The Appendages of Isotelus.

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.

Fig. 7.—A restored section across the thorax ofNeolenus serratus, showing the probable form of attachment of the appendages, their relation to the ventral membrane, and the jaw-like endobases of the coxopodites.

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.

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

Nathorstia transitansWalcott.

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 outlineof the body, the position of the eye, and even a trace of spines about the pygidium (in the figure) are similar to those ofNeolenus, and I would venture the suggestion thatNathorstia transitansis a recently moultedNeolenus 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.

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 suggestsNeolenus. 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 asMolaria spinifera, and the trilobites." In what respect it is transitional does not appear.

Formation and locality: Same as that ofNeolenus serratus. One nearly complete specimen and a few fragments were found.

HISTORICAL.

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

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

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

In the same year Woodward (1884, p. 162, fig. 1-3) reproduced all of Mickleborough's figures, and suggested that the last seven pairs of appendages on the pygidium ofCalymeneandIsoteluswere probably "lamelliform branchiferous appendages, as inLimulusand 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 ofIsotelus 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 nameIsotelus covingtonensis?Foerste (not Ulrich). He advances the view, which I am unable to share, that the cylindrical appearance of the segments of the appendages ofIsotelusis due to post-mortem changes.

Isotelus latusRaymond.

(pl. 10, fig. 1.)

Illustrated:Asaphus platycephalusBillings, 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 latusRaymond, 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.

Isotelus latusRaymond, 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 ofIsotelus maximus, and the segments, while of less diameter than the coxopodites, do not show so great a contrast tothem 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 inIsotelus 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 maximusLocke.

(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 ofIsoteluswith 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. InTriarthrusand 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.[1]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.

[1]If this specimen had the same proportions as specimens ofIsotelus maximusfrom Toronto, the total length would be only 174 mm. The cephalon would be about 52 mm. long, the thorax 58 mm., and the pygidium about 64 mm. long.

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.

(Textfig. 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 ofIsotelus maximusfrom Toronto, Ontario.

Isotelus gigasDekay.

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 calledAsaphus platycephalus, is, in all probability, anIsotelus gigas. Woodward says of it:

I was at once attracted by a specimen ofAsaphus, 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 thejointed palpusof 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, asApus,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.

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.

Fig. 9.—A restored composite ofIsotelus maximusandI. latus. The exopodites are left out because entirely unknown. Drawn by Doctor Elvira Wood. Natural size.

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 gigasDekay. Ordovician, Trenton Limestone. N. America, Canada. Descr. H. Woodward, 1870, Q. J. G. S., XXVI, pp. 486-488, text fig. 1, asAsaphus 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.

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 ofIsotelus gigasfrom 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 arenicolaRaymond.

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. InIsotelus gigasthe 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.

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. InIsotelus gigasthe 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.

Triarthrus beckiGreen.

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

(Also seePart 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.—Œhlert, 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.

Specimens ofTriarthrusretaining 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 withLimulusseems not to be as close inTriarthrusas 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 ofTriarthrus, 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 onTriarthrusuntil 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 ofTriarthruswith those ofCalymeneandCeraurus.

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).

Œhlert (1896) gave an excellent summary in French of the work of Beecher and Walcott onTriarthrus, 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.

Fig. 10.—Triarthrus beckiGreen. A new restoration, modified from Professor Beecher's, to incorporate the results of his later work. The inner ends of the endobases are probably too far apart, as it was not discovered until after the drawing had been made that the appendifers projected within the dorsal furrows. Drawn by Doctor Elvira Wood. × about 3.8.

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 fromspecimens ofTriarthrusandCryptolithushe 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 ofPtychoparia, 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 ofTriarthrus. 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 toTriarthrusis on pages 197 to 219.

Moberg (1907), in connection with a specimen ofEurycare angustatumwhich he thought preserved some appendages, described and illustrated some of the appendages ofTriarthrus.

The most recent discussion ofTriarthrus, 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 ofApus. 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 ofTriarthrusis 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 emphasizeone of the important characteristics ofTriarthrusthe regional differentiation of the appendages.

It should be pointed out that althoughTriarthrusis 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 beckiis 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 moundwhose 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 ofTriarthrus, 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.

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