior pair of antennae (c) consists of a peduncle of five joints and a multiarticulate fila- ment. The first two joints are closely incorporated, and bear an olfactory denticle ; but sometimes, as in Talitrus, the denticle is wanting, probably from the peculiarity of its condition of existence, as the Talitri do not live in water. Occasionally the terminal filament has the joints fused together. This numerical decrease is invariably attended with an increase of strength. In some genera of the Hyperina both pairs of antennae assume a rudimentary condition to such an extent that their respective parts cannot be readily defined : some- times even one or both may be absent. The jaws or mandibles (d) are placed between an anterior and posterior lip or labium ; they consist of a pair of curved triangular blades, each furnished with a cutting edge and a grinding tubercle. Within the inci- sive margin there is frequently a second movable plate, formed upon the type of the preceding. Each mandible has very generally, though not universally, an articulated palpus or appendage. The anterior pair of maxillae (e) consist of three or four foliaceous plates, whereas the second pair (/) have but two ; they are extremely deli- cate, and furnished, upon their anterior margins, with plumose hairs, some of which are strengthened into spines of various shapes. Exceptions to the normal forms exist in the Hyperina. The foot-jaws, or pedipalps (siagonopoda, I. O. W., or maxillipedes) (g) t are the posterior pair of appendages 6 AMPHIPODA. attached to the head ; they have some of the joints foli- aceous. They overlap the preceding appendages of the mouth, and act as a protecting operculum. In the Hy- perina they are small, and do not overlap the whole of the buccal apparatus. The two anterior pairs of legs (gnathopoda, h and z), which in the Podophthalmata are reduced in size, and employed as two additional pairs of foot-jaws or pedi- palps, are here developed into arm-like legs, and are attached to the two anterior segments of the body. They are directed forwards, and generally formed upon the same type, the posterior being the larger ; but to this general rule there are several exceptions. The sixth joint (or propodos, e) is generally enlarged into a hand in both pairs, against the inferior margin of which the seventh joint (or dactylos, 7) doubles back, as a finger against the palm, and impinging against it, gives to the organ a prehensile capability. Sometimes the fifth joint (or carpus, 5), and also the fourth (or meros, 4), are infe- riorly produced, so as to assist in prehension. These ap- pendages seldom attain the form of the analogous chelae in the higher orders ; QalUsoma, Chelura, and one or two others, being the few exceptions to this very general law. All the legs have the first joint * (or coxa, i) developed into a large and squamiform plate, which covers a con- siderable portion of the second joint, and protects the branchial organs (figure *, i" and i'" in p. 2,), as well as the ova and embryos while confined within the incuba- tory pouch during the period of gestation. In the four * By preceding writers, the series of scale-like plates at the sides of the body have been regarded as the homologues of the epimera of the thoracic segments of the Insecta. Mr. Spence Bate, however, considers them as the first joints of the legs, thus dilated for special purposes in the economy of the animals, an opinion which has been accepted by Professors Huxley, Kinahan, and others. (I. 0. W.) The reader will observe that we employ the term joint for a portion of a limb, and articulation for the connecting hinge. GENERAL CHARACTERS. 7 anterior (h, i, k, I), and in some genera the fifth (w,) pairs of limbs, these plates are much larger than in the three or two posterior pairs ; but to compensate for this diminu- tion of size, the three posterior (m, n t o), with few generic exceptions, have the second joint produced posteriorly into a large and squamiform plate. In Caprella and the allied genera, the first joint of each leg is fused with the body of the animal, and is never shaped like scales. The five posterior pairs of legs (pereiopoda, k-o) are the walking appendages ; they homologize with the ten legs in the Decapoda, and as efficiently fulfil their design. Like them, they consist of seven joints; but, unlike them, all articulate in planes vertical to the body of the animal, having no lateral movements. The two ante- rior pairs of walking legs are directed forwards, and the three posterior are directed backwards. Thus the seven pairs of legs constitute three distinct series, generally differing from each other in their proportions, size, and direction, the first and second pairs being subcheliferous, the third and fourth porrected, and the fifth, sixth, and seventh pairs directed backwards. The branchiae (i") consist of a series of vesicles ; a single sac being attached to, and pendent from, the first joint of all the legs except the first, and in the males of some genera also the last. The ova are nurtured within a pouch formed by a series of foliaceous plates (i'") one of which is attached to the first joint of the four anterior pairs of legs in females. In this pouch the embryo continues until it has arrived at a period when there is but a slight dis- tinction in form between it and the parent, except in the Hyperina, as has been shown by Milne-Edwards and Gosse, where the form of the young animal differs considerably from that of the parent. Posterior to the legs used for walking, are three pairs 8 AMPHIPODA. of flexible appendages (1st, 2nd, and 3rd pleopoda, jo, q, r), consisting of a base and two multiarticulate fila- ments or rami fringed with plumose cilia. These are used in swimming, and, powerful for such a purpose, they propel the animal rapidly through the water. Succeed- ing to these are three pairs of appendages (4th, 5th, and 6th pleopoda, s,t 9 v, uropoda, I. O. W.), situated upon the inferior lateral margins of their respective segments. They consist of a single-jointed base, supporting two uniarticulate branches, inflexible and styliform, fringed with spines or hairs. The posterior pair vary consider- ably in form ; in some genera they are furnished with hooks, whilst in others they assume a foliaceous character, circumstances which render them valuable in the recog- nition of species. These appendages are also powerful organs of propulsion. By folding the tail beneath the body, and suddenly striking it out again, those animals which exist in the water, as well as those which live on the shore, are enabled to dart or leap to a considerable distance. In Caprella and its near allies, the whole of the appendages of the tail are absent, or present only in a rudimentary and altered condition. The terminal segment of the animal (telson, z] is represented by an imperfect or rudimentary appendage or scale. From the great variety of shapes which this appendage assumes, it becomes a valuable aid in the determination of genera. The typical form may be con- sidered to be that of an acute-angled triangular scale, the apex being rounded off. Sometimes it is divided into two, as in Gammarus ; again, it is deeply cleft; in one genus it is represented by a hook only ; in some, it is broad, flat, and foliaceous, in others it is cylindrical, the intestinal canal terminating at its extremity. Compared with the podophthalmatous Crustaceans, the animals forming the present order are of small size, the GENERAL CHARACTERS. 9 great majority being less than an inch long, and none exceeding thrice that length. Like all the productions of nature of diminished size, the number of indivi- duals of different species far exceeds that of the larger- sized Crustacea. With few exceptions, there is but little external difference of form between the opposite sexes ; the males, however, contrary to the ordinary rule in the Annulosa, being often larger than the females. There is also but little difference in form between the young and adult individuals of the several species, except where marked characters exist, such as the enlarged form of the hands, or spines on the different segments of the body, which increase in size as the animals become older. The species occur in temperate or high latitudes in greater proportion than in tropical climates. Thus it will be perceived that among the Amphipoda there is a considerable variety of form, some keeping closer to the typical idea of the Order, while others vary more or less considerably. It is therefore desirable, both for clearness of expression and in order to obtain a better knowledge of the whole, that we should arrange together those which more nearly assimilate to each other ; whereas others, which vary in a greater or less degree, should be grouped according to their respective details. In the works of Leach, Latreille, Milne-Edwards, Kroyer, Dana, Zaddach, Liljeborg, and Bruzelius, various modes of classification of these animals have been pro- posed. Based upon a consideration of these various ar- rangements, as well as upon the structure and respective habits of the different animals (resulting from a consider- able observation of their economy and modes of life), and having had the advantage of studying the types of Montagu, Leach, Phipps, M. -Edwards, and others, we have drawn up the following tabular distribution of the Order : 10 GENERAL CLASSIFICATION. I II l4IJl*IE4l tt fin fifi OQHJ ^ 't Ira P^ 2 a^ rSrs ^ 111 ^ && ft o o si i cc-fi ' ! I o AMPIilPODA. 11 Group NOBMALIA. This group comprises the typical Amphipoda, and in- cludes the whole of the order except those forms which have some of the appendages absent, or have the first joint of the legs absorbed into the body of the animal. It is synonymous with the order of Amphipoda of Latreille, Milne-Edwards, &c., and is separated into two divisions namely, GAMMARINA and HYPERINA. Division GAMMARINA. Distinguished from the other division of the group by the smallness of the eyes, the normal form of the antennae, the size of the foot-jaws, and the largeness of the squa- miform plates of the legs. This division corresponds with Milne-Edwards's family of Crevettines, and also with that of Gammaridea of Dana. It is subdivided into VAGANTIA and DOMICOLA. Subdivision VAGANTIA. The VAGANTIA have always the inferior antennae ter- minating in a flexible multiarticulate flagellum, and the posterior pair of caudal appendages are fringed with fine hairs or short spines, and are never furnished with hooks. They construct no abodes to dwell in, but wander from place to place, swimming or walking in the sea, and hopping or wriggling along upon their sides on land. This subdivision represents Milne-Edwards's tribe of Sauteurs. It consists of the two tribes SALTATORIA and NATATQRIA, each consisting of a single family. Tribe SALTATORIA. In this tribe the posterior caudal appendages are never longer than the two preceding pairs. The hairs upon the 12 ORCHESTIIDJ1. animal are short and stiff. The mode of progression, when out of the water, is by leaps. The animals are generally terrestrial or subaquatic. This tribe contains but a single family. Fam. ORCHESTIID^. The superior antennae are never longer than the inferior, and never furnished with a secondary appendage. The mandibles are without a palpiform appendage. The squamiform joints of the legs are largely developed. The caudal appendages are very short and stout, and the pos- terior pair only possess a single branch. The following vignette represents the Hoe Gate, Ply- mouth, the house in which Dr. Leach was born. TALITRUS. 13 A MPHIP DA . ORCHESTIID+E. SALTATORIA. Genus TALITRUS, LATREILLE. Generic character. Superior antennas short and rudimentary. Inferior antennae with the basal joints fused into the facial \vall of the cephalon. Mandibular palpi obsolete. Maxillipedes not unguiculate. First pair of gnathopoda simple ; second pair small and feeble. Coxae of third pair of pereiopoda as deep as the coxse of the second, and divided into two equal lobes. Telson rudimentary.* THE eyes are near the top of the head. The superior antennae are very short, not reaching to the extremity of the second free joint of the inferior. Inferior antennae with the two basal joints absorbed into the frontal wall" of the head. Fingers of the foot-jaws not unguiculate. First pair of legs not having a subchelate hand in either sex : second pair of legs smaller than the first pair, and imperfectly subchelate in both sexes. Coxae of the fifth pair of legs subequally bilobed, and nearly as deep as the coxae of the preceding pair. Middle scale of the tail rudimentary or single. The genus Talitrus was first proposed by Latreille in his "Precis" (1796), under the name of GammaruSj for the reception of the Amphipoda with short upper an- tennae ; the remainder, with longer upper antennae, being arranged in his genus Carcinus. The name Talitrus itself * The structural terms employed in the short generic and specific characters of the AmpUpoda, are those proposed by Mr. Spence Bate in his " Report on the British Edriophthalma," published in the Reports of the British Asso- ciation for 1855. In the text the ordinary English names of the various parts are adopted, as given in the Table of External Organs in page 3. 14 ORCHESTI1DJE. first appears in the year 1802, both in the third volume of Latreille's " Histoire generale des Crustaces et Insectes," and in the second volume of Bosc's " Hist. nat. des Crustaces," the latter writer giving Latreille the credit of the invention. This must be borne in mind, because Latreille, in his "Genera Crust, et Ins.," vol. i., 1806, refers the genus Talitrus to Bosc as its author. In the last-mentioned work we find the genus, according to the views of its founder, to be as extensive as our family Or- chestiidae (which it would consequently have been more correct to have named, after the present genus, Talitridse), embracing the whole of the saltatorial species. Subse- quently Leach separated the species with the first pair of legs cheliferous under the name of Orchestia. In this he has been followed by all subsequent writers. Milne- Edwards, Dana, Desmarest, and others, however, intro- duced into this genus those species which have the second pair of hands as large as in the males of Orchestia ; but Nicolet* has very justly separated them from Talitrus, under the generic name of Orchestoidea. Brandt f has likewise done the same, but, without being aware of what Nicolet had proposed, has given to the same genus the name of Megalorchestia, which Stimpson has followed. Accepting this latter separation of the species into two genera, Talitrus appears to be peculiar to the European coasts and the southern shores of the Mediterranean. The species T. brevicornis of Edwards and T. Novi-Zealandi