The Crustacea to which the Trilobites show some resemblance are the families Apodidae and Branchipodidae of the Order Phyllopoda (see pp. [19]–36). The Trilobita agree with those families in having a large but variable number of trunk-segments, in the possession of a large labrum (hypostome), and in the occurrence of gnathobases on the thoracic appendages; also the foliation of some of the trunk-appendages is somewhat similar. The points of difference, however, are considerable; thus the cephalic appendages are much more specialised in the Apodidae and Branchipodidae than in the Trilobita; in the latter all, with the exception of the antennae, are distinctly biramous, and whilst the basal joints were masticatory the distal parts appear to have been locomotor organs. The appendages of the trunk also differ considerably; in the Trilobita all are clearly biramous, those of the thorax having a schizopodal form. In the possession of a single pair of antennae the Trilobita differ from other Crustacea; but in some forms of Apus the second pair of antennae may be rudimentary or even absent.

There are still other features which characterise the Trilobita: thus the eyes are borne on free cheeks, and differ in structure from those of Phyllopods. The broad pygidium formed of fused segments and without terminal fulcra is quite unlike the slender-jointed abdomen of Apus and Branchipus; and whilst in the Trilobites all the segments bear appendages, in the Phyllopods some, at any rate, of the posterior segments are devoid of appendages. The distinct division of the body into an axial and pleural region is not seen in Phyllopods, and is probably a character of some importance, since it occurs in the great majority of Trilobites, including all the early forms.

The existence of some relationship between the Trilobita and the Leptostraca (Phyllocarida) has been maintained by Professor G. H. Carpenter.[[195]] He points out that some of the earliest Trilobites, such as Holmia kjerulfi (Fig. [148]), possess nearly the same number of segments as Nebalia (Fig. [76], p. 111), and that in the latter genus the cephalic appendages, especially the mandibles and maxillae, are less specialised than in Apus, and consequently differ less from those of Trilobites than do the appendages of the Apodidae. Further, in another genus of the Leptostraca, Paranebalia, the biramous thoracic legs, in which both endopodite and exopodite are elongate, approach those of Trilobites more nearly than do the thoracic legs of Apus.

The view[[196]] that some connexion may exist between the Isopoda and the Trilobita seems to have been based on the similar dorso-ventral flattening of the body, its division into three regions—head, thorax, and abdomen—and the presence of sessile eyes. Beyond this it is difficult to find any resemblance; whilst the differences, such as the variable number of thoracic segments and their biramous appendages in Trilobites, are important.

At present, then, we can only conclude that the Trilobita are more primitive than any other Crustacea, and that their resemblance to some of the Phyllopoda is sufficient to make it probable that they had some ancestral connexion;[[197]] the possibility of such a relationship receives some support from the presence in the Lower Cambrian rocks of Protocaris, a genus of the Phyllopoda which resembles Apus.[[198]] The primitive characters of Trilobites are the variable and often large number of segments in the thorax and pygidium; the presence of a pair of appendages on every segment except the anal; the biramous form of all except the first pair of appendages; and the lack of specialisation shown by the appendages, especially those of the head.

The classification of Trilobites is due largely to the work of Barrande and Salter, and the families defined by those authors have been, in the main, generally adopted. But the phylogenetic relationship of the families has still, to a large extent, to be established. Salter[[199]] arranged the families in four groups, but did not claim that that classification was entirely natural. His groups with the families included in each are:—

1. Agnostini. Without eyes or facial suture. Agnostidae.

2. Ampycini. Facial sutures obscure, or submarginal, or absent. Eyes often absent. Trinucleidae.

3. Asaphini. Facial sutures ending on the posterior margin. Acidaspidae, Lichadidae, Harpedidae, Calymenidae, Paradoxidae, Conocephalidae, Olenidae, Asaphidae, Bronteidae, and Proëtidae.

4. Phacopini. Facial sutures ending on the lateral margins. Eyes well developed. Phacopidae, Cheiruridae, and Encrinuridae.