the Week of Proper 28 / Ordinary 33
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Bible Encyclopedias
Crustacea
1911 Encyclopedia Britannica
or a very large division of the animal kingdom, comprising the familiar crabs, lobsters, crayfish, shrimps and prawns, the sandhoppers and woodlice, the strangely modified barnacles and the minute water-fleas. Besides these the group also includes a multitude of related forms which, from their aquatic habits and generally inconspicuous size, and from the fact that they are commonly neither edible nor noxious, are little known except to naturalists and are undistinguished by any popular names. Collectively, they are ranked as one of the classes forming the sub-phylum Arthropoda, and their distinguishing characters are discussed under that heading. It will be sufficient here to define them as Arthropoda for the most part of aquatic habits, having typically two pairs of antenniform appendages in front of the mouth and at least three pairs of post-oral limbs acting as jaws.
As a matter of fact, however, the range of structural variation within the group is so wide, and the modifications due to parasitism and other causes are so profound, that it is almost impossible to frame a definition which shall be applicable to all the members of the class. In certain parasites, for instance, the adults have lost every trace not only of Crustacean but even of Arthropodous structure, and the only clue to their zoological position is that afforded by the study of their development. In point of size also the Crustacea vary within very wide limits. Certain waterfleas (Cladocera) fall short of one-hundredth of an inch in total length; the giant Japanese crab (Macrocheira) can span over Io ft. between its outstretched claws.
The habits of the Crustacea are no less diversified than their structure. Most of them inhabit the sea, but representatives of all the chief groups are found in fresh water (though the Cirripedia have hardly gained a footing there), and this is the chief home of the primitive Phyllopoda. A terrestrial habitat is less common, but the widely-distributed land Isopoda or woodlice and the land-crabs of tropical regions have solved the problem of adaptation to a subaerial life.
Swimming is perhaps the commonest mode of locomotion, but numerous forms have taken to creeping or walking, and the robber-crab (Birgus latro) of the Indo-Pacific islands even climbs palm-trees. None has the power of flight, though certain pelagic Copepoda are said to leap from the surface of the sea like flying-fish. Apart from the numerous parasitic forms, the only Crustacea which have adopted a strictly sedentary habit of life are the Cirripedia, and here, as elsewhere, profound modifications of structure have resulted, leading ultimately to a partial assumption of the radial type of symmetry which is so often associated with a sedentary life.
Many, perhaps the majority, of the Crustacea are omnivorous or carrion-feeders, but many are actively predatory in their habits, and are provided with more or less complex and efficient instruments for capturing their prey, and there are also many planteaters. Besides the sedentary Cirripedia, numbers of the smaller forms, especially among the Entomostraca, subsist on floating particles of organic matter swept within reach of the jaws by the movements of the other limbs.
Symbiotic association with other animals, in varying degrees of interdependence, is frequent. Sometimes the one partner affords the other merely a convenient means of transport, as in the case of the barnacles which grow on, or of the gulf-weed crab which clings to, the carapace of marine turtles. From this we may pass through various grades of " commensalism," like that of the hermit-crab with its protective anemones, to the cases of actual parasitism. The parasitic habit is most common among the Copepoda and Isopoda, where it leads to complex modifications of structure and life-history. Perhaps the most complete degeneration is found in the Rhizocephala, which are parasitic on other Crustacea. In these the adult consists of a simple saccular body containing the reproductive organs and attached by root-like filaments which ramify throughout the body of the host and serve for the absorption of nourishment (fig. r).
Many of the larger species of Crustacea are used as food by man, the most valuable being the lobster, which is caught in large quantities on both sides of the North Altantic. Perhaps the most important of all Crustacea, however, with respect to the part which they play in the economy of nature, are the minute pelagic Copepoda, of which incalculable myriads form an important constituent of the " plankton " in all the seas of the globe. It is on the plankton that a great part of the higher animal life of the sea ultimately depends for food. The Copepoda live upon the diatoms and other important microscopic vegetable life at the surface of the sea, and in their turn serve as food for fishes and other larger forms and thus, indirectly, for man himself.
Historical Sketch
In common with most branches of natural history, the science of Carcinology may be traced back to its beginnings in the writings of Aristotle. It received additions FIG.
A, Group of Peltogaster socialis on the abdomen of a small hermitcrab; in one of them the fasciculately ramified roots, r, in the liver of the crab are shown (Fritz Muller).
B, Young of Sacculina purpurea with its roots. Magn. 5 diam. (Fritz Muller).
of varying importance at the hands of medieval and later naturalists, and first began to assume systematic form under the influence of Linnaeus. The application of the morphological method to the Crustacea may perhaps be dated from the work of J. C. Fabricius towards the end of the 18th century.
In the first quarter of the Loth century important advances in classification were made by P. A. Latreille, W. E. Leach and others, and J. Vaughan Thompson demonstrated the existence of metamorphosis in the development of the higher Crustacea. A new epoch may be said to begin with H. Milne-Edwards' classical Histoire naturelle des crustaces (1834-1840). It is noteworthy that even at this late date the Cirripedia (Thyrostraca) were still excluded from the Crustacea, though Darwin's Monograph (1851-1854) was soon to make them known with a wealth of anatomical and systematic detail such as was available, at that time, for few other groups of Crustacea. About the same period three authors call for special mention, W. de Haan, J. D. Dana and H. Krdyer. The new impulse given to biological research by the publication of the Origin of Species bore fruit in Fritz Muller's Filr Darwin, in which an attempt was made to reconstruct the phylogenetic history of the class. The same line of work was followed in the long series of important memoirs from the pen of K. F. W. Claus, and noteworthy contributions were made, among many others, by A. Dohrn, Ray Lankester and Huxley. In more recent years the long and constantly increasing list of writers on Crustacea contains no name more honoured than that of the veteran G. O. Sars of Christiania.
Morphology. 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 vII. 18 a ?Ark 7 O Z B I.
G FIG. 2. - Abdominal Somite of a Lobster, separated and viewed from in front. t, tergum; s, sternum; p1, pleuron.
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.
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 ep 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." 1, labrum, or upper lip.
m, metastoma, or lower lip.
I, eyes.
2, antennule (the arrow points to the opening of the socalled auditory organ).
3, antenna.
4, mandible.
5, maxillula (or first maxilla).
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.
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 FIG. 4. - Diagram of an Amphipod. (After Spence Bate and Westwood.) C, cephalon. Ab, abdomen.
Th, thorax. (Only seven of the The numbers appended to the eight thoracic somites are somites do not correspond to the visible, the first being fused enumeration adopted in the text.
with the cephalon.) 21 is the telson.
eyestalks 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.
A character which recurs in the most diverse groups of the Crustacea, and which is probably to be regarded as a primitive attribute FIG. 5. - Phyllopoda and Phyllocarida.
1, Ceratiocarispapilio, U. Silurian, head showing the labrum and Lanark. mouth-parts.
4, larva of Apus cancriformis. 2, Nebalia bipes (one side of 5, Branchipus stagnalis: a, adult carapace removed). female; b, first larval stage (Nauplius); c, second larval 3, Lepidurus Angassi: a, dorsal stage.
aspect; b, ventral aspect of 6, Nauplius of Artemia salina. 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 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.
last thoracic limb of male. In 10 and the arrows indicate the genital apertures.
13, sterna of the thoracic somites, from within.
14, third abdominal somite, with appendages or " swimmerets." 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 Stomato 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.
poda (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.
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). 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 swimmingbranch 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 gnatho FIG. 7. - Squilla mantis bases tends to show that the primitive (Stomatopoda), showing the type of appendage was more complex last four thoracic (leg-bearthan the simple biramous limb, and ing) somites free from the some authorities have regarded the carapace. leaf-like appendages of the Phyllo poda 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 A pus, 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.
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 FIG. 8. - Tanais dubius (?) Kr. magnified 25 times, 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,1, m) are the rudiments of the lamellae which subsequently form the brood-cavity. (Fritz Miller.) 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 importC ance can in most cases be attached to it. A peculiar modification is found in the trunk-limbs of the Cirri pedia (fig. 9), in which both FIG. 9. - A, Balanus (young), side rami are multiarticulate view with cirri protruded. B, Upper and filiform and fringed surface of same; valves closed. C, with long bristles. When Highly magnified view of one of the protruded from the opening cirri. (Morse.) 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.
