The Appendages, Anatomy, and Relationships of Trilobites Part 14

Take now the trilobites. There is no trace of the median pustule in the protaspis of any form, and in many primitive trilobites it is absent. It appears first as a long spine in certain families, and later becomes vestigial and disappears. Very few trilobites of Silurian and later times show it at all.

In the particular case of the Trinucleidae, which were burrowers, the spine is present on only the oldest and most primitive of the group, a form which has only a most rudimentary fringe. It is obvious from the large size of the pygidium in the larval trinucleid that this family is derived from a group of free swimmers. _Trinucleoides reussi_ was perhaps in the transitional stage, just leaving the swimming mode of life, and belonged to a group which had not developed any other "statocyst" than the median spine. Among the later Trinucleidae the spine became a vestigial tubercle, and in some cases entirely disappeared. A similar history can be traced in the Cheiruridae, starting from some such forms as the American Lower Ordovician _Nieszkowskia_ (_N. perforator_ p. ex.).

Another example of a median spine instead of a tubercle is in Goldius rhinoceros (Barrande). Since this species is not from the oldest Goldius-bearing rocks, but from the Lower Devonian, it does not follow what seems to be the general rule, but makes an interesting exception.

Goldius rhinoceros (Barrande) (Supplement, 1872, pl. 9, figs. 12, 13) has the median tubercle elevated into a stubby, recurved spine very suggestive of the horn of a rhinoceros. Since the eyes of this species are very well developed, there seems no especial reason for the elevation of a parietal eye, and the example certainly does not support that interpretation.

3. This tubercle is essentially similar to other tubercles on the median line of cephalon, thorax, and even pygidium. This has been discussed sufficiently under section 1 above, but it may perhaps be justifiable to point out that in some of the Agnostidae there is a median tubercle on both s.h.i.+elds, and since it has not yet been demonstrated beyond question which s.h.i.+eld is the cephalon, to say which one is a parietal eye and which one is a tubercle is impossible.

In other words, the parietal eye can not be differentiated from any other tubercle except by its position.

4. One of the as yet unexplained features of the protaspis of trilobites is the absence of the "nauplius eye." Beecher (1897 B, p.

40) explained this on the ground of the extremely small size of the protaspis and the imperfection of the preservation. If the median tubercle were really a median eye, it should be present in the protaspis and the earlier stages of the ontogeny, even if not in the adult, and should certainly appear before the compound eyes. (In _Limulus_, however, the compound eyes appear first.) The median eye has not so far been seen in any young trilobite in any stage previous to that in which compound eyes are present. The full ontogeny is not known of any species with compound eyes in which the median tubercle is present in the adult, but theoretically the median eye should be most prominent in the young of just those primitive trilobites about whose development most is known.

NERVOUS SYSTEM.

There has been a rather general impression among students of trilobites that the eye-lines, which should be differentiated from the genal caeca, denote the course of the optic nerves, but no other evidence of the nervous system has been found, save the so-called nervures which have been discussed above. In _Apus_ the nerves leading to the eyes come off from the anterior ganglion or "brain" and run directly to the eyes. If conditions were similar in the trilobites, the "brain" was beneath the anterior glabellar lobe, provided, of course, that the eye-lines do indicate the course of the optic nerve.

The ontogenetic history of the eye-lines of trilobites with compound eyes is instructive, and has already been discussed by Lindstroem (1901, pp. 12-25), but he did not cite the case of _Ptychoparia_, which is particularly interesting, because in this genus both eye-lines and "nervures" are present. Beecher (1895 C, p. 171, pl. 8, figs. 5-7) has shown that in _Ptychoparia kingi_ the eye-lines of a specimen in the metaprotaspis stage run forward at a low angle with the glabella, while in the adult their course is nearly at right angles to it. They have therefore swung through an arc of at least 60 and in so doing have had ample opportunity to become coincident with the primary trunks of the genal caeca. Once that was accomplished, it is quite likely that the one fold in the sh.e.l.l would continue to house both structures. In other trilobites, there is a similar backward progression of the eye-lines.

As would be expected, the ventral ganglia and the longitudinal cords left no trace in the test. Since each segment has appendages, there was probably a continuous chain of ganglia back to the posterior end of the pygidium.

VARIOUS GLANDS.

_Dermal glands._--The surface of many trilobites is "ornamented" with pustules and spines which on sectioning are nearly always found to be hollow, and in many cases have a fine opening at the tip. While it is generally believed that the purpose of these spines was protective, yet it is possible that many of them were merely outgrowths which increased the area through which the respiratory function could be carried on. It will be recalled that most of the smooth trilobites are punctate, some of them very conspicuously so, and the spines and pustules of ornamented trilobites may merely subserve the same function as the pores of smooth ones.

If the spines were protective, it would not be surprising if some of them, hollow and open at the top, were poisonous also, and had glands at the base. These are, however, purely matters of speculation so far.

_Renal excretory organs._--Nothing has been seen of any such organs, unless the genal caeca may possibly be of that nature. The main trunks of these always lead to the sides of the anterior glabellar lobe, which is not the point of attachment of either antennae or biramous limbs, so that there seems little chance that they will bear this interpretation.

_Reproductive organs._--Nothing is yet positively known about the reproductive organs or the position of their external openings. If the "exites" of _Neolenus_ could be interpreted as brood-pouches, which does not seem probable, then the genital openings were located near the base of some pair of anterior thoracic appendages.

_The Panderian Organs: Internal Gills or Poison Glands?_

At a meeting of the Mineralogical Society at St. Petersburg, Volborth (1857) announced that Doctor Pander had two years before discovered certain organs on the lower side of the doublure of the pleural lobes of the thorax of a specimen of _Asaphus expansus_. These organs were oval openings in the doublure, one near the posterior margin of the cephalon, and one on each thoracic segment of the half-specimen figured by Volborth in 1863. They were explained by Volborth and by Eichwald (1860, 1863) as the points of attachment of appendages.

Billings (1870) described and figured the "Panderian organs" of "_Asaphus platycephalus_" and stated that he had seen them in _Asaphus_ [_Ogygites_] _canadensis_ and _A. megistos_ [_Isotelus maximus_] as well. He thought some sort of organ was attached to them, but could not suggest its function. Woodward (1870) thought that the openings were "only the fulcral points on which the pleurae move."

Their position outside the fulcra shows that this explanation is impossible.

So far as I am aware, the Panderian organs have been seen only in the Asaphidae. Barrande figured them in "_Ogygia_" [_Hemigyraspis_]

_desiderata_ (1872) and Schmidt in two species of _Pseudasaphus_. They seem to occupy the same position in Bohemian, Russian, and American specimens. There is always one pair of openings on each thoracic segment, and one pair in line with them on the posterior margin of the cephalon. They occur near the anterior margin of the segment, and near the inner end of the doublure. In some cases they are surrounded by a ventrally projecting rim, while in others they have a thin edge. There seem to be no markings on the interior of the sh.e.l.l which are connected with them.

While thinking over the trilobites in connection with the origin of insects, it occurred to me that these hitherto unexplained Panderian organs might possibly be openings to internal gills and that the Asaphidae might have been tending toward an amphibious existence.

On mentioning this to Doctor R. V. Chamberlin of the Museum of Comparative Zoology, he called my attention to the possibility that they might be openings similar to those of the repugnatorial glands of Diplopoda. While no definite decision as to the function can be made, the explanation offered by Doctor Chamberlain seems more plausible than my own, and has suggested still a third, namely, that they might be the openings of poison glands.

If one were to argue that these apertures are really connected with respiration, it might be pointed out that they are ventral in position, while the _foramina repugnatoria_ are always dorsal or lateral, even in diplopods with broad lateral expansions. If offensive secretions were poured out beneath a concave sh.e.l.l like that of a trilobite, they would be so confined as to be but slightly effective against an enemy. This would indicate that if these openings were the outlets of glands, the substance secreted might be a poison used to render prey helpless. On the other hand, openings to gills are normally ventral in position, and if the pleural lobes were folded down against the body, they would be brought very close to the bases of the legs.

A further curious circ.u.mstance is that so far no traces of exopodites have been found on _Isotelus_. The endopodites of both _Isotelus latus_ and _I. maximus_ are fairly well preserved in the single known specimen of each, yet no authentic traces of exopodites have been found with them. Moreover, Walcott sliced specimens of _Isotelus_ from Trenton Falls and found only endopodites. It may also be recalled that the finding of the specimen of _Isotelus arenicola_ at Britannia and the tracks which I attributed to it, suggested to me that it was a sh.o.r.e-loving animal (1910). It offers a field for further inquiry, whether the Asaphidae may not have had internal gills, and whether some primitive member of the family may not have given rise to tracheate arthropods.

[Ill.u.s.tration: Fig. 28. Side view of a specimen of _Isotelus gigas_ Dekay, from which the test of the pleural lobes has been broken to show the position of the Panderian organs. Natural size. Specimen in the Museum of Comparative Zoology.]

The explanation of the Panderian organs as openings of poison glands is less radical than the one just set forth, and so possibly lies nearer the truth. One would expect poison glands to lie at the bases of fangs, and so they do in specialized animals like chilopods and scorpions, but the trilobites may have had the less effective method of pouring out the poison from numerous glands. The purpose may have been merely to paralyze the brachiopod or pelecypod which was incautious enough to open its sh.e.l.l in proximity to the asaphid.

MUSCULATURE.

This is a field which is rather one for investigation than for exposition. Very little has been done, though probably much could be.

The chief obstacle to a clearer understanding of the muscular system lies in the difficulty of getting at the inner surface of the test without obscuring the faint impressions in the process.

There exist in the literature a number of references to scars of attachment of muscles, and any study of the subject should of course begin by the collection of such data. I shall at this time refer to only a few observations on the subject.

The structure and known habits of trilobites make it obvious that strong flexor and extensor muscles must have been present, and some trace of them and of their points of attachment should be found. It is likely that their proximal ends were tough tendons. The muscles holding up the heart and alimentary ca.n.a.l would be less likely to reveal their presence by scars, but there must have been at least one pair of strong muscles extending from the under side of the head across to the hypostoma. Judging from the method of attachment, the muscles moving the limbs were short ones, chiefly within the segments of the legs themselves.

_Flexor Muscles._

Since the majority of trilobites had the power of enrollment, and seem also to have used the pygidia in swimming, the flexors must have been important muscles. Beecher (1902, p. 170) appears to have been the only writer to point out any tangible evidence of their former presence. Walcott (1881, p. 199) had shown that the ventral membrane was reinforced in each segment by a slightly thickened transverse arch. Beecher showed that on this thickened arch in _Triarthrus_, _Isotelus_, _Ptychoparia_, and _Calymene_, there are low longitudinal internal ridges or folds. One of these is central, and there is a pair of diagonal ridges on either side. Beecher interpreted these ridges as separating the strands of the flexor muscles, and believed that a line of median ridges divided a pair of longitudinal muscles, while the outer ridges showed the place of attachment of the pair of strands which was set off to each segment. He did not discuss the question as to where the anterior and posterior ends were attached. In trilobites with short pygidia, the attachment would probably have been near the posterior end, and it is possible that the two scars beneath the doublure and back of the last appendifers in _Ceraurus_ may indicate the point of attachment in that genus.

There is as yet no satisfactory evidence as to where the anterior ends of the flexors were attached. In _Apus_ these muscles unite in an entosternal sinewy ma.s.s above the mouth, but no evidence of any similar ma.s.s has been found in the trilobites and it is likely that the muscles were anch.o.r.ed somewhere on the test of the head.

_Extensor Muscles._

The exact position of these muscles has not been previously discussed.

The interior of the dorsal test shows no such apodemes as are found on the mesosternites, but, as I have shown in the discussion of the alimentary ca.n.a.l of _Calymene_ and _Ceraurus_, there is an opening on either side of the axial lobe between the dorsal test and the abdominal sheath, and it is entirely probable that an extensor muscle pa.s.sed through each of these. The abdominal sheath extends only along the posterior region of the glabella and the anterior part of the thorax, and probably served to protect the soft organs from the strain of the heavy muscles. The extensors (see fig. 29) probably lay along the top of the axial lobe on either side of the median line of the thorax to the pygidium, where they appear to have been attached chiefly on the under side of the anterior ring of the axial lobe, although strands probably continued further back. This is above and slightly in front of the fulcral points on the pleura, and meets the mechanical requirements. _Ceraurus_ (Walcott, 1875, and 1881, p. 222, pl. 4, fig. 5) shows a pair of very distinct scars on the under side of the first ring of the pygidium, and in many other trilobites (_Illaenus_, _Goldius_, etc.) distinct traces of muscular attachment can be seen in this region, even from the exterior. The anterior ends were probably attached by numerous small strands to the top of the glabella, and, princ.i.p.ally, to the neck-ring.

On enrolling, the sternites of all segments are pulled forward and the tergites backward. In straightening out, the reverse process takes place. The areas available for muscular attachment are so disposed as to indicate longitudinal flexor and extensor muscles rather than short muscles extending from segment to segment. Indeed, the tenuity of the ventral membrane is such as to preclude the possibility of enrollment by the use of muscles of that sort, while powerful longitudinal flexors could have been anch.o.r.ed to cephalon and pygidium. The strongly marked character of the neck-ring of trilobites is probably to be explained as due to the attachment of the extensor muscles, rather than to its recent incorporation in the cephalon. The same is true of the anterior ring on the pygidium.

[Ill.u.s.tration: Fig. 29. Restoration of a part of the internal organs of _Ceraurus pleurexanthemus_ as seen from above. At the sides are the extensor muscles, and in the middle, the heart overlying the alimentary ca.n.a.l. Drawn by Doctor Elvira Wood, under the supervision of the author.]

_Possible preservations of extensors and flexors in Ceraurus_.--Among Doctor Walcott's sections are four slices which I should not like to use in proving the presence of longitudinal muscles, but which may be admitted as corroborative evidence. Two of these transverse sections (Nos. 114 and 199) show a dorsal and a ventral pair of dark spots in positions which suggest that they represent the location of the dorsal and ventral muscles, while two others (Nos. 131 and 140) show only the upper pair of spots. The chief objection to this interpretation is that it is difficult to imagine how the muscles could be so replaced that they happen to show in the section. Both the sections showing all four spots are evidently from the anterior part of the thorax, as they show traces of the abdominal sheath, which seems to be squeezed against the inside of the axial lobe, with the muscles (?) forced out to the sides. The ventral pair lie just inside the appendifers, but even if they are sections of muscles, all four are probably somewhat out of place.

_Hypostomial Muscles._

The hypostoma fits tightly against the epistoma, or the doublure when the epistoma is absent, but in no trilobite has it ever been seen ankylosed to the dorsal test, and its rather frail connection therewith is evidenced by the relative rarity of specimens found with it in position. That the hypostoma was movable seems very probable, and that it was held in place by muscles, certain. The maculae were always believed to be muscle scars until Lindstroem (1901, p. 8) announced the discovery by Liljevall of small granules on those of _Goldius polyactin_ (Angelin). These were interpreted as lenses of eyes by Lindstroem, who tried to show that the maculae of all trilobites were functional or degenerate eyes. Most palaeontologists have not accepted this explanation, and since the so-called eyes cover only a part of the surface of the maculae, it is still possible to consider the latter as chiefly muscle-scars.

In Lindstroem's summary (1901, pp. 71, 72) it is admitted that the globular lenses are found only in _Bronteus_ (_Goldius_) (three Swedish species only) and _Cheirurus spinulosus_ Nieszkowski, while the prismatic structure supposed to represent degenerate eyes was found in eleven genera (Asaphidae, Illaenidae, Lichadidae). All of these are forms with well developed eyes, and Lindstroem himself points out that the appearance of actual lenses in the hypostoma was a late development, long after the necessity for them would appear to have pa.s.sed.

The use of the hypostoma has been discussed by Bernard (1892, p. 240) and extracts from his remarks are quoted:

The earliest crustacean-annelids possessed large labra or prostomia projecting backward, still retained in the Apodidae and trilobites.

This labrum almost necessitated a very deliberate manner of browsing. The animal would creep along, and would have to run some way over its food before it could get it into its mouth, the whole process, it seems to us, necessitating a number of small movements backwards and forwards. Small living prey would very often escape, owing to the fact that the animal's mouth and jaws were not ready in position for them when first perceived. The labrum necessitates the animal pa.s.sing forwards over its prey, then darting backward to follow it with its jaws. We here see how useful the gnathobases of _Apus_ must be in catching and holding prey which had been thus pa.s.sed over. Indeed the whole arrangement of the limbs of _Apus_ with the sensory endites forms an excellent trap to catch prey over which the labrum has pa.s.sed.

In alcoholic specimens of _Apus_ the labrum is not in a horizontal plane, as it is in most well preserved trilobites, but is tipped down at an angle of from 30 to 45, and the big mandibles lie under it. It has considerable freedom of motion and is held in place by muscles which run forward and join the under side of the head near its posterior margin. It seems entirely possible that the hypostoma of the trilobite had as much mobility as the labrum of _Apus_, and that byopening downward it brought the mouth lower and nearer the food. It will be recalled that the hypostomata of practically all trilobites are pointed at the posterior margin, there being either a central point or a pair of p.r.o.ngs. By dropping down the hypostoma until the point or p.r.o.ngs rested on or in the substratum, and sending food forward to the mouth by means of the appendages, a trilobite could make of itself a most excellent trap, and if the animal could dart backward as well as forward, the hypostoma would be still more useful.

There is no reason to suppose that they could not move backward, and the "pygidial antennae" of _Neolenus_ indicate that animals of that genus at least did so. This habit of dropping down the hypostoma would also permit the use of those anterior gnathobases which seem too far ahead of the mouth in the trilobites with a long hypostoma.

For actual evidence on this point, it is necessary to have recourse once more to Doctor Walcott's exceedingly valuable slices. From such sections of _Ceraurus_ as his Nos. 100, 106, 108, 170, and 173, it is evident that the hypostoma of that form could be dropped considerably without disrupting the ventral membrane (fig. 30). Sections of _Calymene_ already published (Walcott 1881, pl. 5, figs. 1, 2) show the hypostoma turned somewhat downward, and the slices themselves show sections of the anterior pair of gnathobases beneath the hypostoma.

When the hypostoma was horizontal, these gnathobases were crowded out at the sides.

[Ill.u.s.tration: Fig. 30.--Longitudinal section of cephalon of _Ceraurus pleurexanthemus_, to show position of the mouth and folds of the ventral membrane between the glabella and the hypostoma. The test is in solid black and the part within the ventral membrane dotted. From a photographic enlargement. Specimen 169. 3.9.]