Chapter X: Part I: The Appendages of Trilobites (3)

The exopodites of course furnish the chief difficulty in interpretation. Doctor Walcott finds two sets of structures attached to the coxopodite, a long, slender, spiral exopodite, and a short, broad epipodite with a fringe of long setæ. Since he has given the same interpretation for _Calymene_, _Ceraurus_, and _Acidaspis_, I have considered the question of all three together on a preceding page (p. 48), and given my reasons for regarding both structures as due to sections in different directions across setiferous exopodites.

Sections like those shown in figures 11, 13, and 14 of plate 27 (1918) happen to be cut in or near the plane of the setæ of an exopodite, and so show hairs of considerable length. Such sections are, as would be expected, very few in number, while sections like those shown on figures 4, 5, 7, and 9 of plate 27, which cut the setæ more nearly at right angles, are very common. Slices which give any definite idea of the form of the shaft of the exopodite are exceedingly rare. Perhaps the most satisfactory one is No. 23 (pl. 3, fig. 3, 1881), which shows the proximal part of a long, slender, unsegmented shaft, with the bases of a number of slender setæ. The organ is not complete, as would be inferred from the published figure, but the section cuts diagonally across it, and the total length is unknown. It is directed forward, like the exopodites of Neolenus, but whether or not this is a natural position is yet to be learned.

The proximal, non-setiferous portion of the exopodite is evidently at an angle with the setiferous part. Another similar exopodite is apparently shown by specimen 29 (pl. 3, fig. 9, 1881), which has a similar angulated shaft and just a trace of the bases of the setæ.

_Pygidial Appendages._

That appendages were present under the pygidium is shown by longitudinal sections, but nothing is known of the detail of structure.

_Relation of Hypostoma to Cephalon in Calymene._

In _Calymene_ the shape of the hypostoma bears little relation to the shape of the glabella, and it is relatively smaller, both shorter and narrower, than in Ceraurus. In shape, neglecting the side lappets at the front, it is somewhat rectangular, but rounded at the back, where it is bifurcated by a shallow notch. The anterior edge has a narrow flange all across, which is turned at almost right angles to the plane of the appendage, and which fits against the doublure of the free cheeks at the sides and against the epistoma in the middle. The side lappets show on their inner (upper) surface shallow pits, one on each lappet, which fit over projections that on the dorsal surface show as deep pits in the bottom of the dorsal furrows in front of the anterior glabellar furrows. The appendifers on the head in _Calymene_ take the form of curving projections of shell underneath the glabellar and neck furrows, and owing to the narrowness of the hypostoma, all these are visible from the ventral side, even with it in position. This shield extends back about 0.6 of the length of the cephalon, and to a point a little behind the second glabellar furrow from the back of the head.

In Doctor Walcott's restoration of _Calymene_ he has represented all four pairs of biramous appendages as articulating back of the posterior end of the hypostoma. I think his sections indicate that the gnathobases of two pairs of these appendages rested alongside or beneath it, and in particular, the longitudinal sections (1881, pl. 5) would appear to show that the mouth was some distance in advance of its posterior end.

_Restoration of Calymene._

(Text fig. 16.)

From what has been said above, it is evident that for a restoration of the appendages of _Calymene_ considerable dependence must be placed upon analogy with other trilobites. Nothing is positively known of the antennules, the exopodites of the cephalon, or any appendages, other than coxopodites, of the pygidium, but all were probably present. It is inferred from the slices that the first two pairs of cephalic appendages were poorly developed, the endopodites short and very slender, the coxopodites lying parallel to the sides of the hypostoma and nearly or quite functionless. The gnathites of the last two pairs of cephalic appendages are large, closely approximated at their inner ends, and bear small tooth-like spines. The endopodites are probably somewhat better developed than the anterior ones and more like those on the thorax.

The coxopodites of the thorax appear to have had nearly cylindrical endobases which tapered inward. The endopodites were slender, tapering gradually outward, and probably did not extend beyond the dorsal test. Small spines were present on the distal end of each segment. Each exopodite had a long, slender, unsegmented shaft, to which were attached numerous long, overlapping, flattened setæ. The shaft may have been angulated near the proximal end, and may have been directed somewhat forward and outward as in Neolenus, but the evidence on this point is unsatisfactory. The number of pairs of appendages is that determined by Walcott from longitudinal sections, namely, four pairs on the cephalon beside the antennules, thirteen pairs in the thorax, and nine pairs on the pygidium.

=Calymene= sp. ind.

(pl. 6, figs. 4, 5.)

Illustrated: Walcott, Bull. Mus. Comp. Zool., Harvard Coll., vol.
8, 1881, pl. 6, figs. 5a, b;--Proc. Biol. Soc. Washington, vol. 9,
1894, pl. 1, fig. 10;--Geol. Mag., dec. 4, vol. 1, 1894, pl. 8,
fig. 10;--Smithson. Misc. Coll., vol. 67, 1918, pl. 36, figs. 1, 2,
2a-d.--Milne-Edwards, Ann. Sci. Nat., Zoologie, ser. 6, vol. 12,
1881; pl. 12, figs. 44a, b.

In the United States National Museum there is a thin piece of limestone, about 3 inches square, which has on its surface eight jointed objects that have been called legs of trilobites. Two of these were figured by Walcott (1881, pl. 6, fig. 5). The slab contains specimens of _Dalmanella_ and _Cryptolithus_, in addition to the appendages of trilobites, and is said by Doctor Ulrich to have come from the tipper part of the Point Pleasant formation (Trenton) on the bank of the Ohio River below Covington, Kentucky.

The specimens are all endopodites of long slender form, similar to those of _Triarthrus_, but since that genus does not occur in the Point Pleasant, it is necessary to look upon some other trilobite as the former possessor of these organs. Both _Isotelus_ and _Calymene_ occur at this horizon, and as the specimens obviously do not belong to _Isotelus_ or _Cryptolithus_, it is probable that they were formerly part of a _Calymene_.

All the endopodites are of chitinous material, and the various specimens show, according to the perfection of their preservation, from four to six segments. The endopodite as a whole tapers but slightly outward, and the individual segments are of nearly equal length. They appear to be but little crushed, and are oval in section, with a crimped anterior and posterior margin. One or two show a median longitudinal ridge, such as is seen in some appendages of _Triarthrus_. Each segment is parallel-sided, with a slight expansion at the distal end, where the next segment fits into it.

Under the heading "Ordovician Crustacean Leg," Walcott (1918, p. 154, pl. 36, figs. 1,2) has recently redescribed these specimens, and thinks that they do not belong to _Calymene_, nor, indeed, to any trilobite. He concludes that they were more like what one would expect in an isopod. Passing over the fact that the oldest isopod now known is Devonian, the fossils in question seem to me quite trilobite-like. Walcott says:

The legs are associated with fragments of _Calymene meeki_ but it
is not probable that they belong to that species; if they did, they
are unlike any trilobite leg known to me. The very short coxopodite
and basopodite are unknown in the trilobites of which we have the
legs, as they are fused into one joint forming the long protopodite
in the trilobite. The distal joint is also unlike that of the
trilobite legs known to us.

A great deal of Doctor Walcott's difficulty probably arises from his homology of the coxopodite of the trilobite with the protopodite of the higher Crustacea. The coxopodite of the trilobite is not fused with the basipodite, this latter segment always remaining free. Indeed, Walcott himself says of _Neolenus_ (1918, p. 128):

Each thoracic leg (endopodite) is formed of a large elongate
proximal joint (protopodite), four strong joints each about 1.5
times as long as wide (basopodite, ischiopodite, meropodite and
carpopodite); two slender elongate joints (propodite and
dactylopodite) and a claw-like, more or less tripartite
termination.

Walcott's drawing (pl. 36, fig. 1) is a composite one, and while it shows eight segments, I was not able to count more than seven on any of the specimens themselves. In regard to the terminal segment, the dactylopodite of the limb shown in his plate 36, figure 2, is unusually long, and a comparison with other photographs published on the same plate shows that such long segments are unusual.

Proof that these are appendages of a _Calymene_ is of course wanting, but there is no particular reason so far to say that they are not.

_Measurements:_ Two of the more complete specimens, each showing six segments, are each 8 mm. long.

Somewhat similar to the specimens from Covington are the ones described by Eichwald (1825, p. 39, 1860, pl. 21), the specimens being from the Silurian of Gotland. The figure copied by Walcott (1881, pl. 6, fig. 4) has never been looked upon as entirely satisfactory evidence of the nature of the specimen, and so far as I know, the fossil has not been seen by any modern investigator.

=Ceraurus pleurexanthemus= Green.

(pl. 11; text figs. 12, 17-19, 21, 22, 24, 29, 30.)

Illustrated: Walcott, Ann. Lye. Nat. Hist. New York, vol. II, 1875,
pl. 11;--31st Ann. Rept. New York State Mus. Nat. Hist, 1879, pl.
1, fig. 3;--Bull. Mus. Comp. Zool., Harvard Coll., vol. 8, 1881,
pl. 1, figs. 1-5; pl. 2, figs. 1-4, 6-8; pl. 3, figs. 2, 4-7; pl.
4, figs. 1, 2, 4-6, 8; pl. 6, fig. 3; Smithson. Misc. Coll., vol.
67, 1918, pl. 26, figs. 8, 14, 15; pl. 27, figs. 1-3, 5a, 6-9, 12
(not _Calymene_), (not 15, _Calymene_); pl. 28, figs. 1-5; pl. 34,
fig. 1; pl. 35, fig. 7.--Milne-Edwards, Ann. Sci. Nat., Zoologie,
ser. 6, vol. 12, 1881, pl. 10, figs. 1-18.--Bernard, The Apodidæ,
1892, text figs. 46, 51.

_Cephalic Appendages._

No trace of antennules has yet been found.

I find only three sections cut through the plane of the hypostoma of Ceraurus which show anything of the cephalic appendages, and no one of them is very satisfactory. The best is No. 22, the one figured by Walcott (pl. 3, fig. 2, 1881; pl. 27, fig. 12, 1918), but one should remember that this section is not actually cut in the plane of the hypostoma but is a slice diagonally through the head, cutting through one eye and the posterior end of the hypostoma. It shows what seem to be the coxopodites of the second, third, and fourth pairs of cephalic appendages, the exopodites of the third and fourth pairs, and the metastoma. If this interpretation is correct, the first pair of gnathites lay alongside the hypostoma or under its edge, and were feebly developed, the second pair were attached in front of the tip of the hypostoma, curved back close to it, and their inner ends reached the sides of the metastoma. The third and fourth pairs were back of the metastoma, the third pair was stronger than the second, and the fourth probably like the third.

Specimen 92 shows traces of the slender endopodites belonging to the cephalon, but no details. Specimen 22 shows on one side exopodites (epipodites of Walcott) belonging to the third and fourth cephalic appendages. That belonging to the third shows some long setæ and a trace of the shaft, while the one on the fourth appendage (third coxopodite) has a portion of a broad shaft and a number of long setæ. It should again be remembered that the slice does not cut through the plane of the exopodite, but across it at a low angle, so that a part but not all of the shaft is shown. On the other side of this slice there is a fairly good section of one of the thoracic exopodites. It is, however, turned around in the opposite direction from the others, as would be expected in an enrolled specimen.

Specimens 4 and 5 (pl. 1, figs. 4, 5, 1881) are slices cut diagonally through the head of Ceraurus, in front of the posterior tip of the hypostoma. They show fragments of endopodites and exopodites which may be interpreted as practically identical in form with those of the thorax. Due to the diagonal plane in which the section is cut, slice 5 shows the coxopodites of two pairs of appendages, one lying nearer the median cavity than the other. It is extremely difficult to visualize the interpretation of such sections.

_Thoracic Appendages._

A transverse section through a thoracic segment (No. 128, our fig. 17) shows the relation of coxopodite to appendifer to be the same as in _Calymene_, the upper side of the coxopodite having a notch a little outward from the middle. After seeing that specimen, it is possible to understand slice No. 168, which shows longitudinal sections through a number of coxopodites of the thorax, with fragments of both exopodites and endopodites articulated at the distal ends. These and longitudinal vertical sections like No. 18 (pl. 2, fig. 8, 1881) show that the endobases taper inward, and the general uniformity in width in sections taken at various angles indicates that the coxopodites were not greatly flattened.

A unique slice (No. 111, pl. 2, fig. 2, 1881; pl. 27, fig. 1, 1918; our fig. 18) shows a nearly complete thoracic endopodite, and above it a part of the proximal end of the exopodite of the same segment. When one considers that out of over two thousand sections only this one shows the six successive segments of an endopodite, one realizes how futile it is to expect that dozens of the equally slender "spirals" should be cut so as to show practically all their turns.

This endopodite is slender, all the segments have nearly the same length and diameter, though there is a slight taper outward, each segment is expanded distally for the articulation of the next, and there are small spines on the distal ends of some of them. There is probably a terminal spine present, though it is neither so long nor so plainly visible as in Walcott's photograph.

The exopodite on this same specimen was evidently cut diagonally across near the setiferous edge, showing a section through the shaft and the bases of seven setæ (fig. 18). This section is so exactly what would be obtained by cutting similarly an exopodite of either Neolenus or _Triarthrus_ that it should in itself dispose of the "spiral-exopodite" theory.

Several sections have already been illustrated showing sections across the setæ of the exopodites (pl. 3, figs. 4-6, 1881; pl. 27, figs. 3, 4, 9, 1918), and similar sections are not uncommon. Only a very few, however, show sections in the plane of the exopodite. If only No. 111, described above, were known, it would be inferred that the exopodite had a slender shaft as in _Calymene_, but another good slice, No. 80 (fig. 12, ante) shows that the blade was rather broad, though not so broad as in Neolenus. The other specimen is No. 22, which has already been discussed. The thoracic exopodite of this specimen has been very incorrectly figured by Walcott, as it shows no such palmate shaft as he has indicated, but a long blade-like one is outlined, though its entire width is not actually shown.

_Pygidial Appendages._

Sections 14 and 18 (pl. 2, figs. 4, 8, 1881) prove the presence under the pygidium of three pairs of appendages, the coxopodites and fragments of endopodites of which are shown. Nothing is known of the exopodites.

_Relation of Hypostoma to Cephalon._

In Ceraurus the body portion and posterior end of the hypostoma are roughly oval, about as wide as the glabella at its broadest part, and the posterior edge extends back to within 0.5 to 1 mm. of the neck furrow. The posterior pair of appendifers are behind the hypostoma, while the second pair are in front of its posterior end but escape being covered by it on account of its oval shape. At the anterior end the hypostoma is widened by the presence of two side lappets which extend beyond the boundaries of the glabella. In both Ceraurus and Cheirurus the anterior edge of the hypostoma fits against the doublure at the anterior margin of the head and the epistoma is either entirely absent or is so narrow as not to be seen in specimens in the ordinary state of preservation. A section across the cephalon of _Ceraurus pleurexanthemus_ at the horizon of the eyes shows the sides of the hypostoma fitting closely against the sides of the glabella (Walcott's pl. 1, fig. 1). Further back on the head it is not in contact with the dorsal test, and the gnathobases extend beneath it.

Restoration of _Ceraurus pleurexanthemus_. (pl. 11; text fig. 19.)

The restoration of the appendages of _Ceraurus pleurexanthemus_ is a tentative one, based upon a careful study of the translucent sections prepared by Doctor Walcott. In no case among these sections is the actual test of any appendage preserved, and the real form of each part is generally obscured by the crystallization of the calcite which fills the spaces formerly occupied by animal matter.

No section shows anything which can be identified as any part of the antennules, so that these organs have been supplied from analogy with _Triarthrus_.

There are undoubtedly four pairs of biramous Cephalic appendages, but their points of attachment are not so obvious. There are two pairs of conspicuous appendifers on the posterior part of the cephalon and another pair almost concealed by the hypostoma. It is probable that the appendages of the cephalon were not attached directly beneath them, as the four pairs have to be placed within the space occupied by the three pairs of appendifers. As the mouth is in front of the posterior end of the hypostoma, the gnathites of the first pair of biramous appendages may have extended beneath that organ, or they may have lain beside it, and only become functional when the hypostoma was dropped down in the feeding position. The second pair of gnathites reached just to the tip of the hypostoma, and the other two pairs seemingly curved backward behind it.

The points of attachment on the thorax, as shown clearly in sections, were directly beneath the lower ends of the appendifers. The endopodites were long enough to reach to or a little beyond the outer extremities of the pleural spines, while the exopodites were apparently somewhat shorter. Each endopodite consisted of six short, fairly stout segments, each with at least two spines on the somewhat expanded distal ends. The exact form of the exopodites could not be made out. The shaft was apparently rather short, unsegmented, and fairly broad. The setæ appear from the sections to have been more or less blade-shaped and to have overlapped, as do those of the exopodites of _Cryptolithus_. Judging from their position in the sections, the setæ not only bordered the posterior side of the shaft, but radiated out from the end as well.

The pygidium shows three pairs of functional appendifers, hence three pairs of appendages have been supplied. There is a fourth pair of rudimentary appendifers, but as they are beneath the doublure they could not have borne ambulatory appendages.

The Appendages of Acidaspis trentonensis Walcott.

(pl. 6, fig. 6.)

A single individual of _Acidaspis trentonensis_, obtained from the same locality and horizon as the specimens of _Triarthrus_ and _Cryptolithus_, when cleaned from the ventral side shows a number of poorly preserved endopodites which seem very similar in shape and position to those of _Triarthrus_. One endopodite on the right side of the head and the first five on the right side of the thorax are the best shown. All are slender, are directed first forward at an angle of about 45 with the axis, then, except in the case of the cephalic appendage, turn backward on a gentle curve and extend a little distance beyond the margin of the test, but not as far as the tips of the lateral spines of the thoracic segments.

The individual segments of the endopodites can not be seen clearly enough to make any measurements. On the fourth and fifth endopodites of the thorax, some of the segments seem to be broad and triangular as in _Triarthrus_. All that can be seen indicates that _Acidaspis_ had appendages entirely similar to those of _Triarthrus_, but perhaps not quite so long, as they seem not to have projected beyond the limits of the lateral spines. There are no traces of antennules nor, unfortunately, of exopodites.

_Measurements:_ Length 8 mm.

Walcott (1881, p. 206) stated that his sections had shown the presence in this species of legs "both cephalic and thoracic" and also the "spiral branchiæ." His specimens were from the Trenton at Trenton Falls, New York.

The Appendages of Cryptolithus.

=Cryptolithus tessellatus= Green.

(pl. 6, fig. 7; pls. 7-9; text figs. 20, 25, 45, 46.)

(See also Part IV.)

Illustrated: Beecher, Amer. Jour. Sci., vol. 49, 1895, pl. 3.

When Professor Beecher wrote his short article on the "Structure and Appendages of _Trinucleus_" (1895), he had only three specimens showing appendages. In his later work he cleaned several more, so that there are now thirteen specimens of _Trinucleus_ = _Cryptolithus_ available for study, though some of these do not show much detail. In his last and unpublished study, Beecher devoted the major part of his attention to this genus, and summarized his findings in the drawings which he himself made of the best individuals (text figs. 45, 46). Valiant (1901) stated that he had found a _Trinucleus_ with antennæ in the Frankfort shale south of Rome, New York. The specimen has not been figured.

None of the specimens shows much more of the appendages of the cephalon than, the hypostoma and the antennules, so that we are still in ignorance about the mouth parts.

The most striking characteristics of the appendages are as follows: the antennules are long, and turn backward instead of forward; none of the limbs projects beyond the margin of the dorsal test; the exopodites extend beyond the endopodites, reaching very nearly to the margin of the test; the endopodites are not stretched out at right angles to the axis, but the first three segments have a forward and outward direction as in _Triarthrus_, while the last four turn back abruptly so that they are parallel to the axis; the limbs at the anterior end of the thorax are much more powerful than the others; the dactylopodites of the endopodites show a fringe of setæ instead of three spines as in _Triarthrus_ and _Neolenus_. All these would, as Beecher has already suggested, seem to be adaptations to a burrowing habit of life, the antennules being turned backward and the other appendages kept within the shelter of the dorsal test in order to protect them, and the anterior endopodites enlarged and equipped with extra spines to make them more efficient digging and pushing organs.

_Restoration of Cryptolithus._

(Text fig. 20.)

It should be definitely understood that the present figure is a restoration and not a drawing of a specimen, and that there are many points in the morphology of _Cryptolithus_ about which no information is available, especially about the appendages under the central portion of the cephalon. The information afforded by all the figures published in this memoir is combined here. As gnathites are preserved on none of the specimens, those represented in the figure are purely conventional.

A person who is acquainted only with _Cryptolithus_ preserved in shale, or with figures, usually has a very erroneous idea of the fringe It is not a flat border spread out around the front of the head, but stands at an angle about 45 in uncrushed specimens of most species. When viewed from the lower side, there is a single outer, concentric row of the cup-shaped depressions, bounded within by a prominent girder. This row is in an approximately horizontal plane, while the remainder of the doublure of the fringe rises steeply into the hollow of the cephalon. Since the front of the hypostoma is attached to this doublure, it stands high up within the vault and under the glabella. Two specimens, Nos. 231 and 233, show something of the hypostoma, and they are the only ones known of any American trinucleid. That of specimen 233, the better preserved, is very small, straight across the front, and oval behind. It seems that it is abnormally small in this specimen and I should not be surprised if in other specimens it should be found to be larger.

In the Bohemian _Trinucleoides reussi_ (Barrande), the oldest of the trinucleids, the hypostoma is very commonly present, and is of the proper size to just cover the cavity of the glabella, seen from the lower side, and has, toward the anterior end, side flaps which reach out under the prominent glabellar lobes. This large size of the hypostoma would cause the antennules to be attached outside the dorsal furrows, and the position in which they are attached in the American species of _Cryptolithus_ may be explained as an inherited one, since with the small hypostoma they might have been within the glabella, as in _Triarthrus_.

The antennules are seen in three specimens, and in all cases are directed backward. The particular course in which they are drawn in the restoration is purely arbitrary. The second pair of cephalic appendages are represented as directed downward and forward, since in one or two specimens fragments of forward-pointing endopodites were seen near the front of the cephalon, and because in other trilobites the second pair of appendages is always directed forward. The remaining three pairs have a more solid basis in observed fact, for the two or three specimens retaining fragmentary remains of them indicate that they turn backward like those on the thorax, and that the individual segments are longer and more nearly parallel-sided than those of the more posterior appendages. The gnathites of all the cephalic appendages are admittedly purely hypothetical. None of the specimens shows them. As drawn, they are singularly inefficient as jaws, but if, as is suggested by the casts of the intestines of trinucleids found in Bohemia, these trilobites were mud-feeders, inefficient mouth-parts would be quite in order.

The appendages of the thorax and pygidium can fortunately be taken quite directly from the photographs of the dorsal and ventral sides of well preserved specimens. There is of course a question as to the number and the exact form of those on the pygidium, but I think the present restoration is fairly well justified by the specimens. As would be expected from the narrow axial lobe, the gnathobases of the coxopodites are short and small.

Summary on the Ventral Anatomy of Trilobites.

COMPARISON OF APPENDAGES OF DIFFERENT GENERA.

Since the appendages of _Triarthrus_, _Cryptolithus_, _Neolenus_, _Calymene_, and _Ceraurus_ are now known with some degree of completeness, those of _Isotelus_ somewhat less fully, and something at least of those of _Ptychoparia_, _Kootenia_, and _Acidaspis_, these forms being representatives of all three orders and of seven different families of trilobites, it is of some interest to compare the homologous organs of each.

All in which the various appendages are preserved prove to have a pair of antennules, four pairs of biramous limbs on the cephalon, as many pairs of biramous limbs as there are segments in the thorax, and a variable number of pairs on the pygidium, with, in the case of _Neolenus_ alone, a pair of tactile organs at the posterior end. Each limb, whether of cephalon, thorax, or pygidium, consists of a coxopodite, which is attached on its dorsal side to the ventral integument and supported by an appendifer, an exopodite, and an endopodite. The exopodite is setiferous, and the shaft is of variable form, consisting of one, two, or numerous segments. The endopodite always has six segments, the distal one armed with short movable spines.

_Coxopodite._

The coxopodite does not correspond to the protopodite of higher Crustacea, the basipodite remaining as a separate entity. The inner end of the coxopodite is prolonged into a flattened or cylindrical process, which on the cephalon is more or less modified to assist in feeding, and so becomes a gnathobase or gnathite. The inner ends of the coxopodites of the thorax and pygidium are also prolonged in a similar fashion, but are generally somewhat less modified. These organs also undoubtedly assisted in carrying food forward to the mouth, but since they probably had other functions as well, I prefer to give them the more non-committal name of endobases.

In _Triarthrus_ and _Neolenus_ the endobases are flattened and taper somewhat toward the inward end. In _Isotelus_, _Calymene_ and _Ceraurus_, they appear to have been cylindrical. In other genera they are not yet well known. In all cases, particularly about the mouth, they appear to have been directed somewhat backward from the point of attachment. As it is supposed that these organs moved freely forward and backward, the position in which they occur in the best preserved fossils should indicate something of their natural position when muscles were relaxed.

_Cephalon._

_Antennules._--Antennules are known in _Triarthrus_, _Cryptolithus_, _Neolenus_, and _Ptychoparia_. In all they are long, slender, and composed of numerous segments, which are spiniferous in _Neolenus_, and very probably so in the other genera.

In _Triarthrus_, _Neolenus_, and _Ptychoparia_ they project ahead of the cephalon, emerging quite close together under the front of the glabella, one on either side of the median line. In _Cryptolithus_ they turn backward beneath the body, but since only three or four specimens are known which retain them, it is possible that other specimens would show that these organs were capable of being turned forward as well as backward. The proximal ends of the antennules being ball-like, it is probable, as Doctor Faxon has suggested to me, that these "feelers" had considerable freedom of motion. The antennules of _Triarthrus_ are apparently somewhat less flexible than those of the other genera, and have a double curvature that is seen among the others only in Ptychoparia. The proximal end of an antennule in _Triarthrus_ is a short cylindrical shaft, apparently articulating in a sort of ball-and-socket joint. The proximal end in the other genera is still unknown. The points of attachment in _Triarthrus_ seem to be under the inner part of the second pair of glabellar furrows. In _Cryptolithus_ they appear to be beside the anterior lobe of the glabella under what have long been known as the antennal pits. In the other genera the location is not definitely known, but in _Neolenus_ it seems to be under the dorsal furrows near the anterior end of the glabella. Viewed from the under side, the point of attachment is probably always beside the middle or anterior part of the hypostoma, just behind the side wings.

_Paired biramous appendages._--Behind the antennules all the appendages except those on the anal segment are biramous, consisting of a coxopodite with an inward-directed endobase and an outward-directed pair of branches, the exopodite above, and the six-jointed endopodite beneath. The basipodite really bears the exopodite, but the latter also touches the coxopodite. This structure has been seen in _Triarthrus_, _Cryptolithus_, _Neolenus_, _Kootenia_, _Calymene_, _Ceraurus_, and _Ptychoparia_. In _Triarthrus_, _Neolenus_, _Acidaspis_, _Ptyclioparia_, and Kootenia, the appendages extend beyond the margins of the dorsal test. In _Cryptolithus_ and _Isotelus_ none (other than antennules) does so. In _Isotelus_ and _Acidaspis_ only the endopodites have been seen. In _Triarthrus_, _Calymene_, _Ceraurus_, and _Neolenus_ there are four pairs of appendages behind the antennules. The other genera probably had the same number, but the full structure of the under part of their cephala is not known. In _Triarthrus_ the endopodites of the cephalon are slender, the individual segments parallel-sided, the inner ones flattened, the outer ones cylindrical in section. They project slightly beyond the edge of the cephalon when fully extended, and each terminates in three small spines. In _Cryptolithus_ the endopodites of the cephalon are longer than those of the thorax, but with the possible exception of the first pair, are bent backward at the carpopodite, and do not ordinarily project beyond the brim of the test. In _Neolenus_ the endopodites of the cephalon are rather thick and wide, but are long, project forward, and extend beyond the brim. The individual segments are flattened, probably compressed oval in section. The terminal segment of each is furnished with three strong spines at its distal end. In _Calymene_ and _Ceraurus_ the endopodites appear to consist of slender segments which are oval or circular in section. In _Calymene_ Walcott believed the three distal segments of the last endopodites of the head to be greatly enlarged, giving these appendages a paddle-like form similar to some of the appendages of eurypterids. The evidence for this does not seem to me to be good. The cephalic endopodites of _Isotelus_ are entirely similar to those of the thorax, and are rather short, consisting of a series of short cylindrical segments which do not taper greatly toward the distal end. The endopodites of the cephalon of _Acidaspis_, _Kootenia_, and _Ptychoparia_ are still unknown.

The exopodites of the cephalon seem in all known cases (_Triarthrus_, _Cryptolithus_, _Neolenus_, and Ceraurus) to be like those of the thorax. They point more directly forward in most cases, project beyond the margin of the head normally only in Triarthrus, and usually occupy the region under the cheeks (fixed and free).

The endobases of the coxopodites of the appendages of the cephalon probably in all cases function as mouth-parts (gnathites), and are especially modified for this purpose in Triarthrus, being flattened, shoe-shaped in outline, and so arranged that they work over one another in a shearing fashion. While the more anterior of the coxopodites are attached in front of the posterior tip of the hypostoma, the gnathites of Triarthrus bend backward so that all are behind the hypostoma. In _Calymene_ and _Ceraurus_, two or three pairs of the gnathites are back of the hypostoma, and one or more pairs may be beside or under the hypostoma. In these genera the mouth is probably in front of the tip of the upper lip. In _Isotelus_, the mouth seems to have been situated in the notch between the two branches of the hypostoma, and the gnathites of two or three pairs of the appendages probably worked under its forks. Since the length of the hypostoma differs in the various species of _Isotelus_, there would be a variable number of gnathites projecting under its forks, according to the species. In this genus the gnathites are of the same long form, cylindrical in cross-section, as the endobases of the thoracic segments, but each is bowed back considerably from the point of attachment.

The gnathites of _Neolenus_ are like the endobases of the thorax, but broader. The great length of the hypostoma makes it probable that the mouth was far back and that some of the gnathites were in front of it. The gnathites of _Cryptolithus_ are unknown. Professor Beecher in his drawing shows some fragments with toothed ends near the hypostoma, and it may be that they are inner ends of gnathites, but I see nothing to substantiate such an interpretation. If, as some suppose, _Cryptolithus_ was a mud feeder, the gnathites were probably poorly developed. Of the gnathites of _Kootenia_, _Ptychoparia_, and _Acidaspis_ also nothing is known.

_Thorax._

In each genus there is a pair of appendages for each segment of the thorax. When the axial lobe is narrow, the endobases of the coxopodites are small and short (_Cryptolithus_, _Ceraurus_, _Calymene_). When the axial lobe is wide, the endobases are long and stout (_Isotelus_, _Triarthrus_). The exopodites always lie above and in front of the corresponding endopodites. In Triarthrus the two branches are of practically equal length. In _Cryptolithus_ the exopodites are much the longer. In _Neolenus_, _Calymene_, _Ceraurus_, _Kootenia_, and _Ptychoparia_, the exopodites are shorter than the endopodites.

The exopodites in Triarthrus consist of a proximal shaft, succeeded by numerous short segments, and ending distally in a long, grooved, somewhat spatula-shaped segment. Along the anterior margin of the shaft there are many small spines. Along the posterior margin there are numerous flattened setæ, which all lie in one plane and which seem to be more or less united to one another like the barbs of a feather. The setæ are short, not much longer than the width of one of the thoracic segments, and point backward and outward. In _Cryptolithus_ the shaft does not seem to be made up of small segments, and is narrow, with a decided backward curve. The setæ are considerably longer and much more flattened than in Triarthrus. In _Calymene_ the state of preservation does not allow a very full knowledge of the exopodites, but they appear to have a slender, unjointed shaft and short and delicate setæ. The coiled branches of the exopodites as described by Walcott seem to me to be only ordinary Triarthrus-like organs, and this, as I understand from Schuchert, was also the view of Beecher. In _Ceraurus_ the exopodite seems to have been somewhat paddle-shaped, expanded at the distal end, and to have had rather thick, blade-like setæ.

The exopodite of _Neolenus_ is decidedly leaf-like, and reminds one somewhat of the exites of some of the phyllopods. The shaft is a broad unsegmented blade. The setæ are slender, delicate, flattened, and a little longer than the width of the shaft. The exopodites of this genus point forward all along the body. In _Kootenia_ the exopodites are like those of _Neolenus_, but with a narrower shaft. The exopodites of _Ptychoparia_ appear to be very much like those of Triarthrus, but the shaft is probably not segmented.

The endopodites of the thorax of _Triarthrus_, _Cryptolithus_, and _Acidaspis_ show progressive modification from front to back in the broadening of the individual segments and the assumption by them of a triangular form. Not only do the individual segments become more triangular from front to back, but more of the segments of each endopodite become triangular. This modification has so far been seen in these three genera only. The individual segments, except the distal ones, seem to be flattened in all these genera. The distal end of the terminal segment of each endopodite of _Triarthrus_ bears three small movable spines, and each of the segments usually bears three or more spines, located in sockets along the dorsal surface and at the anterior distal angle of each segment. The endopodite of _Cryptolithus_ is bent backward at the carpopodite and this segment is always thickened. At the distal end of the dactylopodite there is a tuft of spines, the triangular segments have tufts of spines on their posterior corners, and there are groups of spines also in the neighborhood of the articulations.

The endopodites of _Ceraurus_, _Calymene_, and _Isotelus_ are all relatively slender, the segments are parallel-sided, and there seems to be no particular modification from front to back of the thorax. The endopodites of _Isotelus_ are short, the entire six segments of one being but little longer than the coxopodite of the same appendage. The segments of the endopodites of _Neolenus_ are mostly short and wide, and at the distal end of the terminal segment there are three stout spines. In _Kootenia_ the endopodites are long and very slender. The endopodites of Ptychoparia are too poorly preserved to show details, and those of the thorax of _Acidaspis_ likewise reveal little structure, but they seem to have the triangular modification, and to turn back somewhat sharply at about the position of the carpopodite.

_Pygidium._

Beecher showed that in _Triarthrus_ there was a pair of appendages on the pygidium for every segment of which it is composed except the last or anal segment (protopygidium). Walcott has since shown that in _Neolenus_ this segment bears a pair of cerci, and Beecher's drawings show that in his later studies he recognized a spinous plate, the possible bearer of cerci, on the anal segment of _Triarthrus_. The appendages of the anal segment have not yet been seen on other species of trilobites.

The appendages of the pygidium do not show any special modifications, but seem in all cases to be similar to those of the posterior part of the thorax. In _Cryptolithus_ all the pygidial appendages are short and remain beneath the cover of the dorsal test, while in _Triarthrus_ and _Neolenus_ they extend behind it.

In the latter genus the endopodites of the pygidial appendages appear to be practically identical in form with those of the thorax, the individual segments being perhaps a little more nearly square in outline. Like those of the thorax, the segments of the pygidial endopodites bear numerous short spines. The caudal cerci are richly segmented, slightly flexible, spinous tactile organs. They are symmetrically placed, nearly straight when in their natural position, and make an angle of about 75 with one another. They appear to be attached to a narrow rim-like plate which seems to fit in just ahead of the doublure of the pygidium, or perhaps over it.

In _Ceraurus_, _Calymene_, and _Isotelus_, the endopodites of the pygidium are similar to those of the thorax, but seemingly more slender, with less well developed coxopodites, and with, in the last-named genus, slender cylindrical segments. Exopodites are not known on the pygidia of any of these genera, but since they are present and like those of the thorax in _Triarthrus_, _Cryptolithus_, _Neolenus_, and _Ptychoparia_, there is little reason to think that they were absent in _Ceraurus_ or _Calymene_, though there is some question about _Isotelus_.

The limbs are largest and longest on the anterior part of the thorax of a trilobite, and diminish regularly in length and strength to the posterior end of the pygidium. This regular gradation shows, as Beecher was the first to point out, that the growing point of the trilobites is, as in other arthropods, in front of the anal segment. New _free_ segments are introduced into the thorax at the anterior end of the pygidium, and this has led to some confusion between the growing point and the place of introduction of free segments.

If a new segment were introduced at a moult in front of the pygidium, that segment would probably have less fully developed appendages than those adjacent to it, and so make a break in the regular succession. The condition of the appendages corroborates the evidence derived from the ontogeny of the pygidium, and proves that the new segments are introduced at the same growing point as in other Arthropoda.

_Caudal Rami._

Bernard, who believed that the Crustacea had been derived through an _Apus_-like ancestor (1892, pp. 20, 85, 274), pointed out that four or less than four anal cirri were to be expected. Two well developed cirri and two rudimentary ones are present in _Apus_, and they are also to be found in other phyllopods and some isopods. It is, however, characteristic of the Crustacea as a whole to lack appendages on the anal segment. Caudal cirri (cerci) are much more freely developed in the hexapods than in the Crustacea, particularly in the more primitive orders, Palæodictyoptera, Apterygota, Archiptera, and Neuroptera. They are supposed, in this case, to be modified limbs, and therefore not homologous with the bristles on the anal segment of an annelid. Doctor W. M. Wheeler of the Bussey Institution has kindly allowed me to quote the following excerpt from a letter to me, as expressing the opinion of one who has made an extensive study of the embryology of insects:

I would say that I have no doubt that the cerci of insects are
directly inherited from the insect ancestors. They are always
highly developed in the lower insects, and only absent or vestigial
in a few of the most highly specialized orders such as the
Hemiptera, Diptera, and Hymenoptera. I have further no doubt
concerning their being originally ambulatory in function. They are
certainly not developed independently in insects. Embryologically
they arise precisely like the legs, and each cercus contains a
diverticulum of the mesoblastic somite precisely as is the case
with the ambulatory legs and mouth parts.

The "pygidial antennæ" seem to be as fully developed in _Neolenus_ as in any of the other arthropods, and may suggest a common ancestry of the phyllopods, isopods, and hexapods, in the trilobites. They were doubtless tactile organs, and while the evidence is chiefly negative, it would seem that they proved useless, and were lost early in the phylogeny of this group. Possibly the use of the pygidium as a swimming organ proved destructive to them.

HOMOLOGY OF THE CEPHALIC APPENDAGES WITH THOSE OF OTHER CRUSTACEA.

The head of the typical crustacean bears five pairs of appendages, namely, the antennules, antennas, mandibles, and first and second maxillæ, or, as they are more properly called, the maxillulæ and maxillæ.

As Beecher has pointed out, the "antennæ" of the trilobites, on account of their pre-oral position and invariably uniramous character, are quite certainly to be correlated with the antennules.

The second pair of appendages, the first pair of biramous ones, Beecher homologized with the antennæ of other crustaceans, and that homology has been generally accepted, though Kingsley (1897) suggested that it was possible that no representatives of the true antennæ were present.

In preparing the restorations in the present study, the greatest difficulty has been to adjust the organs about the mouth. In _Triarthrus_, numerous specimens show that without question there are four pairs of gnathites back of the hypostoma, and that all four belong to the cephalon. In forms with a long hypostoma, however, there was no room on the cephalon for the attachment of four pairs of gnathites, neither were there enough appendifers to supply the requisite fulcra. At first I supposed I had solved the difficulty by assuming the mouth to be in front of the posterior tip of the hypostoma, as it really is in Ceraurus and _Calymene_, and allowing the gnathites to play under the hypostoma as Walcott (1912) has shown that they do in _Marrella_. Finally, when I came to study in greater detail the slices of _Calymene_ and _Ceraurus_, they seemed to show that the anterior one or two pairs of appendages became degenerate and under-developed. This was probably a specialization due to the great development of the hypostoma in trilobites, that organ being much more prominent in this than in any other group. As the hypostoma lengthened to accommodate the increasing size of sub-glabellar organs (stomach, heart, etc.), the mouth migrated backward, leaving the anterior appendages ahead of it, with their gnathobases, at least, functionless. That such migration has taken place, even in Triarthrus, is shown by the fact that the points of articulation of the first biramous appendages are pre-oral, and it is more obviously true of _Ceraurus_. Correlated with the weakening of the appendages on the lower surface is the loss of glabellar furrows on the upper surface. The glabellar furrows mark lines of infolding of the test to form the appendifers and other rugosities for the attachment of tendons and muscles. It is conceivable that this migration backward of the mouth began very early in the history of the race, and that even before Cambrian times, the antennæ, probably originally biramous appendages like those on the remainder of the body, had dwindled away and become lost. If this is the case, then the first pair of biramous appendages of _Triarthrus_ would be mandibles, the second pair maxillulæ, and the third pair maxillæ.

There remain the last pair of cephalic appendages, and they bring up the whole head problem of the trilobites. Beecher has stated (1897 A, p. 96) his conviction that the head of the trilobite is made up of five segments, representing the third, fourth, fifth, sixth, and seventh neuromeres of the theoretical crustacean. As a matter of fact, he really made up the head of seven segments, since he stated that the first neuromere was represented by the hypostoma and the second by the epistoma and free cheeks.

Jaekel (1901, p. 157) nearly agreed with Beecher, but made eight segments, as he saw five segments in the glabella of certain trilobites. In his table (p. 165) he has listed the segments with their appendages as follows: 1. Acron, with hypostoma; 2, rostrum (epistoma), with free cheeks; 3, first frontal lobe, with (?) antennules; 4, second frontal lobe, with antennæ; 5, mandibles; 6, first, or pre-maxillæ; 7, second maxillæ; 8, occipital segment with maxillipeds.

Jaekel refused to believe that the antennæ of trilobites were really entirely simple, and so homologized them with the antennæ and not the antennules of other Crustacea. In this he was obviously incorrect, but it accounts for his homology of the remainder of the cephalic appendages.

It is, at present, impossible to demonstrate the actual number of somites in the cephalon of the trilobite, but I believe that Beecher was correct in holding that the glabella was composed of four segments. There are, it is true, a number of trilobites (Mesonacidæ, Paradoxidæ Cheiruridæ, etc.) which show distinctly four pairs of glabellar furrows, indicating five segments in the glabella. This is, however, probably due to a secondary division of the first lobe.

The correspondence of the five segments on the dorsal side with the five pairs of appendages makes it unlikely that any pair of limbs has been lost. The condition in _Marrella_, where a trilobite-like cephalon bears five pairs of appendages, the second pair of which are tactile antennæ, is favorable to the above interpretation. In spite of the apparent degeneration of the first two pairs of appendages in _Calymene_, no limbs are actually missing, and if some are dropped out in the later trilobites it would not affect the homology of those now known. I therefore agree with Beecher in homologizing the appendages, pair for pair, with those of the higher Crustacea.

FUNCTIONS OF THE APPENDAGES.

_Antennules._

The antennules were obviously tactile organs, probably freely movable in most trilobites, but in the case of Triarthrus perhaps rather rigid, judging from the great numbers of specimens which show the characteristic sigmoid curve made familiar by Professor Beecher's restoration. The proximal end of the shaft of each antennule of Triarthrus is hemispheric and doubtless fitted into a socket, thus suggesting great mobility of the whole organ. In spite of this, I have seen no specimens in which they did not turn in toward each other and cross the anterior margin very near the median line. In front of the margin, various specimens show evidence of flexibility, but from the proximal end to the margin the position is the same in all specimens.

In all the few specimens of _Cryptolithus_ retaining the antennules, these organs are turned directly backward, but it is entirely within the range of probabilities that while its burrowing habits made this the more usual position, the animal had the power of turning them around to the front when they could be used to advantage in that direction.

_Exopodites._

It has been the opinion of most observers that the exopodites of trilobites were swimming organs, while others have thought that they functioned also in aerating the blood. To the present writer it seems probable that the chief function was that of acting as gills, for which the numerous thin, flattened or blade-like setæ are particularly adapted. That they were also used in swimming is of course possible, but that was not their chief function. It should be remembered that the exopodites are always found dorsal to or above the endopodites, and in a horizontal plane. For use in swimming it would have been necessary to rotate each exopodite into a plane approximately perpendicular to or at least making a considerable angle with the dorsal test. In this position, the exopodites would have been thrust down between the endopodites, and one would expect to find some specimens in which a part at least of the exopodites were ventral to the endopodites. Specimens in this condition have not yet been seen among the fossils. To avoid having the exopodites and endopodites intermingled in this way, the animal would have to bring all the endopodites together along the axial line in a plane approximately perpendicular to the dorsal test, in which case the exopodites would be free to act as swimming organs. The fact that the setæ of an exopodite stay together like the barbs on a feather would of course tend to strengthen the idea that the exopodites could be used in swimming, but that is not the only possible explanation of this condition. The union of the basipodite and exopodite shows that the two branches of the appendage acted together. Every movement of one affected the other, and the motion of the endopodites in either swimming or crawling produced a movement of the exopodites which helped to keep up a circulation of water, thus insuring a constant supply of oxygen.