On the maculæ of the hypostoma.
We shall now turn our attention to the visual organs which Liljevall discovered on the hypostoma of Bronteus. In doing this, we may bear in mind, that the genera in which we really have found lenses on the maculæ are relatively few, but that we shall review the maculæ in all genera, which we have been able to examine, and try to show that even most of these may, although in an inferior degree, have acted as visual organs. At the same time some more details shall be given about the cephalic eyes for comparing them with the hypostomic ones.
Acidaspis Murch.
The hypostoma is of a peculiar type, deviating from that of the other genera, squarish or rectangularly transverse, entirely without terrace lines and no maculæ proper. Barrande has given no less than nine different samples of these hypostomas.
Acidaspis crenata Emmr.
There are certainly no maculæ of the same sort as in so many other trilobites and the interior side of the hypostoma does not bear the least traces of impressions which might be taken as the reverse of the maculæ and still more less as muscular impressions. In fact, the total absence of such in Acidaspis militates against the interpretation of the macula impressions as muscular scars. When the maculæ fail, also the so called muscular impressions fail. Moreover we are able to see still clearer in this matter through what we know about the structure and position of the extremely similar hypostoma of Apus, of which we have given a description in the end of this memoir. This skeletal part of Apus is without any connection with the surroundings, excepting at its anterior margin, and is consequently movable in a direction outwards and upwards. Along that anterior margin it is fixed to the outside of the ventricle through three pair of muscles, three muscles on each side of the margin. Zaddach De Apodis cancriformis anatome (pl. II fig. XIV p. 68).
For the rest the marks of the attachment of the muscles are as a rule in the Crustacea, at least those of the head, elevated small platforms, so in the Trilobites, of which I have excellent specimens in a Bumastus and others. On the inside of the head of Limulus they are faintly elevated patches. The small hollows on the inside of the hypostoma formed by the maculæ are indeed the sockets in which the soft parts of these more or less developed hypostomic eyes were sheltered. But there is still a feature in the hypostoma of Acidaspis which merits our attention and which perhaps may have a significance akin to that of the maculæ.
The hypostoma is square, with two short pointed wings, one on each side of the slightly bent anterior margin and likewise two smaller ones at the corners of the posterior margin. A groove follows on a short distance the lateral margins and the posterior margin and disappears a little below the anterior one. In the same direction, distally, though a little more inside and unconnected with them there are two small grooves, the bottom of which consists of a shell substance of different colour and structure than the other parts. Having been a little ground and seen in transmitted light it exhibits the shape of a club and a homogenous yellow spot, tapering posteriorly and swelling out distally (Pl. I fig. 4). It must be left an open question whether these maculæ share in the nature of visual organs as the quite different maculæ of the other trilobites, but it may be possible that it is so. It must, however, be remarked that there are two types of hypostoma in the genus Acidaspis as shown by the illustrations of Barrande. One has the small grooves, possibly all sheltering the claviform maculæ, disposed as in the now described A. crenata. This group embraces five species of the Bohemian Silurian formation. The other group of three species again has the hypostoma of the same quadratic or rectangular shape, but the two short grooves, which may be expected to contain the maculæ, are placed midways between the anterior and the posterior margins, nearly as the maculæ bearing grooves of other trilobites. We have however not had material for pursuing our researches in this genus, the other species of the Swedish Acidaspidæ being unknown as to their hypostoma.
It may here be added an observation concerning the ornamentation of the exterior surface of the hypostoma of Ac. crenata. It is covered by a great number of diminutive circular or oblong wartlets occupying the whole surface excepting the lower third of the central field just above the posterior groove which is smooth. These wartlets seen through transmitted light (Pl. I fig. 5) show in their interior something like a peculiar black spiculum rising from a bifid rootlet and confined within the wartlet and not extruding from it. In a longitudinal section (Pl. I fig. 6) the spicula perforate the wartlets reaching through their whole length. As in Calymmene these interior pseudo-spicula are tubes, filled with iron-pyrites. It is probable that these tubes were once bearing setæ and quite as in Apus formed a fur of bristles.
The structure of the cephalic eyes (Pl. 1 f. 1-2) is prismatic, but the separate prisms are rather short and broad. Their lower or interior end is convex. The separating lines between the single prisms are not always distinct.
Agnostus Dalm.
Agnostus glandiformis Ang. Although there is not the slightest evidence of eyes in this the largest of its genus, nor any free hypostoma hitherto has been found, we may here give a little account of our researches into this species. The scantiness of material forms a chief obstacle to our knowledge. Only three entire, rolled up specimens have been found, and it is by sectioning and preparing such that any hope can be entertained to gain reliable results.
A rolled up specimen from Andrarum Scania was sectioned lengthwise. The tail-piece closed tight against the head-shield, so there had been little chance for foreign matter to penetrate into the interior which, however, is filled. Close below the cephalic shield there is a remarkable structure, mostly resembling an elongated intestine with swellings joined by more narrow ducts and anteriorly the coherence is interrupted. As in crustaceans and Arthropods in general the stomach and the intestine are situated on the dorsal side of the body, there is nothing unlikely in assuming that this in reality may be the remnants of the intestine. This may also be compared with the observation made by Vollborth in his memoir »Ueber die mit glatten Rumpfgliedern versehenen Trilobiten», 1863, p. 46, tab. 1 fig. 12 where the heart-tube probably is delineated. Barrande has also given figures of what he considers as the intestine in Trinucleus.
Below this organ in Agnostus, there lies a section of a vaulted calcareous plate with its convexity turned against the dorsal side of the head shield, that is to say quite the reverse what might have been expected if it had been the hypostoma in its true position. This may, however, not be any objection against considering it as a sectioned hypostoma, loosened from its connection with the cephalic shield, disturbed in its original position and turned round, when the shell became filled with mud. At the distal end there is a much distinct duplicature.
Asaphus Brongn.
The wellknown hypostoma of this genus has the same characteristic shape, though more pronounced, as in Ptychopyge, with its posterior margin deeply indented, so as to form two large, pointed lobes. There is properly only one median field, surrounded by flat, lateral borders, continuing down into the posterior lobes, from which it is separated through a shallow groove. The macula are situated in this groove, on each side of the inferior border of the central field. They are more or less prominent, but whatever their form may be, their surface is always entirely smooth, lying well circumscribed amidst the surrounding terrace lines.
The following list enumerates all the species in this genus, of which previous authors have delineated the tubercles, though they in the descriptions only in very few instances have mentioned their presence.
A. acuminatus Nieczkowski, »Zusätze zur Monogr. der Trilobiten 1859», tab. I, f. 6. Maculæ most prominent, but no mention made of them in the description.
A. (Isotelus) canalis J. Hall, Pal. N. Y. I, pl. 4 bis, f. 18-19. Though this and a following species probably on account of the deviating conformation of their body, belong to a different genus or subgenus, I mention them along with the Asaphi, as there is the greatest similarity in their hypostoma. Fragment of an interior cast; still more strange is the fragment showing part of the interior surface and the duplicature.
A. expansus L. Br. I pl. VII f. 3, BR. II pl. I f. 2, 2 c. In the two first figures there are no tubercles nor maculæ marked, in the last figure there are distinct tubercles.
A. fallax A. pl. XXVIII f. 3, c. BR. II pl. I, f. 3, both nearly congruent.
A. (Isotelus) gigas J. Hall, Pal. N. Y. 1, pl. 60, f. 7 g, pl. 66 f. 5, the inside of an entire, uncommonly large hypostoma gives a general good view with the macula; which are large and evident. He mentions the maculæ as »two circular spots». »These probably indicate the points for the attachment of muscles and tendons upon the inside,» he adds.
A. ingens Barr. Novák II, pl. I, f. 7, a good figure showing two semiglobular maculæ. Barrande's figures pl. 33, f. 7, 8 are not distinct.
A. ludibundus Tqt. Br. II, pl. 1, f. 7. Two semilunar sulci on the cast of the interior surface, being only the posterior borders of the maculæ which are a little more raised and distinct than the rest.
A. nobilis Barr. pl. 32, f. 6. Indistinct traces of maculæ. In the figure 6, pl. 31 representing the hypostoma of a young specimen, there are no tubercles at all. If the figures were to be relied upon, it might be assumed, that the tubercles appear at a more mature age.
A. Powisii Salter, pl. 23, f. 6. Two semilunar, narrow tubercular maculæ, their interior apices converging towards the anterior border of the hypostoma.
A. raniceps A., pl. XXVIII, f. 2 c., BR. II, pl. 1, f. 4. A good figure.
A. raniceps var. maxima Br. II, pl. 1, f. 6. Semilunar impressions on a cast and thus far incomplete.
A. striatus Boeck. To this belongs probably A. expansus M. Sars »Ueber einige neue oder unvollständig bekannte Trilobiten» in Isis 1835 p. 333, and especially p. 340 and the following (Bemerkungen über die untere Seite von einigen Trilobiten). Sars there gives a very good description of the exterior side, where he also mentions the two maculæ, pl. IX, f. 9 a, b. (»2 kleine Knoten»), Br. I, pl. VIII, f. 4 a, and Br. II, pl. I, f. 9. These latter figures differ in so far, that the former has the maculæ excavated or rather tubular below, in the latter again they are regular.
A. trinucleorum Br. II, pl. I, f. 16 a large specimen with linear maculæ arranged rectangularly in respect to the longitudinal axis of the hypostoma and both on the same level.
A. tyrannus Murch. Salter in Mem. Geol. Survey, Brit. Foss. Dec. II, 1849 pl. V, f. 4, excellent figure. On p. 2 he says: »there is an oval circumscribed tubercle at the origin of each (fork) most distinct on the inner surface». This figure is again reproduced in the »Monograph», pl. 22, f. 6, where it is said »Two linear tubercles with their interior apices converging towards the posterior margin of the hypostoma».
Asaphus sp., probably A. raniceps, Pompeckj, Trilobitenfauna Ost- and West-Preussens, taf. VI, f. 7. p. 80 »an seinem Hinterrande liegen zwei kleine, flache Höckerchen, die als Reste des Hinterlappens aufzufassen sind»(!?).
We shall now describe the Asaphi which we have been able to examine more in detail.
Asaphus expansus L.
In the numerous specimens of which we have examined the hypostoma, there are always small, nearly circular maculæ which by their lighter colour are marked out from the surrounding smooth space of the inferior sinus of the lateral grooves where they are situated. They do not rise at all above the surrounding surface from which they are separated only by a fine, scarcely perceptible line. On the interior surface of the hypostoma they are better seen, and the enclosing line is deeper and more distinct. Their surface is there quite as smooth and even as on the exterior side of the hypostoma. They do not consequently in the least manner resemble muscular impressions as observable on the inside of the glabella of other trilobites. These are on the contrary elevated above the surface and finely striated in various ways.
The peculiar nature of these macula is revealed through the various sections we have made. In pl. I fig. 15 a vertical section across the entire hypostoma is represented. The sectioned, dark maculæ (a) lie on both sides of the faintly curved central field, in the sinus of the lateral grooves, and the strongly developed terrace lines continue sidewards just a little on the interior side, where the duplicature turns round. As seen in a magnified vertical section (fig. 16) the maculæ consist of horizontal, whitish, straight lines, probably lines of successive growth, and these are crossed rectangularly by more irregular whitish lines separated from each other through dark spaces. The whole thus gains the aspect of a dark surface cancellated by white lines. This reticulated or spongious macula is enclosed as to its superior as well as to its inferior part in the compact and homogenous test of the hypostoma and occluded from the influence of the light. Its value as a visual organ consequently is insignificant. In a horizontal section (fig. 17) the structure is still more bewildering. There the whole macula is a confused, spongious white mass with dark spaces between the white meshes, and only at one side, the left one, some obscure indications as of polygones are visible. It would indeed have been impossible to interprete what this means, had not the study of the cephalic eye given a clue thereof.
The eye of this species is covered with a delicate perfectly smooth and glossy integument (f. 12) which is reposing immediately on the prismatic lenses of the cornea. In a few instances, depending on the colour or the state of preservation of that integument the lenses are indistinctly translucent. Owing to their state of preservation their aspect is greatly variable. They are in many instances, as seen in fig. 9, six-sided or some rhombic, foursided or even quadratic (f. 8, 10). They are all of the same length, 0,2 mm. along the surface proper of the eye, but are lengthened to 0,3 mm. towards the border of the eyes, f. 12, which will be described further on; they are of equal breadth, amounting to 0,066 mm. Their inferior ends are slightly convex or nearly plane when well preserved, else, when as often is the case, disintegrated as to be scooped out and vaulted. Their interior structure, as revealed by sections, shows a cylindrical core, f. 8-10, composed of concentric strata. This cylinder fills nearly the whole interior space of the prisms, there being, however, in many instances a compact dark mass between the cylinder and the walls of the prisms. There are also sections in which the prisms are filled with a uniformly black mass without any concentric structure. In longitudinal sections the outlines of the individual prisms are not clearly discernible (f. 11). There are longitudinal, white lines of varying thickness with lateral irregular offshoots, which may join with those from opposite walls, and give the interior a sort of spongious or cancellate appearance.
Towards both sides of the ocular surface, towards the superior and inferior side, a change sets in as to the shape of the lenses, as best seen in horizontal sections (figs. 8, 10) they are lengthened and become more and more indistinct, and at last in the upper and lower marginal zones pass over into a reticulate, spongious mass, which seen in a longitudinal section presents almost the same aspect as in the regular prismatic surface of the eye. Fig. 11 b, a being the visual field. It is, however, more densely reticulate, but a prismatic arrangement is quite as much evident as in the ocular surface proper. In the rule the passage from the prismatic surface to the reticulate is gradual, the prisms becoming by and by irregular in their outline and diminishing in size (fig. 12, a the eye proper, b the border zone), but there are also instances where the distinction between these two fields is sharp and without any gradual transition. In the inferior reticulate zone there are generally some oblongue, funnelshaped pits. I am uncertain whether they are to be regarded as regular parts of the eye structure or rather as burrows of some parasite. They do not continue deep down.
If we now compare the reticulate zone of the eye with the maculæ of the hypostoma, for instance the vertical section fig. 16 with fig. 11 b, we find the most complete identity in structure. In the same manner the horizontal section of the border zone of the eye, fig. 8 a, fig. 10 b, and of the macula, fig. 17 are similar. The same chaotic, spongious mass in both, with some tendency to form prisms more evident in the longitudinal sections, where the same reticulate structure with predominant white, longitudinal streaks is so palpable. That there is a complete identity in structure between the two, the macula of the hypostoma and the border zones of the eye, is as evident as anything can be, but as to the functional identity or what this function may have been it is difficult to decide anything with certainty, at least it seems to me that the capacity of vision must in both have been far more restricted than in the eye proper. They rather give the impression as of rudimentary visual organs.
Asaphus raniceps Dalm.
The maculæ of the hypostoma are placed obliquely in the saline manner as in the preceding species. They are more prominent and the oblong macula, with the longest diameter of 1,2 mm. is on the top of a little mound and surrounded by a fine, elevated marginal line (fig. 23). Its somewhat convex surface is entirely smooth and if sectioned horizontally exhibits the same sort of spongy texture as A. expansus (pl. I f. 24). In a vertical section (fig. 25) the macula does not occupy so large a space as in As. expansus, but rather lies as a lenticular disk in the hypostomic test closer to the superior surface than the inferior. With sufficiently high power the same sort of pillars, divided by horizontal strata is seen. Upon the whole the vertical section is not so clearly developed. The reticulate zone of the eye is more definitely separated front the prismatic zone than in A. expansus. There can be no doubt that there is a correspondence in this species between the structure of the macula! and the spongious zone of the eyes.
Asaphus cornigerus Schloth. (A. Kowalewskyi Lawrow).
Of this strange species with its enormous eyestalks, more than 2 centimeters in length I have through the kindness of Akademiker Friedr. Schmidt in St. Petersburg had occasion to study some specimens.
The rather large maculæ, obovate-circular, are like those of several other Esthonian species oriented inwards and upwards instead of inwards and downwards as in the Swedish species described. As the specimens have suffered through corrosion of the surface there are only faint traces of a marginal line. Their microscopic structure is badly preserved, and they look pale and transparent with only few indications of the spongious texture. The lenses of the cephalic eye are nearly square prisms sometimes with a slight approach to hexaedral pillars. There is no clear transition into a spongious or reticulate border zone. But this may depend upon the bad preservation.
Asaphus fallax Dalm.?
It is very difficult to distinguish this species or rather variety from A. expansus, but if we have found genuine specimens, there are some points in the shape of eyes and maculæ which make it different. The hypostoma (f. 21) is rather more broad and the duplicature is large with an upturned margin. The little macula is placed on the top of a smooth rounded elevation and enclosed by an elevated rim.
The cephalic eves are rather short, regular hexaedral prisms (f. 18-20) and as shown in fig. 18 change into irregular squares near the border of the eve.
Asaphus sp.
from Brunsby kanal, Segerstad parish, isle of Öland. The maculæ are large pale whitish and of an uncommonly fine-meshed reticulation, well limited from the surrounding hypostoma.
Asaphus sp.
from the islet of Sandö, north of Gotland. Although we cannot give any account of its hypostoma, not having had sufficient material, the structure of the eyes is so peculiar that it seems worthy of being recorded. The integument is extremely thin and transparent and the subjacent lenses are clearly seen, and through their impact on the thin integument they make this to stand out in a very low relief above them (fig. 27). These lenses are uncommonly short, forming at the surface rather oblong, slightly hexaedral prisms with a narrow interspace between them. In a section lower down they have the shape of hollow, white rings filled with black mud and in a longitudinal section the white walls of the lenses look like short pointed spikes and interiorly they are completely empty. We here find also the same gradual change from regular cones to the spongious border zone as in the above mentioned species f. 29. The border zone is finely reticulated.
Asaphus (Isotelus) gigas J. Hall.
The enormous hypostoma of this giant resembles in a high degree that of the Asaphi, but is at the same time the most evident verification of the experience that the hypostoma per se cannot be regarded as the sole criterion for determinating the generic affinities of different species. The whole structure of this trilobite in other respects gives it a quite independent position, distinct, from Asaphus.
We have had at our disposal several specimens of the large hypostoma, the dimensions of the largest, fragmentary in its anterior margin, being as follows: breadth 48 mm. length 41 mm., probably 47 mm. when entire, breadth of each of the posterior lobes at their bases 19 mm., length of the same 23 mm. The maculæ which are placed on the flat surface of the hypostoma without being at all elevated, are prominent through their great size and their pale, whitish colour. They are somewhat oblongue having their longest axis directed inwards and downwards. They attain a diameter of four millimeters. Horizontally sectioned, f. 2, they show the spongious texture and vertically the quasi prismatic reticulate structure richly developed, f. 3. The pillars are very distinct and continue without interruption all through the macula and in the interstices there are traverses joining between two pillars or partially filling the darkish interspaces, thus giving the whole the aspect of some »tabulate» coral sectioned. By the inspection of the horizontal section alone, presenting the irregular spongy texture it would have been impossible to imagine the ordinated arrangement which the vertical section reveals to us. The eyes consist of regular hexaedral prisms, and there is a very sharp boundary line between them and the spongious zone, which is very narrow and without distinct separating lines joins with the cephalic test. For the rest there are in all probability at least two different species sent from America under the name of Isotelus gigas, of which only the largest, almost like a Homalonotus, and the hypostoma of which has been partly delineated in this memoir, seems to be the real one. They differ both in the shape of the hypostoma as well as in other respects.
Barrandia Mac Coy.
According to Salter Monogr. pl. 19 fig. 9 his B. Portlocki, of which there is a fragmentary and broken hypostoma, shows feeble traces of oblique macula, nearly in the fashion of Asaphus. In the description at page 139 is told about »the usual pair of tubercles».
Bronteus Goldfuss.
Barrande delineates the maculæ on the hypostoma of
B. palifer, Tab. 45 fig. 17,
B. planus, Tab. 48 fig. 7, and
in the supplementar volume of
B. rhinoceros, Tab. 9 f. 16,
B. furcifer, Tab. 11 fig. 16, but there is not the slightest indication of granulation on any of them, nor is there in the descriptions, generic or specific, the least mention made of the tubercular maculæ. It is also remarkable that in the works of Angelin and Novák where several hypostomas belonging to species of this genus are delineated, not a single one shows these tubercles. We shall now continue the descriptions of the Swedish Brontei, already begun with Br. polyactin in the introductory part of this memoir.
Bronteus irradians Lindstr.
has a hypostoma that much resembles that of Br. polyactin. In its general shape it is similar and the two concentric grooves with the two maculæ placed in the same way, just below the superior groove. These maculæ are much larger than in Br. polyactin, nearly thrice their size. They are also more ovate or rather like a bean, the smooth surface is larger and the granulated spot restricted to a more narrow space forming an oblique patch. The granules or lenses are also individually larger than in the allied species, double their size or 0,06 millimeters. We have not succeeded in making sections of the cephalic eye nor of the maculæ.
Br. platyactin Angelin.
The hypostoma has a transversally triangular form, and is divided only in two fields through a shallow semicircular groove near the posterior margin. The two maculæ are situated above the groove near its superior sinuses. They are elongated, fig. 17, elliptic with the narrow pointed end directed outwards and the broad rounded end inwards. The chief surface is scooped out as a shallow depression. The granulated spot is situated on the broader end and covering it completely. The relatively large lenses are arranged in five regular rows, the uppermost one being the longest. On the interior surface of the hypostoma there are the corresponding sockets of both maculæ with smooth surface. The horizontal sections of the granules figs. 18, 19 present the image of white rings in close contact, without, however, to occasion a prismatic structure, a dark interspace lying between each ring. These lenses are filled with a dark mass, and in some the same sort of radiated structure is perceptible as in the lenses of the cephalic eye. In horizontal sections of the cephalic eye the lenses approach the polyedral shape. In another section near to the surface of another specimen the lenses are decidedly hexaedral. The vertical sections, fig. 14, reveal their real nature as lenses where they lie as a string of beads with a dark nucleus enclosed within a thin whitish shell. They are covered by a thin membranous lining. When seen in transmitted light the lenses proper are dark, and the shell white and in reflected light the lenses are lighter than the rest.
Br. laticauda Angel.
Lower Silurian from Dalecarlia. The hypostoma, f. 10, is of a broad clypeate shape, the anterior margin rounded without any large projecting wings. On the exterior surface there are two grooves parallel with the rounded inferior margin. As in Br. polyactin the two elliptic tubercles are situated on the inferior edge or slope of the upper groove and they are deepened by a shallow depression as in Br. platyactin. The granulated spot situated along the posterior margin of the macula deviates much in shape from that of the other species. It consists of a long and narrow stripe ending in a fine point outwards and widening inwards, where it is rounded, forming thus a claviform, curved elevation. Around the granulated area the maculæ are quite smooth, and show in a horizontal section an irregular structure of tiny black dots nearly resembling the structure of the hypostomic shell. We have not been able to obtain any good vertical section, but by casts of the interior side of the tubercles it is found that the granules form polyedric facets like those of the eyes though perhaps not so regular. There is a specimen with the granules intact, and the polyedric shape is then not so distinct.
Sections of the eyes elucidate the structure, which partially is obscure in other species of Bronteus. There is a thin membranous coating covering the subjacent well formed lenses (fig. 8). When this membrane in some instances has been peeled off the lenses lie bare (fig. 9). These have a dark nucleus and in some instances it seems as if there were two. In a vertical section they look spheroidal, in a horizontal section again they are polyedric, especially when taken somewhat below the surface or near the middle line (fig. 6). In fig. 7, some lenses are delineated in a horizontal section, highly magnified and the corallian appearance is evident. The lenses are well separated by distinct lines, and from their inner tubes whitish reticulations radiate towards the centre.
If we now compare these sections, where the lenses are so distinctly seen, with the more obscure sections of other Brontei, it is evident that the dark points in them are nothing but transformed and deteriorated centres of the lenses. In Br. platyactin they are a little more distinct than in the other species. And it is with these that the granulated spots on the maculæ of the hypostoma are most concordant.
Bronteus sp. indet.
This species, of which only the hypostoma is known, found at Lansa, Gotland, is likely, to judge by that, to be nearly related to Br. polyactin. The shape of this hypostoma is exactly the same and the maculæ are placed in the same position, but they are larger, a little sunk on the blind surface and the group of the lenses is different. Its superior margin forms an ingoing arch and the lenses themselves are not convex, but rounded or slightly polyedric, and separated through thick interspaces.
Bumastus Murchison.
The only recorded hypostoma with maculæ belongs to B. insignis Hall as described by Salter Monogr. p. 208 pl. 27 fig. 7. He says: »A pair of compressed tubercles occurs at the lower third: they are transverse-ovate, and more than their own diameter apart.» This is nearly in concordance with what is seen on the species to be described below.
Although it can in reality be said that the species of Bumastus are the most common of all trilobites in the Silurium of Gotland, in so far that their head pieces and pygidia are met with everywhere in the limestone rocks of the island, the find of entire specimens or of detached hypostomas is amongst the rarest events when collecting there and amongst thousands of fragments, in several places entirely filling large portions of the limestone rock, there have in the Swedish State museum been acquired only three or four hypostomas, which we are now going to describe.
At first we must point out a certain resemblance which prevails between the hypostomas and maculæ in Bumastus and Bronteus. Compare for instance Bum. sulcatus with Br. platyactin and Br. laticauda. The general form is nearly the same in both and the shape of the maculæ almost identical but for the complete want of granulations on those of Bumastus. To this resemblance must be added that there are, as known, forms of trilobites which by some authors have been regarded as Brontei and by others, again, ranked amongst the Illænidæ.
Three species of Bumastus have been recorded as found in the Gotland strata, but of one of these no hypostoma is known. Besides, to judge by the head shields and hypostoma, there are a few new species. To begin with the most common
B. sulcatus Ldm.
The hypostoma is broadly shield-formed and the anterior margin is most characteristic being in its central part elongated in a short evenly rounded projection. The side wings are triangular acuminate, bent a little backwards and towards the interior surface. There is only a single rounded field passing into an evenly bent border, in front of which the projecting, elliptic maculæ lie. A second pair of wings stretches backwards near the posterior margin. We have not obtained any sections of the maculæ.
The cephalic eyes figs. 36-38 are covered with an uncommonly thick stratum of homogenous shell, one and a half times as thick as the subjacent lenses. These are comparatively broad, above passing indistinctly in the covering stratum, below, at their basis, convex. In a horizontal section figs. 33, 34 they exhibit the common hexaedral or pentaedric aspect, their exterior sides being white and confluent without any separating lines. Their interior is dark and from the polygonal sides white lines go in giving the whole the appearance as of a starry, composite coral with its septa. The correspondence in structure with Bronteus laticauda pl. II fig. 7 is striking. A narrow space separates the surface of the eye from a nearly similar patch below the eve on the cheek 35 b, 38 b. But there the position of the composing strata is singularly reversed and the spongious or pseudo-prismatic stratum lies close to the outside and the homogenous stratum on the inside. The white longitudinal lines are well marked out and regular in the vertical section, fig. 38 b, indicating the prismatic structure most clearly.
B. barriensis Murch.
The hypostoma fig. 31 is of so transverse a shape that the breadth much surpasses the height, the former being 6 millim., while the latter attains only 3,5 millim. The anterior margin is so much arcuated and sloping backwards that it forms an obtuse angle. The side wings are triangular and acuminated and from them a thick pad runs backwards and forms the posterior margin. Below the largest field of the exterior surface the maculæ rest on a ridge having a much elongated, vermiform shape and sunk in the surface (fig. 32) a little more than three times longer than broad. In its shape it approaches much to the hypostoma of Bum. insignis Hall as delineated by Salter pl. 27 fig. 7.
In another, undetermined Bumastus (pl. II figs. 47, 48)[40] the hypostoma (from Klints, Othem, isle of Gotland) is of a more elongate obovate form, the anterior margin elevated, rounded, the terrace lines faint, the tubercles are larger than in the former Bumasti, oblong, curved, beanshaped, evenly rounded and smooth.
[40] Probably the same as Bum. sulcatus pl. XII fig. 12 in my paper on the Trilobites of Gotland 1885.
Another of the new species is one that Liljevall has found at Korpklint near Wisby (Pl. II figs. 41-46). The hypostoma is more elongate than transverse with the anterior margin regularly arcuated, by far not so prominent as in B. sulcatus and the former. By a shallow groove the exterior surface is divided in two fields, the anterior one the largest and covered with a few, distantiated, concentric terrace lines. The posterior field is smooth, nearly even and the maculæ lie just in the groove, with their axis oblique to the longitudinal axis of the hypostoma, beanshaped and smooth. The cephalic eyes show in a horizontal section hexaedral white walls enclosing a dark space in which some indistinct radii emanate, fig. 41 a. In the vertical sections the prisms are narrow and elongate and the integument by far not so thick as in Bumastus sulcatus, fig. 42, 43. The border zone of the eye in this (fig. 41) as well as in B. sulcatus, as seen in horizontal sections, agrees with that of the Asaphidæ.
Bumastus sp. inlet.
In a detached piece of limestone, found in Gotland, at Norderstrand, near Wisby, two hypostomas and a free cheek lay embedded along with a coral of the genus Acantholithus and a Bronteus probably belonging to a new species. As the coral has not been found above the uppermost beds of the Lower Silurian it is probable that the trilobites also are derived from the same horizon.
The hypostoma, figured pl. III fig. 1, is broadly tongueshaped, with a short, blunt, triangular wing on each side of the anterior margin. The exterior side is nearly completely occupied by a single large field, which is convex and decorated by sparse terrace lines. Below this a smooth, nearly even plane, reaching to the inferior, elevated border of the hypostoma. The two tubercular macula' are situated exactly on the boundary line between the convex and the plane field. They are elongately ovate, pointed outwards, rounded inwards, with smooth and glossy surface. That portion, on which in the species of Bronteus the granular spot rests, is transversally wrinkled by some faintly elevated, sigmoid lines (fig. 2). It would have been of great interest to investigate the interior structure of these curious maculæ, but the scarcity of the material has prevented our doing so.
The other hypostoma, pl. III fig. 3-5, is larger and has the terrace lines more distantiated and forming more open curves, but for the rest it is of the same shape as the smaller, so that we may not ascribe these small deviations to a specific difference. The maculæ are more elongated, narrow, curved tubercles, thus differing from the next preceding. A section running through its left tubercle does not show any structure, but the shell of the maculæ is extremely thin, thus contrasting strongly with the surrounding thicker shell, fig. 4 a. The cephalic eye, again, remaining on the free cheek found in the same piece of limestone and probably belonging to the same specimen, has its surface well preserved and exhibits semiglobular lenses in low relief, fig. 5.
Calymmene Brongn.
In the following species of this genus hypostomic tubercles have formerly been recorded.
Cal. Blumenbachi Salter Monogr. pl. 8, fig. 9. The identity of the species is dubious.
Cal. duplicata Salter Mon. pl. 9, fig. 22. The tubercles rather more linear and approximated than in the subsequent. Salter calls them »two small transverse lobes ... forming a nearly continuous ridge».
Cal. tuberculata Angel. Tab. XIX fig. 5 c from the inside.
It is remarkable that the eyes or that part of them where the cornea and the lenses should be found, has been destroyed and entirely lost in all specimens of the Calymmene known to us, many hundreds having been searched and every one with a lacuna on that spot. It must have been of an excessive thinness. We know only of one instance, the Bohemian Calym. Arago, in which Barrande has seen the eye intact and provided with a small number of about eleven sphærical lenses.
Calymmene intermedia Ldm.
The clypeiform hypostoma is divided in two fields, one anterior larger, in the centre elevated in a short, blunt knob, and a posterior, bearing the two elevated rounded maculæ, just below the semicircular groove which separates it from the anterior field. These fields are surrounded by a large border which is posteriorly emarginated through a short rounded sinus. The surface is entirely covered by small semiglobular wartlets perforated by a straight pore or duct, which being filled with a black mineral, probably iron pyrites, gives a peculiar and characteristic aspect to the sections. In the same manner the whole mass of the hypostoma is pierced by straight, black lines, also filled up pores, starting from the outside and ending before attaining the inner surface.
The maculæ are prominent, separated from the hypostoma through narrow, distinct grooves, ovate, with the pointed end directed obliquely outwards, against the margins of the hypostoma. They are covered with wartlets and leaving only a little oblong spot free, quite smooth, this being consequently the macula proper. But in a longitudinal section there is not the least distinction between this macula and the other hypostoma, only, that it is free of pores.
Calymmene tuberculata Brünnich.
We have succeeded to prepare a horizontal section of the macula. This is oval, perfectly homogenous and without any structure proper, showing only indistinctly a mottled medley of pale, brownish spots in the clear mass. There is in both maculæ a small dark, angular spot placed in their inferior part, fig. 9.
Centropleura Angelin.
This author has delineated most distinct maculæ in Centr. Steenstrupi pl. III fig. 4, b. two, small elongate darkish spots, near the posterior margin. I have not seen any specimen of this trilobite and have consequently not been able to follow up Angelin's observation.
Chasmops M'Coy.
In his work on the Russian Phacopidæ Friedr. Schmidt has given the figures of Chasmops Eichwaldi and Ch. Wesenbergensis showing, as it were, patches of black maculæ. But as there is no mention of them, and as they are quite unlike the macula? which are described here, I am in doubt of their real nature. We have figured a macula from the hypostoma of a Chasmops, probably Ch. macroura, from a drift block near Rostock. It is elongate and narrow, more than four times longer than broad. Its smooth surface is a little scooped out and no structure is observable.
Chirurus Beyrich.
We find the hypostomic maculæ annotated and delineated as occurring in the following species.
Ch. bimucronatus Salter pl. 5, fig. 5 as two narrow elongated pits, surface seems intact. No mention of them in the letter-press.
Ch. exsul Schmidt pl. VI fig. 9. Indications of the oblong, narrow maculæ.
Ch. macrophthalmus Schm. pl. VII, figs. 1 c, 2. Faint indications of narrow pits.
Ch. Quenstedti Barr. Tab. 42 fig. 3. Maculæ near lower border as two narrow slits.
Ch. spinulosus Schm. pl. VII fig. 10. The lateral pits well visible.
Ch. tumidus Schm. pl. VIII, fig. 22. The pits are short, but present.
The exterior surface of the hypostoma is entirely covered with small wartlets, closely set, of different size in the various species, and spread between them lie a great number of elongate smooth spots dispersed, on the interior side of the hypostoma showing an excavated surface (pl. III fig. 16).
Chirurus (Cyrtometopus) clavifrons Dalm.
The shape of the hypostoma is shown in fig. 21 pl. III. It is regularly shield-formed with posterior margin rounded. A shallow groove running parallel with the margins forms a large, faintly elevated border. The anterior margin projects on both sides in a short, backwards directed, broad and fiat wing, hollowed out with a pit on the front side, and there is a posterior lateral wing, midways between the anterior and posterior margin short, blunt, directed obliquely backwards. The whole exterior surface is covered with diminutive tubercles, the above mentioned smooth spots interspersed on the central disk. A little below the middle there are two lengthened pits, so shallow that they often are not discernible. In a specimen of Chirurus spinulosus Nieczkowski from Estland there are two, almost 3 mm. long dots, one on each side above the maculæ as the fig. 21 shows and in several others of this generic group there are also indications of similar. I cannot compare these dots with anything more than the lengthened spots visible in Acidaspis crenata.
The maculæ are wanting in this species, but the cephalic eyes are well developed. A section of them fig. 19 shows the enormous difference of the shell in the eye and the surroundings where the shell surpasses it many times in thickness. The eye consists in fact only of the ovate, beadlike lenses of which a string is seen sectioned in the figure mentioned. More enlarged (fig. 20) a nucleus is visible in each lens, and in a horizontal section (fig. 18) a little below the surface, where they are more pressed against themselves, they have a polygonal shape. The figure 17 seems to represent lenses that have been much changed interiorly, having only a narrow zone left of the primary structure.
Chirurus glaber Angelin.
As a sample of the shape of the surface of the eyes in this genus, we have given a figure, showing the small rounded lenses.
Chirurus ornatus Dalman.
The cephalic eyes have globular lenses, of more than double the size of those in Chir. clavifrons. The difference between the lenses and the surrounding shell amounts at the highest to thrice the former, while in Ch. clavifrons it is at least seven times as much. In a horizontal section the lenses have a little, darkish irregular nucleus surrounded by a radiated structure (fig. 12).
The maculæ (fig. 14) are slightly concave, oblong, smooth, surrounded by comparatively large granulations.
Chirurus speciosus Dalm.
The exterior surface of the hypostoma is sparingly covered by granules of larger size than in other Chiruri. The maculæ as seen in a cast are prominent near the posterior border, sunk in their centre and surrounded by a narrow distinct border line. In a horizontal section it as not been possible to detect any peculiar structure, only a dark border surrounding the interior clear surface.
Chirurus spinulosus Nieczkowski.
We have had the interior surface of a hypostoma at our disposal from the lower Siluria, of Esthonia (Kuckers C2) belonging to Dr. G. Holm. The macula are well preserved, standing out black on the white surface, oblong, 1,5 millim. in greatest length (fig. 15). There are indications as of a great number of small lenses on these maculæ. Remarkable are the two elongated spots, almost 3 mm. in length, above these maculæ, of which already mention has been made under Ch. clavifrons.
Chirurus sp. indet.
The hypostoma of an undescribed species from Öland. The maculæ are lengthened, ellipsoid and smooth tubercles lying in a groove. The finely granulated surface of the hypostoma shows rare smooth spots interspersed, being on the inside smooth pits, from which again in casts of the hypostoma larger tubercles are moulded. Of the same nature are those that are visible on the nucleus of Chir. speciosus and others. There are still some undescribed Lower Silurian species, for instance one nearly related to Ch. conformis with tubercular maculæ. A large hypostoma from the Leptæna limestone of Dalecarlia, 33 millim. in length, has a longitudinal macula, 2 millim. in length, as seen on the inside of the fragment.
On pl. VI fig. 10 a little hypostoma is delineated that shows some resemblance with that of the Chirurus, but probably belongs to some other, unknown genus. It is Upper Silurian, found at Mulde in Fröjel, Gotland. It is of an elongated ovate shape, with the anterior border faintly arched, the anterior wings broad, truncate. A narrow elevated border surrounds the lateral and posterior margins. The somewhat vaulted surface is covered by fine granulations, and the macula, situated on equal distance from the anterior and posterior margins are ovate, smooth and directed obliquely inwards. There are two minute pointed processes on each side of the lateral borders and one on each side of the pointed posterior margin.
Ctenopyge Linsn.
Ct. spectabilis Br. (I, pl. XII, fig. 12 a).
A small lengthened, apparently smooth hypostoma with two globular tubercular maculæ near the posterior border. No mention made of them in the description of Brögger. Cambrian. I am not, however, quite sure whether this really belongs to a Ctenopyge, as it rather more resembles the type of a Peltura as shown in Pelt. scarabæoides. As Ctenopyge is so nearly related to Sphærophthalmus it could be expected to see its hypostoma of the same type and the maculæ entirely wanting. The eyes of this genus have been described above at page 29.
Cybele Lovén.
Cyb. bellatula Dalman.
We give a new figure of the hypostoma to complete that of Fr. Schmidt, pl. XIII fig. 9. The anterior wings are larger and obtuse, excavated below and the border rim is not so distinct all round as he has figured. There are two pair of oblique, lengthened grooves above each other. The opposite do not, however, join to form a coherent groove across the median field of the hypostoma, which is level between them. No maculæ. Nearer the anterior border close below the wings, there is a small roundish, dark spot. The surface of the hypostoma is granulated.
Cyphaspis Ang.
Cyph. elegantula Ang.
The hypostoma is elongated, anterior margin evenly arched, anterior wings triangular, lateral margins faintly curved as well as the posterior margin, at both sides of which there is one diminutive spine. The posterior wings are short forming a spiny process from the lateral margin. The surface of the hypostoma is smooth with a shallow transverse groove on the inferior moiety and two corresponding lateral impressions below it, one on each side. The elevated lateral borders are longitudinally striated by a few terrace lines. There are no maculæ visible. The eves consist f short hexagonal prisms (figs. 22, 23) nearly resembling those of Proetus.
Dalmanites Emmrich.
In this large genus the hypostoma has been figured in a great number of species. But it is only in the following that we, at least in the figures, if not in the descriptions, are able to detect the maculæ.
D. atavus Barr. Suppl. pl. 5 fig. 14.
D. Calypso J. Hall, vol. VII, pl. XI A. fig. 21 p. 66 »Postero-lateral pits, moderately strong and elongate». There are casts of the two well distinct maculæ.
D. caudatus Brünn.
a. Forbes and Salter in British Organic remains, Dec. II pl. 1, fig. 3 give a good figure, showing the oblique pitlike maculæ. On page 2 is told that »a pair of lateral strong indentations indicate a second furrow above» (»the transverse furrow between the tip»).
b. Salter, pl. 3, figs. 7, 8, bad figures, the description verbally the same as above. What Angelin pl. VIII, fig. 2 c gives as the hypostoma of »Ph. caudata» is in fact the hypostoma of a species of Lichas.
D. Mac Coyi Barr. Supl., pl. 13 f. 32. This magnificent hypostoma belongs to the same group as D. micrurus Hall as to which we may hesitate whether there not be a double pair of maculæ above each other near the posterior border. In this species, however, there is the transverse lower groove and above probably the two maculæ united by a groove so closely as to resemble the lower groove.
D. micrurus J. Hall, Pal. N. Y. III pl. 74 fig. 20. Shows, as it were, a double pair of maculæ, very like D. spinifer Barr.
D. rugosus Barr. pl. 24 f. 23. The two maculæ in the ordinary place, without being united by a groove.
D. socialis Barr. pl. 26 f. 21. Rather of an uncommon shape, not so triangular as in the other species, the two maculæ distinct as narrow slits. P. 553 »Vers l'extrémité du corps central, on aperçoit de chaque côté, près du bord, une impression oblique, alongée et arquée».
D. spinifer Barr. Pl. 25, fig. 20. A large beautifully preserved hypostoma, shows what I think we positively must interpret as a double pair of maculæ close above each other. As stated above there is reason to believe that in other species of Dalmanites there also are indications of four maculæ, though not so evident as in this species. Barrande says nothing about this remarkable feature.
Dalmanites imbricatulus Angelin.
The interesting image, as given by the agglomerated eyes, is represented in the magnified figure 43 Plate III. The lenses lie there separated from each other at much varying distance, some in close contact. The presence of the delicate covering membrane is clearly seen, being of white colour contrasting with the black, glossy lens. It is much lacerated and only preserved around the periphery. The size of the ocelli is on an average 0,5 mm., a little one has 0,3 mm. The surface between the ocelli is most finely granulated. The granules scarcely attaining the fourth of the granules in Dalm. vulgaris.
In a vertical section the regular biconvex lenses are seen to be covered with the extremely thin integument, which is a direct continuation from the test. The test between the lenses is perforated by some longitudinal tubes, as usual in the skeleton of the trilobites. We have not found any hypostoma with the maculæ.
Dalmanites obtusus Ldm.
We have not succeeded in finding any hypostoma, but as the structure of the cephalic eyes is sufficiently well preserved, we here describe it. The lenses seen from the surface look globular, and in a vertical section they are ovate and in a horizontal circular. They are covered by a very thin and delicate membrane, that envelops their superior moiety completely and between the eyes it is enclosed by the surrounding test and growing out from it. In the horizontal section, fig. 45, it looks as a circular frame around the lenses and the membranes of two contiguous lenses sometimes lie so close, that they entirely fill up the interspace between the lenses, leaving only a narrow slit between them marking their boundaries. The space between the lenses, fig. 46, is porous, being perforated by tubes, which continue vertically down through the shell of the cheek.
Dalm. sclerops Dalman.
Commonly this species has been ranged with Phacops, but it is evident that it as to the conformation of its glabella and the head in general is highly discrepant, and in these respects is more concordant with Dalmanites, though still deviating as for instance in the pygidium and some details of the head. Therefore it may be justified with Friedr. Schmidt[41] to consider it as a generic division, though not, as he has it, as an independent genus, but as a subgenus of Dalmanites.