Page:EB1911 - Volume 07.djvu/577

Rh 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.

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 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.

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