StageLengthin mm.Percentageof pygidiumSegmentsin thoraxSegmentsin pygidium10.66300220.7533-1/30331.00460541.503225-651.50253461.75234471.80215382.00176392.501373103.001283113.501193-4124.0011103-4135.0010113145.509122-4156.008133-4166.508143177.007153187.507163197.5061722010.256172
This table shows the rapid increase in the length of the pygidium till the time when the thorax began to be freed, the very rapid decrease during the earlier part of its formation until six segments had been set free, and then a more gradual decrease until the entire seventeen segments had been acquired, after which time the relative length remained constant. From an initial proportion of 30 per cent, it rose to nearly one half the whole length, and then dwindled to a mere 6 per cent, showing conclusively that the thorax grew at the expense of the pygidium.
If this conclusion can be sustained by other trilobites, it indicates that the large pygidium is a more primitive characteristic of a protaspis than is a small one. I have already shown that the pygidium is proportionately larger in the protaspis in the Mesonacidæ, Solenopleuridæ, and Olenidæ, and a glance at Barrande's figures of"Hydrocephalus" carensand"H." saturnoides, both young ofParadoxideswill show that the same process of development goes on in that genus as inSao. There is first an enlargement of the pygidium to a maximum, a rise from 20 per cent to 33 per cent in the case ofH. carensand then, with the introduction of thoracic segments, a very rapid falling off. All of these are, however, trilobites with small pygidia, and it has been a sort of axiom among palæontologists that large pygidia were made up of a number of coalesced segments. While not definitely so stated, it has generally been taken to mean the joining together of segments once free. The asaphid, for instance, has been thought of as descended from some trilobite with rich segmentation, and a body-form like that of aMesonacisorParadoxides.
The appeal to the ontogeny does not give as full an answer to this question as could be wished, for the complete life-history of no trilobite with a large pygidium is yet known. While the answer is not complete, enough can be gained from the study of the ontogeny ofDalmanitesandCyclopygeto show that in these genera also the thorax grows by the breaking down of the pygidium and that no segment is ever added from the thorax to the pygidium. The case ofDalmanites socialisas described by Barrande (1852, p. 552, pl. 26) will be taken up first, as the more complete. The youngest specimen of this species yet found is 0.75 mm. long, the pygidium is distinctly separated from the cephalon, and makes up 25 per cent of the length. This is probably not the form of the shell as it leaves the egg. At this stage there are two segments in the pygidium, but they increase to four when the test is 1 mm. long. The cephalon has also increased in length, however, so that the proportional length is the same. The subjoined table, which is that compiled by Barrande with the proportional length of the pygidium added, is not as complete as could be desired, but affords a very interesting history of the growth of the caudal shield. The maximum proportional length is reached before the introduction of thoracic segments, and during the appearance of the first five segments the size of the pygidium drops from 25 to 15 per cent. Several stages are missing at the critical time between stages 8 and 9 when the pygidium had added three segments to itself and has supplied only one to the thorax. This would appear to have been a sort of resting or recuperative stage for the pygidium, for it increased its own length to 20 per cent, but from this stage up to stage 12 it continued to give up segments to the thorax and lose in length itself. After stage 12, when the specimens were 8 mm. long, no more thoracic segments were added, but new ones were introduced into the pygidium, until it reached a size equal to one fifth the entire length, as compared with one fourth in the protaspis.
StageLengthin mm.Percentageof pygidiumSegmentsin thoraxSegmentsin pygidium10.75250220.75250331.00250441.00221351.25202361.25183371.60154381.60155393.002066103.502076118.001897128.00161151312.00161171419.00181191595.00201111
Since the above was written, Troedsson (1918, p. 57) has described the development ofDalmanites eucentrus, a species found in the Brachiopod shales (Upper Ordovician) of southern Sweden. This species follows a course similar to that ofD. socialis, so that the full series of stages need not be described. The pygidium is, however, of especial interest, for there is a stage in which it shows two more segments than in the adult. Troedsson figures a pygidium 1.28 mm. long which has eight pairs of pleural ribs, while the adult has only six pairs. The ends of all these ribs are free spines, and were the development not known one would say that this was a case of incipient fusion, while as a matter of fact, it is incipient freedom.
A further interest attaches to this case, because of the close relationship betweenD. eucentrusandD. mucronatus. The latter species appears first in theStaurocephalusbeds which underlie the Brachiopod shales, so that in its first appearance it is somewhat the older. The pygidium of the adultD. mucronatusis larger than that ofD. eucentrus, having eight pairs of pleural ribs, the same number as in the young of the latter. In short,D. eucentrusis probably descended fromD. mucronatus, and in its youth passes through a stage in which it has a large pygidium like that species. Once more it appears that the small pygidium is more specialized than the large one.
The full ontogeny ofCyclopygeis not known, but young specimens show conclusively that segments are not transferred from the thorax to the pygidium, but that the opposite occurs. As shown by Barrande (1852) and corroborated by specimens in the Museum of Comparative Zoology, the process is as follows: The third segment of the adult of this species, that is, the fourth from the pygidium, bears a pair of conspicuous cavities on the axial portion. In a young specimen, 7 mm. long, the second segment bears these cavities, but as the thorax has only four segments, this segment is also the second instead of the fourth ahead of the pygidium. The pygidium itself, instead of being entirely smooth, as in the adult state, is smooth on the posterior half, but on the anterior portion has two well formed but still connected segments, the anterior one being more perfect than the other. These are evidently the two missing segments of the thorax, and instead of being in the process of being incorporated in the pygidium, they are in fact about to be cast off from it to become free thoracic segments. In other words, the thorax grows through the degenerationof the pygidium. That the thorax grows at actual expense to the pygidium is shown by the proportions of this specimen. In an adult of this species the pygidium, thorax, and cephalon are to each other as 9:11:13. In the young specimen they are as 10:6:12, the pygidium being longer in proportion both to the thorax and to the cephalon than it would be in the adult.
This conception of the breaking down of the pygidium to form the thorax will be very helpful in explaining many things which have hitherto seemed anomalous. For instance, it indicates that the Agnostidæ, whose subequal shields in early stages have been a puzzle, are really primitive forms whose pygidia do not degenerate; likewise the Eodiscidæ, which, however, show within the family a tendency to free some of the segments. The annelidan Mesonacidæ may not be so primitive after all, and their specialized cephala may be more truly indicative of their status than has previously been supposed.
The facts of ontogeny of trilobites with both small and large pygidia do show that there is a reduction of the relative size of the caudal shield during the growth-stages, and therefore that the large pygidium in the protaspis is probably primitive. The same study also shows that the large pygidium is made up of "coalesced segments" only to the extent that they are potentially free, and not in the sense of fused segments.
WIDTH OF THE AXIAL LOBE.
That the narrow type of axial lobe is more primitive than the wide one has already been demonstrated by the ontogeny of various species, and space need not be taken here to discuss the question. Most Cambrian trilobites have narrow axial lobes even in the adult so that their development does not bring this out very strikingly, though it can be seen in Sao, Ptychoparia, etc., but in Ordovician trilobites such as Triarthrus and especially Isotelus, it is a conspicuous feature.
PRESENCE OR ABSENCE OF A "BRIM."
That the extension of the glabella to the front of the cephalon is a primitive feature is well shown by the development of Sao (Barrande, 1852, pl. 7), Ptychoparia (Beecher, 1895 C, pl. 8), and Paradoxides (Raymond, Bull. Mus. Comp. Zool., vol. 57, 1914), although in the last genus the protaspis has a very narrow brim, the larva during the stages of introduction of new segments a fairly wide one, and most adults a narrow one.
The brim of Sao seems to be formed partly by new growth and partly at the expense of the frontal lobe, for that lobe is proportionately shorter in the adult than in the protaspis. InCryptolithusand probably inHarpes,Harpides, etc., the brim is quite obviously new growth and has nothing to do with the vital organs. Its presence or absence may not have any great significance, but when the glabella extends to the frontal margin, it certainly suggests a more anterior position of certain organs. InSao, the only trilobite in which anything is known of the position of the hypostoma in the young, the posterior end is considerably further forward in a specimen a. 5 mm. long than in one 4 mm. long, thus indicating a backward movement of the mouth during growth, comparable to the backward movement of the eyes.
SEGMENTATION OF THE GLABELLA.
The very smallest specimens ofSaoshow a simple, unsegmented axial lobe, and the same simplicity has been noted in the young of other genera. Beecher considered this asdue to imperfect preservation of the exceedingly small shells, which practically always occur as moulds or casts in soft shale. There is, however, a very general increase in the strength of glabellar segmentation in the early part of the ontogeny of all trilobites whose life history is known, and in some genera, like the Agnostidæ, there is no question of the comparatively late acquisition of glabellar furrows. Even inParadoxides, the furrows appear late in the ontogeny.
Summary.
If absence of eyes on the dorsal surface be primitive, as Beecher has shown, and if the large pygidium, narrow axial lobe, and long unsegmented glabella be primitive, then the known protaspis of the Mesonacidæ and Paradoxidæ is not primitive, that of the Olenidæ is very primitive, and that of the Agnostidæ is primitive except that in one group the axial lobe, when it appears, is rather wide, and in the other a brim is present.
Subsequent development from the simple unsegmented protaspis would appear to show, first, an adaptation to swimming by the use of the pygidium; next, the invagination of the appendifers as shown in the segmentation of the axial lobe indicates the functioning of the appendages as swimming legs; then with the introduction of thoracic segments the assumption of a bottom-crawling habit is indicated. Some trilobites were fully adapted for bottom life, and the pygidium became reduced to a mere vestige in the production of a worm-like body. Other trilobites retained their swimming habits, coupled with the crawling mode of life, and kept or even increased (Isotelus) the large pygidium.
Fig. 35.—A specimen ofWeymouthia nobilis(Ford), collected by Mr. Thomas H. Clark at North Weymouth, Mass. Note the broad smooth shields of this Lower Cambrian eodiscid. × 6.
Fig. 35.—A specimen ofWeymouthia nobilis(Ford), collected by Mr. Thomas H. Clark at North Weymouth, Mass. Note the broad smooth shields of this Lower Cambrian eodiscid. × 6.
In the discussion above I have placed great emphasis on the large size of the primitive pygidium, because, although there is nothing new in the idea, its significance seems to have been overlooked.
If the large pygidium is primitive, then multisegmentation in trilobites can not be primitive but is the result of adaptation to a crawling life. It is annelid-like, but is not in itself to be relied upon as showing relationship to the Chætopoda. Simple trilobites with few segments, like the Agnostidæ, Eodiscidæ etc., were, therefore, properly placed by Beecher atthe base of his classification, and there is now less chance than ever that they can be called degenerate animals.
From the phylogeny of certain groups, such as the Asaphidæ, it is learned that the geologically older members of the family have more strongly segmented anterior and posterior shields than the later ones. That there has been a "smoothing out" is demonstrated by a study of the ontogeny of the later forms. From such examples it has come to be thought that all smooth trilobites are specialized and occupy a terminal position in their genealogical line. This has caused some wonder that smooth agnostids likePhalacroma bibullatumandP. nudumshould be found in strata so old as the Middle Cambrian, and was a source of great perplexity to me in the case ofWeymouthia(Ottawa Nat., vol. 27, 1913) (fig. 35). This is a smooth member of the Eodiscidæ, and, in fact, one of the simplest trilobites known, for while it has three thoracic segments, it shows almost no trace of dorsal furrows or segmentation on cephalon or pygidium, and, of course, no eyes. Following the general rule, I took this to be a smooth-out eodiscid, and was surprised that it should come from the Lower Cambrian, where it is associated withElliptocephalaat Troy, New York, and withCallaviaat North Weymouth, Massachusetts, and where it has lately been found by Kiær associated withHolmiaandKjerulfiaat Tømten, Norway. It now appears it is really in its proper zone, and instead of being the most specialized, is the simplest of the Eodiscidæ.
What appears to be a still simpler trilobite is the form described by Walcott as Naraoia.
Naraoia compactaWalcott.
(Textfig. 36.)
Illustrated: Walcott, Smithson. Misc. Coll., vol. 57, 1912, p. 175, pl. 28, figs. 3, 4.—Cleland, Geology, Physical and Historical, New York, 1916, p. 412, fig. 382 F (somewhat restored).
Illustrated: Walcott, Smithson. Misc. Coll., vol. 57, 1912, p. 175, pl. 28, figs. 3, 4.—Cleland, Geology, Physical and Historical, New York, 1916, p. 412, fig. 382 F (somewhat restored).
This very imperfectly known form is referred by Walcott to the Notostraca on what appear to be wholly inadequate grounds, and while I do not insist on my interpretation, I can not refrain from calling attention to the fact that itcanbe explained as the most primitive of all trilobites. It consists of two subequal shields, the anterior of which shows slight, and the posterior considerable evidence of segmentation. It has no eyes, no glabella, and no thorax, and is directly comparable to a very youngPhalacroma bibullatum(see Barrande 1852, pl. 49, figs. a, b). Walcott states that there is nothing to show how many segments there are in the cephalic shield, but that on one specimen fourteen were faintly indicated on the abdominal covering. The appendages are imperfectly unknown, as no specimen showing the ventral side has yet been described. The possible presence of antennas and three other appendages belonging to the cephalic shield is mentioned, and there are tips of fourteen legs projecting from beneath the side of one specimen. As figured, some of the appendages have the form of exopodites, others of endopodites, indicating that they were biramous.
Naraoiais, so far as now known, possessed of no characteristics which would prevent its reference to the Trilobita, while the presence of a large abdominal as well as a cephalic shield would make it difficult to place in even so highly variable a group as the Branchiopoda. On the other hand, its only exceptional feature as a trilobite is the lack of thorax, and all study of the ontogeny of the group has led us to expect just that sort of a trilobite to be found some day in the most ancient fossiliferous rocks.Naraoiacan, I think,be best explained as a trilobite which grew to the adult state without losing its protaspian form. It was found in the Middle Cambrian of British Columbia.
Even ifNaraoiashould eventually prove to possess characteristics which preclude the possibility of its being a primitive trilobite, it at least represents what I should expect a pre-Cambrian trilobite to look like. What the ancestry of the nektonic primitive trilobite may have been is not yet clear, but all the evidence from the morphology of cephalon, pygidium, and appendages indicates that it was a descendant of a swimming and not a crawling organism.
Since the above was written, the Museum of Comparative Zoology has purchased a specimen of this species obtained from the original locality. The shields are subequal, the posterior one slightly the larger, and the axial lobes are definitely outlined on both. The glabella is about one third the total width, nearly parallel-sided, somewhat pointed at the front. There are no traces of glabellar furrows. The axial lobe of the pygidium is also about one third the total width, extends nearly to the posterior margin, and has a rounded posterior end. The measurements are as follows: Length, 33 mm.; length of cephalon, 16 mm., width, 15 mm.; length of glabella, 11.5 mm., width, 5.5 mm.; length of pygidium, 17 mm., width, 15 mm.; length of axial lobe, 14 mm., width, 5.5 mm.
The species is decidedlyAgnostus-like in both cephalon and pygidium, and were it not so large, might be taken for the young of such a trilobite. The pointed glabella is comparable to the axial lobes of the so called pygidia of the young ofCondylopyge rexandPeronopsis integer(Barrande, Syst. Sil., vol. 1, pl. 49).
The "annelid" theory of the origin of the Crustacea and therefore of the trilobites, originating with Hatschek (1877) and so ably championed by Bernard (1892), has now been a fundamental working hypothesis for some years, and has had a profound influence in shaping thought about trilobites. This hypothesis has, however, its weak points, the principal one being its total inhibition of the workings of that great talisman of the palæontologist, the law of recapitulation. Its acceptance has forced the zoologist to look upon the nauplius as a specially adapted larva, and has caused more than one forced explanation of the protaspis of the trilobite. When so keen a student as Calman says that the nauplius must point in some way to the ancestor of the Crustacea (1909, p. 26), it is time to reëxamine some of the fundamentals. This has been done in the preceding pages and evidence adduced to show that the primitive features of a trilobite indicate a swimming animal, and that the adaptations are those which enabled it to assume a crawling mode of existence. It has also been pointed out that in Naraoia there is preserved down to Middle Cambrian times an animal like that to which ontogeny points as a possible ancestor of the trilobites.Naraoiais not the simplest conceivable animal of its own type, however, for it has built up a pygidium of fourteen or fifteen somites. One would expect to find in Proterozoic sediments remains of similar animals with pygidia composed of only one or two somites, with five pairs of appendages on the cephalon, one or two pairs on the pygidium, a ventral mouth, and a short hypostoma. Anything simpler than this could not, in my opinion, be classed as a trilobite.
What the ancestor of this animal was is mere surmise. It probably had no test, and it may be noted in this connection thatNaraoiahad a very thin shell, as shown by itsstate of preservation, and was in that respect intermediate between the trilobite and the theoretical ancestor. Every analysis of the cephalon of the trilobite shows that it is made up of several segments, certainly five, probably six, possibly seven. Every study of the trilobite, whether of adult, young, or protaspis, indicates the primitiveness of the lateral extensions or pleural lobes. The same studies indicate as clearly the location of the vital organs along the median lobe. These suggestions all point to a soft-bodied, depressed animal composed of few segments, probably with simple marginal eyes, a mouth beneath the anterior margin, tactile organs at one or both ends, with an oval shape, and a straight narrow gut running from anterior mouth to terminal anus. The broad flat shape gives great buoyancy and is frequently developed in the plankton. Inherited by the trilobites, it proved of great use to the swimmers among them.
The known animal which most nearly approaches the form which I should expect the remote ancestor of the trilobites to have had isAmiskwia sagittiformisWalcott (Smithson. Misc. Coll., vol. 57, 1911, p. 112, pl. 22, figs. 3, 4). This "worm" from the Middle Cambrian is similar in outline to the recentSpadella, and is referred by Walcott to the Chætognatha. It has a pair of lateral expansions and a flattened caudal fin, a narrow median alimentary canal, and a pair of rather long simple tentacles. With the exception of a thin septum back of the head, no traces of segmentation are shown.
Some time in the late pre-Cambrian, the pre-trilobite, which probably swam by rhythmic undulations of the body, began to come into occasional contact with a substratum, and two things happened: symmetrically placed, i. e., paired, appendages began to develop on the contact surface, and a test on the dorsal side. The first use of the appendages may have been in pushing food forward to the mouth, and for the greater convenience in catching such material, a fold in front of the mouth may have elongated to form the prototype of the hypostoma. At this time the substratum may not have been the ocean bottom at all, but the animals, still free swimmers, may have alighted at feeding time on floating algæ from the surface of which they collected their food. While the dorsal test was originally jointed at every segment, the undulatory mode of swimming seems to have given way to the method of sculling by means of the posterior end only, or by the use of the appendages, and the anterior segments early became fused together.
The result of the hardening of the dorsal test was of course to reduce to that extent the area available for respiration, and this function was now transferred in part to the limbs, which bifurcated, one branch continuing the food-gathering process and the other becoming a gill. The next step may have been the "discovery" of the ocean bottom and the tapping of an hitherto unexploited supply of food. Upon this, there set in those adaptations to a crawling mode of existence which are so well shown in the trilobite. The crawling legs became lengthened and took on a hardened test, the hypostoma was greatly elongated, pushing the mouth backward, and new segments were added to produce a long worm-like form which could adapt itself to the inequalities of the bottom. That the test of the appendages became hardened later than that of the body is shown by the specimens of Neolenus, in which the dorsal shell as preserved in the shale is thick and solid, while the test of the appendages is a mere film.
The late Proterozoic or very earliest Cambrian was probably the time of the great splitting up into groups. The first development seems to have been among the trilobites themselves, the Hypoparia giving rise to two groups with compound eyes, first the Opisthoparia and later the Proparia. About this same time the Copepoda may have split offfrom the Hypoparia, continuing in the pelagic habitat. At first, most of the trilobites seem to have led a crawling existence, but about Middle Cambrian time they began to go back partially to the ancestral swimming habits, and retained some of the trunk segments to form a larger pygidium. The functional importance of the pygidium explains why it can not be used successfully in making major divisions in classification. Nearly related trilobites may be adapted to diverse methods of life.
EVOLUTION WITHIN THE CRUSTACEA.
The question naturally arises as to whether the higher Crustacea were derived from some one trilobite, or whether the different groups have been developed independently from different stocks. The opinion that all other crustaceans could have been derived from anApus-like form has been rather generally held in recent years, but Carpenter (1903, p. 334) has shown that the leptostracan,Nebalia, is really a more primitive animal thanApus. He has pointed out that in Leptostraca the thorax bears eight pairs of simple limbs with lamelliform exopodites and segmented endopodites, while the abdomen of eight segments has six pairs of pleopods and a pair of furcal processes, so that only one segment is limbless. Contrasted with this are the crowded and complicated limbs of the anterior part of the trunk ofApus, and the appendage-less condition of the hinder portion. Further, a comparison between the appendages of the head ofNebaliaand those ofApusshows that the former are the more primitive. The antennules of Nebalia are elongate, those ofApusgreatly reduced; the mandible ofNebaliahas a long endopodite, and Carpenter points out that from it either the malacostracan mandible with a reduced endopodite or the branchiopodan mandible with none could be derived, but that the former could not have arisen from the latter. The maxillæ ofApusare also much the more specialized and reduced.
Nebaliabeing in all else more primitive thanApus, it follows that the numerous abdominal segments of the latter may well have arisen by the multiplication of an originally moderate number, and the last trace of primitiveness disappears.
It is now possible to add to the results obtained from comparative morphology the testimony of palæontology, already outlined above, and since the two are in agreement, it must be admitted that the modern Branchiopoda are really highly specialized.
As has already been pointed out,Hymenocaris, the leptostracan of the Middle Cambrian, has very much the same sort of appendages as the Branchiopoda of the same age, both being of the trilobite type. Which is the more primitive, and was one derived from the other?
The Branchiopoda were much more abundant and much more highly diversified in Cambrian times than were the Leptostraca, and, therefore, are probably older. Some of the Cambrian branchiopods were without a carapace, and some were sessile-eyed. These were more trilobite-like than Hymenocaris. Many of the Cambrian branchiopods had developed a bivalved carapace, though not so large a one as that of the primitive Leptostraca. The present indications are, therefore, that the Branchiopoda are really older than the Leptostraca, and also that the latter were derived from them. It seems very generally agreed that the Malacostraca are descended from the Leptostraca, and the fossils of the Pennsylvanian supply a number of links in the chain of descent. Thus,Pygocephalus cooperi, with its brood pouches, is believed by Calman (1909, p. 181) to stand at the base of the Peracaridan series of orders, andUronectes,Palæocaris, and the like are Palæozoic representatives of the Syncarida. Others of the Pennsylvanian species appear to tend in the direction of the Stomatopoda,whose true representatives have been found in the Jurassic. The Isopoda seem to be the only group of Malacostraca not readily connected up with the Leptostraca. Their depressed form, their sessile-eyes, and their antiquity all combine to indicate a separate origin for the group, and it has already been pointed out how readily they can be derived directly from the trilobite.
While the Copepoda seem to have been derived directly from the Hypoparia, the remainder of the Crustacea apparently branched off after the compound eyes became fully developed, unless, as seems entirely possible, compound eyes have been developed independently in various groups. Most Crustacea were derived from crawling trilobites (Lower Cambrian or pre-Cambrian Opisthoparia), for they lost the large pygidium, and also the major part of the pleural lobes. In all Crustacea, too, other than the Copepoda and Ostracoda, there is a tendency to lose the exopodites of the antennæ.
These modifications, which produced a considerable difference in the general appearance of the animal, are easily understood. As has been shown in previous pages, the trilobites themselves exhibit the degenerative effect on the anterior appendages of the backward movement of the mouth, and the transformation of a biramous appendage with an endobase into a uniramous antenna is a simple result of such a process. The feeding habits of the trilobites were peculiar and specialized, and it is natural that some members of the group should have broken away from them. In any progressive mode of browsing the hypostoma was a hindrance, so was soon gotten rid of, and the endobases not grouped around the mouth likewise became functionless. The chief factor in the development of the higher Crustacea seems to have been the pinching claw, by means of which food could be conveyed to the mouth. It had the same place in crustacean development that the opposable thumb is believed to have had in that of man.
An intermediate stage between the Trilobita and the higher Crustacea is at last exhibited to us by the wonderful, but unfortunately rather specializedMarrella, already described. It retains the hypostoma and the undifferentiated biramous appendages of the trilobite, but has uniramous antennæ, there are no endobases on the coxopodites of the thoracic appendages, the pygidium is reduced to a single segment, and the lateral lobes of the thorax are also much reduced.Marrellais far from being the simplest of its group, but is the only example which survived even down to Middle Cambrian times of what was probably once an important series of species transitional between the trilobites and the higher Crustacea.
In this theory of the origin of the Crustacea from the Trilobita, the nauplius becomes explicable and points very definitely to the ancestor. According to Calman (1909, p. 23):
The typical nauplius has an oval unsegmented body and three pairs of limbs, corresponding to the antennules, antennas, and mandibles of the adult. The antennules are uniramous, the others biramous, and all three pairs are used in swimming. The antennæ may have a spiniform or hooked masticatory process at the base, and share with the mandibles which have a similar process, the function of seizing and masticating the food. The mouth is overhung by a large labrum or upper lip and the integument of the dorsal surface of the body forms a more or less definite dorsal shield. The paired eyes are as yet wanting, but the median eye is large and conspicuous.
The typical nauplius has an oval unsegmented body and three pairs of limbs, corresponding to the antennules, antennas, and mandibles of the adult. The antennules are uniramous, the others biramous, and all three pairs are used in swimming. The antennæ may have a spiniform or hooked masticatory process at the base, and share with the mandibles which have a similar process, the function of seizing and masticating the food. The mouth is overhung by a large labrum or upper lip and the integument of the dorsal surface of the body forms a more or less definite dorsal shield. The paired eyes are as yet wanting, but the median eye is large and conspicuous.
The large labrum or hypostoma, the biramous character of the appendages, especially of the antennæ, the functional gnathobases on the second and third appendages, and the oval unsegmented shield are all characteristics of the trilobites, and it is interesting to note that all nauplii have the free-swimming habit.
The effect of inheritance and modification through millions of generations is also shown in the nauplius, but rather less than would be expected. The most important modificationis the temporary suppression of the posterior pairs of appendages of the head, so that they are generally developed later than the thoracic limbs. The median or nauplius eye has not yet been found in trilobites, and if it is, as it appears to be, a specialized eye, it has probably arisen since the later Crustacea passed the trilobite stage in their phylogeny.
The oldest Crustacea, other than trilobites, so far known are the Branchiopoda and Phyllocarida described by Walcott and discussed above. It is important to note that while the former have already achieved such modified characteristics that they have been referred to modern orders, they retain the trilobite-like limbs and some of them still have well developed pleural lobes.
Calman (1909, p. 101) says of the Copepoda:
On the hypothesis that the nauplius represents the ancestral type of the Crustacea, the Eucopepoda would be regarded as the most primitive existing members of the class, retaining as they do, naupliar characters in the form of the first three pairs of appendages and in the absence of paired eyes and of a shell-fold. As already indicated, however, it is much more probable that they are to be regarded as a specialized and in some respects degenerate group which, while retaining, in some cases, a very primitive structure of the cephalic appendages, has diverged from the ancestral stock in the reduction of the number of somites, the loss of the paired eyes and the shell-fold, and the simplified form of the trunk-limbs.
On the hypothesis that the nauplius represents the ancestral type of the Crustacea, the Eucopepoda would be regarded as the most primitive existing members of the class, retaining as they do, naupliar characters in the form of the first three pairs of appendages and in the absence of paired eyes and of a shell-fold. As already indicated, however, it is much more probable that they are to be regarded as a specialized and in some respects degenerate group which, while retaining, in some cases, a very primitive structure of the cephalic appendages, has diverged from the ancestral stock in the reduction of the number of somites, the loss of the paired eyes and the shell-fold, and the simplified form of the trunk-limbs.
If the Eucopepoda be viewed in the light of the theory of descent here suggested, it is at once seen that while they are modified and specialized, they more nearly approximate the hypothetical ancestor than any other living Crustacea. Compound eyes are absent, and it can not be proved that they were ever present, although Grobben is said to have observed rudiments of them in the development ofCalanus. The "simplified limbs" are the simple limbs of the trilobite, somewhat modified. The absence of the shell-fold and carapace is certainly a primitive characteristic. Add to this the direct development of the small number of segments, and the infolded pleural lobes, and it must be admitted that the group presents more trilobite-like characteristics than any other. It seems very likely that the primitive features were retained because of the pelagic habitat of a large part of the group.
Ruedemann (Proc. Nat. Acad. Sci., vol. 4, 1918, p. 382, pl.) has recently outlined a possible method of derivation of the acorn barnacles from the phyllocarids. Starting from a recentBalanuswith rostrum and carina separated by two pairs of lateralia, he traces back throughCalophragmuswith three pairs of lateralia toProtobalanusof the Devonian with five pairs. Still older is the newly discoveredEobalanusof the upper Ordovician, which also has five pairs of lateralia but the middle pair is reversed, so that when the lateralia of each side are fitted together, they form a pair of shields like those ofRhinocaris, separated by the rostrum and carina, which are supposed to be homologous with the rostrum and dorsal plate of the Phyllocarida. Ruedemann suggests that the ancestral phyllocarid attached itself by the head, dorsal side downward, and the lateralia were developed from the two valves of the carapace during its upward migration, to protect the ventral side exposed in the new position.
This theory is very ingenious, but has not been fully published at the time of writing, and it seems very doubtful if it can be sustained.
Summary.
The salient points in the preceding discussion should be disentangled from their setting and put forward in a brief summary.
It is argued that the ancestral arthropod was a short and wide pelagic animal of few segments, which so far changed its habits as to settle upon a substratum. As a result of change in feeding habits, appendages were developed, and, due perhaps to physiological change induced by changed food, a shell was secreted on the dorsal surface, covering the whole body. Such a shell need not have been segmented, and, in fact, the stiffer the shell, the more reason for development of the appendages. Activity as a swimming and crawling animal tended to break up the dorsal test into segments corresponding to those of the soft parts, and, by adaptation, a floating animal became a crawling one, with consequent change from a form like that ofNaraoiato one likePædeumias. (See figs.36-40.) A continuation of this line of development by breaking up and loss of the dorsal test led through forms similar toMarrellato the Branchiopoda of the Cambrian, in which not only is there great reduction in the test, but also loss of appendages. The origin of the carapace is still obscure, but Bernard (1892, p. 214, fig. 48) has already pointed out that some trilobites, Acidaspidæ particularly, have backward projecting spines on the posterior margin of the cephalon, which suggest the possibility of the production of such a shield, and inMarrellasuch spines are so extravagantly developed as almost to confirm the probability of suchorigin. In this line of development two pairs of tactile antennæ were produced, while the anomomeristic character of the trilobite was retained. From similar opisthoparian ancestors there were, however, derived primitive Malacostraca retaining biramous antennæ, but with a carapace and reduced pleural lobes and pygidium. From this offshoot were probably derived the Ostracoda, the Cirripedia, and the various orders of the Malacostraca, with the possible exception of the Isopoda. I have suggested independent origins of the Copepoda and Isopoda, but realize the weighty arguments which can be adduced against such an interpretation.
Fig. 36.—Naraoia compactaWalcott. An outline of the test, after Walcott. Natural size.
Fig. 37.—Pagetia clytiaWalcott. An eodiscid with compound eyes. After Walcott. × 5.
Fig. 38.—Asaphiscus wheeleriMeek. A representative trilobite of the Middle Cambrian of the Pacific province. After Meek. × 1/2.
Fig. 39.—Pædeumias robsonensisBurling. Restored from a photograph published by Burling. × 1/4.
Fig. 40.—Robergiasp. Restored from fragments found in the Athens shale (Lower Middle Ordovician), at Saltville, Va. Natural size.
It is customary to speak of the Crustacea and Trilobita as having had a common ancestry, rather than the former being in direct line of descent from the latter, but when it can be shown that the higher Crustacea are all derivable from the Trilobita, and that they possess no characteristics which need have been inherited from any other source than that group, it seems needless to postulate the evolution of the same organs along two lines of development.
I can not go into the question of which are more primitive, sessile or stalked eyes, but considering the various types found among the trilobites, one can but feel that the stalked eyes are not the most simple. While no trilobite had movable stalked eyes, it is possible to homologize free cheeks with such structures. They always bear the visual surface, and, in certain trilobites (Cyclopyge), the entire cheek is broken up into lenses. Since a free cheek is a separate entity, it is conceivable that it might lie modified into a movable organ.
EVOLUTION OF THE MEROSTOMATA.
It has been pointed out above that the Limulava (Sidneyia,Amiella,Emeraldella) have certain characteristics in common with the trilobites on the one hand and the Eurypterida on the other. These relationships have been emphasized by Walcott, who derives the Eurypterida through the Limulava and the Aglaspina from the Trilobita. The Limulava may be derived from the Trilobita, but indicate a line somewhat different from that of the remainder of the Crustacea. In this line the second cephalic appendages do not become antennæ. and the axial lobe seems to broaden out, so that the pleural lobes become an integral part of the body. As in the modern Crustacea, the pygidium is reduced to the anal plate, and this grows out into a spine-like telson.
From the Limulava to the Eurypterida is a long leap, and before it can be made without danger, many intermediate steps must be placed in position. The direct ancestor of the Eurypterida is certainly not to be seen in the highly specializedSidneyia, and probably not inEmeraldella, but it might be sought in a related form with a few more segments. The few species now known do suggest the beginning of a grouping of appendages about the mouth, a suppression of appendages on the abdomen, and a development of gills on the thorax only. Further than that the route is uncertain.
Clarke and Ruedemann, whose recent extensive studies give their opinion much weight, seem fully convinced that the Merostomata could not have been derived from the Trilobita, but are rather inclined to agree with Bernard that the arachnids and the crustaceans were derived independently from similar chætopod annelids (1912, p. 148).
The greater part of their work was, however, finished before 1910, and although they refer to Walcott's description of the Limulava (1911), they did not have the advantage of studying the wonderful series of Crustacea described by him in 1912. While the evidenceis far from clear, it would appear that the discovery of animals with the form of Limiting and the eurypterids and the appendages of trilobites means something more than descent from similar ancestors. Biramous limbs of the type found in the trilobites would probably not be evolved independently on two lines, even if the ancestral stocks were of the same blood.
The Aglaspidæ, as represented byMolariaandHabeliain the Middle Cambrian, are quite obvious closely related to the trilobites easily derived from them, and retain numerous of their characteristics. That they are not trilobites is, however, shown by the presence of two pairs of antennæ, the absence of facial sutures, and the possession of a spine-like telson.
The Aglaspidæ have always been placed in the Merostomata, and nearer the Limulidæ than the Eurypterida. The discovery of appendages does not at all tend to strengthen that view, but indicates rather that they are true Crustacea which have not given rise to any group now known. The exterior form is, however,Limulus-like, and since it is known from ontogeny that the ancestor of that genus was an animal with free body segments, there is still a temptation to try to see in the Aglaspidæ the progenitors of the limulids.
The oldest knownLimulus-like animal other than the Aglaspidæ isNeolimulus falcatusWoodward (Geol. Mag., dec. 1, vol. 5, 1868, p. 1, pl. 1, fig. 1). The structure of the head of this animal is typically limuloid, with simple and compound eyes and even the ophthalmic ridges. Yet, curiously enough, it shows what in a trilobite would be considered the posterior half of the facial suture, running from the eye to the genal angle. The body is composed of eight free segments with the posterior end missing.Belinurus, from the Mississippian and Pennsylvanian, has a sort of pygidium, the posterior three segments being fused together, andPrestwichiaof the Pennsylvanian has all the segments of the abdomen fused together. So far as form goes, a very good series of stages can be selected, from the Aglaspidæ of the Cambrian throughNeolimulusto the Belinuridæ of the late Palæozoic and the Limulidæ of the Mesozoic to recent. Without much more knowledge of the appendages than is now available, it would be quite impossible to defend such a line. It is, however, suggestive.
EVOLUTION OF THE "TRACHEATA."
The trilobites were such abundant and highly variable animals, adapting themselves to various methods of life in the sea, that it appears highly probably that some of them may have become adapted to life on the land. The ancestors of the Chilopoda, Diplopoda, and Insecta appear to have been air-breathing animals as early as the Cambrian, or at latest, the Ordovician. Since absolutely nothing is yet known of the land or even of the fresh-water life of those periods, nothing can now be proved.
In discussing the relationship of the trilobites to the various tracheate animals, I have pointed out such palæontologic evidence as I have been able to gather. Studies in the field of comparative morphology do not fall within my province. I only hope to have made the structure of the trilobite a little more accessible to the student of phylogenies.
SUMMARY ON LINES OF DESCENT.
In order to put into graphic and concise form the suggestions made above, it is necessary to define and give names to some of the groups outlined. The hypothetical ancestorneed not be included in the classification and for reasons of convenience may be referred to merely as the Protostracean.
The group of free-swimming trilobites without thoracic segments was probably a large one, and within it there were doubtless considerable variations and numerous adaptations. While the only known animal which could possibly be referred to this group,Naraoia, is blind, it is entirely possible that other species had eyes, and that the cephala and pygidia were variously modified. For this reason and because of the lack of all thoracic segments, it seems better to erect a new order rather than merely a family for the group, andNektaspia(swimming shields) may be suggested. The only known family is Naraoidæ Walcott, which must be redefined.
MarrellaandHabeliaare types of Crustacea which can neither be placed in the Trilobita nor in any of the established subclasses of the Eucrustacea. They represent a transitional group, the members of which are, so far as known, adapted to the crawling mode of life, though it may prove that there are also swimmers which can be classified with them. To this subclass the nameHaplopodamay be applied, the feet being simple.
The two known families, Marrellidæ Walcott and Aglaspidæ Clarke, belong to different orders, the second having already the name Aglaspina Walcott. The nameMarrellinamay therefore be used for the other.
ForSidneyia, Walcott proposed the new subordinal name Limulava, placing it under the Eurypterida. WhileSidneyia,Emeraldella, andAmiellamay belong to the group that gave rise to the Eurypterida, they are themselves Crustacea, and a place must be found for them in that group. The possession of only one pair of antennæ prevents their reception by the Haplopoda, and allies them to the Trilobita, but the modifications of the trunk and its appendages keep them out of that subclass, and a new one has to be erected for them. This may be known as theXenopoda, in allusion to the strange appendages ofSidneyia.