PART I.. The Appendages of Trilobites.

THE APPENDAGES OF TRILOBITES.

Terminology.

The terminology employed in the succeeding pages is essentially the same as that used by Beecher, with two new terms added. Beecher assigned to the various segments of the limbs the names suggested by Huxley, but sometimes used the name protopodite instead of coxopodite for the proximal one. It is obvious that he did not use protopodite in the correct sense, as indicating a segment formed by the fusion of the coxopodite and basipodite. The usage employed here is shown in figure 1.

The investigation of Ceraurus showed that the appendages were supported by processes extending downward from the dorsal test, and on comparison with other trilobites it appeared that the same was true in Calymene, Cryptolithus, Neolenus, and other genera. Thin sections showed that these processes were formed by invagination of the test beneath the dorsal and glabellar furrows. While these processes are entirely homologous with the entopophyses of Limulus, I have chosen to apply the name appendifer to them in the trilobites.

The only other new term employed is the substitution of endobase for gnathobase in speaking of the inner prolongation of a coxopodite of the trunk region. The term gnathobase implies a function which can not in all cases be proved.

The individual portions of which the limbs are made up are called segments, and the articulations between them, joints. Such a procedure is unusual, but promotes clearness.

The Appendages of Neolenus.

HISTORICAL.

The first mention of Neolenus with appendages preserved was in Doctor Walcott's paper of 1911, in which two figures were given to show the form of the exopodites in comparison with the branchiæ of the eurypterid-like Sidneyia. In 1912, two more figures were presented, showing the antennules, exopodites, and cerci. The specimens were found in the Burgess shale (Middle Cambrian) near Field, in British Columbia. This shale is exceedingly fine-grained, and has yielded a very large fauna of beautifully preserved fossils, either unknown or extraordinarily rare elsewhere. It was stated in this paper (1912 A) that trilobites, with the exception of Agnostus and Microdiscus, were not abundant in the shale.

In discussing the origin of the tracks known as Protichnites, Walcott presented four figures of Neolenus with appendages, and described the three claw-like spines at the tip of each endopodite.

Three new figures of the appendages were also contributed to the second edition of the Eastman-Zittel "Text-book of Paleontology" (1913, p. 701). Later (1916, pl. 9) there was published a photograph of a wonderful slab, bearing on its surface numerous Middle Cambrian Crustacea. Several of the specimens of Neolenus showed appendages.

Finally, in 1918, appeared the "Appendages of Trilobites," in which the limbs of Neolenus were fully described and figured (p. 126), and a restoration presented. Organs previously unknown in trilobites, epipodites and exites, attached to the coxopodites, were found.

=Neolenus serratus= (Rominger).

(Text fig. 2-8.)

Illustrated: Walcott, Smithson. Misc. Coll., vol. 57, 1911, p. 20, pl. 6, figs. 1, 2 (exopodites of thorax and cephalon);--Ibid., vol. 57, 1912, p. 191, pl. 24, figs. 1, la (antennules, caudal rami, and endopodites of thorax);--Ibid., vol. 57, 1912, p. 277, pl. 45, figs. 1-4 (antennules, endopodites of cephalon and thorax, caudal rami);--Text-book of Paleontology, edited by C. R. Eastman, 2d ed., vol. 1. 1913, p. 701, fig. 1343 (exopodites), p. 716, fig. 1376 (abdominal appendages), fig. 1377 (appendages of thorax and pygidium);--Ann. Rept. Smithson. Inst. for 1915, 1916, pl. 9;--Smithson. Misc. Coll., vol. 67, 1918, pp. 126-131 et al., pl. 14, fig. 1; pls. 15-20; pl. 21, fig. 6; pls. 22, 23; pl. 31 (restoration); pl. 34, fig. 3 (restored section); pl. 35, fig. 4; pl. 36, fig. 3 (hypostoma).

The following description of the appendages of Neolenus is summarized from Walcott's paper of 1918, and from a study of the eight specimens mentioned below.

Cephalon.

The antennules are long, slender, and flexible, and lack the formal double curvature so characteristic of those of Triarthrus. There are short fine spines on the distal rims of the segments of the proximal half of each, thus giving great sensitiveness to these organs. In the proximal portion of each, the individual segments are short and wider than long, and in the distal region they are narrow and longer than wide.

There are four pairs of biramous cephalic appendages, which differ only very slightly from the appendages of the thorax. All are of course excessively flattened, and they are here described as they appear.

The coxopodites, shown for the first time in Walcott's paper of 1918, are broad, longer than wide, and truncated on the inner ends, where they bear short, stout, unequal spines similar to those along the anterior margin. The gnathobases are but slightly modified to serve as mouth parts, much less so than in Triarthrus, but the coxopodites of the cephalon are shorter and wider than those of the thorax.

At the distal end of the coxopodite arise the endopodite and exopodite. The endopodite consists of six segments, the distal ones, propodite and dactylopodite, more slender than the others, the last bearing three terminal spines. The first endopodite is shorter than the others and slightly more slender (pl. 16, fig. 1) and the anterior appendages turn forward more or less parallel to the sides of the hypostoma (pl. 22). The basipodite, ischiopodite, meropodite, and carpopodite are, in their flattened condition, roughly rectangular, only a little longer than wide, taper gradually distally, each bears small spines on the outer rim, and some of the proximal ones usually have a row along the margin.

The exopodites of the cephalon, as of the body of Neolenus, are very different from those of any other trilobite whose appendages were previously known. As shown in the photographs (pl. 20, fig. 2; pl. 22), each exopodite consists of a single long, broad, leaf-like blade, not with many segments as in Triarthrus, but consisting of a large basal and small terminal lobe. It bears on its outer margin numerous relatively short, slender, flat setæ. The long axes of the exopodites point forward, and the setæ are directed forward and outward. They stand more nearly at right angles to the shaft on the cephalic exopodites than on those of the thorax. This same type of broad-bladed exopodite is also found on the thorax and pygidium.

The number of functional gnathobases on the cephalon is unknown. That four endopodites were present on one side is shown pretty clearly by specimen 58591 (pl. 16, fig. 3) and while no more than two well preserved exopodites have been seen on a side, there probably were four. Specimen 65513 (pl. 16, fig. 1) shows gnathobases on the second and third appendages of that individual as preserved, but there is no positive evidence that these are really the second and third appendages, for they are obviously displaced. The hypostoma of Neolenus is narrow but long, several specimens showing that it extended back to the horizon of the outer ends of the last pair of glabellar furrows. It is not as wide as the axial lobe, so that, while gnathobases attached beneath the first pair of furrows would probably not reach back to the posterior end of the hypostoma, they might lie parallel to it and not extend beneath. It seems possible, then, that there were four pairs of endobases but that the second rather than the first pair served as mandibles, as seems to be the case in Ceraurus.

Thorax.

The thorax of Neolenus consists of seven segments, and the appendages are well shown (pl. 17, fig. 1; pl. 18, figs. 1, 2; pl. 20, fig. 1.), The endopodites of successive segments vary but little, all are slender but compact, and consist of a long coxopodite with six short, rather broad segments beyond it. In the figures, the endopodites extend some distance in a horizontal direction beyond the edges of the dorsal test, as many as four segments being in some cases visible, but measurements show that the appendages tended to fall outward on decay of the animal. The dactylopodites are provided with terminal spines as in Triarthrus. The coxopodites are long, straight, and slender. They are well shown on only one specimen (pl. 18), where they are seen to be as wide as the basipodite, and the endobases are set with spines on the posterior and inner margins. They are so long that those on opposite sides must have almost met on the median line. The segments of the endopodites are mostly but little, if any, longer than broad, and at the distal end each shows two or more spines. The propodite and dactylopodite are notably more slender than the others. The exopodites of the thorax are broad and flat, and each shaft has two distinct parts with different kinds of setæ. The posterior edge of the proximal lobe is fringed with a slender, flat, overlapping hairs which are a little longer than the width of the lobe, and stand at an angle of about 60 degrees with the direction of the axis of the appendage. The outer lobe is at an angle with the main one, and has short, very fine setæ oh the margin. One or two specimens show some evidence of a joint between the inner and outer lobes, but in the great majority of cases they seem to be continuous; if originally in two segments, they have become firmly united. The exopodites of the thorax, like those of the cephalon, are directed diagonally forward and outward. (pl. 21, fig. 6; pl. 22.)

Pygidium.

The pygidium of Neolenus serratus is large, and usually shows five rings on the axial lobe and four pairs of ribs on the sides. There are five pairs of biramous appendages belonging to this shield, and behind these a pair of jointed cerci. That the number of abdominal appendages should correspond to the number of divisions of the axial lobe rather than to the number of ribs on the pleural lobes is of interest, and in accord with other trilobites, as first shown by Beecher.

The endopodites of the pygidium have the same form as those of the thorax, are long, and very much less modified than those of any other trilobite whose appendages are known. On some specimens, they extend out far beyond the dorsal test, so that nearly all the segments are visible (pl. 17, fig. 3; pl. 18; pl. 19; pl. 20, fig. 1), but in these cases are probably displaced. The segments are short and wide, the whole endopodite tapering gradually outward. The dactylopodite bears terminal spines, and the individual segments also have outward-directed spines.

The cerci appear to have been long, slender, very spinose organs much like the antennules, but stiff rather than flexible. They are a little longer than the pygidium (pl. 17, figs. 1, 2), and seem to be attached to a plate on the under surface of the posterior end and in front of the very narrow doublure. The precise form of this attachment can not be determined from the published figures. They bear numerous fine spines (pl. 17, fig. 3).

Epipodites and Exiles.

Doctor Walcott has found on several specimens of Neolenus remains of organs which he interprets as epipodites and exites attached to the coxopodites. A study of the specimens has, however, convinced me that both the large and small epipodites are really exopodites, and that the exites are badly preserved and displaced coxopodites. Detailed explanation of this interpretation is given below in the description of the several specimens involved.

Description of Individual Specimens.

Doctor Walcott was kind enough to send me eight of the more important specimens of Neolenus figured by him, and since my interpretation of them does not agree in all respects with his, I have thought it fairer to the reader to present here rather full notes explaining the position I have taken. I understand that since I communicated my interpretation of the epipodites and exites to him, Doctor Walcott has submitted the specimens to several palæontologists, who consider that epipodites are really present. Since I am not able to convince myself that their conclusion is based upon sound evidence, I give here my own interpretation. There is of course, no a priori reason why trilobites should not have had epipodites.

Specimen No. 58589.

Illustrated: Walcott, Smithson. Misc. Coll., vol. 57, 1912, pl. 45, fig. 2;--Zittel-Eastman Text-book of Paleontology, vol. 1, 1913, fig. 1377;--Smithson. Misc. Coll., vol. 67, 1918, pl. 18, fig. 1; pl. 20, fig. 1.

This is one of the most important of the specimens, as it shows the coxopodites of three thoracic limbs and the well preserved endopodites of six thoracic and five pairs of pygidial appendages.

The appendages are all shifted to the left till the articular socket of the coxopodite is about 8 mm. outside of its proper position. The endopodites extend a corresponding amount beyond the edge of the dorsal test and are there so flattened that they are revealed as a mere impression. The coxopodites, which are beneath the test, seem to have been somewhat protected by it, and while hopelessly crushed, are not flattened, but rather conformed to the ridges and grooves of the thorax.

The coxopodite of the appendage of the last thoracic segment is best preserved. It is rectangular, about one third as wide as long, with a slight notch in the posterior margin near the outer end. The inner end is obliquely truncated and shows about ten sharp spines which do not appear to be articulated to the segment, but rather to be direct outgrowths from it. There are similar spines along the posterior margin, but only two or three of what was probably once a continuous series are now preserved. On the opposite margin of the coxopodite from the slight depression mentioned above, there is a slight convexity in the outline, which is better shown and explained by the coxopodite just in front of this. That basal segment has the same form as the one just described, but as its posterior margin is for the greater part of its length pushed under the one behind it, the spines are not shown. On the posterior margin, two-thirds of the length from the proximal end, there is a shallow notch, and corresponding to it, a bulge on the anterior side. From analogy with Ceraurus and Calymene it becomes plain that the notch and bulge represent the position of the socket where the coxopodite articulated with the appendifer. Since these structures have not been shown in previous illustrations, a drawing giving my interpretation of them is here inserted (fig. 2). It is evident from the position of the notch that the row of spines was on the dorsal (inner) side of the coxopodite and that the truncation was obliquely downward and outward.

The endopodite of the last thoracic appendage is well preserved and may be described as typical of such a leg in this part. The basipodite is as wide as the coxopodite, and it and the three succeeding segments, ischiopodite, meropodite, and carpopodite, are all parallel-sided, not expanded at the joints, and decrease regularly in width. The propodite and dactylopodite are also parallel-sided, but more slender than the inner segments, and on the end of the dactylopodite there are four little spines, three of them--one large and two small--articulated at the distal end, and the fourth projecting from the posterior outer angle. Each segment has one or more spines on the outer articular end, and the ischiopodite has several directed obliquely outward on the posterior margin. All of the four proximal segments show a low ridge parallel to and near the anterior margin, and several endopodites of the pygidium have a similar ridge and a row of spines along the posterior margin of some of the segments. These features indicate that the segments in question were not cylindrical in life, but compressed. From the almost universal location of the spines on the posterior side of the limbs as preserved, it seems probable that in the natural position the segments were held in a plane at a high angle with the horizontal, the ridge was dorsal and anterior and the row of spines ventral and posterior. Because the spines on the endobases are dorsal it does not follow that those on the endopodites were, for the position of the coxopodite in a crushed specimen does not indicate the position of the endopodite of even the same appendage.

The endopodites of the pygidium are similar to the one just described, except that some of them have spines on the posterior margin of the segments, and a few on the right side have extremely fine, faintly visible spines on the anterior side. The specimen shows fragments of a few exopodites, but nothing worth describing. In the middle of the right pleural lobe there is a small organ which Walcott has interpreted as a small epipodite. It is oval in form, broken at the end toward the axial lobe, and has exceedingly minute short setæ on the posterior margin. From analogy with other specimens, it appears to me to be the outer end of an exopodite.

Measurements: The entire specimen is about 64 mm. long and 52 mm. wide at the genal angles. The thorax is about 41 mm. wide (disregarding the spines) at the seventh segment, and the axial lobe about 13 mm. wide at the same horizon. The measurements of the individual segments of the seventh left thoracic limb are:

Coxopodite, 9 mm. long, 3 mm. wide, the middle of the notch 8 mm. from the inner end, measured along the bottom, and 6 mm. measured along the top. Basipodite, 5 mm. long, 3 mm. wide Ischiopodite, 4 " " 3 " " Meropodite, 3.5 " " 2.5 " " Carpopodite, 3.5 " " 2 " " Propodite, 3 " " 1.25 " " Dactylopodite, 2 " " 1.25 " "

The five distal segments of the last pygidial endopodite are together 10.5 mm. long. The whole six segments of the endopodite of the third thoracic segments are together 21 mm. long. The distance from the appendifer of the third segment to the outer end of the spine is 17 mm. From the center of the notch in the coxopodite to the outer end is 1.5 mm., which, added to the length of the endopodite, 21 mm., makes a distance of 22.5 mm. from the appendifer to the tip of the dactylopodite, showing that if projected straight outward, the endopodites of the thorax would project 5.5 mm. beyond the test, including spines.

The distance across the axial lobe from appendifer to appendifer on the seventh thoracic segment is 12.5 mm. Measured along the top of the coxopodite, it is 6 mm. from the middle of the notch to the inner end, and measured along the bottom it is 8 mm. From the truncated form of the ends it is evident that the coxopodites extended inward and downward from the appendifers, and with the dimensions given above, the inner toothed ends would practically meet on the median line.

Measurements on the appendages of the pygidia show that on this specimen they extend back about twice as far beyond the edge of the pygidium as they should, all being displaced.

Specimen No. 65514.

Illustrated: Walcott, Smithson. Misc. Coll., vol. 67, 1918, pl. 19, figs. 1-3.

This specimen is so twisted apart that it is not possible to determine to what segments the appendages belong, but it exhibits the best preserved exopodites I have seen. The best one is just in front of the pygidium on the matrix, and shows a form more easily seen than described (our fig. 3). There is a broad, flat, leaf-like shaft, the anterior side of which follows a smooth curve, while in the curve on the posterior side, which is convex backward, there is a re-entrant, setting off a small outer lobe whose length is about one third the length of the whole. This lobe seems to be a continuation of the shaft, and the test of the whole is wrinkled and evidently very thin. The main and distal lobes of the shaft both bear numerous delicate setæ, but those of the outer lobe are much shorter and finer than those on the main portion. The latter are flattened and blade-like.

The anterior edge of the shaft shows a narrow stiffening ridge and the setæ are but little longer than its greatest width. The second segment of the pygidium has another exopodite like this one, but shows faintly the line between the two lobes, as though there were two segments.

This specimen also shows some very well preserved endopodites, but they differ in no way from those described from specimen No. 58589. Walcott mentions two large epipodites projecting from beneath the exopodites. I judge that he has reference to the distal lobes of the exopodites, but as these are continuous with the main shaft, there can be no other interpretation of them than that which I have given above.

Measurements: The pygidium is 19 mm. long (without the spines) and about 34 mm. wide at the front. The exopodites show faintly beneath the pygidial shield, but their proximal ends are too indistinct to allow accurate measurement. Apparently they were just about long enough to reach to the margin of the shield. The best preserved one, that of the second segment in the pygidium, is about 11 mm. long, 2.5 mm. wide at the widest; the distal lobe is 2.5 mm. long, and the longest setæ of the main lobe 3.5 mm. long. The pleural lobe of the pygidium is just 11 mm. wide at this point.

The endopodites project from 8 to 12 mm. beyond the pygidium, showing about four segments.

The thoracic exopodite described above is 11 mm. long and 2.75 mm. wide at the widest part. The distal lobe is 3.5 mm. long and 2.25 mm. wide, and the longest setæ on the main lobe 3 mm. long.

Specimen No. 65519.

Illustrated: Walcott, Zittel-Eastman Text-book of Paleontology, vol. 1, 1913, fig. 1343;--Smithson. Misc. Coll., vol. 67, 1918, pl. 21, fig. 6.

This specimen is somewhat difficult to study but is very valuable as showing the natural position of the exopodites of the anterior part of the thorax. Walcott's figures are excellent and show the broad leaf-like shafts, the distal lobes with the re-entrant angles in the posterior margin, and the long fine setæ of the main lobes. None of the distal lobes retains its setæ. All extend back to the dorsal furrows, but the proximal ends are not actually shown.

The specimen is especially important because it shows the same distal lobes as specimen No. 65514, and demonstrates that they are a part of the exopodite and not of any other structure.

Measurements: The exopodite belonging to the fourth thoracic segment is 23 mm. long and 4 mm. wide at the widest part. The longest setæ are 7 mm. in length.

Specimen No. 65520.

Illustrated: Walcott, Smithson. Misc. Coll., vol. 67, 1918, pl. 20, fig. 2; pl. 22, fig. 1.

This is a practically entire specimen, on two blocks, one showing the interior of the shell, and the other the one figured by Walcott, a cast of the interior. The first shows the low rounded appendifers at the anterior angle of each axial tergite. They are almost entirely beneath the dorsal furrows and do not project so far into the axial lobe as those of Ceraurus and Calymene. In fact, only those at the anterior end of the thorax project inward at all. As expected, there are five pairs on the pygidium. The cephalon is unfortunately so exfoliated that the appendifers there are not preserved. The doublure of the pygidium is extremely narrow.

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 setæ. 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 setæ 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.

Illustrated: 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 setæ or the bases of them, and the distal lobe faint traces of much finer ones. The form, and the setæ 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.

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 setæ. The same stout spines that negate the above possible explanation at once suggest that they are coxopodites (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 coxopodite 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 coxopodite, 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 coxopodite (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.

Illustrated: 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 coxopodites. 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 coxopodites 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 coxopodite. 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.

Measurements: The cephalon is 24 mm. long and about 60 mm. wide. The coxopodite 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.

=Nathorstia transitans= Walcott.

Illustrated: 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 associated 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-shelled" condition. Even if not a Neolenus, it is probable, from the state of preservation, that it is an animal which had recently cast its shell.

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 assume 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 setæ. 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 maxillæ." 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 illustrated 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, 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.

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:

The club-shaped bodies lying within the axis are the gnathobases attached at the sides of the axis; the curved members extending outward from the gnathobases are the endopodites; the longitudinal ridges in the ventral membrane between the inner ends of the gnathobases are the buttresses and apodemes of the mesosternites; the slender oblique rod-like bodies shown in the right pleural region in Walcott's figure are portions of the fringes of the exopodites.

In 1910, Mr. W. C. King of Ottawa, Ontario, found at Britannia, a few miles west of Ottawa, the impression in sandstone of the under surface of a large specimen of Isotelus arenicola, described on a later page (p. 39).

Finally (1918, p. 133, pl. 24, figs. 3, 3a; pl. 25), Walcott has redescribed the specimen from Ohio, presenting a new and partially restored figure. He refers also to the specimen from Ottawa under the name Isotelus covingtonensis? Foerste (not Ulrich). He advances the view, which I am unable to share, that the cylindrical appearance of the segments of the appendages of Isotelus is due to post-mortem changes.

=Isotelus latus= Raymond.

(pl. 10, fig. 1.)

Illustrated: Asaphus platycephalus Billings, Quart. Jour. Geol. Soc., London, vol. 26, 1870, pl. 31, figs. 1-3; pl. 32, figs. 1, 2.--Woodward, Geol. Mag., vol. 8, 1871, pl. 8, figs. 1, 1a.--Gerstäcker, in Bronn's "Klassen u. Ordnungen d. Thier-Reichs," 1879, pl. 49, fig. 1.--von Koenen, N. Jahrb. f. Min., etc., vol. 1, 1880, pl. 8, fig. 8.--Milne-Edwards, Ann. Sci. Nat., Zoologie, ser. 6, vol. 12, 1881, pl. 12, fig. 45.

Isotelus latus Raymond, Bull. Victoria Mem. Mus., Geol. Survey Canada, No. 1, 1913, p. 45 (species named).

Isotelus covingtonensis? Walcott (not Foerste), Smithson. Misc. Coll., vol. 67, 1918, p. 134.

Knowledge of the appendages of this species is derived from the specimen which Billings described in 1870. It was found in the Trenton, probably the Middle Trenton, near Ottawa, Ontario, and is preserved in the Victoria Memorial Museum at Ottawa.

Viewed from the upper surface, it shows a large part of the test, but is broken along the sides, so that parts of the free cheeks, considerable of the pleural lobes of the thorax, and one side of the pygidium are missing. Viewed from the lower surface, the appendages are practically confined to the cephalon and thorax.

A short time before his death, Professor Beecher had this specimen and succeeded in cleaning away a part of the matrix so that the appendages show somewhat more clearly than in Billings' time, but they are not so well preserved as on the Mickleborough specimen, found in Ohio somewhat later.

The hypostoma is in place and well preserved; the posterior points are but 3 mm. in advance of the posterior margin of the cephalon. Behind the hypostoma there are only two pairs of cephalic appendages, the first of which is represented by the coxopodite and a trace of the endopodite. The outer end of the coxopodite is close to the outer margin of one of the prongs of the hypostoma and about 3 mm. in front of its posterior end. The gnathobase curves backward and inward, and appears to pass under the tip of the hypostoma. There were probably two appendages in front of this, whose gnathobases projected under the hypostoma, but the specimen shows nothing of them unless it be that one small fragment about 2 mm. back of the center is really a part of a gnathobase.

The specimen retains only the coxopodite and basipodite of the posterior cephalic appendage on the left side. The coxopodite is long and apparently cylindrical, the cross-section being of uniform diameter throughout the length. The inner portion is nearly straight, while the outer part is curved gently forward.

It is possible to make out remains of eight pairs of appendages on the thorax, some of them represented by coxopodites only, but most with more or less poorly preserved endopodites as well. No exopodites are visible. The coxopodites of the thorax seem to be of the same form as the last one on the cephalon, but slightly less curved. All are long and heavy, and there seems to be no decrease in size toward the pygidium. The endopodites are very imperfectly shown. They seem to be longer than those of Isotelus maximus, and the segments, while of less diameter than the coxopodites, do not show so great a contrast to them as do those of that species. The direction of the endopodites is diagonally forward, and the outer portions do not appear to be curved backward as in Isotelus maximus. It would appear also that the endopodites were nearly or quite long enough to reach the outer margin of the dorsal test. On no endopodite can more than three segments be definitely distinguished, but the longest ones are the most obscurely segmented.

No appendages are preserved on the pygidium, but at one side of the median groove there are two projections which may be processes to which the appendages were attached.

Measurements: Total length of specimen, 109 mm. Probable length when complete, 116 mm. Length of cephalon, 40 mm.; width at genal angles, restored, about 62 mm. (Billings' restoration). Width of doublure of front of cephalon on median line, 17 mm.; length of hypostoma, 20 mm. Length of coxopodite of last appendage on left side of cephalon, 10.5 mm.; length of basipodite of the same appendage, 5 mm. Diameter of coxopodite, 2 mm.; diameter of basipodite, 1.5 mm. Length of coxopodite on left side of the second segment of the thorax, 11 mm.; diameter, about 2.5 mm. Length of basipodite of the same, 5 mm.; diameter, about 1.5 mm. Length of ischiopodite, 3.5 mm.; diameter, about 1.5 mm. Length of meropodite, 2.5 mm. (this may be less than the total length as the segment is not completely exposed.) Distance between proximal ends of gnathobases of the fifth thoracic segment, about 7 mm. Distance between outer ends of the coxopodites of the first thoracic segment (estimated from measurements on the left side), 27 mm Distance apart of the dorsal furrows at the first thoracic segment, 27 mm. Length of the longest exopodite which can be traced, about 20 mm.

=Isotelus maximus= Locke.

(pl. 10, fig. 2.)

Illustrated: Mickleborough, Jour. Cincinnati Soc. Nat. Hist., vol. 6, 1883, p. 200, figs. 1-3 (endopodites and coxopodites). Walcott, Science, vol. 3, 1884, p. 279, fig. 1 (endopodites, coxopodites, and traces of exopodites). Woodward, Geol. Mag., dec. 3, vol. 1, 1884, p. 162, figs. 1-3 (copies of Mickleborough's figures). Bernard, The Apodidæ, 1892, text fig. 49. Beecher, Amer. Jour. Sci., vol. 13, 1902, p. 169, pl. 5. figs. 5, 6 (outline from one of Mickleborough's figures and an original figure). Walcott, Smithson. Misc. Coll., vol. 67, 1918, p. 133, pl. 24, figs. 3, 3a; pl. 25, fig. 1.

This specimen, which conies from the Richmond strata 2 miles north of Oxford, Ohio, is the best preserved of the specimens of Isotelus with appendages which has so far been found. The individual consists of two parts, the actual specimen, and the impression of the ventral side.

To describe it I am using very skillfully made plaster reproductions of both parts, presented to the Museum of Comparative Zoology by Doctor Charles D. Walcott, and presumably made after he cleaned the specimen as described in Science (1884). I have also an enlarged photograph (pl. 10, fig. 2) which seems to have been made after some later period of cleaning, probably by Professor Beecher, and I have examined the original specimens in Washington.

Viewed from the dorsal side, it is seen that the individual is very imperfect, the greater part of the cephalon being removed by a diagonal break which cuts off the anterior third of the left eye and extends to the front of the second thoracic segment on the right side. The ends of the pleura of both sides of the thorax are broken away, as are also the greater parts of the pleural lobes and the posterior end of the pygidium. On the ventral side, merely the posterior tips of the hypostoma remain, but the distal ends of the appendages were so far within the outer margin that the appendagiferous area is quite fully retained.

The most conspicuous feature of this specimen is the presence of nine pairs of large coxopodites behind the hypostoma, and of the remains of ten pairs of endopodites, making in all ten pairs of appendages which are easily seen. The apportionment of these segments to cephalon, thorax, and pygidium is not agreed upon by the people who have examined the specimens, but if one remembers that it is the outer and not the inner end of the coxopodite which articulates with the appendifer, it at once becomes evident that the first two pairs of appendages on the specimen are the last two pairs belonging to the cephalon, and that the next eight pairs are those of the thorax.

The impressions of fourteen pairs of coxopodites are readily counted on the pygidium, and as Doctor Walcott noted sixteen pairs on the actual specimens, his number was probably correct.

Cephalon.

Projecting the line of the back of the cephalon through from the dorsal side, it is found that the posterior tips of the hypostoma are 7 mm. in front of the posterior margin of the cephalon, and that the points of attachment of the posterior pair of cephalic appendages (the second pair shown on the specimen) are just within the posterior margin. The gnathobases of this pair of appendages extend back some distance beneath the thorax, and so give the impression that they belong to that part of the body. So far as can be determined, the cephalic appendages do not differ in any way from those of the thorax. On the mould of the ventral surface, just outside of the lateral edge of the right lobe of the hypostoma, is the somewhat imperfectly shown impression of the endopodite of the third cephalic appendage. The point of junction of the endopodite and coxopodite is about 2 mm. in front of the tip of the adjacent branch of the hypostoma, and the gnathobase is curved around just behind it. This accounts for three of the pairs of cephalic appendages. The second cephalic appendages must have thrust their gnathobases under the prongs of the hypostoma, and the endopodites were probably close to its edge. No trace of this pair appears on the specimen.

Thorax.

The thoracic appendages are the best preserved of any, and show the large coxopodites and the more slender endopodites which do not extend to the outer margin of the test. The latter extend forward and outward for about one half their length, then turn backward in a graceful curve.

Walcott's figure in Science shows hair-like markings on the under side of the right half of the thorax. These were interpreted by both Walcott and Beecher as fringes of the exopodites, but since the setæ of those organs on all other trilobites are always above the endopodites, while these are represented as below them, it would seem doubtful if this interpretation can be sustained. Furthermore, I find no trace of them on either cast or mould, and the actual specimen does not now show them.

Pygidium.

The coxopodites and endopodites of the pygidium seem to be similar to those on the thorax, but both are shorter and more slender, and the former decrease in length rapidly toward the posterior end. As mentioned above, it is not perfectly plain how many appendages are present, but I have accepted Doctor Walcott's count of sixteen pairs. Of the endopodites only the barest traces are seen, and of exopodites nothing.

One point of considerable interest in this specimen is the thickness, as it probably gives some measure of the space occupied by the animal. In Triarthrus and other trilobites from Rome, New York, the appendages are pressed directly against the dorsal test, but in this specimen a considerable space intervenes between the plane of the appendages and the shell. Between the central furrow and the inner surface of the dorsal test at the anterior end of the thorax is a distance of 13 mm. and under the dorsal furrows the thickness is about 7 or 8 mm., no accurate measurement being possible in the present state of the specimen.

Measurements: Length of specimen on median line, 121 mm.; probable original length, about 195 mm. (Walcott's restoration). Length of thorax, 58 mm. Width of axial lobe at the first thoracic segment, 45 mm.; total width as preserved, 92 mm.; width as estimated from the mould of the ventral surface, no mm.; Walcott's restoration, 105 mm.

Length of coxopodite of fourth left cephalic appendage, about 18 mm.; diameter, about 2.5 mm. Length of coxopodite of last left cephalic appendage, about 18.5 mm. Distance apart of inner ends of gnathobases of fourth cephalic appendages, about 4 mm. Distance apart of inner ends of endobases of first thoracic segment, about 6 mm. Distance apart of outer ends of coxopodites of first thoracic segment, about 43 mm.

Length of coxopodite of seventh left thoracic appendage 16 mm., diameter about 3.5 mm.; length of basipodite of the endopodite of the same appendage 6 mm.; diameter about 2 mm.; length of ischiopodite 5 mm.; length of meropodite 4.5 mm.; length of carpopodite 4.5 mm.; length of propodite 3 mm.; length of dactylopodite 2.75 mm.; total length of endopodite 25.75 mm.

Length of coxopodite of fourth left thoracic appendage 20 mm., diameter 4 mm.; length of five proximal joints of the endopodite 25 mm.; diameter of basipodite about 2 mm.

RESTORATION OF ISOTELUS.

(Text fig. 9.)

The exopodites have been omitted from this restoration since nothing is known of their actual form. The chief reason for the figure is to contrast the greatly developed coxopodites of the posterior part of the cephalon and thorax with those of other trilobites. The antennules and first two pairs of biramous appendages of the cephalon are more or less hypothetical, and less is known of the appendages of the pygidium than is shown here. The restoration is based somewhat upon Walcott's figure in Science. The outline is that of a specimen of Isotelus maximus from Toronto, Ontario.

=Isotelus gigas= Dekay.

Illustrated: Woodward, Quart. Jour. Geol. Soc., London, vol. 26, 1870, text fig. 1; Geol. Mag., dec. 3, vol. 1. 1884, p. 78, text fig. Milne-Edwards, Ann. Sci. Nat, Zoologie, ser. 6, vol. 12, 1881, pl. 12, fig. 46. Walcott, Bull. Mus. Comp. Zool., Harvard Coll., vol. 8, 1881, pl. 2, fig. 9; Geol. Mag., dec. 4, vol. 1, 1894, pl. 8, fig. 9; Proc. Biol. Soc. Washington, vol. 9, 1894, pl. 1, fig. 9.

The specimen in the British Museum which Woodward called Asaphus platycephalus, is, in all probability, an Isotelus gigas. Woodward says of it:

I was at once attracted by a specimen of Asaphus, from the Black Trenton Limestone (Lower Silurian), which has been much eroded on its upper surface, leaving the hypostoma and what appear to be the appendages belonging to the first, second, and third somites, exposed to view, united along the median line by a longitudinal ridge. The pseudo-appendages, however, have no evidence of any articulations. But what appears to me to be of the highest importance, as a piece of additional information afforded by the Museum specimen, is the discovery of what I believe to be the jointed palpus of one of the maxillæ, which has left its impression upon the side of the hypostoma--just, in fact, in that position which it must have occupied in life, judging by other Crustaceans which are furnished with an hypostoma, as Apus, Serolis, etc.

The palpus is 9 lines in length, the basal joint measures 3 lines, and is 2 lines broad, and somewhat triangular in form.

There appear to be about 7 articulations in the palpus itself, above the basal joint, marked by swellings upon its tubular stem, which is 1 line in diameter.

Desiring to know more of this individual, I wrote to Doctor Bather and was surprised to learn that the specimen which was the basis of Woodward's observations is so badly preserved as to be of no real value. With his permission, I append a note made by Doctor Bather some years ago when selecting fossils to be placed on exhibition:

Asaphus gigas Dekay. Ordovician, Trenton Limestone. N. America, Canada. Descr. H. Woodward, 1870, Q. J. G. S., XXVI, pp. 486-488, text fig. 1, as Asaphus platycephalus. Coll. and presd. J. J. Bigsby, 1851. Regd. I 14431.

This specimen is in the Brit. Mus. Geol. Dept. I 14431. The supposed hypostome is exceedingly doubtful; it lies dorsad of the crushed glabellar skeleton. The "appendage" is merely the edge of a part in the head-shield; the maxilla is some calcite filling, between two such laminæ.

13 Sept. 1911. (Signed) F. A. BATHER.

Walcott figured a slice of Isotelus gigas from Trenton Falls, New York, which shows a few fragments of appendages, but is of particular importance because it shows the presence of well developed appendifers beneath the axial lobe.

=Isotelus arenicola= Raymond.

Illustrated: Ottawa Nat, vol. 24, 1910, p. 129, pl. 2, fig. 5.

The following quotations from my paper are inserted here to complete the record of appendage-bearing specimens:

A rather remarkable specimen of this species was found by W. C. King, Esq., on the shore of Lake Deschenes at Britannia [near Ottawa, Ontario]. This specimen is an impression of the lower surface of the trilobite, and shows a longitudinal ridge corresponding to the central furrow along the axis of the ventral side of the animal, ten pairs of transverse furrows, and the impression of the hypostoma. The doublure of the pygidium has also left a wide smooth impression, but in the cephalic region the hypostoma is the only portion of which there are any traces remaining. The specimen was found on a waterworn surface of the beach, partially covered by shingle....

The transverse furrows are the impressions left by the gnathobases of the basal joints of the legs. They were evidently long and very heavy, but the specimen has been so abraded that all details are obscured. The first six pairs of impressions are longer and deeper than the four behind. The first eight pairs seem to pertain to the thoracic appendages, while the last two belong to the pygidium. From the posterior tips of the hypostoma to the first gnathobases of which traces are present there is a distance of about 22 mm. without impressions. In Isotelus gigas the hypostoma normally extends back to the posterior margin of the cephalon, so that it seems that in this specimen the impressions of the first two pairs of gnathobases under the thorax may not have been preserved. In that case, the six pairs of strong impressions may represent the last six pairs of thoracic segments, and the pygidium might begin with the first of the fainter ones.

Horizon and locality: From the sandstone near the base of the Aylmer (Upper Chazy) formation at Britannia, west of Ottawa, Ontario. Specimen in the Victoria Memorial Museum, Geological Survey of Canada, Ottawa.

The Appendages of Triarthrus.

=Triarthrus becki= Green.

(Pls. 1-5; pl. 6, figs. 1-3; text figs. 1, 10, 11, 33, 42.)

(Also see Part IV.)

Illustrated: Matthew, Amer. Jour. Sci., vol. 46, 1893, pl. 1, figs. 1-7;--Trans. N. Y. Acad. Sci., vol. 12, pl. 8, figs. 1-7.--Beecher, Amer. Jour. Sci., vol. 46, 1893, text figs. 1-3;--Amer. Geol., vol. 13, 1894, pl. 3;--Amer. Jour. Sci., vol. 47, pl. 7, text fig. 1;--Amer. Geol., vol. 15, 1895, pls. 4, 5;--Ibid., vol. 16, 1895, pl. 8, figs. 12-14; pl. 10. fig. 1;--Amer. Jour. Sci., vol. 1, 1896, pl. 8; Geol. Mag., dec. 4, vol. 3, 1896, pl. 9;--Eastman-Zittel Text-book of Paleontology, vol. 1, 1900, text figs. 1267-1269;--2d ed., 1913, fig. 1375; Studies in Evolution, 1901, reprint of all previous figs.;--Amer. Jour. Sci., vol. 13, 1902, pl. 2, figs. 1-5; pl. 3, fig. 1; pl. 4, fig. 1; pl. 5, figs. 2-4;--Geol. Mag., dec. 10, vol. 9, 1902, pls. 9-11, text figs. 1-3.--Walcott, Proc. Biol. Soc. Washington, vol. 9, 1894, pl. 1 figs. 1-6;--Geol. Mag., dec. 4, vol. 1, 1894, pl. 8;--Smithson. Misc. Coll., vol. 67, 1918, pl. 29, figs. 1-11; pl. 30, figs. 17-20; pl. 32; pl. 34, figs. 4-7; pl. 35, fig. 5.--Bernard, Quart. Jour. Geol. Soc., London, vol. 50, 1894, text figs. 11, 12.--Oehlert, Bull. Soc. Géol. France, ser. 3, vol. 24, 1896, text figs. 1-17, 34.--Jaekel, Zeits. d. d. geol. Gesell., vol. 53, 1901, text fig. 24. Moberg, Geol. Fören. Förhandl., vol. 29, pl. 5, 1907, pl. 4, fig. 2; pl. 5, fig. 1.--Handlirsch, Foss. Insekten, 1908, text fig. 6.--Tothill, Amer. Jour. Sci., vol. 42, 1916, p. 380, text fig. 5.--Crampton, Jour. N. Y. Entomol. Soc., vol. 24, 1917, pl. 2, fig. 20.

Historical.

Specimens of Triarthrus retaining appendages were first obtained by Mr. W. S. Valiant from the dark carbonaceous Utica shale near Rome, New York, in 1884, but no considerable amount of material was found until 1892. The first specimens were sent to Columbia University, and were described by Doctor W. D. Matthew (1893). This article was accompanied by a plate of sketches, showing for the first time the presence of antennules in trilobites and indicating something of the endopodites and exopodites of the appendages of the cephalon, thorax, and pygidium. Specimens had not yet been cleaned from the lower side, so that no great amount could then be learned of the detailed structure. Matthew concluded that "The homology with Limulus seems not to be as close in Triarthrus as in the forms studied by Mr. Walcott; but the characters seem to be of a more comprehensive type, approaching the general structure of the other Crustacea rather than any special form."

Professor Beecher's first paper, dated October 9, 1893, merely mentioned the fact that the Yale University Museum had obtained material from Valiant's locality, but was quickly followed by a paper read before the National Academy of Sciences on November 8, and published in December, 1893. This paper described particularly the thoracic appendages.

This was followed in January (1894 A) by an article in which some information about the mode of occurrence of the specimens was added, and in April (1894 B), the limbs of the pygidium were described and figured. The determination of the structure of the appendages of the head evidently presented some difficulty, for the article describing this portion of the animal did not appear until the next February (1895 A). This cleared up the ventral anatomy of Triarthrus, and was followed by a short article (1896 A) accompanied by a restoration of the trilobite showing all the appendages.

This ended Professor Beecher's publications on Triarthrus until his final paper in 1902, although he contributed some of his results and figures to his chapter on the trilobites in the Eastman-Zittel Text-book of Paleontology in 1900.

The discovery of these excellent specimens had of course excited very great interest. Doctor Walcott also studied a number of specimens from Valiant's locality, and published in 1894, with some original figures, the results of his comparison of the appendages of Triarthrus with those of Calymene and Ceraurus.

In his article on the "Systematic Position of the Trilobites," Bernard (1894) used the results of Professor Beecher's studies of 1893, and also quoted the papers by Matthew (1893) and Walcott (1894), though the article by the latter appeared too late to be used except for a note added while Bernard's paper was in press. A final footnote quoted from Professor Beecher's paper of April, 1894 (1894 B).

Oehlert (1896) gave an excellent summary in French of the work of Beecher and Walcott on Triarthrus, with reproductions of many of their figures.

Valiant (1901) in a non-technical article described his long search for trilobites with antennas. The discovery of the wonderful pyritized trilobites at Cleveland's Glen near Rome was not the result of a lucky accident, but the culmination of eight years of labor in a locality especially selected on account of the fineness of grain of the shale.

After 1896, Professor Beecher turned his attention largely to the problem of the classification of trilobites, and while he continued the arduous task of cleaning the matrix from specimens of Triarthrus and Cryptolithus he did not again publish upon the subject of appendages until forced to do so by the doubts cast by Jaekel (1901) upon the validity of his earlier conclusions. Because of certain structures which he thought he had interpreted correctly from a poorly preserved specimen of Ptychoparia, Jaekel came to the conclusion that Beecher's material was not well preserved. Professor Beecher would have taken much more kindly to aspersions upon his opinions than to any slight upon his beloved trilobites, and his article on the "Ventral Integument of Trilobites" of 1902 was designed not only as an answer to Jaekel, but also to show by means of photographs the unusually perfect state of preservation of the specimens of Triarthrus. This article, like so many describing the appendages of trilobites, beginning with Matthew's, was published in two places (Beecher 1902).

Most of Beecher's papers, except the last one, were reprinted in the volume entitled "Studies in Evolution," published by Charles Scribner's Sons at the time of the Yale Bicentennial in 1901. The part pertaining particularly to Triarthrus is on pages 197 to 219.

Moberg (1907), in connection with a specimen of Eurycare angustatum which he thought preserved some appendages, described and illustrated some of the appendages of Triarthrus.

The most recent discussion of Triarthrus, with some new figures, is by Walcott (1918, p. 135, pls. 29, 30). He gives a summary of Beecher's work with numerous quotations. The principal original contribution is a discussion of the form and shape of the appendages before they were flattened out in the shale. He found also what he thought might possibly be the remains of epipodites on three specimens, one of which he illustrated with a photograph. I have seen nothing which could be interpreted as such an organ in the many specimens I have studied.

A point in which Walcott differs from Beecher in the interpretation of specimens is in regard to the development of the endopodites of small pygidia. Beecher (1894 B, pl. 7, fig. 3) illustrated a series of endopodites which he likened to the endites of a thoracic limb of Apus. Doctor Walcott finds that specimens in the United States National Museum show slender endopodites all the way to the back of the pygidium, and thinks that Beecher mistook a mass of terminal segments of exopodites for a series of endopodites. On careful examination, however, the specimen shows, as Beecher indicated, a series of endopodites in undisturbed condition (No. 222, our pl. 4, fig. 5).

Restoration of Triarthrus.

One of the more important points noted in the later studies of Triarthrus is that the gnathites of the cephalic appendages are much less like the endobases under the thorax than Beecher earlier thought, and showed in his restored figures and in his model. The four gnathites of each side are curved, flattened, not club-shaped, and so wide and so close together that they overlap one another. The metastoma is somewhat larger and more nearly circular than Beecher's earlier preparations led him to suppose.

The restoration here presented is modified only slightly from the one designed by Professor Beecher, and the modifications are taken principally from figures published by him. The gnathites are drawn in form more like that shown by the specimens and his figures in the American Geologist (1895 A), and the metastoma is taken from one of the specimens. On the thorax the chief modification is in the addition of a considerable number of spines to the endopodites. In spite of the trivial character of most of these changes, they emphasize one of the important characteristics of Triarthrus the regional differentiation of the appendages.

It should be pointed out that although Triarthrus is usually considered to be a very primitive trilobite, its appendages are more specialized than those of any of the others known. This is shown in their great length, the double curvature of the antennules, the differentiation of four pairs of endobases on the cephalon as gnathites, and the flattening of the segments of the posterior endopodites. These departures from the uniformity existing among the appendages of the other genera lead one to question whether the genus is really so primitive as has been supposed.

Relation of the Cephalic Appendages to the Markings on the Dorsal Surface of the Glabella.

Triarthrus becki is usually represented as having four pairs of glabellar furrows, but the two pairs at the front are exceedingly faint and the first of them is hardly ever visible, though that it does exist is proved by a number of authentic specimens. The neck furrow is narrow and sharply impressed, continuing across the glabella with a slightly backward curvature. In front of it are two pairs of linear, deeply impressed furrows which in their inward and backward sweep are bowed slightly forward, the ends of the corresponding furrows on opposite sides nearly meeting along the crest of the glabella. In front of these, near the median line, is a pair of slight indentations, having the appearance and position of the inner ends of a pair of furrows similar to those situated just behind them.

In front of and just outside this pair are the exceedingly faint impressions of the anterior pair of furrows, these, as said above, being but seldom seen. They are short, slightly indented linear furrows which have their axes perpendicular to the axis of the cephalon, and do not connect with each other or with the dorsal furrows. The latter are narrow, sharply impressed, and merge into a circumglabellar furrow at the front. In front of the circumglabellar furrow is a very narrow rounded ridge, but the anterior end of the glabella is very close to the margin of the cephalon.

Specimen No. 214, which was cleaned from the dorsal side, shows the posterior tip of the hypostoma, apparently in its natural position, 3.5 mm. back from the anterior margin. The entire length of the cephalon is 6 mm., so that the hypostoma reaches back slightly over one half the length (0.583). The greater part of it has been cleaned off, and one sees the proximal portions of the antennules, which are apparently attached just at the sides of the hypostoma, 2.5 mm. apart and 2.25 mm. back from the anterior edge of the cephalon. This position is distinctly within the outline of the glabella and corresponds approximately to the location of the second pair of glabellar furrows. Specimens 214, 215, 216, 217, and 219 all seem to show the same location for the bases of the antennules. Specimen 220 is the one in which the basal shafts are best preserved and the points of attachment seem to be further apart in it than in any of the others. This specimen is 38 mm. long, and the bases of the antennules are 5.5 mm. apart and 4 mm. behind the anterior margin. As the specimen is cleaned from the ventral side, the dorsal furrows do not show distinctly, but another specimen of about the same size (No. 228, 38.5 mm. long) has the dorsal furrows 8 mm. apart 4 mm. back of the anterior margin.

On the same slab with specimens 209 and 210 there is an individual which, although retaining the test, has had the proximal ends of the antennules so pressed against it that the course of the one on the left side is readily visible. It originates in a small oval mound whose posterior margin impinges upon the third glabellar furrow near the middle of its course, and just outside the outer end of the second glabellar furrow. The cephalon of this specimen is 5 mm. long, and the point of origin of the left antennule is 2.75 mm. in front of the posterior margin and 0.75 mm. from the dorsal furrow.

It is therefore evident that the antennules in this species are not attached beneath the dorsal furrows, but within them and opposite the second pair of glabellar furrows.

All cephalic appendages behind the antennules are attached somewhat within the dorsal furrows, the first pair as far forward as the antennules and the last pair apparently under the anterior edge of the neck ring. They do not appear to correspond in position to the posterior glabellar furrows and neck ring, being more crowded. The last pair is attached to appendifers beneath the nuchal segment, and the first pair beneath the third glabellar furrows. There are no depressions on the dorsal surface corresponding to the points of attachment of the mandibles.

Anal Plate.

Professor Beecher, during his first studies of Triarthrus, found no appendages pertaining to the anal segment, but later evidently came upon a spinose anal plate which he caused to be figured. The specimen (No. 201) on which this appendage is preserved is cleaned from the dorsal side, and the anal plate is a small, bilaterally symmetrical, nearly semicircular structure margined with small spines. Specimen 202 also shows the same plate (pl. 5, fig. 6), but it is imperfectly preserved. It has a large perforation in the anterior half. Both of these specimens are in the Yale University Museum.

The anal plate is especially well shown by specimen 65525 in the United States National Museum (fig. 11). This specimen is from Rome, New York, and two photographs of it have been published by Walcott (1918, pl. 29, fig. 6; pl. 30, fig. 19). It is developed from the dorsal side, and the anal plate is displaced, so that it projects behind the end of the pygidium. It is semicircular in shape, with a hemispheric mound at the middle of the anterior half. Two furrows starting from the anterior edge on either side of the mound border its sides, and, uniting back of it, continue as an axial furrow to the posterior margin. The mound is perforated for the opening of the posterior end of the alimentary canal. The lateral borders of the plate bear five pairs of short, symmetrically placed spines. The plate is 1 mm. wide and 0.5 mm. long, and the entire trilobite is 11.5 mm. long.

THE APPENDAGES OF PTYCHOPARIA.

=Ptychoparia striata= (Emmrich).

Illustrated: Jaekel, Zeits. d. d. geol. Gesell., 1901, vol. 53,