The segmentation of the body is Malacostracan, save that two extra segments are present in the abdomen, and the paired compound eyes are borne upon stalks. The eight thoracic limbs are all very similar; they are built on the typical biramous plan, and each carries a bract; they have been compared, owing to their flattened, expanded shape, to the foliaceous limbs of the Phyllopods. The abdominal appendages are also biramous. The heart is greatly elongated, stretching through thorax and abdomen; there are present both the antennary excretory glands characteristic of adult Malacostraca and the maxillary glands characteristic of adult Entomostraca, and both the posterior and anterior livers characteristic of the two Orders respectively are present. This combination of characters justifies the belief that Nebalia represents a primitive form, standing to some extent in an intermediate position between Entomostraca and Malacostraca, but it may be doubted if the special relationship to the Phyllopoda, claimed on the strength of the foliaceous appearance of the thoracic limbs, can be legitimately pressed.
Nebalia shows the clearest signs of relationship to the other primitive Malacostraca, and especially to the Mysidae, which it resembles not only in general form and in the essentially biramous character of its appendages, but also in many embryological points and in the similarity in development of the brood-pouch.[[86]]
A large number of very ancient palaeozoic fossils are known which are placed provisionally with Nebalia in the Division Phyllocarida, and some of these are no doubt closely related to the existing isolated genus. Hymenocaris from the Cambrian.
SERIES 2. EUMALACOSTRACA.
Before entering on a description of the members of this Series it is necessary to introduce and justify a new scheme of classification which has been proposed by Dr. W. T. Calman. This scheme necessitates the abandonment of the old Order Schizopoda, and also ignores the distinction which used to be considered fundamental between the sessile-eyed Crustacea (Edriophthalmata) and the stalk-eyed forms (Podophthalmata).
The old group of Schizopoda, to which Nebalia and the isolated form Anaspides, to be considered later, are undoubtedly related, represent very clearly the stem-forms from which the various branches of the Malacostracan stock diverge. No doubt they are themselves specialised in many directions, since they are a dominant group in present day seas, but their organisation is fundamentally of a primitive type. We see this especially in the comparative absence of fusion or reduction of the segments of the body externally and of the nervous system internally, and in the simple undifferentiated character of the trunk-limbs, all of which conform to the primitive biramous type. The most anterior thoracic limbs of the Schizopods are of particular interest. In the higher Malacostraca three of these limbs are usually turned forwards towards the mouth to act as maxillipedes, and the most anterior of all, the first maxillipede, is apt, especially in the Decapoda, to take on a flattened foliaceous form owing to the expansion of the basal segments to act as gnathobases (see Fig. [1], A, p. 10). Now this appendage in the Schizopods preserves its typical biramous character, and resembles the succeeding thoracic limbs, but in many of the species the basal joints show a tendency to be produced into biting blades (Fig. [1], E, p. 10), thus indicating the first step in the evolution of the foliaceous first maxillipede of the Decapoda. The primitive character of the Schizopods is also indicated by the fact that most of the Decapoda with uniramous limbs on the five hinder thoracic segments pass through what is known as the “Mysis stage” in development, when these limbs are biramous, the exopodites being subsequently lost in most cases.
The “Schizopoda” include a very large number of pelagic Crustacea of moderate size, which superficially appear to resemble one another very closely. The slender, elongated body, the presence of biramous limbs on all the thoracic and abdominal segments, and the possession of a single row of gills at the bases of the thoracic limbs, are, generally speaking, typical of the families Mysidae, Lophogastridae, Eucopiidae, and Euphausiidae, which go to make up the old Order Schizopoda.
It has, however, been pointed out first by Boas,[[87]] and subsequently by Hansen and Calman,[[88]] that the Euphausiidae are in many respects distinct from the other three families, and agree with the Decapoda, while the Eucopiidae, Lophogastridae, and Mysidae agree with the Cumacea, Isopoda, and Amphipoda.
It has, therefore, been suggested by these authors that the classification of the Malacostraca should be revised, and Calman (loc. cit.) has brought forward the following scheme:—
The division Peracarida, including the Eucopiidae, Lophogastridae, and Mysidae (= Mysidacea), the Cumacea, Isopoda, and Amphipoda, is characterised by the fact that when a carapace is present it leaves at least four of the thoracic segments free and uncoalesced: by the presence of a brood-pouch formed from the oostegites on the thoracic limbs of the female: by the elongated heart: by the few and simple hepatic caeca: by the filiform spermatozoa: and by the direct method of development without a complicated larval metamorphosis. The biting face of the mandible has a movable joint, the “lacinia mobilis.”[[89]]