It is not desirable to occupy the limited space of this article by a full description of the limbs and segments of Limulus and Scorpio. The reader is referred to the complete series of figures here given, with their explanatory legends (figs 12, 13, 14, 15). Certain matters, however, require comment and explanation to render the comparison intelligible. The tergites, or chitinized dorsal halves of the body rings, are fused to form a “prosomatic carapace,” or carapace of the prosoma, in both Limulus and Scorpio (see figs. 7 and 8). This region corresponds in both cases to six somites, as indicated by the presence of six pairs of limbs. On the surface of the carapace there are in both animals a pair of central eyes with simple lens and a pair of lateral eye-tracts, which in Limulus consist of closely-aggregated simple eyes, forming a “compound” eye, whilst in Scorpio they present several separate small eyes. The microscopic structure of the central and the lateral eyes has been shown by Lankester and A.G. Bourne (5) to differ; but the lateral eyes of Scorpio were shown by them to be similar in structure to the lateral eyes of Limulus, and the central eyes of Scorpio to be identical in structure with the central eyes of Limulus (see below).Fig.4.—Ventral surface of the same entosternum as that drawn in fig. 3. Letters as in fig. 3 with the addition of NC, neural canal or foramen.(After Lankester,loc. cit.)Fig.5.—Entosternum of one of the mygalomorphous spiders; ventral surface. Ph.N., pharyngeal notch. The posterior median process with its repetition of triangular segments closely resembles the same process in Limulus.(From Lankester,loc. cit.)Fig.6.—Dorsal surface of the same entosternum as that drawn in fig. 5. Ph.N., pharyngeal notch.After Lankester,loc. cit.Following the prosoma is a region consisting of six segments (figs. 14 and 15), each carrying a pair of plate-like appendages in both Limulus and Scorpio. This region is called the mesosoma. The tergites of this region and those of the following region, the metasoma, are fused to form a second or posterior carapace in Limulus, whilst remaining free in Scorpio. The first pair of foliaceous appendages in each animal is the genital operculum; beneath it are found the openings of the genital ducts. The second pair of mesosomatic appendages in Scorpio are known as the “pectens.” Each consists of an axis, bearing numerous blunt tooth-like processes arranged in a series. This is represented in Limulus by the first gill-bearing appendage. The leaves (some 150 in number) of the gill-book (see figure) correspond to the tooth-like processes of the pectens of Scorpio. The next four pairs of appendages (completing the mesosomatic series of six) consist, in both Scorpio and Limulus, of a base carrying each 130 to 150 blood-holding, leaf-like plates, lying on one another like the leaves of a book. Their minute structure is closely similar in the two cases; the leaf-like plates receive blood from the great sternal sinus, and serve as respiratory organs. The difference between the gill-books of Limulus and the lung-books of Scorpio depends on the fact that the latter are adapted to aerial respiration, while the former serve for aquatic respiration. The appendage carrying the gill-book stands out on the surface of the body in Limulus, and has other portions developed besides the gill-book and its base; it is fused with its fellow of the opposite side. On the other hand, in Scorpio, the gill-book-bearing appendage has sunk below the surface, forming a recess or chamber for itself, which communicates with the exterior by an oval or circular “stigma” (fig. 10,stg). That this in-sinking has taken place, and that the lung-books or in-sunken gill-books of Scorpio really represent appendages (that is to say, limbs or parapodia) is proved by their developmental history (seefigs. 17 and 18). They appear at first as outstanding processes on the surface of the body.The exact mode in which the in-sinking of superficial outstanding limbs, carrying gill-lamellae, has historically taken place has been a matter of much speculation. It was to be hoped that the specimen of the Silurian scorpion (Palaeophonus) from Scotland, showing the ventral surface of the mesosoma (fig. 49), would throw light on this matter; but the specimen recently carefully studied by the writer and Pocock reveals neither gill-bearing limbs nor stigmata. The probability appears to be against an actual introversion of the appendage and its lamellae, as was at one time suggested by Lankester. It is probable that such an in-sinking as is shown in the accompanying diagram has taken place (fig. 15); but we are yet in need of evidence as to the exact equivalence of margins, axis, &c., obtaining between the lung-book of Scorpio and the gill-book of Limulus. Zoologists are familiar with many instances (fishes, crustaceans) in which the protective walls of a water-breathing organ or gill-apparatus become converted into an air-breathing organ or lung, but there is no other case known of the conversion of gill processes themselves into air-breathing plates.Fig.7.—Diagram of the dorsal surface ofLimulus polyphemus.oc, Lateral compound eyes.oc′, Central monomeniscous eyes.PA, Post-anal spine.I to VI, The six appendage-bearing somites of the prosoma.VII, Usually considered to be the tergum of the genital somite, but suggested by Pocock to be that of the otherwise suppressed praegenital somite.VIII to XIII, The six somites of the mesosoma, each with a movable pleural spine and a pair of dorsal entopophysis or muscle-attaching ingrowths.XIV to XVIII, The confluent or unexpressed six somites of the metasoma.[According to the system of numbering explained in the text, if VII is the tergum of the praegenital somite (as is probable) it should be labelledPrgwithout any number, and the somites VIII to XIII should be lettered 1 to 6, indicating that they are the six normal somites of the mesosoma; whilst XV to XVIII should be replaced by the numbers 7 to 12—an additional suppressed segment (making up the typical six) being reckoned to the metasomatic fusion.](From Lankester,Q.J. Micr Set. vol. xxi., 1881.)Fig.8.—Diagram of the dorsal surface of a scorpion to compare with fig. 7. Letters and Roman numerals as in fig. 7, excepting that VII is here certainly the tergum of the first somite of the mesosoma—the genital somite—and isnota survival of the embryonic praegenital somite. The anus (not seen) is on the sternal surface.(From Lankester,loc. cit.)The identification of the lung-books of Scorpio with the gill-books of Limulus is practically settled by the existence of the pectens in Scorpio (fig. 14, VIII) on the second mesosomatic somite. There is no doubt thattheseare parapodial or limb appendages, carrying numerous imbricated secondary processes, and therefore comparable in essential structure to the leaf-bearing plates of the second mesosomatic somite of Limulus. They have remained unenclosed and projecting on the surface of the body, as once were the appendages of the four following somites. But they have lost their respiratory function. In non-aquatic life such an unprotected organ cannot subserve respiration. The “pectens” have become more firmly chitinized and probably somewhat altered in shape as compared with their condition in the aquatic ancestral scorpions. Their present function in scorpions is not ascertained. They are not specially sensitive under ordinary conditions, and may be touched or even pinched without causing any discomfort to the scorpion. It is probable that they acquire special sensibility at the breeding season and serve as “guides” in copulation. The shape of the legs and the absence of paired terminal claws in the SilurianPalaeophonus(see figs. 48 and 49) as compared with living scorpions (see fig. 10) show that the early scorpions were aquatic, and we may hope some day in better-preserved specimens than the two as yet discovered, to find the respiratory organs of those creatures in the condition of projecting appendages serving aquatic respiration somewhat as in Limulus, though not necessarily repeating the exact form of the broad plates of Limulus.It is important to note that the series of lamellae of the lung-book and the gill-book correspondexactlyin structure, the narrow, flat blood-space in the lamellae being interrupted by pillar-like junctions of the two surfaces in both cases (see Lankester (4)), and the free surfaces of the adjacent lamellae being covered with a very delicate chitinous cuticle which is drawn out into delicate hairs and processes. The elongated axis which opens at the stigma in Scorpio and which can be cleared of soft, surrounding tissues and coagulated blood so as to present the appearance of a limb axis carrying the book-like leaves of the lung is not really, as it would seem to be at first sight, the limb axis. That is necessarily a blood-holding structure and is obliterated and fused with soft tissues of the sternal region so that the lamellae cannot be detached and presented as standing out from it. The apparent axis or basal support of the scorpion’s lung-books shown in the figures, is a false or secondary axis and merely a part of the infolded surface which forms the air-chamber. The maceration of the soft parts of a scorpion preserved in weak spirit and the cleaning of the chitinized in-grown cuticle give rise to the false appearance of a limb axis carrying the lamellae. The margins of the lamellae of the scorpion’s lung-book, which arelowermostin the figures (fig. 15) and appear to be free, are really those which are attached to the blood-holding axis. The true free ends are those nearest the stigma.Passing on now from the mesosoma we come in Scorpio to the metasoma of six segments, the first of which is broad whilst the rest are cylindrical. The last is perforated by the anus and carries the post-anal spine or sting. The somites of the metasoma carry no parapodia. In Limulus the metasoma is practically suppressed. In the allied extinct Eurypterines it is well developed, and resembles that of Scorpio. In the embryo Limulus (fig. 42) the six somites of the mesosoma are not fused to form a carapace at an early stage, and they are followed by three separately marked metasomatic somites; the other three somites of the metasoma have disappeared in Limulus, but are represented by the unsegmented prae-anal region. It is probable that we have in the metasoma of Limulus a case of the disappearance of once clearly demarcated somites. It would be possible to suppose, on the other hand, that new somites are only beginning to make their appearance here. The balance of various considerations is against the latter hypothesis. Following the metasoma in Limulus, we have as in Scorpio the post-anal spine—in this case not a sting, but a powerful and important organ of locomotion, serving to turn the animal over when it has fallen upon its back. The nature of the post-anal spine has been strangely misinterpreted by some writers. Owen (7) maintained that it represented a number of coalesced somites, regardless of its post-anal position and mode of development. The agreement of the grouping of the somites, of the form of the parapodia (appendages, limbs) in each region, of the position of the genital aperture and operculum, of the position and character of the eyes, and of the powerful post-anal spines not seen in other Arthropods, is very convincing as to the affinityof Limulus and Scorpio. Perhaps the most important general agreement of Scorpio compared with Limulus and the Eurypterines is the division of the body into the three regions (or tagmata)—prosoma, mesosoma and metasoma—each consisting of six segments, the prosoma having leg-like appendages, the mesosoma having foliaceous appendages, and the metasoma being destitute of appendages.Fig.9.—Ventral view of the posterior carapace or meso-metasomatic (opisthospmatic) fusion ofLimulus polyphemus. The soft integument and limbs of the mesosoma have been removed as well as all the viscera and muscles, so that the inner surface of the terga of these somites with their entopophyses are seen. The unsegmented dense chitinous sternal plate of the metasoma (XIII to XVIII) is not removed. Letters as in fig. 7.After Lankester,loc. cit.)Fig.10.—Ventral view of a scorpion,Palamnaeus indus, de Geer, to show the arrangement of the coxae of the limbs, the sternal elements, genital plate and pectens.M, Mouth behind the oval median camerostome.I, The chelicerae.II, The chelae.III to VI, the four pairs of walking legs.VIIgo, The genital somite or first somite of the mesosoma with the genital operculum (a fused pair of limbs).VIIIp, The pectiniferous somite.IXstgto XIIstg, the four pulmonary somites.met, The pentagonal metasternite of the prosoma behind all the coxae.x, The sternum of the pectiniferous somite.y, The broad first somite of the metasoma.Fig.11.—Third leg ofLimulus polyphemus, showing the division of the fourth segment of the leg by a groove S into two, thus giving seven segments to the leg as in scorpion.(From a drawing by Pocock.)In 1893, some years after the identification of the somites of Limulus with those of Scorpio, thus indicated, had been published, zoologists were startled by the discovery by a Japanese zoologist, Kishinouye (8), of a seventh prosomatic somite in the embryo of Limulus longispina. This was seen in longitudinal sections, as shown in fig. 19. The simple identification of somite with somite in Limulus and Scorpio seemed to be threatened by this discovery. But in 1896 Dr August Brauer of Marburg (9) discovered in the embryo of Scorpio a seventh prosomatic somite (see VII PrG, figs. 17 and 18), or, if we please so to term it, apraegenitalsomite, hitherto unrecognized. In the case of Scorpio this segment is indicated in the embryo by the presence of a pair of rudimentary appendages, carried by a well-marked somite. As in Limulus, so in Scorpio, this unexpected somite and its appendages disappear in the course of development. In fact, more or less complete “excalation” of the somite takes place. Owing to its position it is convenient to term the somite which is excalated in Limulus and Scorpio “the praegenital somite.” It appears not improbable that the sternal plates wedged in between the last pair of legs in both Scorpio and Limulus, viz. the pentagonal sternite of Scorpio (fig. 10) and the chilaria of Limulus (see figs. 13 and 20), may in part represent in the adult the sternum of the excalated praegenital somite. This has not been demonstrated by an actual following out of the development, but the position of these pieces and the fact that they are (in Limulus) supplied by an independent segmental nerve, favours the view that they may comprise the sternal area of the vanished praegenital somite. This interpretation, however, of the “metasternites” of Limulus and Scorpio is opposed by the coexistence in Thelyphonus (figs. 55, 57 and 58) of a similar metasternite with a complete praegenital somite. H.J. Hansen (10) has recognized that the “praegenital somite” persists in a rudimentary condition, forming a “waist” to the series of somites in the Pedipalpi and Araneae. The present writer is of opinion that it will be found most convenient to treat this evanescent somite as something special, and not to attempt to reckon it to either the prosoma or the mesosoma. These will then remain as typically composed each of six appendage-bearing somites-the prosoma comprising in addition the ocular prosthomere.1When the praegenital somite or traces of it are present it should not be called “the seventh prosomatic” or the “first mesosomatic,” but simply the “praegenital somite.” The first segment of the mesosoma of Scorpio and Limulus thus remains the first segment, and can be identified as such throughout the Eu-arachnida, carrying as it always does the genital apertures. But it is necessary to remember, in the light of recent discoveries, that the sixth prosomatic pair of appendages is carried on the seventh somite of the whole series, there being two prosthomeres or somites in front of the mouth, the first carrying the eyes, the second the chelicerae; also that the first mesosomatic or genital somite is not the seventh or even the eighth of the whole senes of somites which have been historically present, but is the ninth, owing to the presence or to the excalation of a praegenital somite. It seems that confusion and trouble will be best avoided by abstaining from the introduction of the non-evident somites, the ocular and the praegenital, into the numerical nomenclature of the component somites of the three great body regions. We shall, therefore, ignoring the ocular somite, speak of the first, second, third, fourth, fifth and sixth leg-bearing somites of the prosoma, and indicate the appendages by the Roman numerals, I, II, III, IV, V, VI, and whilst ignoring the praegenital somite we shall speak of the first, second, third, &c., somite of the mesosoma or opisthosoma (united mesosoma and metasoma) and indicate them by the Arabic numerals.There are a number of other important points of structure besides those referring to the somites and appendages in which Limulus agrees with Scorpio or other Arachnida and differs from other Arthropoda. The chief of these are as follows:—1.The Composition of the Head(that is to say, of the anterior part of the prosoma)with especial Reference to the Region in Front of the Mouth.—It appears (seeArthropoda) that there is embryological evidence of the existence of two somites in Arachnida which were originally post-oral, but have become prae-oral by adaptational shifting of the oral aperture. These forwardly-slipped somites are called “prosthomeres.” The first of these has, in Arachnids as in other Arthropods, its pair of appendages represented by the eyes. The second has for its pair of appendages the small pair of limbs which in all living Arachnids is either chelate or retrovert (as in spiders), and is known as the chelicerae. It is possible, as maintained by some writers (Patten and others), that the lobes of the cerebral nervous mass in Arachnids indicate a larger number of prosthomeres as having fused in this region, but there is noembryologicalevidence at present which justifies us in assuming the existence in Arachnids of more than two prosthomeres. The position of the chelicerae of Limulus and of the ganglionic nerve-masses from which they receive their nerve-supply, is closely similar to that of the same structures in Scorpio. The cerebral mass is in Limulus more easily separated by dissection as a median lobe distinct from the laterally-placed ganglia of the chelceral somite than is the case in Scorpio, but the relations are practically the same in the two forms. Formerly it was supposed that in Limulus both the chelicerae and the next following pair of appendages were prosthomerous, as in Crustacea, but the dissections of Alphonse Milne-Edwards (6) demonstratedthe true limitations of the cerebrum, whilst embryological researches have done as much for Scorpio. Limulus thus agrees with Scorpio and differs from the Crustacea, in which there are three prosthomeres—one ocular and two carrying palpiform appendages. It is true that in the lower Crustacea (Apus, &c.) we have evidence of the gradual movement forward of the nerve-ganglia belonging to these palpiform appendages. But although in such lower Crustacea the nerve-ganglia of the third prosthomere have not fused with the anterior nerve-mass, there is no question as to the prae-oral position of two appendage-bearing somites in addition to the ocular prosthomere. The Crustacea have, in fact, three prosthomeres in the head and the Arachnida only two, and Limulus agrees with the Arachnida in this respect and differs from the Crustacea. The central nervous systems of Limulus and of Scorpio present closer agreement in structure than can be found when a Crustacean is compared with either. The wide divarication of the lateral cords in the prosoma and their connexion by transverse commissures, together with the “attraction” of ganglia to the prosomatic ganglion group which properly belong to hinder segments, are very nearly identical in the two animals. The form and disposition of the ganglion cells are also peculiar and closely similar in the two. (See Patten (42) for important observations on the neuromeres, &c., of Limulus and Scorpio.)Fig.12.—The prosomatic appendages ofLimulus polyphemus(right) and Scorpio (left),Palamnaeus induscompared. The corresponding appendages are marked with the same Roman numeral. The Arabic numerals indicate the segments of the legs.cox, Coxa or basal segment of the leg.stc, The sterno-coxal process or jaw-like up-growth of the coxa.epc, The articulated movable outgrowth of the coxa, called the epi-coxite (present only in III of the scorpion and III, IV and V of Limulus).ex1, The exopodite of the sixth limb of Limulus.a, b, c, d, Movable processes on the same leg (see for some suggestions on the morphology of this leg, Pocock inQuart. Journ. Micr. Sci.March 1901; see also fig. 50 below and explanation).(From Lankester,loc. cit.)Fig.13.—Diagrams of the metasternitest, with genital operculumop, and the first lamelligerous pair of appendagesga, with uniting sternal elementstof Scorpio (left) and Limulus (right).(From Lankester,loc. cit.)2.The Minute Structure of the Central Eyes and of the Lateral Eyes.—Limulus agrees with Scorpio not only in having a pair of central eyes and also lateral eyes, but in the microscopic structure of those organs, which differs in the central and lateral eyes respectively. The central eyes are “simple eyes,” that is to say, have a single lens, and are hence called “monomeniscous.” The lateral eyes are in Limulus “compound eyes,” that is to say, consist of many lenses placed close together; beneath each lens is a complex of protoplasmic cells, in which the optic nerve terminates. Each such unit is termed an “ommatidium.” The lateral eyes of Scorpio consist of groups of separate small lenses each with its ommatidium, but they do not form a continuous compound eye as in Limulus. The ommatidium (soft structure beneath the lens-unit of a compound eye) is very simple in both Scorpio and Limulus. It consists of a single layer of cells, continuous with those which secrete the general chitinous covering of the prosoma. The cells of the ommatidium are a good deal larger than the neighbouring common cells of the epidermis. They secrete the knob-like lens (fig. 22). But they also receive the nerve fibres of the optic nerve. They are at the same time both optic nerve-end cells, that is to say, retina cells, and corneagen cells or secretors of the chitinous lens-like cornea. In Limulus (fig. 23) each ommatidium has a peculiar ganglion cell developed in a central position, whilst the ommatidium of the lateral eyelets of Scorpio shows small intermediate cells between the larger nerve-end cells. The structure of the lateral eye of Limulus was first described by Grenacher, and further and more accurately by Lankester and Bourne (5) and by Watase; that of Scorpio by Lankester and Bourne, who showed that the statements of von Graber were erroneous, and that the lateral eyes of Scorpio have a single cell-layered or “monostichous” ommatidium like that of Limulus. Watase has shown, in a very convincing way, how by deepening the pit-like set of cells beneath a simple lens the more complex ommatidia of the compound eyes of Crustacea and Hexapoda may be derived from such a condition as that presented in the lateral eyes of Limulus and Scorpio. (For details the reader is referred to Watase (11) and to Lankester and Bourne (5).) The structure of the central eyes of Scorpio and spiders and also of Limulus differs essentially from that of the lateral eyes in having two layers of cells (hence called diplostichous) beneath the lens, separated from one another by a membrane (figs. 24 and 25). The upper layer is the corneagen and secretes the lens, the lower is the retinal layer. The mass of soft cell-structures beneath a large lens of a central eye is called an “ommatoeum.” It shows in Scorpio and Limulus a tendency to segregate into minor groups or “ommatidia.” It is found that in embryological growth the retinal layer of the central eyes forms as a separate pouch, which is pushed in laterally beneath the corneagen layer from the epidermic cell layer. Hence it is in origin double, and consists of a true retinal layer and a post-retinal layer (fig. 24, B), though these are not separated by a membrane. Accordingly the diplostichous ommatoeum or soft tissue of the Arachnid’s central eye should strictly be called “triplostichous,” since the deep layer is itself doubled or folded. The retinal cells of both the lateral and central eyes of Limulus and Scorpio produce cuticular structures on their sides; each such piece is a rhabdomere and a number (five or ten) uniting form a rhabdom (fig. 26). In the specialized ommatidia of the compound eyes of Crustacea and Hexapods the rhabdom is an important structure.2It is a very significant fact that the lateral and central eyes of Limulus and Scorpio not only agree each with each in regard to their monostichous and diplostichous structure, but also in the formation in both classes of eyes of rhabdomeres and rhabdoms in which the component pieces are five or a multiple of five (fig. 26). Whilst each unit of the lateral eye of Limulus has a rhabdom of ten3piecesforming a star-like chitinous centre in section, each lateral eye of Scorpio has several rhabdoms of five or less rhabdomeres, indicating that the Limulus lateral eye-unit is more specialized than the detached lateral eyelet of Scorpio, so as to present a coincidence of one lens with one rhabdom. Numerous rhabdomeres (grouped as rhabdoms in Limulus) are found in the retinal layer of the central eyes also.Fig.14.—The first three pairs of mesosomatic appendages of Scorpio and Limulus compared.VII, The genital operculum.VIII, The pectens of Scorpio and the first branchial plate of Limulus.IX, The first pair of lung-books of Scorpio and the second branchial plate of Limulus.gp, Genital pore.epst, Epistigmatic sclerite.stg, Stigma or orifice of the hollow tendons of the branchial plates of Limulus.(After Lankester,loc. cit.)Whilst Limulus agrees thus closely with Scorpio in regard to the eyes, it is to be noted that no Crustacean has structures corresponding to the peculiar diplostichous central eyes, though these occur again (with differences in detail) inHexapoda. Possibly, however, an investigation of the development of the median eyes of some Crustacea (Apus, Palaemon) may prove them to be diplostichous in origin.3.The so-called“Coxal Glands.”—In 1882 (Proc. Roy. Soc. No. 221) Lankester described under the name “coxal glands” a pair of brilliantly white oviform bodies lying in the Scorpion’s prosoma immediately above the coxae of the fifth and sixth pairs of legs (fig. 27). These bodies had been erroneously supposed by Newport (12) and other observers to be glandular outgrowths of the alimentary canal. They are really excretory glands, and communicate with the exterior by a very minute aperture on the posterior face of the coxa of the fifth limb on each side. When examined with the microscope, by means of the usual section method, they are seen to consist of a labyrinthine tube lined with peculiar cells, each cell having a deep vertically striated border on the surface farthest from the lumen, as is seen in the cells of some renal organs. The coils and branches of the tube are packed by connective tissue and blood spaces. A similar pair of coxal glands, lobate instead of ovoid in shape, was described by Lankester in Mygale, and it was also shown by him that the structures in Limulus called “brick-red glands” by Packard have the same structure and position as the coxal glands of Scorpio and Mygale. In Limulus these organs consist each of four horizontal lobes lying on the coxal margin of the second, third, fourth, and fifth prosomatic limbs, the four lobes being connected to one another by a transverse piece or stem (fig. 28). Microscopically their structure is the same in essentials as that of the coxal glands of Scorpio (13). Coxal glands have since been recognized and described in other Arachnida. In 1900 it was shown that the coxal gland of Limulus is provided with a very delicate thin-walled coiled duct which opens, even in the adult condition, by a minute pore on the coxa of the fifth leg (Patten and Hazen,13A). Previously to this, Lankester’s pupil Gulland had shown (1885) that in the embryo the coxal gland is a comparatively simple tube, which opens to the exterior in this position and by its other extremity into a coelomic space. Similar observations were made by Laurie (17) in Lankester’s laboratory (1890) with regard to the early condition of the coxal gland of Scorpio, and by Bertkau (41) as to that of the spider Atypus. H.M. Bernard (13B) showed that the opening remains in the adult scorpion. In all the embryonic or permanent opening is on the coxa of the fifth pair of prosomatic limbs. Thus an organ newly discovered in Scorpio was found to have its counterpart in Limulus.The name “coxal gland” needs to be carefully distinguished from “crural gland,” with which it is apt to be confused. The crural glands, which occur in many terrestrial Arthropods, are epidermal in origin and totally distinct from the coxal glands. The coxal glands of the Arachnida are structures of the same nature as the green glands of the higher Crustacea and the so-called “shell glands” of the Entomostraca. The latter open at the base of the fifth pair of limbs of the Crustacean, just as the coxal glands open on the coxal joint of the fifth pair of limbs of the Arachnid. Both belong to the category of “coelomoducts,” namely, tubular or funnel-like portions of the coelom opening to the exterior in pairs in each somite (potentially,) and usually persisting in only a few somites as either “urocoels” (renal organs) or “gonocoets” (genital tubes). In Peripatus they occur in every somite of the body. They have till recently been very generally identified with the nephridia of Chaetopod worms, but there is good reason for considering the true nephridia (typified by the nephridia of the earthworm) as a distinct class of organs (see Lankester in vol. ii. chap. in. ofA Treatise on Zoology, 1900). The genital ducts of Arthropoda are, like the green glands, shell glands and coxal glands, to be regarded as coelomoducts (gonocoels). The coxal glands do not establish any special connexion between Limulus and Scorpio, sincetheyalso occur in the same somite in the lower Crustacea, but it is to be noted that the coxal glands of Limulus are in minute structure and probably in function more like those of Arachnids than those of Crustacea.Fig.15.—The remaining three pairs of mesosomatic appendages of Scorpio and Limulus. Letters as in fig. 14.l130 indicates that there are 130 lamellae in the scorpion’s lung-book, whilstl150 indicates that 150 similar lamellae are counted in the gill of Limulus.(After Lankester,loc. cit.)4.The Entosternites and their Minute Structure.—Strauss-Dürckheim (1) was the first to insist on the affinity between Limulus and the Arachnids, indicated by the presence of a free suspended entosternum or plastron or entosternite in both. We have figured here (figs. 1 to 6) the entosternites of Limulus, Scorpio and Mygale. Lankester some years ago made a special study of the histology (3) of these entosternites for the purpose of comparison, and also ascertained the relations of the very numerous muscles which are inserted into them (4). The entosternites are cartilaginous in texture, but they have neither the chemical character nor the microscopic structure of the hyaline cartilage of Vertebrates. They yield chitin in place of chondrin or gelatin—as does also the cartilage of the Cephalopod’s endoskeleton. In microscopic structure they all present the closest agreement with one another. We find a firm, homogeneous or sparsely fibrillated matrix in which are embedded nucleated cells (corpuscles of protoplasm) arranged in rows of three, six or eight, parallel with the adjacent lines of fibrillation.Fig.16.—Diagram to show the way in which an outgrowing gill-process bearing blood-holding lamellae, may give rise, if the sternal body wall sinks inwards, to a lung-chamber with air-holding lamellae.I is the embryonic condition.bs, Blood sinus.L is the condition of outgrowth withgl, gill lamellae.A is the condition of in-sinking of the sternal surface and consequent enclosure of the lamelligerous surface of the appendage in a chamber with narrow orifice—the pulmonary air-holding chamber.pl, Pulmonary lamellae.bs, Blood sinus.(After Kingsley.)Fig.17.—Embryo of scorpion, ventral view showing somites and appendages.sgc, Frontal groove.sa, Rudiment of lateral eyes.obl, Camerostome (upper lip).so, Sense-organ of Patten.PrGabp1, Rudiment of the appendage of the praegenital somite which disappears.abp2, Rudiment of the right half of the genital operculum.abp3, Rudiment of the right pecten.abp4toabp7. Rudiments of the four appendages which carry the pulmonary lamellae.I to VI, Rudiments of the six limbs of the prosoma.VIIPrG, The evanescent praegenital somite.VIII, The first mesosomatic somite or genital somite.IX, The second mesosomatic somite or pectiniferous somite.X to XIII, The four pulmoniferous somites.XIV, The first metasomatic somite.(After Brauer,Zeitsch. wiss. Zool., vol. lix., 1895.)A minute entosternite having the above-described structure is found in the Crustacean Apus between the bases of the mandibles, and also in the Decapoda in a similar position, but in no Crustacean does it attain to any size or importance. On the other hand, the entosternite of the Arachnida is a very large and important featurein the structure of the prosoma, and must play an important part in the economy of these organisms. In Limulus (figs. 1 and 2) it has as many as twenty-five pairs of muscles attached to it, coming to it from the bases of the surrounding limbs and from the dorsal carapace and from the pharynx. It consists of an oblong plate 2 in. in length and 1 in breadth, with a pair of tendinous outgrowths standing out from it at right angles on each side. It “floats” between the prosomatic nerve centres and the alimentary canal. In each somite of the mesosoma is a small, free entosternite having a similar position, but below or ventral to the nerve cords, and having a smaller number of muscles attached to it. The entosternite was probably in origin part of the fibrous connective tissue lying close to the integument of the sternal surface—giving attachment to muscles corresponding more or less to those at present attached to it. It became isolated and detached, why or with what advantage to the organism it is difficult to say, and at that period of Arachnidan development the great ventral nerve cords occupied a more lateral position than they do at present. We know that such a lateral position of the nerve cords preceded the median position in both Arthropoda and Chaetopoda. Subsequently to the floating off of the entosternite the approximation of the nerve cords took place in the prosoma, and thus they were able to take up a position below the entosternite. In the mesosoma the approximation had occurred before the entosternites were formed.Fig.18.—Portion of a similar embryo at a later stage of growth. The praegenital somite, VII PrG, is still present, but has lost its rudimentary appendages;go, the genital operculum, left half; Km, the left pecten;abp4toabp7, the rudimentary appendages of the lung-sacs.(After Brauer,loc. cit.)Fig.19.—Section through an early embryo ofLimulus longispina, showing seven transverse divisions in the region of the unsegmented anterior carapace. The seventh, VII, is anterior to the genital operculum,op, and is the cavity of the praegenital somite which is more or less completely suppressed in subsequent development, possibly indicated by the area marked VII in fig. 7 and by the great entopophyses of the prosomatic carapace.(After Kishinouye,Journ. Sci. Coll. Japan, vol. v., 1892.)In the scorpion (figs. 3 and 4) the entosternite has tough membrane-like outgrowths which connect it with the body-wall, both dorsally and ventrally forming an oblique diaphragm, cutting off the cavity of the prosoma from that of the mesosoma. It was described by Newport as “the diaphragm.” Only the central and horizontal parts of this structure correspond precisely to the entosternite of Limulus: the right and left anterior processes (markedapin figs. 3 and 4, and RAP, LAP, in figs. 1 and 2) correspond in the two animals, and the median lateral processlmpof the scorpion represents the tendinous outgrowths ALR, PLR of Limulus. The scorpion’s entosternite gives rise to outgrowths, besides the great posterior flaps,pf, which form the diaphragm, unrepresented in Limulus. These are a ventral arch forming a neural canal through which the great nerve cords pass (figs. 3 and 4,snp), and further a dorsal gastric canal and arterial canal which transmit the alimentary tract and the dorsal artery respectively (figs. 3 and 4, GC, DR).In Limulus small entosternites are found in each somite of the appendage-bearing mesosoma, and we find in Scorpio, in the only somite of the mesosoma which has a well-developed pair of appendages, that of the pectens, a small entosternite with ten pairs of muscles inserted into it. The supra-pectinal entosternite lies ventral to the nerve cords.In Mygale (figs. 5 and 6) the form of the entosternite is more like that of Limulus than is that of Scorpio. The anterior notch Ph.N. is similar to that in Limulus, whilst the imbricate triangular pieces of the posterior median region resemble the similarly-placed structures of Limulus in a striking manner.It must be confessed that we are singularly ignorant as to the functional significance of these remarkable organs—the entosternites. Their movement in an upward or downward direction in Limulus and Mygale must exert a pumping action on the blood contained in the dorsal arteries and the ventral veins respectively. In Scorpio the completion of the horizontal plate by oblique naps, so as to form an actual diaphragm shutting off the cavity of the prosoma from the rest of the body, possibly gives to the organs contained in the anterior chamber a physiological advantage in respect of the supply of arterial blood and its separation from the venous blood of the mesosoma. Possibly the movement of the diaphragm may determine the passage of air into or out of the lung-sacs. Muscular fibres connected with the suctorial pharynx are in Limulus inserted into the entosternite, and the activity of the two organs may be correlated.5.The Blood and the Blood-vascular System.—The blood fluids of Limulus and Scorpio are very similar. Not only are the blood corpuscles of Limulus more like in form and granulation to those of Scorpio than to those of any Crustacean, but the fluid is in both animals strongly impregnated with the blue-coloured respiratory proteid, haemocyanin. This body occurs also in the blood of Crustacea and of Molluscs, but its abundance in both Limulus and Scorpio is very marked, and gives to the freshly-shed blood a strong indigo-blue tint.Fig.20.—View of the ventral surface of the mid-line of the prosomatic region ofLimulus polyphemus. The coxae of the five pairs of limbs following the chelicerae were arranged in a series on each side between the mouth, M, and the metasternites,mets.sf, The sub-frontal median sclerite.Ch, The chelicerae.cam, The camerostome or upper lip.M, The mouth.pmst, The promesosternal sclerite of chitinous plate, unpaired.mets, The right and left metasternites (corresponding to the similarly placed pentagonal sternite of Scorpio). Natural size.(After Lankester.)Fig.2l.—Development of the lateral eyes of a scorpion.h, Epidermic cell-layer;mes, mesoblastic connective tissue;n, nerves; II, III, IV, V, depressions of the epidermis in each of which a cuticular lens will be formed.(From Korschelt and Heider, after Laurie.)Fig.22.—Section through the lateral eye ofEuscorpius italicus.lens, Cuticular lens.nerv c, Retinal cells (nerve-end cells).rhabd, Rhabdomes.nerv f, Nerve fibres of the optic nerve.int, Intermediate cells (lying between the bases of the retinal cells).(After Lankester and Bourne from Parker and Habwell’sText book of Zoology, Macmillan & Co.)Fig.23.—Section through a portion of the lateral eye of Limulus, showing three ommatidia—A, B and C.hyp, The epidermic cell-layer (so-called hypodermis), the cells of which increase in volume below each lens,l, and become nerve-end cells or retinula-cells,rt; in A, the lettersrhpoint to a rhabdomere secreted by the cellrt;c, the peculiar central spherical cell;n, nerve fibres;mes, mesoblastic skeletal tissue;ch, chitinous cuticle.(From Korschelt and Heider after Watase.)The great dorsal contractile vessel or “heart” of Limulus is closely similar to that of Scorpio; its ostia or incurrent orifices are placed in the same somites as those of Scorpio, but there is one additional posterior pair. The origin of the paired arteries from theheart differs in Limulus from the arrangement obtaining in Scorpio, in that a pair of lateral commissural arteries exist in Limulus (as described by Alphonse Milne-Edwards (6)) leading to a suppression of the more primitive direct connexion of the four pairs of posterior lateral arteries and of the great median posterior arteries with the heart itself (fig. 29). The arterial system is very completely developed in both Limulus and Scorpio, branching repeatedly until minute arterioles are formed, not to be distinguished from true capillaries; these open into irregular swollen vessels which are the veins or venous sinuses. A very remarkable feature in Limulus, first described by Owen, is the close accompaniment of the prosomatic nerve centres and nerves by arteries, so close indeed that the great ganglion mass and its out-running nerves are actually sunk in or invested by arteries. The connexion is not so intimate in Scorpio, but is nevertheless a very close one, closer than we find in any other Arthropods in which the arterial system is well developed,e.g.the Myriapoda and some of the arthrostracous Crustacea. It seems that there is a primitive tendency in the Arthropoda for the arteries to accompany the nerve cords, and a “supra-spinal” artery—that is to say, an artery in close relation to the ventral nerve cords—has been described in several cases. On the other hand, in many Arthropods, especially those which possess tracheae, the arteries do not have a long course, but soon open into wide blood sinuses. Scorpio certainly comes nearer to Limulus in the high development of its arterial system, and the intimate relation of the anterior aorta and its branches to the nerve centres and great nerves, than does any other Arthropod.An arrangement of great functional importance in regard to the venous system must now be described, which was shown in 1883 by Lankester to be common to Limulus and Scorpio. This arrangement has not hitherto been detected in any other class than the Arachnida, and if it should ultimately prove to be peculiar to that group, would have considerable weight as a proof of the close genetic affinity of Limulus and Scorpio.Fig.24.—Diagrams of the development and adult structure of one of the paired central eyes of a scorpion.A, Early condition before the lens is deposited, showing the folding of the epidermic cell-layer into three.B, Diagram showing the nature of this infolding.C, Section through the fully formed eye.h, Epidermic cell-layer.r, The retinal portion of the same which, owing to the infolding, lies betweengl, the corneagen or lens-forming portion, andpr, the post-retinal or capsular portion or fold.l, Cuticular lens.g, Line separating lens from the lens-forming or corneagen cells of the epidermis.n, Nerve fibres.rh, Rhabdomeres.[How the inversion of the nerve-end-cells and their connexion with the nerve-fibres is to be reconciled with the condition found in the adult, or with that of the monostichous eye, has not hitherto been explained.](From Korschelt and Heider.)The great pericardial sinus is strongly developed in both animals. Its walls are fibrous and complete, and it holds a considerable volume of blood when the heart itself is contracted. Opening in pairs in each somite, right and left into the pericardial sinus are large veins, which bring the blood respectively from the gill-books and the lung-books to that chamber, whence it passes by the ostia into the heart. The blood is brought to the respiratory organs in both cases by a great venous collecting sinus having a ventral median position. In both animalsthe wall of the pericardial sinus is connected by vertical muscular bands to the wall of the ventral venous sinus(its lateral expansions around the lung-books in Scorpio) in each somite through which the pericardium passes. There are seven pairs of theseveno-pericardiac vertical musclesin Scorpio, and eight in Limulus (see figs. 30, 31, 32). It is obvious that the contraction of these musclesmust cause a depression of the floor of the pericardium and a rising of the roof of the ventral blood sinus, and a consequent increase of volume and flow of blood to each. Whether the pericardium and the ventral sinus are made to expand simultaneously or all the movement is made by one only of the surfaces concerned, must depend on conditions of tension. In any case it is clear that we have in these muscles an apparatus for causing the blood to flow differentially in increased volume into either the pericardium, through the veins leading from the respiratory organs, or from the body generally into the great sinuses which bring the blood to the respiratory organs. These muscles act so as to pump the blood through the respiratory organs.Fig.25.—Section through one of the central eyes of a young Limulus.L, Cuticular or corneous lens.hy, Epidermic cell-layer.corn, Its corneagen portion immediately underlying the lens.ret, Retinula cells.nf, Nerve fibres.con. tiss, Connective tissue (mesoblastic skeletal tissue).(After Lankester and Bourne,Q. J. Mic. Sci., 1883.)It is not surprising that with so highly developed an arterial system Limulus and Scorpio should have a highly developed mechanism for determining the flow of blood to the respiratory organs. That this is, so to speak, a need of animals with localized respiratory organs is seen by the existence of provisions serving a similar purpose in other animals,e.g.the branchial hearts of the Cephalopoda.The veno-pericardiac muscles of Scorpio were seen and figured by Newport but not described by him. Those of Limulus were described and figured by Alphonse Milne-Edwards, but he called them merely “transparent ligaments,” and did not discover their muscular structure. They are figured and their importance for the first time recognized in the memoir on the muscular and skeletal systems of Limulus and Scorpio by Lankester, Beck and Bourne (4).6.Alimentary Canal and Gastric Glands.—The alimentary canal in Scorpio, as in Limulus, is provided with a powerful suctorial pharynx, in the working of which extrinsic muscles take a part. The mouth is relatively smaller in Scorpio than in Limulus—in fact is minute, as it is in all the terrestrial Arachnida which suck the juices of either animals or plants. In both, the alimentary canal takes a straight course from the pharynx (which bends under it downwards and backwards towards the mouth in Limulus) to the anus, and is a simple, narrow, cylindrical tube (fig. 33). The only point in which the gut of Limulus resembles that of Scorpio rather than that of any of the Crustacea, is in possessing more than a single pair of ducts or lateral outgrowths connected with ramified gastric glands or gastric caeca. Limulus has two pairs of these, Scorpio as many as six pairs. The Crustacea never have more than one pair. The minute microscopic structure of the gastric glands in the two animals is practically identical. The functions of these gastric diverticula have never been carefully investigated. It is very probable that in Scorpio they do not serve merely to secrete a digestive fluid (shown in other Arthropoda to resemble the pancreatic fluid), but that they also become distended by the juices of the prey sucked in by the scorpion—as certainly must occur in the case of the simple unbranched gastric caeca of the spiders.The most important difference which exists between the structure of Limulus and that of Scorpio is found in the hinder region of the alimentary canal. Scorpio is here provided with a single or double pair of renal excretory tubes, which have been identified by earlier authors with the Malpighian tubes of the Hexapod and Myriapod insects. Limulus is devoid of any such tubes. We shall revert to this subject below.Fig.26.A, Diagram of a retinula of the central eye of a scorpion consisting of five retina-cells (ret), with adherent branched pigment cells (pig).B, Rhabdom of the same, consisting of five confluent rhabdomeres.C, Transverse section of the rhabdom of a retinula of the scorpion’s central eye, showing its five constituent rhabdomeres as rays of a star.D, Transverse section of a retinula of the lateral eye of Limulus, showing ten retinula cells (ret), each bearing a rhabdomere (rhab).(After Lankester.)Fig.27.—Diagram showing the position of the coxal glands of a scorpion,Buthus australis, Lin., in relation to the legs, diaphragm (entosternal flap), and the gastric caeca.1 to 6, The bases of the six prosomatic limbs.A, prosomatic gastric gland (sometimes called salivary).B, Coxal gland.C, Diaphragm of Newport = fibrous flap of the entosternum.D, Mesosomatic gastric caeca (so-called liver).E, Alimentary canal.(From Lankester,Q. J. Mic. Sci., vol. xxiv. N.S. p. 152.)7.Ovaries and Spermaries: Gonocoels and Gonoducts.—The scorpion is remarkable for having the specialized portion of coelom from the walls of which egg-cells or sperm-cells are developed according to sex, in the form of a simple but extensive network. It is not a pair of simple tubes, nor of dendriform tubes, but a closed network. The same fact is true of Limulus, as was shown by Owen (7) in regard to the ovary, and by Benham (14) in regard to the testis. This is a very definite and remarkable agreement, since such a reticular gonocoel is not found in Crustacea (except in the male Apus). Moreover, there is a significant agreement in the character of the spermatozoa of Limulus and Scorpio. The Crustacea are—with the exception of the Cirrhipedia—remarkable for having stiff, motionless spermatozoids. In Limulus Lankester found (15) the spermatozoa to possess active flagelliform “tails,” and to resemble very closely those of Scorpio which, as are those of most terrestrial Arthropoda, are actively motile. This is a microscopic point of agreement, but is none the less significant.In regard to the important structures concerned with the fertilization of the egg, Limulus and Scorpio differ entirely from one another.The eggs of Limulus are fertilized in the sea after they have been laid. Scorpio, being a terrestrial animal, fertilizes by copulation. The male possesses elaborate copulatory structures of a chitinous nature, and the eggs are fertilized in the female without even quitting the place where they are formed on the wall of the reticular gonocoel. The female scorpion is viviparous, and the young are produced in a highly developed condition as fully formed scorpions.Fig.28.—The right coxal gland ofLimulus polyphemus, Latr.a2toa5, Posterior borders of the chitinous bases of the coxae of the second, third, fourth and fifth prosomatic limbs.b, Longitudinal lobe or stolon of the coxal gland.c. Its four transverse lobes or outgrowths corresponding to the four coxae.(From Lankester,loc. cit., after Packard.)Differences between Limulus and Scorpio.—We have now passed in review the principal structural features in which Limulus agrees with Scorpio and differs from other Arthropoda. There remains for consideration the one important structural difference between the two animals. Limulus agrees with the majority of the Crustacea in being destitute of renal excretory caeca or tubes opening into the hinder part of the gut. Scorpio, on the other hand, in common with all air-breathing Arthropoda except Peripatus, possesses these tubules, which are often called Malpighian tubes. A great deal has been made of this difference by some writers. It has been considered by them as proving that Limulus, in spite of all its special agreements with Scorpio (which, however, have scarcely been appreciated by the writers in question), really belongs to the Crustacean line of descent, whilst Scorpio, by possessing Malpighian tubes, is declared to be unmistakably tied together with the other Arachnida to the tracheate Arthropods, the Hexapods, Diplopods, and Chilopods, which all possess Malpighian tubes.Fig.29.—Diagram of the arterial system of A, Scorpio, and B, Limulus. The Roman numerals indicate the body somites and the two figures are adjusted for comparison.ce, Cerebral arteries;sp, supra-spinal or medullary artery;c, caudal artery;l, lateral anastomotic artery of Limulus. The figure B also shows the peculiar neural investiture formed by the cerebral arteries in Limulus and the derivation from this of the arteries to the limbs, III, IV, VI, whereas in Scorpio the latter have a separate origin from the anterior aorta.From Lankester, “Limulus an Arachnid.”It must be pointed out that the presence or absence of such renal excretory tubes opening into the intestine appears to be a question of adaptation to the changed physiological conditions of respiration, and not of morphological significance, since a pair of renal excretory tubes of this nature is found in certain Amphipod Crustacea (Talorchestia, &c.) which have abandoned a purely aquatic life. This view has been accepted and supported by Professors Korschelt and Heider (16). An important fact in its favour was discovered by Laurie (17), who investigated the embryology of two species of Scorpio under Lankester’s direction. It appears that the Malpighian tubes of Scorpio are developed from the mesenteron, viz. that portion of the gut which is formed by the hypoblast, whereas in Hexapod insects the similar caecal tubes are developed from the proctodaeum or in-pushed portion of the gut which is formed from epiblast. In fact it is not possible to maintain that the renal excretory tubes of the gut are of one common origin in the Arthropoda. They have appeared independently in connexion with a change in the excretion of nitrogenous waste in Arachnids, Crustacea, and the other classes of Arthropoda when aerial, as opposed to aquatic, respiration has been established—and they have been formed in some cases from the mesenteron, in other cases from the proctodaeum. Their appearance in the air-breathing Arachnids does not separate those forms from the water-breathing Arachnids which are devoid of them, any more than does their appearance in certain Amphipoda separate those Crustaceans from the other members of the class.Further, it is pointed out by Korschelt and Heider that the hinder portion of the gut frequently acts in Arthropoda as an organ of nitrogenous excretion in the absence of any special excretory tubules, and that the production of such caeca from its surface in separate lines of descent does not involve any elaborate or unlikely process of growth. In other words, the Malpighian tubes of the terrestrial Arachnida arehomoplasticwith those of Hexapoda and Myriapoda, and nothomogeneticwith them. We are compelled to take a similar view of the agreement between the tracheal air-tubes of Arachnida and other tracheate Arthropods. They are homoplasts (see18) one of another, and do not owe their existence in the various classes compared to a common inheritance of an ancestral tracheal system.
It is not desirable to occupy the limited space of this article by a full description of the limbs and segments of Limulus and Scorpio. The reader is referred to the complete series of figures here given, with their explanatory legends (figs 12, 13, 14, 15). Certain matters, however, require comment and explanation to render the comparison intelligible. The tergites, or chitinized dorsal halves of the body rings, are fused to form a “prosomatic carapace,” or carapace of the prosoma, in both Limulus and Scorpio (see figs. 7 and 8). This region corresponds in both cases to six somites, as indicated by the presence of six pairs of limbs. On the surface of the carapace there are in both animals a pair of central eyes with simple lens and a pair of lateral eye-tracts, which in Limulus consist of closely-aggregated simple eyes, forming a “compound” eye, whilst in Scorpio they present several separate small eyes. The microscopic structure of the central and the lateral eyes has been shown by Lankester and A.G. Bourne (5) to differ; but the lateral eyes of Scorpio were shown by them to be similar in structure to the lateral eyes of Limulus, and the central eyes of Scorpio to be identical in structure with the central eyes of Limulus (see below).
Following the prosoma is a region consisting of six segments (figs. 14 and 15), each carrying a pair of plate-like appendages in both Limulus and Scorpio. This region is called the mesosoma. The tergites of this region and those of the following region, the metasoma, are fused to form a second or posterior carapace in Limulus, whilst remaining free in Scorpio. The first pair of foliaceous appendages in each animal is the genital operculum; beneath it are found the openings of the genital ducts. The second pair of mesosomatic appendages in Scorpio are known as the “pectens.” Each consists of an axis, bearing numerous blunt tooth-like processes arranged in a series. This is represented in Limulus by the first gill-bearing appendage. The leaves (some 150 in number) of the gill-book (see figure) correspond to the tooth-like processes of the pectens of Scorpio. The next four pairs of appendages (completing the mesosomatic series of six) consist, in both Scorpio and Limulus, of a base carrying each 130 to 150 blood-holding, leaf-like plates, lying on one another like the leaves of a book. Their minute structure is closely similar in the two cases; the leaf-like plates receive blood from the great sternal sinus, and serve as respiratory organs. The difference between the gill-books of Limulus and the lung-books of Scorpio depends on the fact that the latter are adapted to aerial respiration, while the former serve for aquatic respiration. The appendage carrying the gill-book stands out on the surface of the body in Limulus, and has other portions developed besides the gill-book and its base; it is fused with its fellow of the opposite side. On the other hand, in Scorpio, the gill-book-bearing appendage has sunk below the surface, forming a recess or chamber for itself, which communicates with the exterior by an oval or circular “stigma” (fig. 10,stg). That this in-sinking has taken place, and that the lung-books or in-sunken gill-books of Scorpio really represent appendages (that is to say, limbs or parapodia) is proved by their developmental history (seefigs. 17 and 18). They appear at first as outstanding processes on the surface of the body.
The exact mode in which the in-sinking of superficial outstanding limbs, carrying gill-lamellae, has historically taken place has been a matter of much speculation. It was to be hoped that the specimen of the Silurian scorpion (Palaeophonus) from Scotland, showing the ventral surface of the mesosoma (fig. 49), would throw light on this matter; but the specimen recently carefully studied by the writer and Pocock reveals neither gill-bearing limbs nor stigmata. The probability appears to be against an actual introversion of the appendage and its lamellae, as was at one time suggested by Lankester. It is probable that such an in-sinking as is shown in the accompanying diagram has taken place (fig. 15); but we are yet in need of evidence as to the exact equivalence of margins, axis, &c., obtaining between the lung-book of Scorpio and the gill-book of Limulus. Zoologists are familiar with many instances (fishes, crustaceans) in which the protective walls of a water-breathing organ or gill-apparatus become converted into an air-breathing organ or lung, but there is no other case known of the conversion of gill processes themselves into air-breathing plates.
oc, Lateral compound eyes.
oc′, Central monomeniscous eyes.
PA, Post-anal spine.
I to VI, The six appendage-bearing somites of the prosoma.
VII, Usually considered to be the tergum of the genital somite, but suggested by Pocock to be that of the otherwise suppressed praegenital somite.
VIII to XIII, The six somites of the mesosoma, each with a movable pleural spine and a pair of dorsal entopophysis or muscle-attaching ingrowths.
XIV to XVIII, The confluent or unexpressed six somites of the metasoma.
The identification of the lung-books of Scorpio with the gill-books of Limulus is practically settled by the existence of the pectens in Scorpio (fig. 14, VIII) on the second mesosomatic somite. There is no doubt thattheseare parapodial or limb appendages, carrying numerous imbricated secondary processes, and therefore comparable in essential structure to the leaf-bearing plates of the second mesosomatic somite of Limulus. They have remained unenclosed and projecting on the surface of the body, as once were the appendages of the four following somites. But they have lost their respiratory function. In non-aquatic life such an unprotected organ cannot subserve respiration. The “pectens” have become more firmly chitinized and probably somewhat altered in shape as compared with their condition in the aquatic ancestral scorpions. Their present function in scorpions is not ascertained. They are not specially sensitive under ordinary conditions, and may be touched or even pinched without causing any discomfort to the scorpion. It is probable that they acquire special sensibility at the breeding season and serve as “guides” in copulation. The shape of the legs and the absence of paired terminal claws in the SilurianPalaeophonus(see figs. 48 and 49) as compared with living scorpions (see fig. 10) show that the early scorpions were aquatic, and we may hope some day in better-preserved specimens than the two as yet discovered, to find the respiratory organs of those creatures in the condition of projecting appendages serving aquatic respiration somewhat as in Limulus, though not necessarily repeating the exact form of the broad plates of Limulus.
It is important to note that the series of lamellae of the lung-book and the gill-book correspondexactlyin structure, the narrow, flat blood-space in the lamellae being interrupted by pillar-like junctions of the two surfaces in both cases (see Lankester (4)), and the free surfaces of the adjacent lamellae being covered with a very delicate chitinous cuticle which is drawn out into delicate hairs and processes. The elongated axis which opens at the stigma in Scorpio and which can be cleared of soft, surrounding tissues and coagulated blood so as to present the appearance of a limb axis carrying the book-like leaves of the lung is not really, as it would seem to be at first sight, the limb axis. That is necessarily a blood-holding structure and is obliterated and fused with soft tissues of the sternal region so that the lamellae cannot be detached and presented as standing out from it. The apparent axis or basal support of the scorpion’s lung-books shown in the figures, is a false or secondary axis and merely a part of the infolded surface which forms the air-chamber. The maceration of the soft parts of a scorpion preserved in weak spirit and the cleaning of the chitinized in-grown cuticle give rise to the false appearance of a limb axis carrying the lamellae. The margins of the lamellae of the scorpion’s lung-book, which arelowermostin the figures (fig. 15) and appear to be free, are really those which are attached to the blood-holding axis. The true free ends are those nearest the stigma.
Passing on now from the mesosoma we come in Scorpio to the metasoma of six segments, the first of which is broad whilst the rest are cylindrical. The last is perforated by the anus and carries the post-anal spine or sting. The somites of the metasoma carry no parapodia. In Limulus the metasoma is practically suppressed. In the allied extinct Eurypterines it is well developed, and resembles that of Scorpio. In the embryo Limulus (fig. 42) the six somites of the mesosoma are not fused to form a carapace at an early stage, and they are followed by three separately marked metasomatic somites; the other three somites of the metasoma have disappeared in Limulus, but are represented by the unsegmented prae-anal region. It is probable that we have in the metasoma of Limulus a case of the disappearance of once clearly demarcated somites. It would be possible to suppose, on the other hand, that new somites are only beginning to make their appearance here. The balance of various considerations is against the latter hypothesis. Following the metasoma in Limulus, we have as in Scorpio the post-anal spine—in this case not a sting, but a powerful and important organ of locomotion, serving to turn the animal over when it has fallen upon its back. The nature of the post-anal spine has been strangely misinterpreted by some writers. Owen (7) maintained that it represented a number of coalesced somites, regardless of its post-anal position and mode of development. The agreement of the grouping of the somites, of the form of the parapodia (appendages, limbs) in each region, of the position of the genital aperture and operculum, of the position and character of the eyes, and of the powerful post-anal spines not seen in other Arthropods, is very convincing as to the affinityof Limulus and Scorpio. Perhaps the most important general agreement of Scorpio compared with Limulus and the Eurypterines is the division of the body into the three regions (or tagmata)—prosoma, mesosoma and metasoma—each consisting of six segments, the prosoma having leg-like appendages, the mesosoma having foliaceous appendages, and the metasoma being destitute of appendages.
M, Mouth behind the oval median camerostome.
I, The chelicerae.
II, The chelae.
III to VI, the four pairs of walking legs.
VIIgo, The genital somite or first somite of the mesosoma with the genital operculum (a fused pair of limbs).
VIIIp, The pectiniferous somite.
IXstgto XIIstg, the four pulmonary somites.
met, The pentagonal metasternite of the prosoma behind all the coxae.
x, The sternum of the pectiniferous somite.
y, The broad first somite of the metasoma.
In 1893, some years after the identification of the somites of Limulus with those of Scorpio, thus indicated, had been published, zoologists were startled by the discovery by a Japanese zoologist, Kishinouye (8), of a seventh prosomatic somite in the embryo of Limulus longispina. This was seen in longitudinal sections, as shown in fig. 19. The simple identification of somite with somite in Limulus and Scorpio seemed to be threatened by this discovery. But in 1896 Dr August Brauer of Marburg (9) discovered in the embryo of Scorpio a seventh prosomatic somite (see VII PrG, figs. 17 and 18), or, if we please so to term it, apraegenitalsomite, hitherto unrecognized. In the case of Scorpio this segment is indicated in the embryo by the presence of a pair of rudimentary appendages, carried by a well-marked somite. As in Limulus, so in Scorpio, this unexpected somite and its appendages disappear in the course of development. In fact, more or less complete “excalation” of the somite takes place. Owing to its position it is convenient to term the somite which is excalated in Limulus and Scorpio “the praegenital somite.” It appears not improbable that the sternal plates wedged in between the last pair of legs in both Scorpio and Limulus, viz. the pentagonal sternite of Scorpio (fig. 10) and the chilaria of Limulus (see figs. 13 and 20), may in part represent in the adult the sternum of the excalated praegenital somite. This has not been demonstrated by an actual following out of the development, but the position of these pieces and the fact that they are (in Limulus) supplied by an independent segmental nerve, favours the view that they may comprise the sternal area of the vanished praegenital somite. This interpretation, however, of the “metasternites” of Limulus and Scorpio is opposed by the coexistence in Thelyphonus (figs. 55, 57 and 58) of a similar metasternite with a complete praegenital somite. H.J. Hansen (10) has recognized that the “praegenital somite” persists in a rudimentary condition, forming a “waist” to the series of somites in the Pedipalpi and Araneae. The present writer is of opinion that it will be found most convenient to treat this evanescent somite as something special, and not to attempt to reckon it to either the prosoma or the mesosoma. These will then remain as typically composed each of six appendage-bearing somites-the prosoma comprising in addition the ocular prosthomere.1When the praegenital somite or traces of it are present it should not be called “the seventh prosomatic” or the “first mesosomatic,” but simply the “praegenital somite.” The first segment of the mesosoma of Scorpio and Limulus thus remains the first segment, and can be identified as such throughout the Eu-arachnida, carrying as it always does the genital apertures. But it is necessary to remember, in the light of recent discoveries, that the sixth prosomatic pair of appendages is carried on the seventh somite of the whole series, there being two prosthomeres or somites in front of the mouth, the first carrying the eyes, the second the chelicerae; also that the first mesosomatic or genital somite is not the seventh or even the eighth of the whole senes of somites which have been historically present, but is the ninth, owing to the presence or to the excalation of a praegenital somite. It seems that confusion and trouble will be best avoided by abstaining from the introduction of the non-evident somites, the ocular and the praegenital, into the numerical nomenclature of the component somites of the three great body regions. We shall, therefore, ignoring the ocular somite, speak of the first, second, third, fourth, fifth and sixth leg-bearing somites of the prosoma, and indicate the appendages by the Roman numerals, I, II, III, IV, V, VI, and whilst ignoring the praegenital somite we shall speak of the first, second, third, &c., somite of the mesosoma or opisthosoma (united mesosoma and metasoma) and indicate them by the Arabic numerals.
There are a number of other important points of structure besides those referring to the somites and appendages in which Limulus agrees with Scorpio or other Arachnida and differs from other Arthropoda. The chief of these are as follows:—
1.The Composition of the Head(that is to say, of the anterior part of the prosoma)with especial Reference to the Region in Front of the Mouth.—It appears (seeArthropoda) that there is embryological evidence of the existence of two somites in Arachnida which were originally post-oral, but have become prae-oral by adaptational shifting of the oral aperture. These forwardly-slipped somites are called “prosthomeres.” The first of these has, in Arachnids as in other Arthropods, its pair of appendages represented by the eyes. The second has for its pair of appendages the small pair of limbs which in all living Arachnids is either chelate or retrovert (as in spiders), and is known as the chelicerae. It is possible, as maintained by some writers (Patten and others), that the lobes of the cerebral nervous mass in Arachnids indicate a larger number of prosthomeres as having fused in this region, but there is noembryologicalevidence at present which justifies us in assuming the existence in Arachnids of more than two prosthomeres. The position of the chelicerae of Limulus and of the ganglionic nerve-masses from which they receive their nerve-supply, is closely similar to that of the same structures in Scorpio. The cerebral mass is in Limulus more easily separated by dissection as a median lobe distinct from the laterally-placed ganglia of the chelceral somite than is the case in Scorpio, but the relations are practically the same in the two forms. Formerly it was supposed that in Limulus both the chelicerae and the next following pair of appendages were prosthomerous, as in Crustacea, but the dissections of Alphonse Milne-Edwards (6) demonstratedthe true limitations of the cerebrum, whilst embryological researches have done as much for Scorpio. Limulus thus agrees with Scorpio and differs from the Crustacea, in which there are three prosthomeres—one ocular and two carrying palpiform appendages. It is true that in the lower Crustacea (Apus, &c.) we have evidence of the gradual movement forward of the nerve-ganglia belonging to these palpiform appendages. But although in such lower Crustacea the nerve-ganglia of the third prosthomere have not fused with the anterior nerve-mass, there is no question as to the prae-oral position of two appendage-bearing somites in addition to the ocular prosthomere. The Crustacea have, in fact, three prosthomeres in the head and the Arachnida only two, and Limulus agrees with the Arachnida in this respect and differs from the Crustacea. The central nervous systems of Limulus and of Scorpio present closer agreement in structure than can be found when a Crustacean is compared with either. The wide divarication of the lateral cords in the prosoma and their connexion by transverse commissures, together with the “attraction” of ganglia to the prosomatic ganglion group which properly belong to hinder segments, are very nearly identical in the two animals. The form and disposition of the ganglion cells are also peculiar and closely similar in the two. (See Patten (42) for important observations on the neuromeres, &c., of Limulus and Scorpio.)
cox, Coxa or basal segment of the leg.
stc, The sterno-coxal process or jaw-like up-growth of the coxa.
epc, The articulated movable outgrowth of the coxa, called the epi-coxite (present only in III of the scorpion and III, IV and V of Limulus).
ex1, The exopodite of the sixth limb of Limulus.
a, b, c, d, Movable processes on the same leg (see for some suggestions on the morphology of this leg, Pocock inQuart. Journ. Micr. Sci.March 1901; see also fig. 50 below and explanation).
2.The Minute Structure of the Central Eyes and of the Lateral Eyes.—Limulus agrees with Scorpio not only in having a pair of central eyes and also lateral eyes, but in the microscopic structure of those organs, which differs in the central and lateral eyes respectively. The central eyes are “simple eyes,” that is to say, have a single lens, and are hence called “monomeniscous.” The lateral eyes are in Limulus “compound eyes,” that is to say, consist of many lenses placed close together; beneath each lens is a complex of protoplasmic cells, in which the optic nerve terminates. Each such unit is termed an “ommatidium.” The lateral eyes of Scorpio consist of groups of separate small lenses each with its ommatidium, but they do not form a continuous compound eye as in Limulus. The ommatidium (soft structure beneath the lens-unit of a compound eye) is very simple in both Scorpio and Limulus. It consists of a single layer of cells, continuous with those which secrete the general chitinous covering of the prosoma. The cells of the ommatidium are a good deal larger than the neighbouring common cells of the epidermis. They secrete the knob-like lens (fig. 22). But they also receive the nerve fibres of the optic nerve. They are at the same time both optic nerve-end cells, that is to say, retina cells, and corneagen cells or secretors of the chitinous lens-like cornea. In Limulus (fig. 23) each ommatidium has a peculiar ganglion cell developed in a central position, whilst the ommatidium of the lateral eyelets of Scorpio shows small intermediate cells between the larger nerve-end cells. The structure of the lateral eye of Limulus was first described by Grenacher, and further and more accurately by Lankester and Bourne (5) and by Watase; that of Scorpio by Lankester and Bourne, who showed that the statements of von Graber were erroneous, and that the lateral eyes of Scorpio have a single cell-layered or “monostichous” ommatidium like that of Limulus. Watase has shown, in a very convincing way, how by deepening the pit-like set of cells beneath a simple lens the more complex ommatidia of the compound eyes of Crustacea and Hexapoda may be derived from such a condition as that presented in the lateral eyes of Limulus and Scorpio. (For details the reader is referred to Watase (11) and to Lankester and Bourne (5).) The structure of the central eyes of Scorpio and spiders and also of Limulus differs essentially from that of the lateral eyes in having two layers of cells (hence called diplostichous) beneath the lens, separated from one another by a membrane (figs. 24 and 25). The upper layer is the corneagen and secretes the lens, the lower is the retinal layer. The mass of soft cell-structures beneath a large lens of a central eye is called an “ommatoeum.” It shows in Scorpio and Limulus a tendency to segregate into minor groups or “ommatidia.” It is found that in embryological growth the retinal layer of the central eyes forms as a separate pouch, which is pushed in laterally beneath the corneagen layer from the epidermic cell layer. Hence it is in origin double, and consists of a true retinal layer and a post-retinal layer (fig. 24, B), though these are not separated by a membrane. Accordingly the diplostichous ommatoeum or soft tissue of the Arachnid’s central eye should strictly be called “triplostichous,” since the deep layer is itself doubled or folded. The retinal cells of both the lateral and central eyes of Limulus and Scorpio produce cuticular structures on their sides; each such piece is a rhabdomere and a number (five or ten) uniting form a rhabdom (fig. 26). In the specialized ommatidia of the compound eyes of Crustacea and Hexapods the rhabdom is an important structure.2It is a very significant fact that the lateral and central eyes of Limulus and Scorpio not only agree each with each in regard to their monostichous and diplostichous structure, but also in the formation in both classes of eyes of rhabdomeres and rhabdoms in which the component pieces are five or a multiple of five (fig. 26). Whilst each unit of the lateral eye of Limulus has a rhabdom of ten3piecesforming a star-like chitinous centre in section, each lateral eye of Scorpio has several rhabdoms of five or less rhabdomeres, indicating that the Limulus lateral eye-unit is more specialized than the detached lateral eyelet of Scorpio, so as to present a coincidence of one lens with one rhabdom. Numerous rhabdomeres (grouped as rhabdoms in Limulus) are found in the retinal layer of the central eyes also.
VII, The genital operculum.
VIII, The pectens of Scorpio and the first branchial plate of Limulus.
IX, The first pair of lung-books of Scorpio and the second branchial plate of Limulus.
gp, Genital pore.
epst, Epistigmatic sclerite.
stg, Stigma or orifice of the hollow tendons of the branchial plates of Limulus.
Whilst Limulus agrees thus closely with Scorpio in regard to the eyes, it is to be noted that no Crustacean has structures corresponding to the peculiar diplostichous central eyes, though these occur again (with differences in detail) inHexapoda. Possibly, however, an investigation of the development of the median eyes of some Crustacea (Apus, Palaemon) may prove them to be diplostichous in origin.
3.The so-called“Coxal Glands.”—In 1882 (Proc. Roy. Soc. No. 221) Lankester described under the name “coxal glands” a pair of brilliantly white oviform bodies lying in the Scorpion’s prosoma immediately above the coxae of the fifth and sixth pairs of legs (fig. 27). These bodies had been erroneously supposed by Newport (12) and other observers to be glandular outgrowths of the alimentary canal. They are really excretory glands, and communicate with the exterior by a very minute aperture on the posterior face of the coxa of the fifth limb on each side. When examined with the microscope, by means of the usual section method, they are seen to consist of a labyrinthine tube lined with peculiar cells, each cell having a deep vertically striated border on the surface farthest from the lumen, as is seen in the cells of some renal organs. The coils and branches of the tube are packed by connective tissue and blood spaces. A similar pair of coxal glands, lobate instead of ovoid in shape, was described by Lankester in Mygale, and it was also shown by him that the structures in Limulus called “brick-red glands” by Packard have the same structure and position as the coxal glands of Scorpio and Mygale. In Limulus these organs consist each of four horizontal lobes lying on the coxal margin of the second, third, fourth, and fifth prosomatic limbs, the four lobes being connected to one another by a transverse piece or stem (fig. 28). Microscopically their structure is the same in essentials as that of the coxal glands of Scorpio (13). Coxal glands have since been recognized and described in other Arachnida. In 1900 it was shown that the coxal gland of Limulus is provided with a very delicate thin-walled coiled duct which opens, even in the adult condition, by a minute pore on the coxa of the fifth leg (Patten and Hazen,13A). Previously to this, Lankester’s pupil Gulland had shown (1885) that in the embryo the coxal gland is a comparatively simple tube, which opens to the exterior in this position and by its other extremity into a coelomic space. Similar observations were made by Laurie (17) in Lankester’s laboratory (1890) with regard to the early condition of the coxal gland of Scorpio, and by Bertkau (41) as to that of the spider Atypus. H.M. Bernard (13B) showed that the opening remains in the adult scorpion. In all the embryonic or permanent opening is on the coxa of the fifth pair of prosomatic limbs. Thus an organ newly discovered in Scorpio was found to have its counterpart in Limulus.
The name “coxal gland” needs to be carefully distinguished from “crural gland,” with which it is apt to be confused. The crural glands, which occur in many terrestrial Arthropods, are epidermal in origin and totally distinct from the coxal glands. The coxal glands of the Arachnida are structures of the same nature as the green glands of the higher Crustacea and the so-called “shell glands” of the Entomostraca. The latter open at the base of the fifth pair of limbs of the Crustacean, just as the coxal glands open on the coxal joint of the fifth pair of limbs of the Arachnid. Both belong to the category of “coelomoducts,” namely, tubular or funnel-like portions of the coelom opening to the exterior in pairs in each somite (potentially,) and usually persisting in only a few somites as either “urocoels” (renal organs) or “gonocoets” (genital tubes). In Peripatus they occur in every somite of the body. They have till recently been very generally identified with the nephridia of Chaetopod worms, but there is good reason for considering the true nephridia (typified by the nephridia of the earthworm) as a distinct class of organs (see Lankester in vol. ii. chap. in. ofA Treatise on Zoology, 1900). The genital ducts of Arthropoda are, like the green glands, shell glands and coxal glands, to be regarded as coelomoducts (gonocoels). The coxal glands do not establish any special connexion between Limulus and Scorpio, sincetheyalso occur in the same somite in the lower Crustacea, but it is to be noted that the coxal glands of Limulus are in minute structure and probably in function more like those of Arachnids than those of Crustacea.
4.The Entosternites and their Minute Structure.—Strauss-Dürckheim (1) was the first to insist on the affinity between Limulus and the Arachnids, indicated by the presence of a free suspended entosternum or plastron or entosternite in both. We have figured here (figs. 1 to 6) the entosternites of Limulus, Scorpio and Mygale. Lankester some years ago made a special study of the histology (3) of these entosternites for the purpose of comparison, and also ascertained the relations of the very numerous muscles which are inserted into them (4). The entosternites are cartilaginous in texture, but they have neither the chemical character nor the microscopic structure of the hyaline cartilage of Vertebrates. They yield chitin in place of chondrin or gelatin—as does also the cartilage of the Cephalopod’s endoskeleton. In microscopic structure they all present the closest agreement with one another. We find a firm, homogeneous or sparsely fibrillated matrix in which are embedded nucleated cells (corpuscles of protoplasm) arranged in rows of three, six or eight, parallel with the adjacent lines of fibrillation.
Fig.16.—Diagram to show the way in which an outgrowing gill-process bearing blood-holding lamellae, may give rise, if the sternal body wall sinks inwards, to a lung-chamber with air-holding lamellae.
I is the embryonic condition.
bs, Blood sinus.
L is the condition of outgrowth withgl, gill lamellae.
A is the condition of in-sinking of the sternal surface and consequent enclosure of the lamelligerous surface of the appendage in a chamber with narrow orifice—the pulmonary air-holding chamber.
pl, Pulmonary lamellae.
bs, Blood sinus.
(After Kingsley.)
Fig.17.—Embryo of scorpion, ventral view showing somites and appendages.
sgc, Frontal groove.
sa, Rudiment of lateral eyes.
obl, Camerostome (upper lip).
so, Sense-organ of Patten.
PrGabp1, Rudiment of the appendage of the praegenital somite which disappears.
abp2, Rudiment of the right half of the genital operculum.
abp3, Rudiment of the right pecten.
abp4toabp7. Rudiments of the four appendages which carry the pulmonary lamellae.
I to VI, Rudiments of the six limbs of the prosoma.
VIIPrG, The evanescent praegenital somite.
VIII, The first mesosomatic somite or genital somite.
IX, The second mesosomatic somite or pectiniferous somite.
X to XIII, The four pulmoniferous somites.
XIV, The first metasomatic somite.
(After Brauer,Zeitsch. wiss. Zool., vol. lix., 1895.)
A minute entosternite having the above-described structure is found in the Crustacean Apus between the bases of the mandibles, and also in the Decapoda in a similar position, but in no Crustacean does it attain to any size or importance. On the other hand, the entosternite of the Arachnida is a very large and important featurein the structure of the prosoma, and must play an important part in the economy of these organisms. In Limulus (figs. 1 and 2) it has as many as twenty-five pairs of muscles attached to it, coming to it from the bases of the surrounding limbs and from the dorsal carapace and from the pharynx. It consists of an oblong plate 2 in. in length and 1 in breadth, with a pair of tendinous outgrowths standing out from it at right angles on each side. It “floats” between the prosomatic nerve centres and the alimentary canal. In each somite of the mesosoma is a small, free entosternite having a similar position, but below or ventral to the nerve cords, and having a smaller number of muscles attached to it. The entosternite was probably in origin part of the fibrous connective tissue lying close to the integument of the sternal surface—giving attachment to muscles corresponding more or less to those at present attached to it. It became isolated and detached, why or with what advantage to the organism it is difficult to say, and at that period of Arachnidan development the great ventral nerve cords occupied a more lateral position than they do at present. We know that such a lateral position of the nerve cords preceded the median position in both Arthropoda and Chaetopoda. Subsequently to the floating off of the entosternite the approximation of the nerve cords took place in the prosoma, and thus they were able to take up a position below the entosternite. In the mesosoma the approximation had occurred before the entosternites were formed.
In the scorpion (figs. 3 and 4) the entosternite has tough membrane-like outgrowths which connect it with the body-wall, both dorsally and ventrally forming an oblique diaphragm, cutting off the cavity of the prosoma from that of the mesosoma. It was described by Newport as “the diaphragm.” Only the central and horizontal parts of this structure correspond precisely to the entosternite of Limulus: the right and left anterior processes (markedapin figs. 3 and 4, and RAP, LAP, in figs. 1 and 2) correspond in the two animals, and the median lateral processlmpof the scorpion represents the tendinous outgrowths ALR, PLR of Limulus. The scorpion’s entosternite gives rise to outgrowths, besides the great posterior flaps,pf, which form the diaphragm, unrepresented in Limulus. These are a ventral arch forming a neural canal through which the great nerve cords pass (figs. 3 and 4,snp), and further a dorsal gastric canal and arterial canal which transmit the alimentary tract and the dorsal artery respectively (figs. 3 and 4, GC, DR).
In Limulus small entosternites are found in each somite of the appendage-bearing mesosoma, and we find in Scorpio, in the only somite of the mesosoma which has a well-developed pair of appendages, that of the pectens, a small entosternite with ten pairs of muscles inserted into it. The supra-pectinal entosternite lies ventral to the nerve cords.
In Mygale (figs. 5 and 6) the form of the entosternite is more like that of Limulus than is that of Scorpio. The anterior notch Ph.N. is similar to that in Limulus, whilst the imbricate triangular pieces of the posterior median region resemble the similarly-placed structures of Limulus in a striking manner.
It must be confessed that we are singularly ignorant as to the functional significance of these remarkable organs—the entosternites. Their movement in an upward or downward direction in Limulus and Mygale must exert a pumping action on the blood contained in the dorsal arteries and the ventral veins respectively. In Scorpio the completion of the horizontal plate by oblique naps, so as to form an actual diaphragm shutting off the cavity of the prosoma from the rest of the body, possibly gives to the organs contained in the anterior chamber a physiological advantage in respect of the supply of arterial blood and its separation from the venous blood of the mesosoma. Possibly the movement of the diaphragm may determine the passage of air into or out of the lung-sacs. Muscular fibres connected with the suctorial pharynx are in Limulus inserted into the entosternite, and the activity of the two organs may be correlated.
5.The Blood and the Blood-vascular System.—The blood fluids of Limulus and Scorpio are very similar. Not only are the blood corpuscles of Limulus more like in form and granulation to those of Scorpio than to those of any Crustacean, but the fluid is in both animals strongly impregnated with the blue-coloured respiratory proteid, haemocyanin. This body occurs also in the blood of Crustacea and of Molluscs, but its abundance in both Limulus and Scorpio is very marked, and gives to the freshly-shed blood a strong indigo-blue tint.
sf, The sub-frontal median sclerite.
Ch, The chelicerae.
cam, The camerostome or upper lip.
M, The mouth.
pmst, The promesosternal sclerite of chitinous plate, unpaired.
mets, The right and left metasternites (corresponding to the similarly placed pentagonal sternite of Scorpio). Natural size.
lens, Cuticular lens.
nerv c, Retinal cells (nerve-end cells).
rhabd, Rhabdomes.
nerv f, Nerve fibres of the optic nerve.
int, Intermediate cells (lying between the bases of the retinal cells).
The great dorsal contractile vessel or “heart” of Limulus is closely similar to that of Scorpio; its ostia or incurrent orifices are placed in the same somites as those of Scorpio, but there is one additional posterior pair. The origin of the paired arteries from theheart differs in Limulus from the arrangement obtaining in Scorpio, in that a pair of lateral commissural arteries exist in Limulus (as described by Alphonse Milne-Edwards (6)) leading to a suppression of the more primitive direct connexion of the four pairs of posterior lateral arteries and of the great median posterior arteries with the heart itself (fig. 29). The arterial system is very completely developed in both Limulus and Scorpio, branching repeatedly until minute arterioles are formed, not to be distinguished from true capillaries; these open into irregular swollen vessels which are the veins or venous sinuses. A very remarkable feature in Limulus, first described by Owen, is the close accompaniment of the prosomatic nerve centres and nerves by arteries, so close indeed that the great ganglion mass and its out-running nerves are actually sunk in or invested by arteries. The connexion is not so intimate in Scorpio, but is nevertheless a very close one, closer than we find in any other Arthropods in which the arterial system is well developed,e.g.the Myriapoda and some of the arthrostracous Crustacea. It seems that there is a primitive tendency in the Arthropoda for the arteries to accompany the nerve cords, and a “supra-spinal” artery—that is to say, an artery in close relation to the ventral nerve cords—has been described in several cases. On the other hand, in many Arthropods, especially those which possess tracheae, the arteries do not have a long course, but soon open into wide blood sinuses. Scorpio certainly comes nearer to Limulus in the high development of its arterial system, and the intimate relation of the anterior aorta and its branches to the nerve centres and great nerves, than does any other Arthropod.
An arrangement of great functional importance in regard to the venous system must now be described, which was shown in 1883 by Lankester to be common to Limulus and Scorpio. This arrangement has not hitherto been detected in any other class than the Arachnida, and if it should ultimately prove to be peculiar to that group, would have considerable weight as a proof of the close genetic affinity of Limulus and Scorpio.
A, Early condition before the lens is deposited, showing the folding of the epidermic cell-layer into three.
B, Diagram showing the nature of this infolding.
C, Section through the fully formed eye.
h, Epidermic cell-layer.
r, The retinal portion of the same which, owing to the infolding, lies betweengl, the corneagen or lens-forming portion, andpr, the post-retinal or capsular portion or fold.
l, Cuticular lens.
g, Line separating lens from the lens-forming or corneagen cells of the epidermis.
n, Nerve fibres.
rh, Rhabdomeres.
The great pericardial sinus is strongly developed in both animals. Its walls are fibrous and complete, and it holds a considerable volume of blood when the heart itself is contracted. Opening in pairs in each somite, right and left into the pericardial sinus are large veins, which bring the blood respectively from the gill-books and the lung-books to that chamber, whence it passes by the ostia into the heart. The blood is brought to the respiratory organs in both cases by a great venous collecting sinus having a ventral median position. In both animalsthe wall of the pericardial sinus is connected by vertical muscular bands to the wall of the ventral venous sinus(its lateral expansions around the lung-books in Scorpio) in each somite through which the pericardium passes. There are seven pairs of theseveno-pericardiac vertical musclesin Scorpio, and eight in Limulus (see figs. 30, 31, 32). It is obvious that the contraction of these musclesmust cause a depression of the floor of the pericardium and a rising of the roof of the ventral blood sinus, and a consequent increase of volume and flow of blood to each. Whether the pericardium and the ventral sinus are made to expand simultaneously or all the movement is made by one only of the surfaces concerned, must depend on conditions of tension. In any case it is clear that we have in these muscles an apparatus for causing the blood to flow differentially in increased volume into either the pericardium, through the veins leading from the respiratory organs, or from the body generally into the great sinuses which bring the blood to the respiratory organs. These muscles act so as to pump the blood through the respiratory organs.
L, Cuticular or corneous lens.
hy, Epidermic cell-layer.
corn, Its corneagen portion immediately underlying the lens.
ret, Retinula cells.
nf, Nerve fibres.
con. tiss, Connective tissue (mesoblastic skeletal tissue).
It is not surprising that with so highly developed an arterial system Limulus and Scorpio should have a highly developed mechanism for determining the flow of blood to the respiratory organs. That this is, so to speak, a need of animals with localized respiratory organs is seen by the existence of provisions serving a similar purpose in other animals,e.g.the branchial hearts of the Cephalopoda.
The veno-pericardiac muscles of Scorpio were seen and figured by Newport but not described by him. Those of Limulus were described and figured by Alphonse Milne-Edwards, but he called them merely “transparent ligaments,” and did not discover their muscular structure. They are figured and their importance for the first time recognized in the memoir on the muscular and skeletal systems of Limulus and Scorpio by Lankester, Beck and Bourne (4).
6.Alimentary Canal and Gastric Glands.—The alimentary canal in Scorpio, as in Limulus, is provided with a powerful suctorial pharynx, in the working of which extrinsic muscles take a part. The mouth is relatively smaller in Scorpio than in Limulus—in fact is minute, as it is in all the terrestrial Arachnida which suck the juices of either animals or plants. In both, the alimentary canal takes a straight course from the pharynx (which bends under it downwards and backwards towards the mouth in Limulus) to the anus, and is a simple, narrow, cylindrical tube (fig. 33). The only point in which the gut of Limulus resembles that of Scorpio rather than that of any of the Crustacea, is in possessing more than a single pair of ducts or lateral outgrowths connected with ramified gastric glands or gastric caeca. Limulus has two pairs of these, Scorpio as many as six pairs. The Crustacea never have more than one pair. The minute microscopic structure of the gastric glands in the two animals is practically identical. The functions of these gastric diverticula have never been carefully investigated. It is very probable that in Scorpio they do not serve merely to secrete a digestive fluid (shown in other Arthropoda to resemble the pancreatic fluid), but that they also become distended by the juices of the prey sucked in by the scorpion—as certainly must occur in the case of the simple unbranched gastric caeca of the spiders.
The most important difference which exists between the structure of Limulus and that of Scorpio is found in the hinder region of the alimentary canal. Scorpio is here provided with a single or double pair of renal excretory tubes, which have been identified by earlier authors with the Malpighian tubes of the Hexapod and Myriapod insects. Limulus is devoid of any such tubes. We shall revert to this subject below.
A, Diagram of a retinula of the central eye of a scorpion consisting of five retina-cells (ret), with adherent branched pigment cells (pig).
B, Rhabdom of the same, consisting of five confluent rhabdomeres.
C, Transverse section of the rhabdom of a retinula of the scorpion’s central eye, showing its five constituent rhabdomeres as rays of a star.
D, Transverse section of a retinula of the lateral eye of Limulus, showing ten retinula cells (ret), each bearing a rhabdomere (rhab).
1 to 6, The bases of the six prosomatic limbs.
A, prosomatic gastric gland (sometimes called salivary).
B, Coxal gland.
C, Diaphragm of Newport = fibrous flap of the entosternum.
D, Mesosomatic gastric caeca (so-called liver).
E, Alimentary canal.
7.Ovaries and Spermaries: Gonocoels and Gonoducts.—The scorpion is remarkable for having the specialized portion of coelom from the walls of which egg-cells or sperm-cells are developed according to sex, in the form of a simple but extensive network. It is not a pair of simple tubes, nor of dendriform tubes, but a closed network. The same fact is true of Limulus, as was shown by Owen (7) in regard to the ovary, and by Benham (14) in regard to the testis. This is a very definite and remarkable agreement, since such a reticular gonocoel is not found in Crustacea (except in the male Apus). Moreover, there is a significant agreement in the character of the spermatozoa of Limulus and Scorpio. The Crustacea are—with the exception of the Cirrhipedia—remarkable for having stiff, motionless spermatozoids. In Limulus Lankester found (15) the spermatozoa to possess active flagelliform “tails,” and to resemble very closely those of Scorpio which, as are those of most terrestrial Arthropoda, are actively motile. This is a microscopic point of agreement, but is none the less significant.
In regard to the important structures concerned with the fertilization of the egg, Limulus and Scorpio differ entirely from one another.The eggs of Limulus are fertilized in the sea after they have been laid. Scorpio, being a terrestrial animal, fertilizes by copulation. The male possesses elaborate copulatory structures of a chitinous nature, and the eggs are fertilized in the female without even quitting the place where they are formed on the wall of the reticular gonocoel. The female scorpion is viviparous, and the young are produced in a highly developed condition as fully formed scorpions.
a2toa5, Posterior borders of the chitinous bases of the coxae of the second, third, fourth and fifth prosomatic limbs.
b, Longitudinal lobe or stolon of the coxal gland.
c. Its four transverse lobes or outgrowths corresponding to the four coxae.
Differences between Limulus and Scorpio.—We have now passed in review the principal structural features in which Limulus agrees with Scorpio and differs from other Arthropoda. There remains for consideration the one important structural difference between the two animals. Limulus agrees with the majority of the Crustacea in being destitute of renal excretory caeca or tubes opening into the hinder part of the gut. Scorpio, on the other hand, in common with all air-breathing Arthropoda except Peripatus, possesses these tubules, which are often called Malpighian tubes. A great deal has been made of this difference by some writers. It has been considered by them as proving that Limulus, in spite of all its special agreements with Scorpio (which, however, have scarcely been appreciated by the writers in question), really belongs to the Crustacean line of descent, whilst Scorpio, by possessing Malpighian tubes, is declared to be unmistakably tied together with the other Arachnida to the tracheate Arthropods, the Hexapods, Diplopods, and Chilopods, which all possess Malpighian tubes.
It must be pointed out that the presence or absence of such renal excretory tubes opening into the intestine appears to be a question of adaptation to the changed physiological conditions of respiration, and not of morphological significance, since a pair of renal excretory tubes of this nature is found in certain Amphipod Crustacea (Talorchestia, &c.) which have abandoned a purely aquatic life. This view has been accepted and supported by Professors Korschelt and Heider (16). An important fact in its favour was discovered by Laurie (17), who investigated the embryology of two species of Scorpio under Lankester’s direction. It appears that the Malpighian tubes of Scorpio are developed from the mesenteron, viz. that portion of the gut which is formed by the hypoblast, whereas in Hexapod insects the similar caecal tubes are developed from the proctodaeum or in-pushed portion of the gut which is formed from epiblast. In fact it is not possible to maintain that the renal excretory tubes of the gut are of one common origin in the Arthropoda. They have appeared independently in connexion with a change in the excretion of nitrogenous waste in Arachnids, Crustacea, and the other classes of Arthropoda when aerial, as opposed to aquatic, respiration has been established—and they have been formed in some cases from the mesenteron, in other cases from the proctodaeum. Their appearance in the air-breathing Arachnids does not separate those forms from the water-breathing Arachnids which are devoid of them, any more than does their appearance in certain Amphipoda separate those Crustaceans from the other members of the class.
Further, it is pointed out by Korschelt and Heider that the hinder portion of the gut frequently acts in Arthropoda as an organ of nitrogenous excretion in the absence of any special excretory tubules, and that the production of such caeca from its surface in separate lines of descent does not involve any elaborate or unlikely process of growth. In other words, the Malpighian tubes of the terrestrial Arachnida arehomoplasticwith those of Hexapoda and Myriapoda, and nothomogeneticwith them. We are compelled to take a similar view of the agreement between the tracheal air-tubes of Arachnida and other tracheate Arthropods. They are homoplasts (see18) one of another, and do not owe their existence in the various classes compared to a common inheritance of an ancestral tracheal system.
PRO, Prosoma.
dpm, Dorso-plastral muscle.
art, Lateral artery.
tsm1, Tergo-sternal muscle (labelleddvin fig. 31) of the second (pectiniferous) mesosomatic somite; this is the most anterior pair of the series of six, none are present in the genital somite.
tsm4, Tergo-sternal muscle of the fifth mesosomatic somite.
tsm6, Tergo-sternal muscle of the enlarged first metasomatic somite.
Per, Pericardium.
VPM1to VPM7, The series of seven pairs of veno-pericardiac muscles (labelledpvin fig. 31).
Conclusions arising from the Close Affinity of Limulus and Scorpio.—When we consider the relationships of the various classes of Arthropoda, having accepted and established the fact of the close genetic affinity of Limulus and Scorpio, we are led to important conclusions. In such a consideration we have to make use not only of the fact just mentioned, but of three important generalizations which serve as it were as implements for the proper estimation of the relationships of any series of organic forms. First of all there is the generalization that the relationships of the various forms of animals (or of plants) to one another is that of the ultimate twigs of a much-branching genealogical tree. Secondly, identity of structure in two organisms does not necessarily indicate that the identical structure has been inherited from an ancestor common to the two organisms compared (homogeny), but may be due to independent development of a like structure in two different lines of descent (homoplasy). Thirdly, those members of a group which, whilst exhibiting undoubted structural characters indicative of their proper assignment to that group, yet are simpler than and inferior in elaboration of their organization to other members of the group, are not necessarily representatives of the earlier and primitive phases in the development of the group—but are very often examples of retrogressive change or degeneration. The second and third implements of analysis above cited are of the nature of cautions or checks. Agreements are notnecessarilydue to common inheritance; simplicity is notnecessarilyprimitive and ancestral.
On the other hand, we must not rashly set down agreements as due to “homoplasy” or “convergence of development” if we find two or three or more concurrent agreements. The probability is against agreement being due to homoplasy when the agreement involves a number of really separate (not correlated) coincidences. Whilst the chances are in favour of someonehomoplastic coincidence or structural agreement occurring between some member or other of a large groupaand some member or other of a large groupb, the matter is very differentwhen by such an initial coincidence the two members have been particularized. The chances against these two selected members exhibitinganotherreally independent homoplastic agreement are enormous: let us say 10,000 to 1. The chances against yet another coincidence are a hundred million to one, and against yet one more “coincidence” they are the square of a hundred million to one. Homoplasy can only be assumed when the coincidence is of a simple nature, and is such as may be reasonably supposed to have arisen by the action of like selective conditions upon like material in two separate lines of descent.4
So, too, degeneration is not to be lightly assumed as the explanation of a simplicity of structure. There is a very definite criterion of the simplicity due to degeneration, which can in most cases be applied. Degenerative simplicity is never uniformly distributed over all the structures of the organism. It affects many or nearly all the structures of the body, but leaves some, it may be only one, at a high level of elaboration and complexity. Ancestral simplicity is more uniform, and does not co-exist with specialization and elaboration of a single organ. Further: degeneration cannot be inferred safely by the examination of an isolated case; usually we obtain a series of forms indicating the steps of a change in structure—and what we have to decide is whether the movement has been from the simple to the more complex, or from the more complex to the simple. The feathers of a peacock afford a convenient example of primitive and degenerative simplicity. The highest point of elaboration in colour, pattern and form is shown by the great eye-painted tail feathers. From these we can pass by gradual transitions in two directions, viz. either to the simple lateral tail feathers with a few rami only, developed only on one side of the shaft and of uniform metallic coloration—or to the simple contour feathers of small size, with the usual symmetrical series of numerous rami right and left of the shaft and no remarkable colouring. The one-sided specialization and the peculiar metallic colouring of the lateral tail feathers mark them as the extreme terms of a degenerative series, whilst the symmetry, likeness of constituent partsinter se, and absence of specialized pigment, as well as the fact that they differ little from any average feather of birds in general, mark the contour feather as primitively simple, and as the starting-point from which the highly elaborated eye-painted tail feather has gradually evolved.
Applying these principles to the consideration of the Arachnida, we arrive at the conclusion that the smaller and simpler Arachnids are not the more primitive, but that the Acari or mites are, in fact, a degenerate group. This was maintained by Lankester in 1878 (19), again in 1881 (20); it was subsequently announced as a novelty by Claus in 1885 (21). Though the aquatic members of a class of animals are in some instances derived from terrestrial forms, the usual transition is from an aquatic ancestry to more recent land-living forms. There is no doubt, from a consideration of the facts of structure, that the aquatic water-breathing Arachnids, represented in the past by the Eurypterines and to-day by the sole survivor Limulus, have preceded the terrestrial air-breathing forms of that group. Hence we see at once that the better-known Arachnida form a series, leading from Limulus-like aquatic creatures through scorpions, spiders and harvest-men, to the degenerate Acari or mites. The spiders are specialized and reduced in apparent complexity, as compared with the scorpions, but they cannot be regarded as degenerate since the concentration of structure which occurs in them results in greater efficiency and power than are exhibited by the scorpion. The determination of the relative degree of perfection of organization attained by two animals compared is difficult when we introduce, as seems inevitable, the question of efficiency and power, and do not confine the question to the perfection of morphological development. We have no measure of the degree of power manifested by various animals—though it would be possible to arrive at some conclusions as to how that “power” should be estimated. It is not possible here to discuss that matter further. We must be content to point out that it seems that the spiders, the pedipalps, and other large Arachnids have not been derived from the scorpions directly, but have independently developed from aquatic ancestors, and from one of these independent groups—probably through the harvest-men from the spiders—the Acari have finally resulted.
d, Chelicera.
ch, Chela.
cam, Camerostome.
m, Mouth.
ent, Entosternum.
p, Pecten.
stig1, First pulmonary aperture.
stig4, Fourth pulmonary aperture.
dam, Muscle from carapace to a praeoral entosclerite.
ad, Muscle from carapace to entosternum.
md, Muscle from tergite of genital somite to entosternum (same asdpmin fig. 30).
dv1todv6, Dorso-ventral muscles (same as the series labelledtsmin fig. 30).
pv1topv7, The seven veno-pericardiac muscles of the right side (labelled VPM in fig. 30).
Suc, Suctorial pharynx.
al, Alimentary canal.
Ph, Pharynx.
M, Mouth.
Est, Entosternum.
VS, Ventral venous sinus.
chi, Chilaria.
go, Genital operculum.
br1tobr5, Branchial appendages,
met, Unsegmented metasoma.
Entap4, Fourth dorsal entapophysis of left side.
tsm, Tergo-sternal muscles, six pairs as in Scorpio (labelleddvin fig. 31).
VPM1to VPM8, The eight pairs of veno-pericardiac muscles (labelledpvin fig. 31). VPM1is probably represented in Scorpio, though not marked in figs. 30 and 31.
ps, Muscular suctorial enlargement of the pharynx.
sal, Prosomatic pair of gastric caeca in Scorpio, called salivary glands by some writers.
c1, andc2, The anterior two pairs of gastric caeca and ducts of the mesosomatic region.
c3,c4andc5. Caeca and ducts of Scorpio not represented in Limulus.
M, The Malpighian or renal caecal diverticula of Scorpio.
pro, The proctodaeum or portion of gut leading to anus and formed embryologically by an inversion of the epiblast at that orifice.