The Appendages, Anatomy, and Relationships of Trilobites Part 3

The cast of the interior shows, rather faintly, the exopodites of the right side of the thorax and of the left side of the cephalon, and, still more faintly, the caudal rami and a few pygidial endopodites.

The exopodites on the right side are in what seems to be the customary position, directed obliquely forward and outward, and the tips of their distal lobes project slightly beyond the edge of the test. These lobes were interpreted by Walcott as epipodites, but after comparing them with the terminal lobes of the exopodites of specimens No. 65519 and 65514 I think there can be no doubt that they represent the same structure. The pleura of the individual thoracic segments on this side of the specimen have an unusual appearance, for they are bluntly rounded or obtusely pointed, instead of being spinose.

The interpretation of the appendages of the cephalon is somewhat difficult. At the left of the glabella there are two large exopodites, the anterior of which lies over and partially conceals the other.

These show by their position that they belong to the fourth and fifth cephalic appendages. In front of these lie two appendages which may be either endopodites or exopodites, but which I am inclined to refer to the latter. Both are narrow and shaped like endopodites, but bear on their outer edges close-set fine setae. They also show what might be considered as faint traces of segmentation. If the first of these ran under the end of the exopodite behind it, as shown in Walcott's figure (pl. 22), then it would be necessary to interpret it as an endopodite, but it really continues down between the exopodite and the glabella, and seems to be attached opposite the middle of the eye. The specimen does not indicate clearly whether this appendage is above or below the exopodite behind it, but one's impression is that it is above, in which case it also must be an exopodite. The appendage in front, being similar, is similarly interpreted. If this be correct, then the exopodites of the second and third cephalic appendages are much shorter and narrower than those of the fourth and fifth. All of these appendages are obviously out of position, for the cheek has been pushed forward away from the thorax, though still pivoting on its inner angle at the neck-ring, till the eye has been brought up to the dorsal furrow. In this way the anterior exopodites have been thrust under the glabella and all the appendages have been moved to the right of their original position. The anterior exopodite is very poorly shown, but seems to be articulated in front of the eye. The posterior exopodites are very similar to those on the thorax. The distal lobe is shown only by the second from the last. It has the same form as the distal lobes on the thoracic exopodites, and like them has much finer setae than the main lobe, but it does not stand at so great an angle with the axis of the main lobe, nor yet is it so straight as shown in Walcott's figure.

_Measurements:_ The specimen is about 72 mm. long and 54 mm. wide at the genal angles. The pygidium is 22 mm. long and 37 mm. wide. The doublure is 1.5 mm. wide. The exopodite of the third thoracic segment is 19.5 mm. long. The pleural lobe at this point is 13 mm. wide without the spines and 18.5 mm. wide with them. The third exopodite of the cephalon was apparently about 15 mm. long when complete.

Specimen No. 65515.

Ill.u.s.trated: Walcott, Smithson. Misc. Coll., vol. 67, 1918, pl. 20, figs. 3, 4.

This is a small piece of the axial portion of a badly crushed Neolenus, showing appendages on the left side as viewed from above. On the posterior half there are three large appendages which have the exact form of the exopodites of other specimens. There is a broad, oval, proximal lobe and a distal one at an angle with it. The proximal part of the shaft has fine setae or the bases of them, and the distal lobe faint traces of much finer ones. The form, and the setae so far as they are preserved, are exactly like those of the exopodites on the specimens previously described. (See fig. 4, page 26.) Beneath them there are slender, poorly preserved endopodites.

In front of the exopodites and endopodites lie a series of structures which Walcott has called exites, but for which I can see another explanation. Walcott has shown them as four broad rounded lobes, but his figure must be looked upon as a drawing and not as a photograph, for it has been very much retouched.

For convenience of discussion, these lobes may be called Nos. 1, 2, 3, and 4, the last being the posterior one (fig. 5). This lobe is best shown on the matrix, where the anterior end is seen to be margined by stout spines, while the posterior end lies over the endopodite and under the exopodite behind it. No. 3 is sunk below the level of the others, and only a part of it has been uncovered. Its margin bears strong spines of different sizes. Its full shape can not be made out, but it has neither the shape nor the form of spines shown in figure 3, plate 20 (1918). Lobes 2 and 1 and another lobe in front of 1 seem to form a continuous series and to be part of a single appendage. They are all in one plane, arc so continuous that the joints between them can be made out with difficulty and if they do belong together, can easily be explained.

[Ill.u.s.tration: Fig. 5.--A sketch of the so-called exites of _Neolenus serratus_ (Rominger), to show the form and the character of the spines. 2.]

[Ill.u.s.tration: Fig. 6.--Endopodite of a cephalic appendage of _Neolenus serratus_ (Rominger), showing the very broad c.o.xopodite.

2.]

Before calling these structures new organs not previously seen on trilobites, it is of course necessary to inquire if they can be interpreted as representing any known structures. That they can not be exopodites is obvious, since they are bordered by short stout spines instead of setae. The same stout spines that negate the above possible explanation at once suggest that they are c.o.xopodites (compare fig 6).

At first sight, the so-called exites seem too wide and too rounded to be so interpreted, but if reference be had to the specimens rather than the figures, it will be noted that the only well preserved structure (No. 2) is longer than wide, has spines only on one side and one end, and does not differ greatly from the c.o.xopodite of specimen No. 58589 (pl. 18, 1918). If structures 2, 1, and the segment ahead of 1 are really parts of one appendage, it can only be an endopodite, of which No. 2 is the c.o.xopodite, No. 1 the basipodite, and the next segment the ischiopodite. If one looks carefully, there are no traces of spines on either end of No. 1, but only on the margin. The extreme width of No. 2 is against this interpretation as a c.o.xopodite (see, however, fig. 6), but it may be rolled out very flat, as this is an unusually crushed specimen. No. 2 is 10 mm. long and 6 mm. wide at the widest point. No. 1 is 5 mm. long and 3.5 mm. wide.

The crucial point in this determination is whether 2 and 1 are parts of the same appendage. I believe they are, but others may differ.

Specimen No. 65513.

Ill.u.s.trated: Walcott, Smithson. Misc. Coll., vol. 57, 1912, pl. 45, fig. 3;--Ibid., vol. 67, 1918, pl. 16, figs. 1, 2.

This is nearly all of the right half of an entire specimen, but the only appendages of any interest are those of the cephalon. Five endopodites emerge from beneath that shield, but as all are displaced it is not possible to say how many belong to the head. When held at the proper angle to the light, the second and third from the front show faintly the partial outlines of the c.o.xopodites. The anterior side and end of the best preserved one shows irregular stout spines of unequal sizes, and the inner end is truncated obliquely (fig. 6).

These c.o.xopodites are like those on the thorax of specimen No. 58589, but shorter and wider. This of course suggests that the "exite" No. 2 of specimen No. 65515 may be a cephalic c.o.xopodite. The endopodite of this appendage, like the others on this cephalon, is shorter and stouter than the thoracic or pygidial endopodites of the others described.

[Ill.u.s.tration: Fig. 7.--A restored section across the thorax of _Neolenus serratus_, showing the probable form of attachment of the appendages, their relation to the ventral membrane, and the jaw-like endobases of the c.o.xopodites.]

_Measurements:_ The cephalon is 24 mm. long and about 60 mm. wide. The c.o.xopodite of the third appendage is about 10 mm. long and 5.5 mm.

wide at the widest point. The corresponding endopodite is 19 mm. long and projects 11 mm. beyond the margin, which is about 5 mm. further than it would project were the appendage restored to its proper position.

RESTORATION OF NEOLENUS.

(Text figs. 7, 8.)

This restoration is based upon the information obtained from the studies which have been detailed in the preceding pages, and differs materially from that presented by Doctor Walcott. The appendages are not shown in their natural positions, but as if flattened nearly into a horizontal plane. The metastoma is added without any evidence for its former presence.

The striking features of the appendages are the broad unsegmented exopodites which point forward all along the body, and the strong endopodites, which show practically no regional modification. Although the exopodites have a form which is especially adapted for use in swimming, their position is such as to indicate that they were not so used. The stout endopodites, on the other hand, probably performed the double function of natatory and ambulatory legs.

[Ill.u.s.tration: Fig. 8.--_Neolenus serratus_ (Rominger). A restoration of the ventral surface, with the endopodites omitted from one side, to permit a better exposition of the exopodites. The position and number of the appendages about the mouth are in considerable doubt. Restored by Doctor Elvira Wood under the supervision of the writer. About one-half larger than the average specimen.]

=Nathorstia transitans= Walcott.

Ill.u.s.trated: Walcott, Smithson. Misc. Coll., vol. 57, 1912, pl. 28, fig. 2.

The badly preserved specimen on which this genus and species was based is undoubtedly a trilobite, but for some reason it does not find a place in Walcott's recent article on "Appendages" (1918). The preservation is different from that of the a.s.sociated trilobites, being merely a shadowy impression, indicating a very soft test. The general outline of the body, the position of the eye, and even a trace of spines about the pygidium (in the figure) are similar to those of _Neolenus_, and I would venture the suggestion that _Nathorstia transitans_ is a recently moulted _Neolenus serratus_, still in the "soft-sh.e.l.led" condition. Even if not a Neolenus, it is probable, from the state of preservation, that it is an animal which had recently cast its sh.e.l.l.

Walcott describes such fragments of appendages as remain, as follows:

Head. A portion of what may be an antenna projects from beneath the right anterior margin; from near the left posterolateral angle a large four-jointed appendage extends backward. I a.s.sume that this may be the outer portion of the large posterior appendage (maxilla) of the head.

Thorax. Traces of several slender-jointed thoracic legs project from beneath the anterior segments and back of these on the right side more or less of six legs have been pushed out from beneath the dorsal shield; these are composed of three or four long slender joints; fragments of the three proximal joints indicate that they are shorter and larger and that they have a fringe of fine setae.

Indications of a branchial lobe (gill) are seen in two specimens where the legs are not preserved. This is often the case both among the Merostomata (pl. 29, fig. 3, _Molaria_) and Trilobita (pl. 24, fig. 2, _Ptychoparia_).

Two caudal rami project a little distance beneath the posterior margin of the dorsal shield.

This latter feature of course suggests _Neolenus_. The other appendages are too poorly preserved to allow comparison without seeing the specimen.

The specific name was given "on account of its suggesting a transition between a Merostome-like form, such as _Molaria spinifera_, and the trilobites." In what respect it is transitional does not appear.

Formation and locality: Same as that of _Neolenus serratus_. One nearly complete specimen and a few fragments were found.

The Appendages of Isotelus.

HISTORICAL.

The first specimen of _Isotelus_ with appendages was described orally by Billings before the Natural History Society of Montreal in 1864, and in print six years later (1870, p. 479, pls. 31, 32). The specimen is described in detail on a later page. Billings recognized the remains of eight pairs of legs on the thorax, a pair for each segment, and he inferred from the fact that the appendages projected forward that they were ambulatory rather than natatory organs. He was unable to make out the exact number of the segments in the appendages, but thought each showed at least four or five.

Having examined the individual sent to London by Billings, Woodward (1870, p. 486, fig, 1) reviewed the collection from the American Trenton in the British Museum and found a specimen in the "Black Trenton limestone," from Ottawa, Ontario, in which, alongside the hypostoma, was a jointed appendage, which he described as the "jointed palpus of one of the maxillae." This has always been considered an authentic "find," but I am informed by Doctor Bather that the specimen does not show any real appendage. For further discussion, see under _Isotelus gigas_.

In 1871, Billings' specimen was examined by Professors James D. Dana (1871, p. 320), A. E. Verrill, and Sydney I. Smith, who agreed that the structures identified by Billings as legs were merely semicalcified arches of the membrane of the ventral surface, which opinion seems to have been adopted by zoologists generally in spite of the fact that the most elementary consideration of the structure of the thorax of a trilobite should have shown its falsity. While the curvature of the thoracic segments was convex forward, that of the supposed ventral arches was convex backward, and the supposed arches extended across so many segments as to have absolutely prevented any great amount of motion of the segments of the thorax on each other.

Enrollment, a common occurrence in _Isotelus_, would have been absolutely impossible had any such calcified arches been present.

Walcott, in his study of trilobites in thin section (1881, pp. 192, 206, pl. 2, fig. 9), obtained eleven slices of _Isotelus gigas_ which showed remains of appendages. He figured one of the sections, stating that it "shows the basal joint of a leg and another specimen not ill.u.s.trated gives evidence that the legs extended out beneath the pygidium, as indicated by their basal joints."

The second important specimen of an _Isotelus_ with appendages was found by Mr. James Pugh in strata of Richmond age 2 miles north of Oxford, Ohio, and is now in the U. S. National Museum. It was first described by Mickleborough (1883, p. 200, fig. 1-3). In two successive finds, a year apart, the specimen itself and its impression were recovered. Since I am redescribing the specimen in this memoir (see p. 35), it only remains to state here that Mickleborough interpreted the structures essentially correctly, though not using the same terminology as that at present adopted. His view that the anterior appendages were chelate can not, however, be supported, nor can his idea that the sole appendages of the pygidium were foliaceous branchial organs.

Walcott (1884, p. 279, fig. 1) studied the original specimens and presented a figure which is much more detailed and clear than those of Mickleborough. By further cleaning the specimen he made out altogether twenty-six pairs of appendages. He stated that one of these belonged to the cephalon, nine to the thorax,[1] and the remaining sixteen to the pygidium. He showed that the endopodites of the pygidium were of practically the same form as those on the thorax, and stated that the "leg beneath the thorax of the Ohio trilobite shows seven joints in two instances; the character of the terminal joint is unknown." His figure shows, and he mentions, markings which are interpreted as traces of the fringes of the exopodites.

[Footnote 1: The posterior one of these he believed to have been crowded forward from beneath the pygidium.]

In the same year Woodward (1884, p. 162, fig. 1-3) reproduced all of Mickleborough's figures, and suggested that the last seven pairs of appendages on the pygidium of _Calymene_ and _Isotelus_ were probably "lamelliform branchiferous appendages, as in _Limulus_ and in living Isopoda."

Professor Beecher published, in 1902, an outline taken from Mickleborough's figure of this specimen, to call attention to certain discontinuous ridges along the axial cavity of the anterior part of the pygidium and posterior end of the thorax. These ridges are well shown in Mickleborough's figure, though not in that of Walcott, and their presence on the specimen was confirmed by a study by Schuchert, who contributed a diagrammatic cross-section to Beecher's paper (1902, p. 169, pl. 5, figs. 5, 6). Beecher summarized in a paragraph his interpretation of this specimen: