The Appendages, Anatomy, and Relationships of Trilobites Part 15

[Ill.u.s.tration: Fig. 31.--A copy of Doctor Moberg's figure of _Nileus armadillo_, showing the position of the muscle scars.]

If the hypostoma were used in the manner indicated, the muscles must have been more efficient than those of the labrum of _Apus_, and it is probable that they crossed to the dorsal test. Just where they were attached is an unsolved problem. Barrande (1852, pl. 1, fig. 1) has indicated an anterior pair of scars and a single median one on the frontal lobe of _Dalmanites_ that may be considered in this connection, and also three pairs of scars on the last two lobes of the glabella of _Proetus_ (1852, pl. 1, fig. 7). Moberg (1902, p. 295, pl. 3, figs. 2, 3, text fig. 1) has described in some detail the muscle-scars of a rather remarkable specimen of _Nileus armadillo_ Dalman. While, as I shall point out, I do not agree wholly with Professor Moberg's interpretation, I give here a translation (made for Professor Beecher) of his description, with a copy of his text figure:

The well preserved surface of the sh.e.l.l permits one to note not only the tubercle (t) but a number of symmetrically arranged glabellar impressions. And because of their resemblance to the muscular insertions of recent crustaceans, I must interpret them as such. They appear partly as rounded hollows (k and i), also as elongate straight or curved areas (a, b, c, e, g, h) made up of shallow impressions or furrows about 1 mm. long, sub-parallel, and standing at an angle to the trend of the areas. Impression e is especially well marked, inasmuch as the perpendicular furrows are arranged in a shallow crescentic depression; and impression d shows besides the obscure furrows a number of irregularly rounded depressions. Larger similar ones occur at f, and in part extend over toward g.

The meaning of these impressions, or their myologic significance, could be discussed, although such discussion might rather be termed guessing.

Inner organs, such as the heart and stomach, might have been attached to the sh.e.l.l along impressions a and b. Also along or behind c and h, which both continue into the free cheeks, ligaments or muscular fibers may have been inserted. From d, e, f, and g, muscles have very likely gone out to the cephalic appendages.

Against this it may be urged that impression d is too far forward to have belonged to the first pair of feet. Again, the impression h may in reality represent two confluent muscular insertions, from the first of which, in that case, arose the muscles of the fourth pair of cephalic feet. Were this the case, the muscles of the first pair of cheek feet should be attached at e. And d in turn may be explained as the attachment of the muscles of the antennae, k those of the hypostoma, and from i possibly those of the epistoma. That k is here named as the starting point of the hypostomial muscles and not those of the antennae, depends partly on the a.n.a.logous position of i and partly on the fact that the hypostoma of _Nileus armadillo_ (text figure, in which the outline of the hypostoma is dotted), by reason of it? wing-like border, could not have permitted the antennae to reach forward, but rather only outward or backward.

My own explanation would be that impressions e, f, and g correspond to the glabellar furrows, h the neck furrow, and all four show the places of attachment of the appendifers. Those at d may possibly be connected with the antennae, although I should expect those organs to be attached under the dorsal furrows at the sides of the hypostoma. It will be noted that either b, k, or i correspond well with the maculae of the hypostoma and some or all of them may be the points of attachment of hypostomial muscles. They correspond also with the anterior scars of _Dalmanites_.

EYES.

While I have nothing to add to what has been written about the eyes of trilobites, this sketch of the anatomy would be incomplete without some reference to the little which has been done on the structure of these organs.

Quenstedt (1837, p. 339) appears to have been the first to compare the eyes of trilobites with those of other Crustacea. Johannes Muller had pointed out in 1829 (Meckel's Archiv) that two kinds of eyes were found in the latter group, compound eyes with a smooth cornea, and compound eyes with a facetted coat. Quenstedt cited _Trilobites esmarkii_ Schlotheim (=_Illaenus cra.s.sicauda_ Dalman) as an example of the first group, and _Calymene macrophthalma_ Brongniart (=_Phacops latifrons_ Bronn) for the second. Misreading the somewhat careless style of Quenstedt, Barrande (1852, p. 133) reverses these, one of the few slips to be found in the voluminous writings of that remarkable savant.

Burmeister (1843; 1846, p. 19) considered the two kinds of eyes as essentially the same, and accounted for the conspicuous lenses of Phacops on the supposition that the cornea was thinner in that genus than in the trilobites with smooth eyes.

Barrande (1852, p. 135) recognized three types of eyes in trilobites, adding to Quenstedt's smooth and facetted compound eyes the groups of simple eyes found in Harpes. In his sections of 1852, pl. 3, figs.

15-25, which are evidently diagrammatic, he shows separated biconvex lenses in both types of compound eyes, _Phacops_ and _Dalmanites_ on one hand, and _Asaphus_, _Goldius_, _Acidaspis_, and _Cyclopyge_ on the other. Clarke ( 1888), Exner ( 1891 ) and especially Lindstroem (1901) have since published much more accurate figures and descriptions. The first person to study the eye in thin section seems to have been Packard (1880), who published some very sketchy figures of specimens loaned him by Walcott. He studied the eyes of _Isotelus gigas_, _Bathyurus longispinus_, _Calymene_, and _Phacops_, and decided that the two types of eyes were fundamentally the same.

He also compared them with the eyes of _Limulus_.

Clarke (1888), in a careful study of the eye of _Phacops rana_, found that the lenses were unequally biconvex, the curvature greater on the inner surface. The lens had a circular opening on the inner side, leading into a small pear-shaped cavity. The individual lenses were quite distinct from one another, and separated by a continuation of the test of the cheek.

Exner (1891, p. 34), in a comparison of the eyes of Phacops and _Limulus_, came to the opinion that they were very unlike, and that the former were really aggregates of simple eyes.

Lindstroem (1901, pp. 27-31) came to the conclusion that besides the blind trilobites there were trilobites with two kinds of compound eyes, trilobites with aggregate eyes, and trilobites with stemmata and ocelli. His views may be briefly summarized.

I. Compound eyes.

1. Eyes with prismatic, plano-convex lenses.

"A pellucid, smooth and glossy integument, a direct continuation of the common test of the body, covers the corneal lenses, quite as is the case in so many of the recent Crustacea. The lenses are closely packed, minute, usually hexagonal in outline, flat on the outer and convex on the inner surface. Such eyes are best developed in _Asaphus_, _Illaenus_, _Nileus_, _b.u.mastus_, _Proetus_, etc."

2. Eyes with biconvex lenses.

The surface of the eye is a ma.s.s of contiguous lenses, covered by a thin membrane which is frequently absent from the specimens, due to poor preservation. The lenses are biconvex, and being in contact with one another, are usually hexagonal, although in some cases they nearly retain their globular shape. Such eyes are found in Bury care, _Peltura_, _Sphaeropthalmus_, _Ctenopyge_, _Goldius_, _Cheirurus_, and probably others.

II. Aggregate eyes.

The individual lenses are comparatively large, distinct from one another, each lying in its own socket. There is, however, a thin membrane, which covers all those in any one aggregate, and is a continuation of the general integument of the body. This membrane is continued as a thickened infolding which forms the sockets of the lenses.

Such eyes are known in the Phacopidae only.

III. Stemmata and ocelli.

The stemmata are present only in _Harpes_, where there may be on the summit of the cheek two or three ocelli lying near one another.

Each, viewed from above, is nearly circular in outline, almost hemispheric, glossy and shining. In section they prove to be convex above and flat or slightly concave beneath. The test covers and separates them, as in the case of the aggregate eyes.

The ocelli of the Trinucleidae and _Eoharpes_ are smaller, and the detailed structure not yet investigated.

Lindstroem concludes that so far as its facets or lenses are concerned, the eye of the trilobite shows the greatest a.n.a.logy with the Isopoda, and the least with _Limulus_.

SUMMARY.

The simplest eyes found among the Trilobita are the ocelli. These consist of a Simple thickening of the test to form a convex surface capable of concentrating light. The similarity in position of the paired ocelli of trilobites and the simple eyes of copepods has perhaps a significance.

The schizochroal eyes may well be compared with the aggregate eyes of the chilopods and scorpions. The mere presence of a common external covering is not sufficient to prove this a true compound eye, especially as the covering is merely a continuation of the general test.

The holochroal eyes are of two kinds, one with plano-convex and one with biconvex lenses. The latter would seem to be mechanically the more perfect of the two, and it is worthy of note that the trilobites possessing the biconvex lenses have, in general, much smaller eyes than those with the other type.

If, as some investigators claim, the parietal eye of Crustacea originates by the fusion of two lateral ocelli, trilobites show a primitive condition in lacking this eye, which may have originated through the migration toward the median line of ocelli like those of the Trinucleidae.

s.e.x.

That the s.e.xes were separate in the Trilobita there can be very little doubt, but the study of the appendages has as yet revealed nothing in the way of s.e.xual differences. One of the most important points still to be determined is the location of the genital openings.

In many modern Crustacea, the antennae or antennules are modified as claspers, and it is barely possible that the curious double curvature of the antennules of Triarthrus indicates a function of this sort. The antennules of many specimens have the rather formal double curvature, turning inward at the outer ends, and retain this position of the frontal appendages, no matter what may be the condition of those on the body. Other specimens have the antennules variously displaced, indicating that they are quite flexible. It is conceivable that the individuals with rigid antennules are males, the others females.

It is interesting to note that the antennules of _Ptychoparia permulta_ Walcott (1918, pl. 21, fig. 1) have the same recurved form.

All the specimens of Neolenus, however, show very flexible antennas.

Barrande and Salter laid great stress upon the "forme longue" and "forme large" as indicating male and female. This was based upon the supposition that the female of any animal would probably have a broader test than the male, a hypothesis which seems to be very little supported by fact. In practical application it was found that the apparent difference was so often due to the state of preservation or the confusion of two or more species, that for many years little reference has been made to this supposed s.e.x difference.

EGGS.

In his cla.s.sic work on the trilobites of Bohemia, Barrande described three kinds of spherical and one of capsule-shaped bodies which he considered to be the eggs of trilobites. After a review of the literature and a study of specimens in the collections of the Museum of Comparative Zoology, it can be said that none of these fossils has proved to be a trilobite egg, but that they may be plants. A full account of them will be published elsewhere.

Walcott (1881) and Billings (1870) have described similar bodies within the tests of _Calymene_ and _Ceraurus_, but without showing positive evidence as to their nature.

Methods Of Life.

This is a subject upon which much can be inferred, but little proved.

Without trying to cover all possibilities, it may be profitable to see what can be deduced from what is known of the structure of the external test, the internal anatomy, and the appendages. This can, to a certain extent, be controlled by what is inferred from the strata in which the specimens are found, the state of preservation, and the a.s.sociated animals. (For other details, see the discussion of "Function of the Appendages" in Part I.)

HABITS OF LOCOMOTION.

The methods of locomotion may be deduced with some safety from a study of the appendages, and, as has repeatedly been pointed out, all trilobites could probably swim by their use. This swimming was evidently done with the head directed forward, and could probably be accomplished indifferently well with either the dorsal (gastronectic, Dollo) or the ventral (notonectic) side up. If food were sought on the bottom by means of sight, the animal would probably swim dorsal side up, for by canting from side to side it could see the bottom just as easily as though it were ventral side up, and at the same time it would be in position to drop quickly on the prey. In collecting food at the surface, it might swim ventral side up.

All trilobites could probably crawl by the use of the appendages, and, as has already been pointed out, there are great differences in the adjustment of the appendages to different methods of crawling. Some crawled on their "toes," some by means of the entire endopodites, and some apparently used the c.o.xopodites to push themselves along. That the normal direction of crawling was forward is indicated by the position of the eyes and sensory antennules. There is no evidence that their mechanism was irreversible, however, and the position of the mouth and the shape of the hypostoma indicate that they usually backed into feeding position. The caudal rami of Neolenus were evidently sensory, and the animal was prepared to go in either direction.