The Appendages, Anatomy, and Relationships of Trilobites Part 27

The shaft of the exopodite is made up of numerous short segments which at their anterior outer angles are produced into spines, and which also bear movable spines along the anterior border. As shown in several other specimens, the exopodite ends in a more or less long spoon-shaped segment bearing on its lower surface a broad groove. No setae appear to be attached to this, but both anterior and posterior margins bear numerous small, apparently movable spines. From the groove along the ventral side of the remainder of the exopodite arise numerous long slender filaments which become progressively shorter toward the tip. This specimen shows that they are not cylindrical, but are flattened along opposite faces, at least at their distal ends.

While no connection can be seen between adjacent setae, they seem to stay together like the barbs on a feather.

_Measurements:_ Length, 33 mm., width at back of cephalon, 16 mm.; from front of cephalon to back of hypostoma, 6 mm.

Specimen No. 219 (pl. 2, fig. 6; pl. 4, fig. 4).

Ill.u.s.trated: Amer. Jour. Sci., vol. 13, 1902, pl. 4, fig. 1, pl. 5, fig. 4 (photograph and drawing).

The endopodites of most of the appendages of the thorax are well shown, and occasional portions of exopodites. The c.o.xopodites are long, flattened, and do not taper much. The anterior and posterior edges of the basipodites of the endopodites of the first two segments are approximately parallel, but on the succeeding endopodites the basipodites and ischiopodites are triangular in form, with the apex backward. In successive endopodites toward the posterior end, the angle made by the backward-directed sides of the basipodites becomes increasingly acute, so that in some of the posterior appendages this segment is wider than long. The ischiopodite shows a similar increase of width and angularity on successive segments, and the meropodites and carpopodites also become wider on the posterior segments, and even triangular in outline toward the back of the thorax and on the pygidium.

Along the median portion of the axial lobe the specimen has been cleaned until the inner side of the ventral membrane was reached. Here the test shows on the inner surface at each segment of the thorax a series of low ridges which are roughly parallel to the axial line, but which really converge in an anterior direction. Between the ridges are shallow canoe-shaped depressions, which have the appearance of areas for the insertion of muscles.

_Measurements:_ Length, 31 mm.; width at back of head, 15 mm.; distance, in a straight line, from point of insertion of the right antennule to its tip, 14.25 mm.; it projects 12 mm. beyond the cephalon.

Specimen No. 218 (pl. 6, fig. 3; text fig. 43).

This specimen is a large one, developed from the lower side, but retains only the endopodites of a few appendages. The cephalon and anterior portion of the thorax are missing.

Professor Beecher had a drawing made to show the appendages on the right-hand side of the last two segments of the thorax, seen of course from the ventral side. This drawing shows well the broadening of the basipodite, ischiopodite, and meropodite, while the c.o.xopodite is thick and heavy, and the inner end of the gnathobase somewhat rugose.

Almost every segment of the endopodites has one or more pits for insertion of spines, these being along the anterior or posterior margins. The exopodites lack the setae, but show no unusual features.

[Ill.u.s.tration: Fig. 43.--_Triarthrus becki_ Green. Drawing to represent the writer's interpretation of the appendages of specimen 218. Drawn by Miss Wood. 10.]

Specimen No. 222 (pl. 4, fig. 5).

Ill.u.s.trated: Amer. Jour. Sci., vol. 47, 1894, pl. 7, fig. 3 (drawing).

A small specimen, developed from the lower side, and used by Professor Beecher to ill.u.s.trate the form of the segments of the endopodites of the pygidium. In addition to this, it shows very well the form of the endopodites of the thorax. All of the appendages on the specimen are shifted to the left of their normal position. This specimen differs from most of the others in that the segments of the endopodites do not lie with their greatest width in the horizontal plane, but were embedded vertically, with the posterior edge downward. From this circ.u.mstance they retain their natural shape, and it is seen that they are naturally flattened, with about the same thickness in proportion to length and width as in some of the modern isopods (Serolis, for instance). In even the most anterior of these endopodites (that of the second segment) the ischiopodite, meropodite, and carpopodite are triangular in shape, with the point backward, but in all the endopodites at the anterior end of the thorax, the triangle has a very obtuse angle at the apex, and the base is much longer than the perpendicular. On the other hand, those of the pygidium, which were figured by Beecher, have a number of short wide segments, all wider than long, and, excepting the dactylopodites, triangular in form.

_Measurements:_ Length, 8.75 mm.; width at back of cephalon, about 4 mm.

Specimen No. 230 (pl. 5, fig. 3; text fig. 44).

Ill.u.s.trated: Amer. Jour. Sci., vol. 47, 1894, pl. 7, fig. 2 (drawing); Ibid., vol. 13, 1902, pl. 2, fig. 2.

[Ill.u.s.tration: Fig. 44.--_Triarthrus becki_ Green. Appendages of the posterior part of the thorax and pygidium of specimen 230. Inked by Miss Wood from a tracing made under the direction of Professor Beecher.]

An entire specimen of medium size, developed from the ventral side. It seems to have been the first one to yield to Professor Beecher any satisfactory knowledge of the appendages of the pygidium. There are five endopodites, all on one side, which appear to belong here. The segments in this region are characterized by their short, wide, triangular form. At the apex of each is a small tuft of spines or short hairs, and the ventral surfaces of some of the endopodites show pits for the insertion of spines.

_Measurements:_ Length, 21 mm.; width at back of cephalon, 10 mm.

Cryptolithus tessellatus Green.

Specimen No. 233 (pl. 7, fig. 1; text fig. 45).

This is the best preserved entire specimen. It is developed from the lower side, and shows the hypostoma, antennules, and a few fragmentary appendages of the cephalon, the outer portions of the exopodites of thorax and pygidium on both sides, and the endopodites on the left side.

The hypostoma is imperfectly preserved and is turned completely around, so that the anterior margin is directed backward, and the posterior one is so much in the shadow that it does not show well in any of the photographs. The form is, however, essentially like that of _Trinucleoides reussi_ (Barrande), the only other trinucleid of which the hypostoma is known, except that the border does not extend so far forward along the sides, and it is much smaller.

The antennules are not inserted close to the hypostoma, as in Triarthrus, but at some distance from it, and, as nearly as can be determined, directly beneath the antennal pits which are seen near the front of the glabella in many species of trinucleids.

[Ill.u.s.tration: Fig. 45.--_Cryptolithus tessellatus_ Green. Drawing of specimen 233, made by Professor Beecher. 9. Below are parts of two of the endopodites of specimen 236, showing the interarticular membranes. 41.]

The antennules are long, and are composed of far fewer and longer segments than those of Triarthrus. In this specimen they converge backward, cross each other and at the distal end are more or less intertwined.

As is shown in the drawing and photograph, very little can be learned from this individual about the other appendages of the cephalon. A few fragments of exopodites on either side suggest that these members pointed forward and were much like those in Triarthrus, but nothing conclusive is shown.

The exopodites and endopodites of the left side of the thorax are best preserved. The exopodites are above the endopodites, and only that portion exposed from the ventral side which projects beyond the line at which the endopodites bend backward. The endopodite on the left side of the first thoracic segment is the best preserved. It shows seven segments, the outer ones best. The c.o.xopodite is short and narrow, the basipodite somewhat heavier and longer, while the carpopodite and propodite are the widest and strongest segments. The propodite is triangular and flattened, like the segments on the middle and posterior part of the thorax of Triarthrus. At the inner end of the ischiopodite and meropodite are tufts of spines pointing inward and backward. These are not shown on any of the photographs, but may be seen with the light striking the specimen at the proper angle.

It is not possible to count the exact number of limbs, but one gets the impression that on the left side of this specimen there are twenty-one sets of appendages, six of which of course belong to the thorax. On the thorax and anterior part of the pygidium, successive endopodites show the propodites and dactylopodites becoming progressively more slender and shorter, while the ischiopodites, meropodites and carpopodites become shorter and more triangular, and with increasingly large numbers of short spines on their posterior borders. Back of the fourth endopodite on the pygidium it is not possible to make out the detail, but the appearance is of an endopodite consisting of short broad segments fringed at the back with short spines, the ones at the very posterior end appearing to be exceedingly short and rudimentary.

The exopodites are not so well shown as in some others but the setae are flattened and blade-shaped, and often bear numerous small spines.

_Measurements:_ Length (lacking most of the fringe), 10.5 mm. Width of thorax, 10.5 mm. Length of hypostome, 1.41 mm., width at front, 1.46 mm. The distance from back of fringe to end of antennules is 5.4 mm.

If straightened out, the left antennule would be about 6.1 mm. long.

In the first 3.1 mm., there are only ten segments, so that the average length of a segment is 0.31 mm. The distance from the inner end of the endobase of the first segment of the thorax to the outer end of the meropodite is 2.43 mm., and from that point to the end of the dactylopodite 2.47 mm. making the total length 4.90 mm. These measurements are taken from the photograph. Measurements taken from Professor Beecher's drawing, which was made with the camera-lucida, give a total length of 4.57 mm., the distance to the outer end of the meropodite being 2.3 mm. and thence to the tip of the dactylopodite 2.27 mm. Detailed measurements of the segments, on the photograph, are as follows: c.o.xopodite, 0.321 mm.; basipodite, 0.78 mm.; ischiopodite, 0.68 mm.; meropodite, 0.642 mm.; carpopodite, 0.642 mm.; propodite, 1.01 mm., dactylopodite, 0.825 mm.

Specimen No. 235 (pl. 7, fig. 2; pl. 8, fig. 3; pl. 9, figs. 1, 2).

Ill.u.s.trated: Amer. Jour. Sci., vol. 49, 1895, pl. 3, figs. 5, 6.

Specimens 235 and 236 were originally parts of an entire _Cryptolithus_, but, as Professor Beecher has explained, the specimen was cut in two longitudinally on the median line, and the halves transversely just back of the cephalon, so that each now represents one half of a thorax and pygidium. Both halves have been cleaned from both upper and lower side, a perfectly marvelous piece of work, for the thickness is no greater than that of a thin sheet of paper, and the soft shale of the matrix has a very slight cohesive power.

Both sides of specimen 235 were figured, but the dorsal side was apparently then somewhat less fully developed than at present. On plate 9 are two figures in which specimens 235 and 236 are brought together again, and both dorsal and ventral sides ill.u.s.trated.

On the dorsal side, specimen 235 shows portions of three exopodites which lie in a direction roughly parallel to the outer portions of the endopodites on the lower side, that is, their direction if projected would reach the axis in an acute angle back of the end of the pygidium. The setae stand at right angles to the shaft, and on a portion of it 0.5 mm. long there are seven of them. This is a fragment of an exopodite near the front of the thorax, and the setae, which are flattened, are about 1.63 mm. long.

On the ventral side this same specimen shows incomplete endopodites and exopodites of about seventeen segments, six of which would belong to the thorax and the remainder to the pygidium. The greater part of the appendages belonging to the pygidium are exceedingly small (about 0.15 mm. long) and so incompletely exposed that the structure can not be definitely made out.

The endopodites of the thoracic segments all lack the greater part of their proximal segments and are all of practically the same form. They turn abruptly backward at the outer end of the meropodite, and the carpopodite of each is greatly widened, projects inward and is armed with tufts of spines. The propodite and dactylopodite are wide, flattened, and taper but slightly outward, the dactylopodite bearing on its distal end a tuft of spines. On several of the endopodites, the meropodites are visible and they bear on their inner ends fringes of spines pointing inward. Behind these well preserved appendages the proximal segments of several endopodites are visible, and a regular succession of flattened, oval bodies armed with numerous forward-pointing spines. These latter bodies Professor Beecher took to be leaf-like exopodites, which they certainly resemble, and as they lie beyond the line of endopodites they probably do belong to the outer halves of the appendages.

The exopodites under the thorax are long, the shaft shows numerous short segments, and is in each case bent backward, though not through a right angle. They extend considerably beyond the endopodites. The setae do not diverge from the shaft at a right angle as on the dorsal side of this same specimen, but at an acute angle, indicating that they were not rigid. The individual hairs are broad and blade-shaped, frequently with a linear depression along the median line, perhaps due to collapse of the internal tube.

_Measurements:_ The greatest length of the fragment in its present state is 5 mm. The dactylopodite of the second endopodite (without terminal spines) is 0.18 mm. long, the propodite 0.23 mm. long and 0.15 mm. wide; the carpopodite is 0.24 mm. long and 0.38 mm. wide. All measurements were made on the photographs.

Specimen No. 236 (pl. 7, figs. 3-5; pl. 9, figs. 1, 2; text fig. 45).

The right half of the same thorax and pygidium as specimen No. 235.