The Appendages, Anatomy, and Relationships of Trilobites Part 18

Walcott referred _Burgessia_ and _Waptia_ to new families under the Notostraca, while _Yohoia_ and _Opabina_ were placed with the Anostraca. Except for the development of the carapace, there is a striking similarity between _Waptia_ and _Yohoia_, serving to connect the two groups.

The Branchiopoda were very highly specialized as early as Middle Cambrian time, the carapace of the Notostraca being fully developed and the abdomen limbless. Some (_Burgessia_) had numerous segments, but most had relatively few. The most striking point about them, however, is that so far as is known none of them had phyllopodan limbs. While the preservation is in most cases unsatisfactory, such limbs as are preserved are trilobite-like, and in the case of _Burgessia_ there can be no possible doubt of the structure. Another interesting feature is the retention by _Yohoia_ of vestiges of pleural lobes. The Middle Cambrian Branchiopoda are more closely allied to the Trilobita than are the modern ones, but still the subcla.s.s is not so closely related to that group as has been thought.

Modern _Apus_ is certainly much less like a trilobite than has been supposed, and very far from being primitive. The Branchiopoda of the Middle Cambrian could have been derived from the trilobites by the loss of the pleural lobes, the development of the posterior margin of the cephalon to form a carapace, and the loss of the appendages from the abdominal segments. Modern branchiopods can be derived from those of the Middle Cambrian by the modification of the appendages through the reduction of the endopodite and exopodite and the growth of the endites and exites from the proximal segments.

Carpenter (1903, p. 334), from his study of recent crustaceans, has already come to the conclusion that the Branchiopoda are not the most primitive subcla.s.s, and this opinion is strengthened by evidence derived from the Trilobita and from the Branchiopoda of the Middle Cambrian.

COPEPODA.

The non-parasitic Eucopepoda are in many ways much nearer to the trilobites than any other Crustacea. These little animals lack the carapace, and the body is short, with typically ten free segments and a telson bearing caudal furcae. The head is composed of five segments (if the first thoracic segment is really the fused first and second), is often flattened, and lacks compound eyes. Pleural lobes are well developed, but instead of being flattened as in the trilobite, they are turned down at the sides or even incurved. A labrum is present.

The antennules, antennae, and mandibles are quite like those of trilobites. The antennules are very long and made up of numerous segments. The antennae are biramous, the junction between the c.o.xopodite and basipodite is well marked, and the endopodite consists of only two segments.

The mandibles are said to "retain more completely than in any other Crustacea the form of biramous swimming limbs which they possess in the nauplius." The c.o.xopodites form jaws, while both the reduced endopodite and exopodite are furnished with long setae. The maxillulae are also biramous, but very different in form from those of the trilobite, and the maxillae are phyllopodan.

The first thoracic limb is uniramous and similar to the maxillae, but the five following pairs are biramous swimming legs with c.o.xopodite, basipodite, exopodite, and endopodite. Both the exopodite and endopodite are shorter than in the trilobites, but bear setae and spines.

The last pair of thoracic limbs are usually modified in the male into copulatory organs. In some females they are enlarged to form plates for the protection of the eggs, in others they are unmodified. In still others they are much reduced or disappear. The abdomen is without appendages.

The development in Copepoda is direct, by the addition posteriorly to the larval form (nauplius) of segments, and the appendages remain nearly unmodified in the adult.

Altogether, the primitive Copepoda seem much more closely allied to the Trilobita than any other modern Crustacea, but unfortunately no fossil representative of the subcla.s.s has been found. This is not so surprising when one considers the habits and the habitat of most of the existing species. Many are parasitic, many pelagic in both fresh and marine waters, and many of those living on the bottom belong to the deep sea or fresh water. Most free-living forms are minute, and all have thin tests.

The eyes of copepods are of interest, in that they suggest the paired ocelli of the Harpedidae and Trinucleidae. In the Copepoda there are, in the simplest and typical form of these organs, three ocelli, each supplied with its own nerve from the brain. Two of these are dorsal and look upward, while the third is ventral. In some forms the dorsal ocelli are doubled, so that five in all are present (cf. some species of Harpes with three ocelli on each mound). In some, the cuticle over the dorsal eyes is thickened so as to form a lens, as appears to be the case in the trilobites. These peculiar eyes may be a direct inheritance from the Hypoparia.

ARCHICOPEPODA.

Professor Schuchert has called my attention to the exceedingly curious little crustacean which Handlirsch (1914) has described from the Tria.s.sic of the Vosges. Handlirsch erected a new species, genus, family, and order for this animal, which he considered most closely allied to the copepods, hence the ordinal name. _Euthycarcinus kessleri_, the species in question, was found in a clayey lens in the Voltzia sandstone (Upper Bunter). a.s.sociated with the new crustacean were specimens of _Estheria_ only, but in the Voltzia sandstone itself land plants, fresh and brackish water animals, and occasionally, marine animals are found. The clayey lens seems to have been of fresh or brackish water origin.

All of the specimens (three were found) are small, about 35 mm. long without including the caudal rami, crushed flat, and not very well preserved. The head is short, not so wide as the succeeding segments, and apparently has large compound eyes at the posterior lateral angles. The thorax consists of six segments which are broader than the head or abdomen. The abdomen, which is not quite complete in any one specimen, is interpreted by Handlirsch as having four segments in the female and five in the male. Least satisfactory of all are traces of what are interpreted by the describer as a pair of long stiff unsegmented cerci or stylets on the last segment.

The ventral side of one head shield shows faint traces of several appendages which must have presented great difficulty in their interpretation. A pair of antennules appear to spring from near the front of the lower surface, and the remainder of the organs are grouped about the mouth, which is on the median line back of the center. Handlirsch sees in these somewhat obscure appendages four pairs of biramous limbs, antennae, mandibles, maxillulae, and maxillae, both branches of each consisting of short similar segments, endopodites and exopodites being alike pediform.

Each segment of the thorax has a pair of appendages, and those on the first two are clearly biramous. The endopodites are walking legs made up of numerous short segments (twelve or thirteen according to Handlirsch's drawing), while the exopodite is a long breathing and rowing limb, evidently of great flexibility and curiously like the antennules of the same animal. The individual segments are narrow at the proximal end, expand greatly at the sides, and have a concave distal profile. A limb reminds one of a stipe of _Diplograptus_.

Both branches are spiniferous.

No appendages are actually present on the abdomen, but each segment has a pair of scars showing the points of attachment. From the small size of these, it is inferred that the limbs were poorly developed.

This species is described in so much detail because, if it is a primitive copepod, it has a very important bearing on the ancestry of that group and is the only related form that has been found fossil.

The non-parasitic copepods have typically ten (eleven) free segments, including the telson, and the four abdominal segments are much more slender than the six in front of them. In this respect the agreement is striking, and the presence of five pairs of appendages in the head and six free segments in the thorax is a more primitive condition than in modern forms where the first two thoracic segments are apparently fused (Calman, 1909, p. 73).

The large compound eyes of this animal are of course not present in the copepods, but as vestiges of eyes have been found in the young of _Cala.n.u.s_, it is possible that the ancestral forms had eyes.

The greatest difficulty is in finding a satisfactory explanation of the appendages. The general condition is somewhat more primitive than in the copepods, for all the appendages are biramous, while in the modern forms the maxillipeds are uniramous and the sixth pair of thoracic appendages are usually modified in the male as copulatory organs. In the copepods the modification is in the direction of reduction, both endopodites and exopodites usually possessing fewer segments than the corresponding branches in the trilobites. The endopodite of _Euthycarcinus_, on the contrary, possesses, if Handlirsch's interpretation is correct, twice as many segments as the endopodite of a trilobite. If the Copepoda are descended from the trilobites, as everything tends to indicate, then _Euthycarcinus_ is certainly not a connecting link. The only truly copepodan characteristic of this genus is the agreement in number and disposition of free segments. The division into three regions instead of two, the compound eyes, and the structure of the appendages are all foreign to that group.

With the Limulava fresh in mind, one is tempted to compare _Euthycarcinus_ with that ancient type. The short head and large marginal eyes recall _Sidneyia_, and the grouping of the appendages about the mouth also suggests that genus and _Emeraldella_. In the Limulava likewise there is a contraction of the posterior segments, although it is behind the ninth instead of the sixth. There is no likeness in detail between the appendages of the Limulava and those of _Euthycarcinus_, but the composite claws of _Sidneyia_ show that in this group there was a tendency toward the formation of extra segments.

If this fossil had been found in the Cambrian instead of the Tria.s.sic, it would probably have been referred to the Limulava, and is not at all impossible that it is a descendant from that group. As a connecting link between the Trilobita and Copepoda it is, however, quite unsatisfactory.

OSTRACODA.

The bivalved sh.e.l.l of the Ostracoda gives to this group of animals an external appearance very different from that of the trilobites, but the few appendages, though highly modified, are directly comparable.

The development, although modified by the early appearance of the bivalved sh.e.l.l within which the nauplius lies, is direct. Imperfect compound eyes are present in one family.

The antennules are short and much modified by functioning as swimming, creeping, or digging organs. They consist of eight or less segments.

The antennas are also locomotor organs, and in most orders are biramous. The mandibles are biramous and usually with, but sometimes without, a gnathobase. The maxillulae are likewise biramous but much modified.

The h.o.m.ology of the third post-oral limb is in question, some considering it a maxilla and others a maxilliped. It has various forms in different genera. It is always much modified, but exopodite and endopodite are generally represented at least by rudiments. The fourth post-oral limb is a lobed plate, usually not distinctly segmented, and the fifth a uniramous pediform leg. The sixth, if present at all, is vestigial.

Very little comparison can be made between the Ostracoda and Trilobita, other than in the ground-plan of the limbs, but the presence of biramous antennae is a primitive characteristic.

CIRRIPEDIA.

Like the ostracod, the adult cirriped bears little external resemblance to the trilobite. The form of the nauplius is somewhat peculiar, but it has the typical three pairs of appendages, to which are added in the later metanauplius stages the maxillae and six pairs of thoracic appendages. In the adult, the antennules, which serve for attachment of the larva, usually persist in a functionless condition, while the antennas disappear. The mandibles, maxillulae, and maxillae are simple and much modified to form mouth parts, and the six pairs of thoracic appendages are developed into long, multisegmented, biramous appendages bearing numerous setae which serve for catching prey. Paired eyes are present in later metanauplius stages, but lost early in the development. The relationship to the trilobite evidently is not close.

MALACOSTRACA.

_1. Phyllocarida._

The oldest malacostracans whose appendages are known are species of _Hymenocaris_. One, described as long ago as 1866 by Salter, has what seem to be a pair of antennae and a pair of jaw-like mouth-parts.

Another more completely known species has recently been reported by Walcott (1912 A, p. 183, pl. 31, figs. 1-6). This latter form is described as having five pairs of cephalic appendages: a pair of minute antennules beside the small pedunculated eyes, a pair of large uniramous antennae, slender mandibles and maxillulae, and large maxillae composed of short stout segments. There are eight pairs of biramous thoracic limbs, the exopodites setiferous, the endopodites composed of short wide segments and ending in terminal claw-like spines. These appendages are like those of trilobites.

_Hymcnocaris_ belongs to the great group of extinct ceratocarid Crustacea which are admitted to the lowest of the malacostracan orders, Phyllocarida, because of their resemblance to _Nebalia_, _Paranebalia_, _Nebaliopsis_, and _Nebaliella_, the four genera which are at present living. The general form of the recent and fossil representatives of the order is strikingly similar. The chief outward difference is that in many of the fossils the telson is accompanied by two furcal rami, while in the modern genera it is simple. It now becomes possible to make some comparison between the appendages of _Hymcnocaris_ of the Middle Cambrian and the Nebaliidae of modern seas.

In both there are five pairs of cephalic and eight of thoracic appendages, while those of the abdomen of Hymenocaris are not known.

In both, the antennules are less developed than the antennae. In the Nebaliidae the antennules show evidence of having been originally double (they are obviously so in the embryo), while they are single in _Hymcnocaris_. In both, the antennae are simple. The remaining cephalic organs are too little shown by the specimen from the Middle Cambrian to allow detailed comparison. The mandibles, maxillulae, and maxillae of _Nebalia_ are, however, of types which could be derived from the trilobite.

In three of the genera of the Nebaliidae, the eight pairs of thoracic limbs are all similar to one another, though those of the genera differ. All are biramous. The limbs of _Hymcnocaris_ can apparently be most closely correlated with those of _Nebalia antarctica_, in which the endopodite consists of short flattened segments, and the exopodite is a long setiferous plate. Epipodites are present in both _Nebalia_ and _Hymcnocaris_.

So far as the appendages of _Hymenocaris_ are known, they agree very well with those of the Nebaliidae, and since they are of the trilobite type, it may safely be stated that the Trilobita and Malacostraca are closely related.

_2. Syncarida._

Walcott (1918, p. 170) has compared the limbs of _Neolenus_ with those of the syncarid genera _Anaspides_ and _Koonunga_. These are primitive Malacostraca without a carapace, but as they have a compressed test and _Anaspides_ has stalked eyes, their gross anatomy does not suggest the trilobite. The thoracic appendages are very trilobite-like, since the endopodite has six segments (in _Anaspides_) and a multisegmented setiferous exopodite. The c.o.xopodites, except of the first thoracic segment, do not, however, show endobases, and those which are present are peculiar articulated ones. The cephalic appendages are specialized, and the antennules double as in most of the Malacostraca.

External epipodites are very numerous on the anterior limbs.

This group extends back as far as the Pennsylvanian and had then probably already become adapted to fresh-water life. It may be significant that the Palaeozoic syncarids appear to have lacked epipodites. While differing very considerably from the Trilobita, the Syncarida could have been derived from them.

_3. Isopoda._

Since the earliest times there has been a constant temptation to compare the depressed shields of the trilobites with the similar ones of isopods. Indeed, when _Scrolls_ with its Lichadian body was first discovered about a hundred years ago, it was thought that living trilobites had been found at last. The trilobate body, cephalic shield, sessile eyes, abdominal shield, and pleural extensions make a wonderful parallel. This similarity is, however, somewhat superficial.

The appendages are very definitely segregated in groups on the various regions of the body, and while the pleopods are biramous, the thoracic legs are without exopodites (except in very early stages of development of one genus). The Isopoda arose just at the time of the disappearance of the Trilobita, and there seems a possibility of a direct derivation of the one group from the other. It should be pointed out that while the differences of Isopoda from Trilobita are important, they are all of a kind which could have been produced by the development from a trilobite-like stock. For example: