Crustacea

External Structure: Body.—As in all Arthropoda the body consists of a series of segments or somites which may be free or more or less coalesced together. In its simplest form the exoskeleton of a typical somite is a ring of chitin defined from the rings in front and behind by areas of thinner integument forming moveable joints, and having a pair of appendages articulated to its ventral surface on either side of the middle line. Frequently, however, this exoskeletal somite may be differentiated into various regions. A dorsal and a ventral plate are often distinguished, known respectively as the tergum and the sternum, and the tergum may overhang the insertion of the limb on each side as a free plate called the pleuron. The name epimeron is sometimes applied to what is here called the pleuron, but the word has been used in widely different senses and it seems better to abandon it. The typical form of a somite is well seen, for example, in the segments which make up the abdomen or “tail” of a lobster or crayfish (fig. 2). The posterior terminal segment of the body, on which the opening of the anus is situated, never bears appendages. The nature of this segment, which is known as the “anal segment” or telson (fig. 3, T), has been much discussed, some authorities holding that it is a true somite, homologous with those which precede it. Others have regarded it as representing the fusion of a number of somites, and others again as a “median appendage” or as a pair of appendages fused. Its morphological nature, however, is clearly shown by its development. In the larval development of the more primitive Crustacea, the number of somites, at first small, increases by the successive appearance of new somites between the last-formed somite and the terminal region which bears the anus. The “growing point” of the trunk is, in fact, situated in front of this region, and, when the full number of somites has been reached, the unsegmented part remaining forms the telson of the adult.

Fig. 3.—The Separated Somites and Appendages of the Common Lobster (Homarus gammarus).
C, carapace covering the cephalothorax. Ab, abdominal somites. T, telson, having the uropods or appendages of the last abdominal somite spread out on either side of it, forming the “tail-fan.” l, labrum, or upper lip. m, metastoma, or lower lip. 1, eyes. 2, antennule (the arrow points to the opening of the so-called auditory organ). 3, antenna. 4, mandible.	5, maxillula (or first maxilla). 6, maxilla (second maxilla). 7-9, first, second and third maxillipeds. ex, exopodite. ep, epipodite. g, gill. 10, sixth thoracic limb (second walking-leg) of female. 11, last thoracic limb of male. In 10 and 11 the arrows indicate the genital apertures. 13, sterna of the thoracic somites, from within. 14, third abdominal somite, with appendages or “swimmerets.”

In no Crustacean, however, do all the somites of the body remain distinct. Coalescence, or suppression of segmentation (“lipomerism”), may involve more or less extensive regions. This is especially the case in the anterior part of the body, where, in correlation with the “adaptational shifting of the oral aperture” (see Arthropoda), a varying number of somites unite to form the “cephalon” or head. Apart from the possible existence of an ocular somite corresponding to the eyes (the morphological nature of which is discussed below), the smallest number of head-somites so united in any Crustacean is five. Even where a large number of the somites have fused, there is generally a marked change in the character of the appendages after the fifth pair, and since the integumental fold which forms the carapace seems to originate from this point, it is usual to take the fifth somite as the morphological limit of the cephalon throughout the class. It is quite probable, however, that in the primitive ancestors of existing Crustacea a still smaller number of somites formed the head. The three pairs of appendages present in the “nauplius” larva show certain peculiarities of structure and development which seem to place them in a different category from the other limbs, and there is some ground for regarding the three corresponding somites as constituting a “primary cephalon.” For practical purposes, however, it is convenient to include the two following somites also as cephalic.

Fig. 4.—Diagram of an Amphipod. (After Spence Bate and Westwood.)
C, cephalon. Th, thorax. (Only seven of the eight thoracic somites are visible, the first being fused with the cephalon.)	Ab, abdomen. The numbers appended to the somites do not correspond to the enumeration adopted in the text. 21 is the telson.

A remarkable feature found only in the Stomatopoda is the reappearance of segmentation in the anterior part of the cephalic region. Whether the movably articulated segments which bear the eye-stalks and the antennules in this aberrant group correspond to the primitive head somites or not, their distinctness is certainly a secondarily acquired character, for it is not found in the larvae, nor in any of the more primitive groups of Malacostraca.

The body proper is usually divisible into two regions to which the names thorax and abdomen are applied. Throughout the whole of the Malacostraca the thorax consists of eight and the abdomen of six somites (fig. 4), and the two regions are sharply distinguished by the character of their appendages. In the various groups of the Entomostraca, on the other hand, the terms thorax and abdomen, though conveniently employed for purposes of systematic description, do not imply any homology with the regions so named in the Malacostraca. Sometimes they are applied, as in the Copepoda, to the limb-bearing and limbless regions of the trunk, while in other cases, as in the Phyllopoda, they denote, respectively, the regions in front of and behind the genital apertures.

Fig. 5.—Phyllopoda and Phyllocarida.
1, Ceratiocaris papilio, U. Silurian, Lanark. 2, Nebalia bipes(one side of carapace removed). 3, Lepidurus Angassi: a, dorsal aspect; b, ventral aspect of head showing the labrum and mouth-parts.	4, larva of Apus cancriformis. 5, Branchipus stagnalis: a, adult female; b, first larval stage (Nauplius); c, second larval stage. 6, Nauplius of Artemia salina.

A character which recurs in the most diverse groups of the Crustacea, and which is probably to be regarded as a primitive attribute of the class, is the possession of a carapace or shell, arising as a dorsal fold of the integument from the posterior margin of the head-region. In its most primitive form, as seen in the Apodidae (fig. 5, 3) and in Nebalia (fig. 5, 2), this shell-fold remains free from the trunk, which it envelops more or less completely. It may assume the form of a bivalve shell entirely enclosing the body and limbs, as in many Phyllopoda (fig. 6) and in the Ostracoda. In the Cirripedia it forms a fleshy “mantle” strengthened by shelly plates or valves which may assume a very complex structure. In many cases, however, the shell-fold coalesces with some of the succeeding somites. In the Decapoda (fig. 3), this coalescence affects only the dorsal region of the thoracic somites, and the lateral portions of the carapace overhang on each side, enclosing a pair of chambers within which lie the gills. The arrangement is similar in Schizopoda and Stomatopoda (fig. 7), except that the coalescence does not usually involve the posterior thoracic somites, several of which remain free, though they may be overlapped by the carapace.

From Morse’s Zoology.

Fig. 6.—Estheria, sp.; D from Dubuque, Iowa; (e) the eye. L from Lynn, Massachusetts (nat. size). S presents a highly magnified section of one of the valves to show the successive moults. B an enlarged portion of the edge of the shell along the back, showing the overlap of each growth.

In the Isopoda and Amphipoda, where, as a rule, all the thoracic somites except the first are distinct (fig. 4), there seems at first sight to be no shell-fold. A comparison with the related Tanaidacea (fig. 8) and Cumacea (or Sympoda), however, leads to the conclusion that the coalescence of the first thoracic somite with the cephalon really involves a vestigial shell-fold, and, indeed, traces of this are said to be observed in the embryonic development of some Isopoda. It seems likely that a similar explanation is to be applied to the coalescence of one or two trunk-somites with the head in the Copepoda, and, if this be so, the only Crustacea remaining in which no trace of a shell-fold is found in the adult are the Anostracous Phyllopoda such as Branchipus (fig. 5, 5).

Fig. 7.—Squilla mantis (Stomatopoda), showing the last four thoracic (leg-bearing) somites free from the carapace.

General Morphology of Appendages.—Amid the great variety of forms assumed by the appendages of the Crustacea, it is possible to trace, more or less plainly, the modifications of a fundamental type consisting of a peduncle, the protopodite, bearing two branches, the endopodite and exopodite. This simple biramous form is shown in the swimming-feet of the Copepoda and Branchiura, the “cirri” of the Cirripedia, and the abdominal appendages of the Malacostraca (fig. 3, 14). It is also found in the earliest and most primitive form of larva, known as the Nauplius. As a rule the protopodite is composed of two segments, though one may be reduced or suppressed and occasionally three may be present. In many cases, one of the branches, generally the endopodite, is more strongly developed than the other. Thus, in the thoracic limbs of the Malacostraca, the endopodite generally forms a walking-leg while the exopodite becomes a swimming-branch or may disappear altogether. Very often the basal segment of the protopodite bears, on the outer side, a lamellar appendage (more rarely, two), the epipodite, which may function as a gill. In the appendages near the mouth one or both of the protopodal segments may bear inwardly-turned processes, assisting in mastication and known as gnathobases. The frequent occurrence of epipodites and gnathobases tends to show that the primitive type of appendage was more complex than the simple biramous limb, and some authorities have regarded the leaf-like appendages of the Phyllopoda as nearer the original form from which the various modifications found in other groups have been derived. In a Phyllopod such as Apus the limbs of the trunk consist of a flattened, unsegmented or obscurely segmented axis or corm having a series of lobes or processes known as endites and exites on its inner and outer margins respectively. In all the Phyllopoda the number of endites is six, and the proximal one is more or less distinctly specialized as a gnathobase, working against its fellow of the opposite side in seizing food and transferring it to the mouth. The Phyllopoda are the only Crustacea in which distinct and functional gnathobasic processes are found on appendages far removed from the mouth. The two distal endites are regarded as corresponding to the endopodite and exopodite of the higher Crustacea, the axis or corm of the Phyllopod limb representing the protopodite. The number of exites is less constant, but, in Apus, two are present, the proximal branchial in function and the distal forming a stiffer plate which probably aids in swimming. It is not altogether easy to recognize the homologies of the endites and exites even within the order Phyllopoda, and the identification of the two distal endites as corresponding to the endopodite and exopodite of higher Crustacea is not free from difficulty. It is highly probable, however, that the biramous limb is a simplification of a more complex primitive type, to which the Phyllopod limb is a more or less close approximation.

Fig. 8.—Tanais dubius (?) Kr. ♀, showing the orifice of entrance (x) into the cavity overarched by the carapace in which an appendage of the maxilliped (f) plays. On four feet (i, k, l, m) are the rudiments of the lamellae which subsequently form the brood-cavity. (Fritz Müller.)

Fig. 9.—A, Balanus (young), side view with cirri protruded. B, Upper surface of same; valves closed. C, Highly magnified view of one of the cirri. (Morse.)

The modifications which this original type undergoes are usually more or less plainly correlated with the functions which the appendages have to discharge. Thus, when acting as swimming organs, the appendages, or their rami, are more or less flattened, or oar-like, and often have the margins fringed with long plumose hairs. When used for walking, one of the rami, usually the inner, is stout and cylindrical, terminating in a claw, and having the segments united by definite hinge-joints. The jaws have the gnathobasic endites developed at the expense of the rest of the limb, the endopodite and exopodite persisting only as sensory “palps” or disappearing altogether. When specialized as bearers of sensory (olfactory or tactile) organs, the rami are generally elongated, many-jointed and flagelliform. This modification is usually only found in the antennules and antennae, but it may exceptionally be found in the appendages of the trunk, as, for instance, in the thoracic legs of some Decapods (e.g. Mastigocheirus). Very often one or other of the appendages may be modified for prehension, the seizing of prey or the holding of a mate. In this case, the claw-like terminal segment may be simply flexed against the preceding in the same way as the blade of a penknife shuts up against the handle. The penultimate segment is often broadened, so that the terminal claw shuts against a transverse edge (fig. 4), or, finally, the penultimate segment may be produced into a thumb-like process opposed to the movable terminal segment or finger, forming a perfect chela or forceps, as, for instance, in the large claws of a crab or lobster. This chelate condition may be assumed by almost any of the appendages, and sometimes it appears in different appendages in closely related forms, so that no very great phylogenetic importance can in most cases be attached to it. A peculiar modification is found in the trunk-limbs of the Cirripedia (fig. 9), in which both rami are multiarticulate and filiform and fringed with long bristles. When protruded from the opening of the shell these “cirri” are spread out to form a casting-net for the capture of minute floating prey.

Gills or branchiae may be developed by parts of an appendage becoming thin-walled and vascular and either expanded into a thin lamella or ramified. Some of the special modifications of branchiae are referred to below.

Special Morphology of Appendages.—In many Crustacea the eyes are borne on stalks which are movably articulated with the head and which may be divided into two or three segments. The view is commonly held that these eye-stalks are really limbs, homologous with the other appendages. In spite of much discussion, however, it cannot be said that this point has been finally settled. The evidence of embryology is decidedly against the view that the eye-stalks are limbs. They are absent in the earliest and most primitive larval forms (nauplius), and appear only late in the course of development, after many of the trunk-limbs are fully formed. In the development of the Phyllopod Branchipus, the eyes are at first sessile, and the lateral lobes of the head on which they are set grow out and become movably articulated, forming the peduncles. The most important evidence in favour of their appendicular nature is afforded by the phenomena of regeneration. When the eye-stalk is removed from a living lobster or prawn, it is found that under certain conditions a many-jointed appendage like the flagellum of an antennule or antenna may grow in its place. It is open to question, however, how far the evidence from such “heteromorphic regeneration” can be regarded as conclusive on the points of homology. The fact that in certain rare cases among insects a leg may apparently be replaced by a wing tends to show that under exceptional conditions similar forms may be assumed by non-homologous parts.

The antennules (or first antennae) are almost universally regarded as true appendages, though they differ from all the other appendages in the fact that they are always innervated from the “brain” (or preoral ganglia), and that they are uniramous in the nauplius larva and in all the Entomostracan orders. As regards their innervation an apparent exception is found in the case of Apus, where the nerves to the antennules arise, behind the brain, from the oesophageal commissures, but this is, no doubt, a secondary condition, and the nerve-fibres have been traced forwards to centres within the brain. In the Malacostraca, the antennules are often biramous, but there is considerable doubt as to whether the two branches represent the endopodite and exopodite of the other limbs, and three branches are found in the Stomatopoda and in some Caridea. In the great majority of Crustacea the antennules are purely sensory in function and carry numerous “olfactory” hairs. They may, however, be natatory as in many Ostracoda and Copepoda, or prehensile, as in some Copepoda. The most peculiar modification, perhaps, is that found in the Cirripedia (Thyrostraca), in the larvae of which the antennules develop into organs of attachment, bearing the openings of the cement-glands, and becoming, in the adult, involved in the attachment of the animal to its support.

The antennae (second antennae) are of special interest on account of the clear evidence that, although preoral in position in all adult Crustacea, they were originally postoral appendages. In the nauplius larva they lie rather at the sides than in front of the mouth, and their basal portion carries a hook-like masticatory process which assists the similar processes of the mandibles in seizing food. In the primitive Phyllopoda, and less distinctly in some other orders, the nerves supplying the antennae arise, not from the brain, but from the circum-oesophageal commissures, and even in those cases where the nerves and the ganglia in which they are rooted have been moved forwards to the brain, the transverse commissure of the ganglia can still be traced, running behind the oesophagus.

The functions of the antennae are more varied than is the case with the antennules. In many Entomostraca (Phyllopoda, Cladocera, Ostracoda, Copepoda) they are important, and sometimes the only, organs of locomotion. In some male Phyllopoda they form complex “claspers” for holding the female. They are frequently organs of attachment in parasitic Copepoda, and they may be completely pediform in the Ostracoda. In the Malacostraca they are chiefly sensory, the endopodite forming a long flagellum, while the exopodite may form a lamellar “scale,” probably useful as a balancer in swimming, or may disappear altogether. A very curious function sometimes discharged by the antennules or antennae of Decapods is that of forming a respiratory siphon in sand-burrowing species.

The mandibles, like the antennae, have, in the nauplius, the form of biramous swimming limbs, with a masticatory process originating from the proximal part of the protopodite. This form is retained, with little alteration in some adult Copepoda, where the biramous “palp” still aids in locomotion. A somewhat similar structure is found also in some Ostracoda. In most cases, however, the palp loses its exopodite and it often disappears altogether, while the coxal segment forms the body of the mandible, with a masticatory edge variously armed with teeth and spines. In a few Ostracoda, by a rare exception, the masticatory process is reduced or suppressed, and the palp alone remains, forming a pediform appendage used in locomotion as well as in the prehension of food. In parasitic blood-sucking forms the mandibles often have the shape of piercing stylets, and are enclosed in a tubular proboscis formed by the union of the upper lip (labrum) with the lower lip (hypostome or paragnatha).

The maxillulae and maxillae (or, as they are often termed, first and second maxillae) are nearly always flattened leaf-like appendages, having gnathobasic lobes or endites borne by the segments of the protopodite. The endopodite, when present, is unsegmented or composed of few segments and forms the “palp,” and outwardly-directed lobes representing the exopodite and epipodites may also be present. These limbs undergo great modification in the different groups. The maxillulae are sometimes closely connected with the “paragnatha” or lobes of the lower lip, when these are present, and it has been suggested that the paragnatha are really the basal endites which have become partly separated from the rest of the appendage.

The limbs of the post-cephalic series show little differentiation among themselves in many Entomostraca. In the Phyllopoda they are for the most part all alike, though one or two of the anterior pairs may be specialized as sensory (Apus) or grasping (Estheriidae) organs. In the Cirripedia (Thyrostraca) the six pairs of biramous cirriform limbs differ only slightly from each other, and in many Copepoda this is also the case. In other Entomostraca considerable differentiation may take place, but the series is never divided into definite “tagmata” or groups of similarly modified appendages. It is highly characteristic of the Malacostraca, however, that the trunk-limbs are divided into two sharply defined tagmata corresponding to the thoracic and abdominal regions respectively, the limit between the two being marked by the position of the male genital openings. The thoracic limbs have the endopodites converted, as a rule, into more or less efficient walking-legs, and the exopodites are often lost, while the abdominal limbs more generally preserve the biramous form and are, in the more primitive types, natatory. These tagmata may again be subdivided into groups preserving a more or less marked individuality. For example, in the Amphipoda (fig. 4) the abdominal appendages are constantly divided into an anterior group of three natatory “swimmerets” and a posterior group of three limbs used chiefly in jumping or in burrowing. In nearly all Malacostraca the last pair of abdominal appendages (uropods) differ from the others, and in the more primitive groups they form, with the telson, a lamellar “tail-fan” (fig. 3, T), used in springing backwards through the water. In the thoracic series it is usual for one or more of the anterior pairs to be pressed into the service of the mouth, forming “foot-jaws” or maxillipeds. In the Decapoda three pairs are thus modified, and in the Tanaidacea, Isopoda and Amphipoda only one. In the Schizopoda and Cumacea the line of division is less sharp, and the varying number of so-called maxillipeds recognized by different authors gives rise to some confusion of terminology in systematic literature.

Gills.—In many of the smaller Entomostraca (Copepoda and most Ostracoda) no special gills are present, and respiration is carried on by the general surface of the body and limbs. When present, the branchiae are generally differentiations of parts of the appendages, most often the epipodites, as in the Phyllopoda. In the Cirripedia, however, they are vascular processes from the inner surface of the mantle or shell-fold, and in some Ostracoda they are outgrowths from the sides of the body. In the primitive Malacostraca the gills were probably, as in the Phyllopoda and in Nebalia, the modified epipodites of the thoracic limbs, and this is the condition found in some Schizopoda. In the Cumacea and Tanaidacea only the first thoracic limb has a branchial epipodite. In the Amphipoda, the gills though arising from the inner side of the bases of the thoracic legs are probably also epipodial in nature. In the Isopoda the respiratory function has been taken over by the abdominal appendages, both rami or only the inner becoming thin or flattened. In the Decapoda the branchial system is more complex. The gills are inserted at the base of the thoracic limbs, and lie within a pair of branchial chambers covered by the carapace. Three series are distinguished, podobranchiae, attached to the proximal segments of the appendages, pleurobranchiae, springing from the body-wall, and an intermediate series, arthrobranchiae, inserted on the articular membrane of the joint between the limb and the body. The podobranchiae are clearly epipodites, or, more correctly, parts of the epipodites, and it is probable that the arthro- and pleurobranchiae are also epipodial in origin and have migrated from the proximal segment of the limbs on to the adjacent body-wall.

Adaptations for aërial respiration are found in some of the land-crabs, where the lining membrane of the gill-chamber is beset with vascular papillae and acts as a lung. In some of the terrestrial Isopoda or woodlice (Oniscoidea) the abdominal appendages have ramified tubular invaginations of the integument, filled with air and resembling the tracheae of insects.

Internal Structure: Alimentary System.—In almost all Crustacea the food-canal runs straight through the body, except at its anterior end, where it curves downwards to the ventrally-placed mouth. In a few cases its course is slightly sinuous or twisted, but the only cases in which it is actually coiled upon itself are found in the Cladocera of the family Lynceidae (Alonidae) and in a single recently-discovered genus of Cumacea (Sympoda). As in all Arthropoda, it is composed of three divisions, a fore-gut or stomodaeum, ectodermal in origin and lined by an inturning of the chitinous cuticle, a mid-gut formed by endoderm and without a cuticular lining, and a hind-gut or proctodaeum, which, like the fore-gut, is ectodermal and is lined by cuticle. The relative proportions of these three divisions vary considerably, and the extreme abbreviation of the mid-gut found in the common crayfish (Astacus) is by no means typical of the class. Even in the closely-related lobster (Homarus) the mid-gut may be 2 or 3 in. long.

In a few Entomostraca (some Phyllopoda and Ostracoda) the chitinous lining of the fore-gut develops spines and hairs which help to triturate and strain the food, and among the Ostracods there is occasionally (Bairdia) a more elaborate armature of toothed plates moved by muscles. It is among the Malacostraca, however, and especially in the Decapoda, that the “gastric mill” reaches its greatest perfection. In most Decapods the “stomach” or dilated portion of the fore-gut is divided into two chambers, a large anterior “cardiac” and a smaller posterior “pyloric.” In the narrow opening between these, three teeth (fig. 10) are set, one dorsally and one on each side. These teeth are connected with a framework of movably articulated ossicles developed as thickened and calcified portions of the lining cuticle of the stomach and moved by special muscles in such a way as to bring the three teeth together in the middle line. The walls of the pyloric chamber bear a series of pads and ridges beset with hairs and so disposed as to form a straining apparatus.

The mid-gut is essentially the digestive and absorptive region of the alimentary canal, and its surface is, in most cases, increased by pouch-like or tubular outgrowths which not only serve as glands for the secretion of the digestive juices, but may also become filled by the more fluid portion of the partially digested food and facilitate its absorption. These outgrowths vary much in their arrangement in the different groups. Most commonly there is a pair of lateral caeca, which may be more or less ramified and may form a massive “hepato-pancreas” or “liver.”

Fig. 10.—Gastric Teeth of Crab and Lobster.

1a, Stomach of common crab, Cancer pagurus, laid open, showing b, b, b, some of the calcareous plates inserted in its muscular coat; g, g, the lateral teeth, which when in use are brought in contact with the sides of the median tooth m; c, c, the muscular coat. 1b′ and 1b″, The gastric teeth enlarged to show their grinding surfaces. 2, Gastric teeth of common lobster, Homarus vulgaris. 3a and 3b, Two crustacean teeth (of Dithyrocaris) from the Carboniferous series of Renfrewshire (these, however, may be the toothed edges of the mandibles).

The whole length of the alimentary canal is provided, as a rule, with muscular fibres, both circular and longitudinal, running in its walls, and, in addition, there may be muscle-bands running between the gut and the body-wall. In the region of the oesophagus these muscles are more strongly developed to perform the movements of deglutition, and, where a gastric mill is present, both intrinsic and extrinsic muscles co-operate in producing the movements of its various parts. The hind-gut is also provided with sphincter and dilator muscles, and these may produce rhythmic expansion and contraction, causing an inflow and outflow of water through the anus, which has been supposed to aid in respiration.

In the parasitic Rhizocephala and in a few Copepoda (Monstrillidae) the alimentary canal is absent or vestigial throughout life.

Circulatory System.—As in the other Arthropoda, the circulatory system in Crustacea is largely lacunar, the blood flowing in spaces or channels without definite walls. These spaces make up the apparent body-cavity, the true body-cavity or coelom having been, for the most part, obliterated by the great expansion of the blood-containing spaces. The heart is of the usual Arthropodous type, lying in a more or less well-defined pericardial blood-sinus, with which it communicates by valvular openings or ostia. In the details of the system, however, great differences exist within the limits of the class. There is every reason to believe that, in the primitive Arthropoda, the heart was tubular in form, extending the whole length of the body, and having a pair of ostia in each somite. This arrangement is retained in some of the Phyllopoda, but even in that group a progressive abbreviation of the heart, with a diminution in the number of the ostia, can be traced, leading to the condition found in the closely related Cladocera, where the heart is a subglobular sac, with only a single pair of ostia. In the Malacostraca, an elongated heart with numerous segmentally arranged ostia is found only in the aberrant group of Stomatopoda and in the transitional Phyllocarida. In the other Malacostraca the heart is generally abbreviated, and even where, as in the Amphipoda, it is elongated and tubular, the ostia are restricted in number, three pairs only being usually present. In many Entomostraca the heart is absent, and it is impossible to speak of a “circulation” in the proper sense of the term, the blood being merely driven hither and thither by the movements of the body and limbs and of the alimentary canal.

A very remarkable condition of the blood-system, unique, as far as is yet known among the Arthropoda, is found in a few genera of parasitic Copepoda (Lernanthropus, Mytilicola). In these there is a closed system of vessels, not communicating with the body-cavity, and containing a coloured fluid. There is no heart. The morphological nature of this system is unknown.

Excretory System.—The most important excretory or renal organs of the Crustacea are two pairs of glands lying at the base of the antennae and of the second maxillae respectively. The two are probably never functional together in the same animal, though one may replace the other in the course of development. Thus, in the Phyllopoda, the antennal gland develops early and is functional during a great part of the larval life, but it ultimately atrophies, and in the adult (as in most Entomostraca) the maxillary gland is the functional excretory organ. In the Decapoda, where the antennal gland alone is well-developed in the adult, the maxillary gland sometimes precedes it in the larva. The structure of both glands is essentially the same. There is a more or less convoluted tube with glandular walls connected internally with a closed “end-sac” and opening to the exterior by means of a thin-walled duct. Development shows that the glandular tube is mesoblastic in origin and is of the nature of a coelomoduct, while the end-sac is to be regarded as a vestigial portion of the coelom. In the Branchiopoda the maxillary gland is lodged in the thickness of the shell-fold (when this is present), and, from this circumstance, it often receives the somewhat misleading name of “shell-gland.” In the Decapoda the antennal gland is largely developed and is known as the “green gland.” The external duct of this gland is often dilated into a bladder, and may sometimes send out diverticula, forming a complex system of sinuses ramifying through the body. The green gland and the structures associated with it in Decapods were at one time regarded as constituting an auditory apparatus.

In addition to these two pairs of glands, which are in all probability the survivors of a series of segmentally arranged coelomoducts present in the primitive Arthropoda, other excretory organs have been described in various Crustacea. Although the excretory function of these has been demonstrated by physiological methods, however, their morphological relations are not clear. In some cases they consist of masses of mesodermal cells, within which the excretory products appear to be stored up instead of being expelled from the body.

Nervous System.—The central nervous system is constructed on the same general plan as in the other Arthropoda, consisting of a supra-oesophageal ganglionic mass or brain, united by circum-oesophageal connectives with a double ventral chain of segmentally arranged ganglia. In the primitive Phyllopoda the ventral chain retains the ladder-like arrangement found in some Annelids and lower worms, the two halves being widely separated and the pairs of ganglia connected together across the middle line by double transverse commissures. In the higher groups the two halves of the chain are more or less closely approximated and coalesced, and, in addition, a concentration of the ganglia in a longitudinal direction takes place, leading ultimately, in many cases, to the formation of an unsegmented ganglionic mass representing the whole of the ventral chain. This is seen, for example, in the Brachyura among the Decapoda. The brain, or supra-oesophageal ganglion, shows various degrees of complexity. In the Phyllopoda it consists mainly of two pairs of ganglionic centres, giving origin respectively to the optic and antennular nerves. The centres for the antennal nerves form ganglionic swellings on the oesophageal connectives. In the higher forms, as already mentioned, the antennal ganglia have become shifted forwards and coalesced with the brain. In the higher Decapoda, numerous additional centres are developed in the brain and its structure becomes extremely complex.

Eyes.—The eyes of Crustacea are of two kinds, the unpaired, median or “nauplius” eye, and the paired compound eyes. The former is generally present in the earliest larval stages (nauplius), and in some Entomostraca (e.g. Copepoda) it forms the sole organ of vision in the adult. In the Malacostraca it is absent in the adult, or persists only in a vestigial condition, as in some Decapoda and Schizopoda. It is typically tripartite, consisting of three cup-shaped masses of pigment, the cavity of each cup being filled with columnar retinal cells. At their inner ends (towards the pigment) these cells contain rod-like structures, while their outer ends are connected with the nerve-fibres. In some cases three separate nerves arise from the front of the brain, one going to each of the three divisions of the eye. In the Copepoda the median eye may undergo considerable elaboration, and refracting lenses and other accessory structures may be developed in connexion with it.

The compound eyes are very similar in the details of their structure (see Arthropoda) to those of insects (Hexapoda). They consist of a varying number of ommatidia or visual elements, covered by a transparent region of the external cuticle forming the cornea. In most cases this cornea is divided into lenticular facets corresponding to the underlying ommatidia.

As has been already stated, the compound eyes are often set on movable peduncles. It is probable that this is the primitive condition from which the sessile eyes of other forms have been derived. In the Malacostraca the sessile eyed groups are certainly less primitive than some of those with stalked eyes, and among the Entomostraca also there is some evidence pointing in the same direction.

Although typically paired, the compound eyes may occasionally coalesce in the middle line into a single organ. This is the case in the Cladocera, the Cumacea and a few Amphipoda.

Mention should also be made of the partial or complete atrophy of the eyes in many Crustacea which live in darkness, either in the deep sea or in subterranean habitats. In these cases the peduncles may persist and may even be modified into spinous organs of defence.

Other Sense-Organs.—As in Arthropoda, the hairs or setae on the surface of the body are important organs of sense and are variously modified for special sensory functions. Many, perhaps all, of them are tactile. They are movably articulated at the base where they are inserted in pits formed by a thinning away of the cuticle, and each is supplied by a nerve-fibril. When feathered or provided with secondary barbs the setae will respond to movements or vibrations in the surrounding water, and have been supposed to have an auditory function. In certain divisions of the Malacostraca more specialized organs are found which have been regarded as auditory. In the majority of the Decapoda there is a saccular invagination of the integument in the basal segment of the antennular peduncle having on its inner surface “auditory” setae of the type just described. The sac is open to the exterior in most of the Macrura, but completely closed in the Brachyura. In the former case it contains numerous grains of sand which are introduced by the animal itself after each moult and which are supposed to act as otoliths. Where the sac is completely closed it generally contains no solid particles, but in a few Macrura a single otolith secreted by the walls of the sac is present. In the Mysidae among the Schizopoda a pair of similar otocysts are found in the endopodites of the last pair of appendages (uropods). These contain each a single concretionary otolith.

Recent observations, however, make it very doubtful whether aquatic Crustacea can hear at all, in the proper sense of the term, and it has been shown that one function, at least, of the so-called otocysts is connected with the equilibration of the body. They are more properly termed statocysts.

Another modification of sensory setae is supposed to be associated with the sense of smell. In nearly all Crustacea the antennules and often also the antennae bear groups of hair-like filaments in which the chitinous cuticle is extremely delicate and which do not taper to a point but end bluntly. These are known as olfactory filaments or aesthetascs. They are very often more strongly developed in the male sex, and are supposed to guide the males in pursuit of the females.

Glands.—In addition to the digestive and excretory glands already mentioned, various glandular structures occur in the different groups of Crustacea. The most important of these belong to the category of dermal glands, and may be scattered over the surface of the body and limbs, or grouped at certain points for the discharge of special functions. Such glands occurring on the upper and lower lips or on the walls of the oesophagus have been regarded as salivary. In some Amphipoda the secretion of glands on the body and limbs is used in the construction of tubular cases in which the animals live. In some freshwater Copepoda the secretion of the dermal glands forms a gelatinous envelope, by means of which the animals are able to survive desiccation. In certain Copepoda and Ostracoda glands of the same type produce a phosphorescent substance, and others, in certain Amphipoda and Branchiura, are believed to have a poisonous function. Possibly related to the same group of structures are the greatly-developed cement-glands of the Cirripedia, which serve to attach the animals to their support.

Phosphorescent Organs.—Many Crustacea belonging to very different groups (Ostracoda, Copepoda, Schizopoda, Decapoda) possess the power of emitting light. In the Ostracoda and Copepoda the phosphorescence, as already mentioned, is due to glands which produce a luminous secretion, and this is the case also in certain members of the Schizopoda and Decapoda. In other cases in the last two groups, however, the light-producing organs found on the body and limbs have a complex and remarkable structure, and were formerly described as accessory eyes. Each consists of a globular capsule pierced at one or two points for the entrance of nerves which end in a central cup-shaped “striated body.” This body appears to be the source of light, and has behind it a reflector formed of concentric lamellae, while, in front, in some cases, there is a refracting lens. The whole organ can be rotated by special muscles. Organs of this type are best known in the Euphausiidae among the Schizopoda, but a modified form is found in some of the lower Decapods.

Reproductive System.—In the great majority of Crustacea the sexes are separate. Apart from certain doubtful and possibly abnormal instances among Phyllopoda and Amphipoda, the only exceptions are the sessile Cirripedia and some parasitic Isopoda (Cymothoidae), where hermaphroditism is the rule. Parthenogenesis is prevalent in the Branchiopoda and Ostracoda, often in more or less definite seasonal alternation with sexual reproduction. Where the sexes are distinct, a more or less marked dimorphism often exists. The male is very often provided with clasping organs for seizing the female. These may be formed by the modification of almost any of the appendages, often the antennules or antennae or some of the thoracic limbs, or even the mandibular palps (some Ostracoda). In addition, some of the appendages in the neighbourhood of the genital apertures may be modified for the purpose of transferring the genital products to the female, as, for instance, the first and second abdominal limbs in the Decapoda. In the higher Decapoda the male is generally larger than the female and has stronger chelae. On the other hand, in other groups the male is often smaller than the female. In the parasitic Copepoda and Isopoda the disparity in size is carried to an extreme degree, and the minute male is attached, like a parasite, to the enormously larger female.

The Cirripedia present some examples of sexual relationships which are only paralleled, in the animal kingdom, among the parasitic Myzostomida. While the great majority are simple hermaphrodites, capable of cross and self fertilization, it was discovered by Darwin that, in certain species, minute degraded males exist, attached within the mantle-cavity of the ordinary individuals. Since these dwarf males pair, not with females, but with hermaphrodites, Darwin termed them “complemental” males. In other species the large individuals have become purely female by atrophy of the male organs, and are entirely dependent on the dwarf males for fertilization. In spite of the opinion of some distinguished zoologists to the contrary, it seems most probable that the separation of the sexes is in this case a secondary condition, derived from hermaphroditism through the intermediate stage represented by the species having complemental males.

The gonads, as in other Arthropoda, are hollow saccular organs, the cavity communicating with the efferent ducts. They are primitively paired, but often coalesce with each other more or less completely. The ducts are present only as a single pair, except in one genus of parasitic Isopoda (Hemioniscus), where two pairs of oviducts are found. Various accessory structures may be connected with the efferent ducts in both sexes. The oviducts may have diverticula serving as receptacles for the spermatozoa (in cases where internal impregnation takes place), and may be provided with glands secreting envelopes or shells around the eggs. The male ducts often have glandular walls, secreting capsules or spermatophores within which the spermatozoa are packed for transference to the female. The terminal part of the male ducts may be protrusible and act as an intromittent organ, or this function may be discharged by some of the appendages, as, for instance, in the Brachyura.

Fig. 11.—Side view of Crab, the abdomen extended and carrying a mass of eggs beneath it; e, eggs. (After Morse.)

The position of the genital apertures varies very greatly in the different groups of the class. They are farthest forward in the case of the female organs of the Cirripedia, where the openings are on the first thoracic (fourth postoral) somite. The most posterior position is occupied by the genital apertures of certain Phyllopoda (Polyartemia), which lie behind the nineteenth trunk-somite. It is characteristic of the Malacostraca that the position of the genital apertures is constantly different in the two sexes, the female openings being on the sixth, and those of the male on the eighth thoracic somite.

Very few Crustacea are viviparous in the sense that the eggs are retained within the body until hatching takes place (some Phyllopoda), but, on the other hand, the great majority carry the eggs in some way or other after their extrusion. In some Phyllopoda (Apus) egg-sacs are formed by modification of certain of the thoracic feet. The eggs are retained between the valves of the shell in some Phyllopoda and in the Cladocera and Ostracoda, and they lie in the mantle cavity in the Cirripedia. In the Copepoda they are agglutinated together into masses attached to the body of the female. Among the Malacostraca some Schizopoda, the Cumacea, Tanaidacea, Isopoda and Amphipoda (sometimes grouped all together as Peracarida) have a marsupium or brood-pouch formed by overlapping plates attached to the bases of some of the thoracic legs. In most of the Decapoda the eggs are carried by the female, attached to the abdominal appendages (fig. 11). A few cases are known in which the developing embryos are nourished by a special secretion while in the brood-chamber of the mother (Cladocera, terrestrial Isopoda).

Embryology.

The majority of the Crustacea are hatched from the egg in a form differing more or less from that of the adult, and pass through a series of free-swimming larval stages. There are many cases, however, in which the metamorphosis is suppressed, and the newly-hatched young resemble the parent in general structure. The relative size of the eggs and the amount of nutritive yolk which they contain are generally much greater in those forms which have a direct development.

The details of the early embryonic stages vary considerably within the limits of the class. They are of interest, however, rather from the point of view of general embryology than from that of the special student of the Crustacea, and cannot be fully dealt with here.

Segmentation is usually of the superficial or centrolecithal type. The hypoblast is formed either by a definite invagination or by the immigration of isolated cells, known as vitellophags, which wander through the yolk and later become associated into a definite mesenteron, or by some combination of these two methods. The blastopore generally occupies a position corresponding to the posterior end of the body. The mesoblast of the cephalic (naupliar) region probably arises in connexion with the lips of the blastopore and consists of loosely-connected cells or mesenchyme. In the region of the trunk, in many cases, paired mesoblastic bands are formed, growing in length by the division of teloblastic cells at the posterior end, and becoming segmented into somites. The existence of true coelom-sacs is somewhat doubtful. The rudiments of the first three pairs of appendages commonly appear simultaneously, and, even in forms with embryonic development, they show differences in their mode of appearance from the succeeding somites. Further, a definite cuticular membrane is frequently formed and shed at this stage, which corresponds to the nauplius-stage of larval development.

Fig. 12.—Nauplius of a Prawn (Penaeus). (Fritz Müller).

The larval metamorphoses of the Crustacea have attracted much attention, and have been the subject of much discussion in view of their bearing on the phylogenetic history of the group. In those Crustacea in which the series of larval stages is most complete, the starting-point is the form already mentioned under the name of nauplius. The typical nauplius (fig. 12) has an oval unsegmented body and three pairs of limbs corresponding to the antennules, antennae and mandibles of the adult. The antennules are uniramous, the others biramous, and all three pairs are used in swimming. The antennae have a spiniform or hooked masticatory process at the base, and share with the mandibles, which have a similar process, the function of seizing and masticating the food. The mouth is overhung by a large labrum or upper lip, and the integument of the dorsal surface of the body forms a more or less definite dorsal shield. The paired eyes are, as yet, wanting, but the unpaired eye is large and conspicuous. A pair of frontal papillae or filaments, probably sensory, are commonly present.

A nauplius larva differing only in details from the typical form just described is found in the majority of the Phyllopoda, Copepoda and Cirripedia, and in a more modified form, in some Ostracoda. Among the Malacostraca the nauplius is less commonly found, but it occurs in the Euphausiidae among the Schizopoda and in a few of the more primitive Decapoda (Penaeidea) (fig. 12). In most of the Crustacea which hatch at a later stage there is, as already mentioned, more or less clear evidence of an embryonic nauplius stage. It seems certain, therefore, that the possession of a nauplius larva must be regarded as a very primitive character of the Crustacean stock.

As development proceeds, the body of the nauplius elongates, and indications of segmentation begin to appear in its posterior part. At successive moults the somites increase in number, new somites being added behind those already differentiated, from a formative zone in front of the telsonic region. Very commonly the posterior end of the body becomes forked, two processes growing out at the sides of the anus and often persisting in the adult as the “caudal furca.” The appendages posterior to the mandibles appear as buds on the ventral surface of the somites, and in the most primitive cases they become differentiated, like the somites which bear them, in regular order from before backwards. The limb-buds early become bilobed and grow out into typical biramous appendages which gradually assume the characters found in the adult. With the elongation of the body, the dorsal shield begins to project posteriorly as a shell-fold, which may increase in size to envelop more or less of the body or may disappear altogether. The rudiments of the paired eyes appear under the integument at the sides of the head, but only become pedunculated at a comparatively late stage.

The course of development here outlined, in which the nauplius gradually passes into the adult form by the successive addition of somites and appendages in regular order, agrees so well with the process observed in the development of the typical Annelida that we must regard it as being the most primitive method. It is most closely followed by the Phyllopods such as Apus or Branchipus, and by some Copepoda.

Fig. 13.—Early Stages of Balanus. (After Spence Bate.)
A, Nauplius. e, Eye. B, Cypris-larva with a bivalve shell and just before becoming attached (represented feet upwards for comparison with E, where it is attached).	C, After becoming attached, side views. D, Later stage, viewed from above. E, Side view, later stage and with cirri extended.
The dots indicate the actual size.

Fig. 14.—Zoea of Common Shore-Crab in its second stage. (Spence Bate.)

r, Rostral spine. s, Dorsal spine. m, Maxillipeds. t, Buds of thoracic feet. a, Abdomen.

In most Crustacea, however, this primitive scheme is more or less modified. The earlier stages may be suppressed or passed through within the egg (or within the maternal brood-chamber), so that the larva, on hatching, has reached a stage more advanced than the nauplius. Further, the gradual appearance and differentiation of the successive somites and appendages may be accelerated, so that comparatively great advances take place at a single moult. In the Cirripedia, for example, the latest nauplius stage (fig. 13, A) gives rise directly to the so-called Cypris-larva (fig. 13, B), differing widely from the nauplius in form, and possessing all the appendages of the adult. Another very common modification of the primitive method of development is found in the accelerated appearance of certain somites or appendages, disturbing the regular order of development. This modification is especially found in the Malacostraca. Even in those which have most fully retained the primitive order of development, as in the Penaeidea and Euphausiidae, the last pair of abdominal appendages make their appearance in advance of those immediately in front of them. The same process, carried further, leads to the very peculiar larva known as the Zoea, in the typical form of which, found in the Brachyura (fig. 14), the posterior five or six thoracic somites have their development greatly retarded, and are still represented by a short unsegmented region of the body at a time when the abdominal somites are fully formed and even carry appendages. The Zoea was formerly regarded as a recapitulation of an ancestral form, but there can be no doubt that its peculiarities are the result of secondary modification. It is most typically developed in the most specialized Decapoda, the Brachyura, while the more primitive groups of Malacostraca, the Euphausiidae, Penaeidea and Stomatopoda, retain the primitive order of appearance of the somites, and, for the most part, of the limbs. At the same time, the tendency to a retardation in the development of the posterior thoracic somites is very general in Malacostracan larvae, and may perhaps be correlated with the fact that in the primitive Phyllocarida the whole thoracic region is very short and the limbs closely crowded together.

Fig. 15.—Nauplius of Tetraclita porosa after the first moult.

(Fritz Müller.)

Besides the nauplius and the zoea there are many other types of Crustacean larvae, distinguished by special names, though, as their occurrence is restricted within the limits of the smaller systematic groups, they are of less general interest. We need only mention the Mysis-stage (better termed Schizopod-stage) found in many Macrura (as, for example, the lobster), which differs from the adult in having large natatory exopodites on the thoracic legs.

Most of the larval forms swim freely at the surface of the sea, and many show special adaptations to this habit of life. As in many other “pelagic” organisms, spines and processes from the surface of the body are often developed, which are probably less important as defensive organs than as aids to flotation. This is well seen in the nauplius of many Cirripedia (fig. 15) and in nearly all zoeae. Perhaps the most striking example is the zoea-like larva of the Sergestidae, known as Elaphocaris, which has an extraordinary armature of ramified spines. The same purpose is probably served by the extreme flattening of the body in the membranous Phyllosoma-larva of the rock-lobsters and their allies (Loricata).