Part 2

The interior surface of the hypostoma shows all the protuberances of the exterior surface (tubercular maculæ, spines and granulations) as shallow pits, the exterior grooves being on the contrary elevated ridges. For the rest the interior surface is smooth and its margins are posteriorly covered with a more or less broad duplicature, from the lateral margins of which the posterior wings project.

The manner in which LILJEVALL became aware of the interesting structural features of which I am going to give an account, is as follows. We were about to describe and delineate some new or not sufficiently known Upper Silurian trilobites from the island of Gotland. In order to draw a specimen of Bronteus polyactin ANGELIN, which is rarely found in the Wenlock and Ludlow strata of Gotland, LILJEVALL was preparing and cleaning its hypostoma. The shape of this hypostoma, as represented in pl. II fig. 22, is clypeiform, tapering towards the posterior margin, which is regularly tongue shaped. The anterior margin is faintly curved and at both sides elongated in a broad, pointed wing. Its exterior surface is divided in three transverse fields, defined through two shallow grooves, the anterior and largest field occupying more than the moiety, the median one is crescent-shaped and narrow, and the posterior one is nearly of the same size and confluent with the lateral borders of the hypostoma, which near the median transverse axis of the latter project in a blunt angle on each side. The surface, for the rest smooth, is covered by concentric, irregular terrace lines. On the inferior edge of the upper groove two small tubercles are situated, one on each side closer to the lateral margins than to the central axis. The size of their longitudinal axis amounts to 0,96 mm. in the shortest and to 1,07 mm. in the longest specimens. This axis is parallel with the groove and consequently oblique to the longitudinal axis of the hypostoma. They are oblong or ellipsoid, their inferior apices bluntly pointed or rounded, varying a little as seen in the three specimens figured, (Pl. II figs. 23, 24, 25). For about two thirds their surface is perfectly smooth or rather glossy, and the lower third is covered with a compact accumulation of small granules. The extension of this peculiar accumulation varies, being somewhat larger in some specimens, and the limit which runs oblique towards the glossy part is more or less curved. Such a granule taken single is of the extremest minuteness, measuring 0,055 millimeter at the highest. Their outline is perfectly circular and semiglobose.

When LILJEVALL saw this group of granules he was at once struck with the perfect, outward similarity between them and the facets of the compound eyes on the upper surface of the head of this same species, and I fully concurred in the same view. It now became of importance to ascertain whether the intimate structure also corresponded in both, and microscopic sections were accordingly prepared. Although not so clear as in other species of this genus a vertical section of a cephalic eye (Pl. II fig. 21) shows a median row of black spots which compared with other sections (Pl. II fig. 9) must be the centres of the badly preserved lenses. This is confirmed by the horizontal section (Pl. II fig. 20) where the well known image of the polyedric facets appeared. In the same manner the granulated spot of the hypostoma shows in a vertical section even more clearly (pl. II fig. 27) the elongate, coherent lenses with a black centre and the polyedric form of the lenses in a horizontal section (Pl. II fig. 26). The assumption of a pair of small adventive eyes on the exterior side of the hypostoma became thus based on the clear evidence of the perfect structural agreement between them and the eyes of the head. But it was necessary to strengthen this evidence through extended researches on the hypostoma of as many trilobite genera as possible and also to study the intimate structure of the cephalic eyes in as many genera as could be accessible.

We shall now give first a general account of the structure of the eyes in the trilobites as far as we have been able to study them, and then proceed to describe the maculæ of the various genera, which we have observed.

With regard to the presence or non existence of visual organs, we must remark that a considerably greater number of genera than those which unanimously have been regarded as blind, also are devoid of eyes though they still by many authors are ranked amongst the oculate ones and we then had better first to make a review of them and thus to eliminate them from the number of the oculate species.

=Blind Trilobites.=

In the chaos of generic forms and in the great disagreement which prevails as to the systematizing of the trilobites of the Cambrian time, there is a thorough revision of them highly needed by a person having access not only to the literature, but also to the original specimens. It is almost impossible in the present state of things to tell with any degree of certainty how many well established genera had been living during that period. Hence the difficulty of fixing the systematic names of many specimens the visual organs of which are to be described.

My researches on the visual organs of the Cambrian trilobites are founded on the Angelinian Collection in the Swedish State Museum together with collections of foreign species, but also largely on the waste European and American literature, though we have to deplore the often occurring inexactitude of the figures, especially in the older works, and constructed or schematized figures in some of the newer ones, which give a quite false notion of the structure. There is no lack of figures to show how it ought to be, according to preconceived notions and, on the other hand, a great scarcity of representations, to show how it really is. In spite of all this there is a sufficiently great number of well established facts to demonstrate the organization of the Cambrian genera.

The trilobites of this division may be called blind only in so far as they have no eyes on the upper surface of the head, but they may have been provided with visual organs, though more imperfect, on the hypostoma as really seems to have been the case with some of them.

According to the structure of the head shield the blind trilobites may be subdivided into the following well characterized groups. These are:

I. Without facial ridge: 1) The Archæan Trilobites.

II. With facial ridge: 1) The Olenellidæ. 2) The Olenidæ and related.

=I. Blind trilobites without facial ridge (= »eye lobe»).=

=Group 1. The Archæan Trilobites.=

In these the head shield is in one piece without any facial suture and facial ridge, and without the least trace of anything that might be called a visual organ and they must consequently be considered as totally blind. In contradistinction to the following groups, excepting the oldest Olenellidæ, the head consists of only three parts, 1) glabella, and 2-3) the two fixed cheeks. These genera range from the oldest zone in which hitherto trilobites have been found, that of Olenellus (Holmia) Kjerulfi, to the zone of Paradoxides Forchhammeri, and some, as Agnostus, even continue as high in the Lower Silurian series as in the Brachiopod schists. Beside Agnostus the other genera are Conocoryphe (seven species in the Swedish Cambrium), Toxotis, Ctenocephalus (?), Elyx, Aneuacanthus, Conophrys and Microdiscus.

»Harpides» breviceps ANG., also belongs here. Anopocare of ANGELIN should also be regarded as one of this group. But it cannot be retained any longer because it is founded on two other, well known forms, being, according to LINNARSSON, Peltura scarabæoides (pl. 27 fig. 1, a in Pal. Scand.) and young specimens of Sphærophthalmus alatus (ibid. figs. 1 & 2).

It is remarkable that some of the Conocoryphidæ have an imperfect facial ridge, to be compared with the commenced one in Sao BARR. I (pl. 7 fig. 9). So the American Con. trilineata and reticulate. WALCOTT U. S. Geol. Survey 10th Rep. pt. I, pl. XCV f. 5 & 6. It is, as it were, arrested in the development and these adult trilobites had stopped, where the larva of Sao was proceeding in its second stage. They are the forerunners of the blind trilobites with facial suture, belonging to the third group. It needs scarcely be mentioned that the genera now enumerated have hardly anything in common, beside the general character of the head, and that real affinity exists only between Agnostus and Microdiscus, and probably also between Conocoryphe, Ctenocephalus and Elyx.

BEECHER asserts that there is a suture in Agnostus, but in vain we have searched for it in numerous well preserved specimens and Dr HOLM also denies its presence. Nor are there any signs of closed up sutures, which also could not possibly be expected in so early a stage of evolution. It may then be taken as well settled that a fundamental character in these the oldest of all known trilobites is the total want of a facial suture and a compactness of the whole head shield which later is broken up in several parts through the disjunction of the free cheeks. In the Lower Silurian formation there are a few genera sharing in the same structure of the head shield, though by no means else related. Such are Dindymene, Areia, Carmon and in the U. Silurian Cromus. The two species forming BARRANDE'S genus Dindymene are so dissimilar that Dind. Friderici Augusti had better to be removed to a new genus and the first described one to be retained as type of the genus Dindymene. The same is the case with Carmon, where the type species C. mutilus is blind and without free cheeks while the other species, known only by its fixed cheeks and glabella is one of the Olenidæ.

=II. Blind trilobites with facial ridge.=

This large division embraces the second and third groups or, with a few exceptions, all the rest of the Cambrian trilobites on account of a feature in the cephalic sculpture common to them all, though widely different as to its first origin in both. What forms the prominent and common characteristic of these two groups is the presence of the facial ridge, which emanates from the basis or the front of the foremost segment of the glabella and in a great variety of different shapes continues backwards near to the posterior border of the head. It has received several names as eye-line, palpebral lobe, ocular ridge, eye-lobe, ocular fillet (MATTHEW). In German it is named Augen-leiste, in French filet (BARRANDE) and in Swedish ögonlist. Some authors make a difference between the more narrow part, calling it eye line, and the thicker posterior node, which they name the palpebral lobe proper.

As this peculiar ridge exists before any facial suture has made its appearance and separated the head shield in five parts, viz. the median glabellar part, the two fixed cheeks and the two free cheeks, and as it occurs in genera which never possessed any facial suture, and where no eye ever was formed, it is not adequate to call it an ocular ridge etc. the more so, as it, at least during a long series of genera succeeding each other, has had no connection whatever with any eye. I therefore propose to call it facial ridge (in swedish faciallist). It occurs on the head of almost all Cambrian trilobites, excepting the archaic ones, and it is retained in the later Cambrian Peltura, Sphærophthalmus etc., which have real, compound eyes, as well as in a few Lower Silurian genera as Triarthrus, Pliomera, Euloma, in the Upper Silurian Arethusina and Acidaspis and most persisting in Harpes, ranging from the Lower Silurian Lower Red Orthoceratite Limestone into the middle Devonian beds.

It presents itself in the most variable shapes, and as it in fact can be followed through its development in the oldest known species, it is suitable to begin its description together with the characteristics peculiar to the oldest or second group in this large subdivision, that of the Olenellidæ.

=Group 1. The Olenellidæ.=

In this group we have two families of different age, the older, less developed the Olenellidæ proper, and the younger the Paradoxidæ. The former consists of the genera Olenellus, Holmia, Mesonacis and Schmidtia, and we shall attend to them first. They have no facial suture and consequently a tripartite head shield like the Archaic ones, no eyes, but there is that strongly developed and most characteristic facial ridge. As seen in Olenellus Thompsoni the crescent like ridge starts as a direct outflow from the base of the first segment of the glabella, and is in direct continuation with it as an integral part. It is roundbacked, regularly curved and at its starting point as broad as one of the posterior glabellar segments. It is regularly semilunate, tapers posteriorly and ends near the occipital segment. From the narrow second segment of the glabella, inside the just described larger ridge, a smaller ridge emanates, broader and flatter than the former, slightly curved, and ends between the third and fourth glabellar segment. Already in Holmia Kjerulfi the ridge is modified. There it is only one ridge, the anterior one, nearly as doubled through a shallow groove running along its back. The second one has dwindled away so as to be seen only as a narrow stripe near the occipital segment and ending outside this in a point. The anterior ridge, represented by HOLM as consisting of two nearly independent parts, is indeed in one piece, though its dorsal groove sometimes is deep, and it is with its total breadth joint to the first glabellar segment. Along its outer edge, where it lies close to the cheek, a narrow slit runs, and I suppose that it is to be considered as the first indication of the forming of a facial suture, which however does not reach longer than the ridge. As in Paradoxides it is so tight to the cheek, that there has been no place for an eye. It is no accidental break, its edges being too regular and unbroken in all specimens. This ridge is by far much shorter than in Olenellus and terminates opposite the third segment.

1. Hypothetical figure, based on fig. 2. represents a stage preceding the next. There are three pair of pleura and three segments of the rhachis.

2. Copy of WALCOTT'S fig. 5 in »Bullet. 1886», pl. XVII. Three pleura at right, two at left, the larger formed through fusion of the two hindmost in fig. 1.

3. Copy of WALCOTT'S fig. 6: the second and third pleura have been united into one.

4. Hypothetical figure as a further development, following upon the stage represented in fig. 3. Fusion completed between all posterior plectra (2, 3, 4), thus forming a single large pleuron composed of the three mentioned.