1911 Encyclopædia Britannica/Arthropoda
ARTHROPODA, a name, denoting the possession by certain animals of jointed limbs, now applied to one of the three sub-phyla into which one of the great phyla (or primary branches) of coelomocoelous animals—the Appendiculata—is divided; the other two being respectively the Chaetopoda and the Rotifera. The word “Arthropoda” was first used in classification by Siebold and Stannius (Lehrbuch der vergleich. Anatomie, Berlin, 1845) as that of a primary division of animals, the others recognized in that treatise being Protozoa, Zoophyta, Vermes, Mollusca and Vertebrata. The names Condylopoda and Gnathopoda have been subsequently proposed for the same group. The word refers to the jointing of the chitinized exo-skeleton of the limbs or lateral appendages of the animals included, which are, roughly speaking, the Crustacea, Arachnida, Hexapoda (so-called “true insects”), Centipedes and Millipedes. This primary group was set up to indicate the residuum of Cuvier’s Articulata when his class Annélides (the modern Chaetopoda) was removed from that embranchement. At the same time C. T. E. von Siebold and H. Stannius renovated the group Vermes of Linnaeus, and placed in it the Chaetopods and the parasitic worms of Cuvier, besides the Rotifers and Turbellarian worms.
The result of the knowledge gained in the last quarter of the 19th century has been to discredit altogether the group Vermes (see Worm), thus set up and so largely accepted by German writers even at the present day. We have, in fact, returned very nearly to Cuvier’s conception of a great division or branch, which he called Articulata, including the Arthropoda and the Chaetopoda (Annélides of Lamarck, a name adopted by Cuvier), and differing from it only by the inclusion of the Rotifera. The name Articulata, introduced by Cuvier, has not been retained by subsequent writers. The same, or nearly the same, assemblage of animals has been called Entomozoaria by de Blainville (1822), Arthrozoa by Burmeister (1843), Entomozoa or Annellata by H. Milne-Edwards (1855), and Annulosa by Alexander M‘Leay (1819), who was followed by Huxley (1856). The character pointed to by all these terms is that of a ring-like segmentation of the body. This, however, is not the character to which we now ascribe the chief weight as evidence of the genetic affinity and monophyletic (uni-ancestral) origin of the Chaetopods, Rotifers and Arthropods. It is the existence in each ring of the body of a pair of hollow lateral appendages or parapodia, moved by intrinsic muscles and penetrated by blood-spaces, which is the leading fact indicating the affinities of these great sub-phyla, and uniting them as blood-relations. The parapodia (fig. 8) of the marine branchiate worms are the same things genetically as the "legs" of Crustacea and Insects (figs. 10 and 11). Hence the term Appendiculata was introduced by Lankester (preface to the English edition of Gegenbaur’s Comparative Anatomy, 1878) to indicate the group. The relationships of the Arthropoda thus stated are shown in the subjoined table:—
The Rotifera are characterized by the retention of what appears in Molluscs and Chaetopods as an embryonic organ, the velum or ciliated prae-oral girdle, as a locomotor and food-seizing apparatus, and by the reduction of the muscular parapodia to a rudimentary or non-existent condition in all present surviving forms except Pedalion. In many important respects they are degenerate—reduced both in size and elaboration of structure.
The Chaetopoda are characterized by the possession of horny epidermic chaetae embedded in the integument and moved by muscles. Probably the chaetae preceded the development of parapodia, and by their concentration and that of the muscular bundles connected with them at the sides of each segment, led directly to the evolution of the parapodia. The parapodia of Chaetopoda are never coated with dense chitin, and are, therefore, never converted into jaws; the primitive "head-lobe" or prostomium persists, and frequently carries eyes and sensory tentacles. Further, in all members of the sub-phylum Chaetopoda the relative position of the prostomium, mouth and peristomium or first ring of the body, retains its primitive character. We do not find in Chaetopoda that parapodia, belonging to primitively post-oral rings or body-segments (called "somites," as proposed by H. Milne-Edwards), pass in front of the mouth by adaptational shifting of the oral aperture. (See, however, 8.)
The Arthropoda might be better called the "Gnathopoda," since their distinctive character is, that one or more pairs of appendages behind the mouth are densely chitinized and turned (fellow to fellow on opposite sides) towards one another so as to act as jaws. This is facilitated by an important general change in the position of the parapodia; their basal attachments are all more ventral in position than in the Chaetopoda, and tend to approach from the two sides towards the mid-ventral line. Very usually (but not in the Onychophora = Peripatus) all the parapodia are plated with chitin secreted by the epidermis, and divided into a series of joints—giving the "arthropodous" or hinged character.
There are other remarkable and distinctive features of structure which hold the Arthropoda together, and render it impossible to conceive of them as having a polyphyletic origin, that is to say, as having originated separately by two or three distinct lines of descent from lower animals; and, on the contrary, establish the view that they have been developed from a single line of primitive Gnathopods which arose by modification of parapodiate annulate worms not very unlike some of the existing Chaetopods. These additional features are the following—(1) All existing Arthropoda have an ostiate heart and have undergone "phleboedesis," that is to say, the peripheral portions of the blood-vascular system are not fine tubes as they are in the Chaetopoda and as they were in the hypothetical ancestors of Arthropoda, but are swollen so as to obliterate to a large extent the coelom, whilst the separate veins entering the dorsal vessel or heart have coalesced, leaving valvate ostia (see fig. 1) by which the blood passes from a pericardial blood-sinus formed by the fused veins into the dorsal vessel or heart (see Lankester’s Zoology, part ii., introductory chapter, 1900). The only exception to this is in the case of minute degenerate forms where the heart has disappeared altogether. The rigidity of the integument caused by the deposition of dense chitin upon it is intimately connected with the physiological activity and form of all the internal organs, and is undoubtedly correlated with the total disappearance of the circular muscular layer of the body-wall present in Chaetopods. (2) In all existing Arthropoda the region in front of the mouth is no longer formed by the primitive prostomium or head-lobe, but one or more segments, originally post-oral, with their appendages have passed in front of the mouth (prosthomeres). At the same time the prostomium and its appendages cease to be recognizable as distinct elements of the head. The brain no longer consists solely of the nerve-ganglion-mass proper to the prostomial lobe, as in Chaetopoda, but is a composite (syncerebrum) produced by the fusion of this and the nerve-ganglion-masses proper to the prosthomeres or segments which pass forwards, whilst their parapodia (= appendages) become converted into eye-stalks, and antennae, or more rarely grasping organs. (3) As in Chaetopoda, coelomic funnels (coelomoducts) may occur right and left as pairs in each ring-like segment or somite of the body, and some of these are in all cases retained as gonoducts and often as renal excretory organs (green glands, coxal glands of Arachnida, not crural glands, which are epidermal in origin); but true nephridia, genetically identical with the nephridia of earthworms, do not occur (on the subject of coelom, coelomoducts and nephridia, see the introductory chapter of part ii. of Lankester’s Treatise on Zoology).
Tabular Statement of the Grades, Classes and Sub-classes of the Arthropoda.—It will be convenient now to give in the clearest form a statement of the larger subdivisions of the Arthropoda which it seems necessary to recognize at the present day. The justification of the arrangement adopted will form the substance of the rest of the present article. The orders included in the various classes are not discussed here, but are treated of under the following titles:—Peripatus (Onychophora), Centipede and Millipede (Myriapoda), Hexapoda (Insecta), Arachnida and Crustacea.
Sub-Phylum ARTHROPODA (of the Phylum Appendiculata).
Grade A. Hyparthropoda (hypothetical forms connecting ancestors of Chaetopoda with those of Arthropoda).
Grade B. Protarthropoda.
- Class Onychophora.
Grade C. Euarthropoda.
- Class 1. Diplopoda.
- Class 2. Arachnida.
- Grade a. Anomomeristica.
- Grade b. Nomomeristica.
- (a) Pantopoda.
- (b) Euarachnida.
- Ex.—Limulus, Scorpio, Mygale, Acarus.
- Grade a. Anomomeristica.
- Class 3. Crustacea.
- Grade a. Entomostraca.
- Ex.—Apus, Branchipus, Cyclops, Balanus.
- Grade b. Malacostraca.
- Ex.—Nebalia, Astacus, Oniscus, Gammarus.
- Grade a. Entomostraca.
- Class 4. Chilopoda.
- Class 5. Hexapoda (syn. Insecta Pterygota).
- Ex.—Locusta, Phryganea, Papilio, Apis, Musca, Cimex, Lucanus, Machilis.
Incertae sedis—Tardigrada, Pentastomidae (degenerate forms).
The Segmentation of the Body of Arthropoda.—The body of the Arthropoda is more or less clearly divided into a series of rings, segments, or somites which can be shown to be repetitions one of another, possessing identical parts and organs which may be larger or smaller, modified in shape or altogether suppressed in one somite as compared with another. A similar constitution of the body is more clearly seen in the Chaetopod worms. In the Vertebrata also a repetition of units of structure (myotomes, vertebrae, &c.)—which is essentially of the same nature as the repetition in Arthropods and Chaetopods, but in many respects subject to peculiar developments—is observed. The name “metamerism” has been given to this structural phenomenon because the “meres,” or repeated units, follow one another in line. Each such “mere” is often called a “metamere.” A satisfactory consideration of the structure of the Arthropods demands a knowledge of what may be called the laws of metamerism, and reference should be made to the article under that head.
The Theory of the Arthropod Head.—The Arthropod head is a tagma or group of somites which differ in number and in their relative position in regard to the mouth, in different classes. In a simple Chaetopod (fig. 2) the head consists of the first somite only; that somite is perforated by the mouth, and is provided with a prostomium or prae-oral lobe. The prostomium is essentially a part or outgrowth of the first somite, and cannot be regarded as itself a somite. It gives rise to a nerve-ganglion mass, the prostomial ganglion. In the marine Chaetopods (the Polychaeta) (fig. 3), we find the same essential structure, but the prostomium may give rise to two or more tactile tentacles, and to the vesicular eyes. The somites have well-marked parapodia, and the second and third, as well as the first, may give rise to tentacles which are directed forward, and thus contribute to form “the head.” But the mouth remains as an inpushing of the wall of the first somite.
The Arthropoda are all distinguished from the Chaetopoda by the fact that the head consists of one or more somites which lie in front of the mouth (now called prosthomeres), as well as of one or more somites behind it (opisthomeres). The first of the post-oral somites invariably has its parapodia modified so as to form a pair of hemignaths (mandibles). About 1870 the question arose for discussion whether the somites in front of the mouth are to be considered as derived from the prostomium of a Chaetopod-like ancestor. Milne-Edwards and Huxley had satisfied themselves with discussing and establishing, according to the data at their command, the number of somites in the Arthropod head, but had not considered the question of the nature of the prae-oral somites. Lankester (2) was the first to suggest that (as is actually the fact in the Nauplius larva of the Crustacea) the prae-oral somites or prosthomeres and their appendages were ancestrally post-oral, but have become prae-oral “by adaptational shifting of the oral aperture.” This has proved to be a sound hypothesis and is now accepted as the basis upon which the Arthropod head must be interpreted (see Korschelt and Heider (3)). Further, the morphologists of the ’fifties appear, with few exceptions, to have accepted a preliminary scheme with regard to the Arthropod head and Arthropod segmentation generally, which was misleading and caused them to adopt forced conclusions and interpretations. It was conceived by Huxley, among others, that the same number of cephalic somites would be found to be characteristic of all the diverse classes of Arthropoda, and that the somites, not only of the head but of the various regions of the body, could be closely compared in their numerical sequence in classes so distinct as the Hexapods, Crustaceans and Arachnids.
The view which it now appears necessary to take is, on the contrary, this—viz that all the Arthropoda are to be traced to a common ancestor resembling a Chaetopod worm, but differing from it in having lost its chaetae and in having a prosthomere in front of the mouth (instead of prostomium only) and a pair of hemignaths (mandibles) on the parapodia of the buccal somite. From this ancestor Arthropods with heads of varying degrees of complexity have been developed characteristic of the different classes, whilst the parapodia and somites of the body have become variously modified and grouped in these different classes. The resemblances which the members of one class often present to the members of another class in regard to the form of the limb-branches (rami) of the parapodia and the formation of tagmata (regions) are not hastily to be ascribed to common inheritance, but we must consider whether they are not due to homoplasy—that is, to the moulding of natural selection acting in the different classes upon fairly similar elements under like exigencies.
The structure of the head in Arthropods presents three profoundly separated grades of structure dependent upon the number of prosthomeres which have been assimilated by the prae-oral region. The classes presenting these distinct plans of head-structure cannot be closely associated in any scheme of classification professing to be natural. Peripatus, the type-genus of the class Onychophora, stands at the base of the series with only a single prosthomere (fig. 4). In Peripatus the prostomium of the Chaetopod-like ancestor is atrophied, but it is possible that two processes on the front of the head (FP) represent in the embryo the dwindled prostomial tentacles. The single prosthomere carries the retractile tentacles as its “parapodia.” The second somite is the buccal somite (II, fig. 4); its parapodia have horny jaws on their ends, like the claws on the following legs (fig. 9), and act as hemignaths (mandibles). The study of sections of the embryo establishes these facts beyond doubt. It also shows us that the neuromeres, no less than the embryonic coelomic cavities, point to the existence of one, and only one, prosthomere in Peripatus, of which the “protocerebrum,” P, is the neuromere, whilst the deuterocerebrum, D, is the neuromere of the second or buccal somite. A brief indication of these facts is given by saying that the Onychophora are “deuterognathous”—that is to say, that the buccal somite carrying the mandibular hemignaths is the second of the whole series.
What has become of the nerve-ganglion of the prostomial lobe of the Chaetopod in Peripatus is not clearly ascertained, nor is its fate indicated by the study of the embryonic head of other Arthropods so far. Probably it is fused with the protocerebrum, and may also be concerned in the history of the very peculiar paired eyes of Peripatus, which are like those of Chaetopods in structure—viz vesicles with an intravesicular lens, whereas the eyes of all other Arthropods have essentially another structure, being “cups” of the epidermis, in which a knob-like or rod-like thickening of the cuticle is fitted as refractive medium.
In Diplopoda (Julus, &c.) the results of embryological study point to a composition of the front part of the head exactly similar to that which we find in Onychophora. They are deuterognathous.
The Arachnida present the first stage of progress. Here embryology shows that there are two prosthomeres (fig. 5), and that the gnathobases of the chelae which act as the first pair of hemignaths are carried by the third somite. The Arachnida are therefore tritognathous. The two prosthomeres are indicated by their coelomic cavities in the embryo (I and II, fig. 5), and by two neuromeres, the protocerebrum and the deuterocerebrum. The appendages of the first prosthomere are not present as tentacles, as in Peripatus and Diplopods, but are possibly represented by the eyes or possibly altogether aborted. The appendages of the second prosthomere are the well-known chelicerae of the Arachnids, rarely, if ever, antenniform, but modified as “retroverts” or clasp-knife fangs in spiders.
The Crustacea (fig. 6) and the Hexapoda (fig. 7) agree in having three somites in front of the mouth, and it is probable, though not ascertained, that the Chilopoda (Scolopendra, &c.) are in the same case. The three prosthomeres or prae-oral somites of Crustacea due to the sinking back of the mouth one somite farther than in Arachnida are not clearly indicated by coelomic cavities in the embryo, but their existence is clearly established by the development and position of the appendages and by the neuromeres.
The eyes in some Crustacea are mounted on articulated stalks, and from the fact that they can after injury be replaced by antenna-like appendages it is inferred that they represent the parapodia of the most anterior prosthomere. The second prosthomere carries the first pair of antennae and the third the second pair of antennae. Sometimes the pair of appendages has not a merely tactile jointed ramus, but is converted into a claw or clasper. Three neuromeres—a proto-, deutero-, and trito-cerebrum—corresponding to those three prosthomeres are sharply marked in the embryo. The fourth somite is that in which the mouth now opens, and which accordingly has its appendages converted into hemignathous mandibles. The Crustacea are tetartognathous.
The history of the development of the head has been carefully worked out in the Hexapod insects. As in Crustacea and Arachnida, a first prosthomere is indicated by the paired eyes and the protocerebrum; the second prosthomere has a well-marked coelomic cavity, carries the antennae, and has the deuterocerebrum for its neuromere. The third prosthomere is represented by a well-marked pair of coelomic cavities and the tritocerebrum (III, fig. 7), but has no appendages. They appear to have aborted. The existence of this third prosthomere corresponding to the third prosthomere of the Crustacea is a strong argument for the derivation of the Hexapoda, and with them the Chilopoda, from some offshoot of the Crustacean stem or class. The buccal somite, with its mandibles, is in Hexapoda, as in Crustacea, the fourth: they are tetartognathous.
The adhesion of a greater or less number of somites to the buccal somite posteriorly (opisthomeres) is a matter of importance, but of minor importance, in the theory and history of the Arthropod head. In Peripatus no such adhesion or fusion occurs. In Diplopoda two opisthomeres—that is to say, one in addition to the buccal somite—are united by a fusion of their terga with the terga of the prosthomeres. Their appendages are respectively the mandibles and the gnathochilarium.
In Arachnida the highest forms exhibit a fusion of the tergites of five post-oral somites to form one continuous carapace united with the terga of the two prosthomeres. The five pairs of appendages of the post-oral somites of the head or prosoma thus constituted all primitively carry gnathobasic projections on their coxal joints, which act as hemignaths: in the more specialized forms the mandibular gnathobases cease to develop.
In Crustacea the fourth or mandibular somite never has less than the two following somites associated with it by the adaptation of their appendages as jaws, and the ankylosis of their terga with that of the prosthomeres. But in higher Crustacea the cephalic “tagma” is extended, and more somites are added to the fusion, and their appendages adapted as jaws of a kind.
|Fig. 8. — Diagram of the somite-appendage or parapodium of a Polychaet Chaetopod. The chaetae are omitted.|
|nr.l1, nr.l2, Neuropodial lobes or endites.|
|nt.l1, nt.l2, Notopodial lobes or exiles.|
|The parapodium is represented with its neural or ventral surface uppermost.|
The Hexapoda are not known to us in their earlier or more primitive manifestations; we only know them as possessed of a definite number of somites arranged in definite numbers in three great tagmata. The head shows two jaw-bearing somites besides the mandibular somite (V, VI, in fig. 7)—thus six in all (as in some Crustacea), including prosthomeres, all ankylosed by their terga to form a cephalic shield. There is, however, good embryological evidence in some Hexapods of the existence of a seventh somite, the supra-lingual, occurring between the somite of the mandibles and the somite of the first maxillae (4). This segment is indicated embryologically by its paired coelomic cavities. It is practically an excalated somite, having no existence in the adult. It is probably not a mere coincidence that the Hexapod, with its two rudimentary somites devoid of appendages, is thus found to possess twenty-one somites, including that which carries the anus, and that this is also the number present in the Malacostracous Crustacea.
The Segmented Lateral Appendages or Limbs of Arthropoda.—It has taken some time to obtain any general acceptance of the view that the parapodia of the Chaetopoda and the limbs of Arthropoda are genetically identical structures; yet if we compare the parapodium of Tomopteris or of Phyllodoce with one of the foliaceous limbs of Branchipus or Apus, the correspondences of the two are striking. An erroneous view of the fundamental morphology of the Crustacean limb, and consequently of that of other Arthropoda, came into favour owing to the acceptance of the highly modified limbs of Astacus as typical. Protopodite, endopodite, exopodite, and epipodite were considered to be the morphological units of the crustacean limb. Lankester (5) has shown (and his views have been accepted by Professors Korschelt and Heider in their treatise on Embryology) that the limb of the lowest Crustacea, such as Apus, consists of a corm or axis which may be jointed, and gives rise to outgrowths, either leaf-like or filiform, on its inner and outer margins (endites and exites). Such a corm (see figs. 10 and 11), with its outgrowths, may be compared to the simple parapodia of Chaetopoda with cirrhi and branchial lobe (fig. 8). It is by the specialization of two “endites” that the endopodite and exopodite of higher Crustacea are formed, whilst a flabelliform exite is the homogen or genetic equivalent of the epipodite (see Lankester, “Observations and Reflections on Apus Cancriformis,” Q. J. Micr. Sci.). The reduction of the outgrowth-bearing “corm” of the parapodium of either a Chaetopod or an Arthropod to a simple cylindrical stump, devoid of outgrowths, is brought about when mechanical conditions favour such a shape. We see it in certain Chaetopods (e.g. Hesione) and in the Arthropod Peripatus (fig. 9). The conversion of the Arthropod’s limb into a jaw, as a rule, is effected by the development of an endite near its base into a hard, chitinized, and often toothed gnathobase (see figs. 10 and 11, en1). It is not true that all the biting processes of the Arthropod limb are thus produced—for instance, the jaws of Peripatus are formed by the axis or corm itself, whilst the poison-jaws of Chilopods, as also their maxillae, appear to be formed rather by the apex or terminal region of the ramus of the limb; but the opposing jaws (= hemignaths) of Crustacea, Arachnida and Hexapoda are gnathobases, and not the axis or corm. The endopodite (corresponding to the fifth endite of the limb of Apus, see fig. 10) becomes in Crustacea the “walking leg” of the mid-region of the body; it becomes the palp or jointed process of anterior segments. A second ramus, the “exopodite,” often is also retained in the form of a palp or feeler. In Apus, as the figure shows, there are four of these “antenna-like” palps or filaments on the first thoracic limb. A common modification of the chief ramus of the Arthropod parapodium is the chela or nipper formed by the elongation of the penultimate joint of the ramus, so that the last joint works on it—as, for instance, in the lobster’s claw. Such chelate rami or limb-branches are independently developed in Crustacea and in Arachnida, and are carried by somites of the body which do not correspond in position in the two groups. The range of modification of which the rami or limb-branches of the limbs of Arthropoda are capable is very large, and in allied orders or even families or genera we often find what is certainly the palp of the same appendage (as determined by numerical position of the segments)—in one case antenniform, in another chelate, in another pediform, and in another reduced to a mere stump or absent altogether. Very probably the power which the appendage of a given segment has of assuming the perfected form and proportions previously attained by the appendage of another segment must be classed as an instance of “homoeosis,” not only where such a change is obviously due to abnormal development or injury, but also where it constitutes a difference permanently established between allied orders or smaller groups, or between the two sexes.
Fig. 9. — Three somite-appendages or parapodia of Peripatus.
|A, A walking leg; p1 to p4, the characteristic “pads”; f, the foot; cl1, cl3, the two claws.|
|B, An oral papilla, one of the second pair of post-oral appendages.|
|C, One of the first post-oral pair of appendages or mandibles; cl1, cl2, the greatly enlarged claws. (Compare A.)|
|The appendages are represented with the neural or ventral surface uppermost.|
The most extreme disguise assumed by the Arthropod parapodium or appendage is that of becoming a mere stalk supporting an eye—a fact which did not obtain general credence until the experiments of Herbst in 1895, who found, on cutting off the eye-stalk of Palaemon, that a jointed antenna-like appendage was regenerated in its place. Since the eye-stalks of Podophthalmate Crustacea represent appendages, we are forced to the conclusion that the sessile eyes of other Crustacea, and of other Arthropoda generally, indicate the position of appendages which have atrophied.
From what has been said, it is apparent that we cannot, in attempting to discover the affinities and divergences of the various forms of Arthropoda, attach a very high phylogenetic value to the coincidence or divergence in form of the appendages belonging to the somites compared with one another.
After Lankester, Q. J. Mic. Sci. vol. xxi., 1881.
|Fig. 10. — The second thoracic (fifth post-oral) appendage of the left side of Apus cancriformis, placed with its ventral or neural surface uppermost to compare with figs. 8 and 9.|
|1, 2,||The two segments of the axis.|
|en2 to en6, The five following “endites.”|
|fl,||The flabellum or anterior exite.|
|br,||The bract or posterior exite.|
The principal forms assumed by the Arthropod parapodium and its rami may be thus enumerated:—
(1) Axial corm well developed, unsegmented or with two to four segments; lateral endites and exites (rami) numerous and of various lengths (certain limbs of lower Crustacea).
(2) Corm, with short unsegmented rami, forming a flattened foliaceous appendage, adapted to swimming and respiration (trunk-limbs of Phyllopods).
(3) Corm alone developed; with no endites or exites, but provided with terminal chitinous claws (ordinary leg of Peripatus), with terminal jaw teeth (jaw of Peripatus), or with blunt extremity (oral papilla of same) (see fig. 9).
(4) Three of the rami of the primitive limb (endites 5 and 6, and exite 1) specially developed as endopodite, exopodite, and epipodite—the first two often as firm and strongly chitinized, segmented, leg-like structures; the original axis or corm reduced to a basal piece, with or without a distinct gnathobase (endite 1)—typical tri-ramose limb of higher Crustacea.
(5) One ramus (the endopodite) alone developed—the original axis or corm serving as its basal joint with or without gnathobase. This is the usual uni-ramose limb found in the various classes of Arthropoda. It varies as to the presence or absence of the jaw-process and as to the stoutness of the segments of the ramus, their number (frequently six, plus the basal corm), and the modification of the free end. This may be filiform or brush-like or lamellate when it is an antenna or palp; a simple spike (walking leg of Crustacea, of other aquatic forms, and of Chilopods and Diplopods); the terminal joint flattened (swimming leg of Crustacea and Gigantostraca); the terminal joint provided with two or with three recurved claws (walking leg of many terrestrial forms—e.g. Hexapoda and Arachnida); the penultimate joint with a process equal in length to the last joint, so as to form a nipping organ (chelae of Crustaceans and Arachnids); the last joint reflected and movable on the penultimate, as the blade of a clasp-knife on its handle (the retrovert, toothed so as to act as a biting jaw in the Hexapod Mantis, the Crustacean Squilla and others); with the last joint produced into a needle-like stabbing process in spiders.
After Lankester, Q. J. Mic. Sci. vol. xxi., 1881.
|Fig. 11. — The first thoracic (fourth post-oral) appendage of Apus cancriformis (right side).|
|Ax1 to Ax4,||the four segments of the axis with muscular bands.|
|En2 to En5, The elongated jointed endites (rami).|
|En6,||The rudimentary sixth endite (exopodite of higher Crustacea).|
|Fl,||The flabellum which becomes the epipodite of higher forms.|
|Br,||The bract devoid of muscles and respiratory in function.|
(6) Two rami developed (usually, but perhaps not always, the equivalents of the endopodite and exopodite) supported on the somewhat elongated corm (basal segment). This is the typical “bi-ramose limb” often found in Crustacea. The rami may be flattened for swimming, when it is “a bi-ramose swimmeret,” or both or only one may be filiform and finely annulate; this is the form often presented by the antennae of Crustacea, and rarely by prae-oral appendages in other Arthropods.
(7) The endopoditic ramus is greatly enlarged and flattened, without or with only one jointing, the corm (basal segment) is evanescent; often the plate-like endopodites of a pair of such appendages unite in the middle line with one another or by the intermediary of a sternal up-growth and form a single broad plate. These are the plate-like swimmerets and opercula of Gigantostraca and Limulus among Arachnids and of Isopod Crustaceans. They may have rudimentary exopodites, and may or may not have branchial filaments or lamellae developed on their posterior faces. The simplest form to which they may be reduced is seen in the genital operculum of the scorpion.
(8) The gnathobase becomes greatly enlarged and not separated by a joint from the corm; it acts as a hemignath or half jaw working against its fellow of the opposite side. The endopodite may be retained as a small segmented palp at the side of the gnathobase or disappear (mandible of Crustacea, Chilopoda and Hexapoda).
(9) The corm becomes the seat of a development of a special visual organ, the Arthropod eye (as opposed to the Chaetopod eye). Its jointing (segmentation) may be retained, but its rami disappear (Podophthalmous Crustacea). Usually it becomes atrophied, leaving the eye as a sessile organ upon the prae-oral region of the body (the eye-stalk and sessile lateral eyes of Arthropoda generally, exclusive of Peripatus).
(10) The forms assumed by special modification of the elements of the parapodium in the maxillae, labium, &c., of Hexapods, Chilopods, Diplopods, and of various Crustacea, deserve special enumeration, but cannot be dealt with without ample space and illustration.
It may be pointed out that the most radical difference presented in this list is that between appendages consisting of the corm alone without rami (Onychophora) and those with more or less developed rami (the rest of the Arthropoda). In the latter class we should distinguish three phases: (a) those with numerous and comparatively undeveloped rami; (b) those with three, or two highly developed rami, or with only one—the corm being reduced to the dimensions of a mere basal segment; (c) those reduced to a secondary simplicity (degeneration) by overwhelming development of one segment (e.g. the isolated gnathobase often seen as “mandible” and the genital operculum).
There is no reason to suppose that any of the forms of limb observed in Arthropoda may not have been independently developed in two or more separate diverging lines of descent.
Branchiae.—In connexion with the discussion of the limbs of Arthropods, a few words should be devoted to the gill-processes. It seems probable that there are branchial plumes or filaments in some Arthropoda (some Crustacea) which can be identified with the distinct branchial organs of Chaetopoda, which lie dorsal of the parapodia and are not part of the parapodium. On the other hand, we cannot refuse to admit that any of the processes of an Arthropod parapodium may become modified as branchial organs, and that, as a rule, branchial out-growths are easily developed, de novo, in all the higher groups of animals. Therefore, it seems to be, with our present knowledge, a hopeless task to analyse the branchial organs of Arthropoda and to identify them genetically in groups.
A brief notice must suffice of the structure and history of the Eyes, the Tracheae and the so-called Malpighian tubes of Arthropoda, though special importance attaches to each in regard to the determination of the affinities of the various animals included in this great sub-phylum.
The Eyes.—The Arthropod eye appears to be an organ of special character developed in the common ancestor of the Euarthropoda, and distinct from the Chaetopod eye, which is found only in the Onychophora where the true Arthropod eye is absent. The essential difference between these two kinds of eye appears to be that the Chaetopod eye (in its higher developments) is a vesicle enclosing the lens, whereas the Arthropod eye is a pit or series of pits into which the heavy chitinous cuticle dips and enlarges knobwise as a lens. Two distinct forms of the Arthropod eye are observed—the monomeniscous (simple) and the polymeniscous (compound). The nerve-end-cells, which lie below the lens, are part of the general epidermis. They show in the monomeniscous eye (see article Arachnida, fig. 26) a tendency to group themselves into “retinulae,” consisting of five to twelve cells united by vertical deposits of chitin (rhabdoms). In the case of the polymeniscous eye (fig. 23, article Arachnida) a single retinula or group of nerve-end-cells is grouped beneath each associated lens. A further complication occurs in each of these two classes of eye. The monomeniscous eye is rarely provided with a single layer of cells beneath its lens; when it is so, it is called monostichous (simple lateral eye of Scorpion, fig. 22, article Arachnida). More usually, by an infolding of the layer of cells in development, we get three layers under the lens; the front layer is the corneagen layer, and is separated by a membrane from the other two which, more or less, fuse and contain the nerve-end-cells (retinal layer). These eyes are called diplostichous, and occur in Arachnida and Hexapoda (fig. 24, article Arachnida).
On the other hand, the polymeniscous eye undergoes special elaboration on its lines. The retinulae become elongated as deep and very narrow pits (fig. 12 and explanation), and develop additional cells near the mouth of the narrow pit. Those nearest to the lens are the corneagen cells of this more elaborated eye, and those between the original retinula cells and the corneagen cells become firm and transparent. They are the crystalline cells or vitrella (see Watase, 7). Each such complex of cells underlying the lenticle of a compound eye is called an “ommatidium”; the entire mass of cells underlying a monomeniscous eye is an “ommataeum.” The ommataeum, as already stated, tends to segregate into retinulae which correspond potentially each to an ommatidium of the compound eye. The ommatidium is from the first segregate and consists of few cells. The compound eye of the king-crab (Limulus) is the only recognized instance of ommatidia in their simplest state. Each can be readily compared with the single-layered lateral eye of the scorpion. In Crustacea and Hexapoda of all grades we find compound eyes with the more complicated ommatidia described above. We do not find them in any Arachnida.
It is difficult in the absence of more detailed knowledge as to the eyes of Chilopoda and Diplopoda to give full value to these facts in tracing the affinities of the various classes of Arthropods. But they seem to point to a community of origin of Hexapods and Crustacea in regard to the complicated ommatidia of the compound eye, and to a certain isolation of the Arachnida, which are, however, traceable, so far as the eyes are concerned, to a distant common origin with Crustacea and Hexapoda through the very simple compound eyes (monostichous, polymeniscous) of Limulus.
Fig. 12.—Diagram to show the derivation of the unit or “ommatidium” of the compound eye of Crustacea and Hexapoda, C, from a simple monomeniscous monostichous eye resembling the lateral eye of a scorpion, A, or the unit of the compound lateral eye of Limulus (see article Arachnida, figs. 22 and 23). B represents an intermediate hypothetical form in which the cells beneath the lens are beginning to be superimposed as corneagen, vitrella and retinula, instead of standing side by side in horizontal series. The black represents the cuticular product of the epidermal cells of the ocular area, taking the form either of lens, cl, of crystalline body, cry, or of rhabdom, rhab; hy, hypodermis or epidermal cells; corn1, laterally-placed cells in the simpler stage, A, which like the nerve-end cells, vit1 and ret1, are corneagens or lens-producing; corn, specialized corneagen or lens-producing cells; vit1, potential vitrella cells with cry1, potential crystalline body now indistinguishable from retinula cells and rhabdomeres; vit, vitrella cell with cry, its contained cuticular product, the crystalline cone or body; ret1, rhab1, retinula cells and rhabdom of scorpion undifferentiated from adjacent cells, vit1; ret, retinula cell; rhab, rhabdom; nf, optic nerve-fibres. (Modified from Watase.)
The Tracheae.—In regard to tracheae the very natural tendency of zoologists has been until lately to consider them as having once developed and once only, and therefore to hold that a group “Tracheata” should be recognized, including all tracheate Arthropods. We are driven by the conclusions arrived at as to the derivation of the Arachnida from branchiate ancestors, independently of the other tracheate Arthropods, to formulate the conclusion that tracheae have been independently developed in the Arachnidan class. We are also, by the isolation of Peripatus and the impossibility of tracing to it all other tracheate Arthropoda, or of regarding it as a degenerate offset from some one of the tracheate classes, forced to the conclusion that the tracheae of the Onychophora have been independently acquired. Having accepted these two conclusions, we formulate the generalization that tracheae can be independently acquired by various branches of Arthropod descent in adaptation to a terrestrial as opposed to an aquatic mode of life. A great point of interest therefore exists in the knowledge of the structure and embryology of tracheae in the different groups. It must be confessed that we have not such full knowledge on this head as could be wished for. Tracheae are essentially tubes like blood-vessels—apparently formed from the same tissue elements as blood-vessels—which contain air in place of blood, and usually communicate by definite orifices, the tracheal stigmata, with the atmosphere. They are lined internally by a cuticular deposit of chitin. In Peripatus and the Diplopods they consist of bunches of fine tubes which do not branch but diverge from one another; the chitinous lining is smooth. In the Hexapods and Chilopods, and the Arachnids (usually), they form tree-like branching structures, and their finest branches are finer than any blood-capillary, actually in some cases penetrating a single cell and supplying it with gaseous oxygen. In these forms the chitinous lining of the tubes is thickened by a close-set spiral ridge similar to the spiral thickening of the cellulose wall of the spiral vessels of plants. It is a noteworthy fact that other tubes in these same terrestrial Arthropoda—namely, the ducts of glands—are similarly strengthened by a chitinous cuticle, and that a spiral or annular thickening of the cuticle is developed in them also. Chitin is not exclusively an ectodermal product, but occurs also in cartilaginous skeletal plates of mesoblastic origin (connective tissue). The immediate cavities or pits into which the tracheal stigmata open appear to be in many cases ectodermic in sinkings, but there seems to be no reason (based on embryological observation) for regarding the tracheae as an ingrowth of the ectoderm. They appear, in fact, to be an air-holding modification of the vasifactive connective tissue. Tracheae are abundant just in proportion as blood-vessels become suppressed. They are reciprocally exclusive. It seems not improbable that they are two modifications of the same tissue-elements. In Peripatus the stigmatic pits at which the tracheae communicate with the atmosphere are scattered and not definite in their position. In other cases the stigmata are definitely paired and placed in a few segments or in several. It seems that we have to suppose that the vasifactive tissue of Arthropoda can readily take the form of air-holding instead of blood-holding tubes, and that this somewhat startling change in its character has taken place independently in several instances—viz. in the Onychophora, in more than one group of Arachnida, in Diplopoda, and again in the Hexapoda and Chilopoda.
The Malpighian Tubes.—This name is applied to the numerous fine caecal tubes of noticeable length developed from the proctodaeal invert of ectodermal origin in Hexapods. These tubes are shown to excrete nitrogenous waste products similar to uric acid. Tubes of renal excretory function in a like position occur in most terrestrial Arthropoda—viz. in Chilopoda, Diplopoda and Arachnida. They are also found in some of the semi-terrestrial and purely aquatic Amphipod Crustaceans. But the conclusion that all such tubes are identical in essential character seems to be without foundation. The Malpighian tubes of Hexapods are outgrowths of the proctodaeum, but those of Scorpion and the Amphipod Crustacea are part of the metenteron or endodermal gut, though originating near its junction with the proctodaeum. Hence the presence or absence of such tubes cannot be used as an argument as to affinity without some discrimination. The Scorpion’s so-called Malpighian tubes are not the same organs as those so named in the other Tracheata. Such renal caecal tubes seem to be readily evolved from either metenteron or proctodaeum when the conditions of the out-wash of nitrogenous waste-products are changed by the transference from aquatic to terrestrial life. The absence of such renal caeca in Limulus and their presence in the terrestrial Arachnida is precisely on a parallel with their absence in aquatic Crustacea and their presence in the feebly branchiate Amphipoda.
Group Characters.—We shall now pass the groups of the Arthropoda in review, attempting to characterize them in such a way as will indicate their probable affinities and genetic history.
Sub-Phylum ARTHROPODA.—The characters of the sub-phylum and those of the associated sub-phyla Chaetopoda and Rotifera have been given above, as well as the general characters of the phylum Appendiculata which comprises these great sub-phyla.
(a) The integument is covered by a delicate soft cuticle (not firm or plated) which allows the body and its appendages great range of extension and contraction.
(b) The paired claws on the ends of the parapodia and the fang-like modifications of these on the first post-oral appendages (mandibles) are the only hard chitinous portions of the integument.
(c) The head is deuterognathous—that is to say, there is only one prosthomere, and accordingly the first and only pair of hemignaths is developed by adaptation of the appendages of the second somite.
(d) The appendages of the third somite (second post-oral) are clawless oral papillae.
(e) The rest of the somites carry equi-formal simple appendages, consisting of a corm or axis tipped with two chitinous claws and devoid of rami.
(f) The segmentation of the body is anomomeristic, there being no fixed number of somites characterizing all the forms included.
(g) The pair of eyes situated on the prosthomere are not of the Euarthropod type, but resemble those of Chaetopods (hence Nereid-ophthalmous).
(h) The muscles of the body-wall and gut do not consist of transversely-striped muscular fibre, but of the unstriped tissue observed also in Chaetopoda.
(i) A pair of coelomoducts is developed in every somite including the prosthomere, in which alone it atrophies in later development.
(j) The ventral nerve-cords are widely separated—in fact, lateral in position.
(k) There are no masses of nerve-cells forming a ganglion (neuromere) in each somite. (In this respect the Protarthropoda are at a lower stage than most of the existing Chaetopoda.)
(l) The genital ducts are formed by the enlargement of the coelomoducts of the penultimate somite.
With the characters of the grade; add the presence within the body of fine unbranched tracheal tubes, devoid of spiral thickening, opening to the exterior by numerous irregularly scattered tracheal pits.
Genera—Eoperipatus, Peripatopsis, Opisthopatus, &c. (See Peripatus.)
Grade C (of the Arthropoda).—Euarthropoda.
(a) Integument heavily plated with firm chitinous cuticle, allowing no expansion and retraction of regions of the body nor change of dimensions, except, in some cases, a dorso-ventral bellows movement. The separation of the heavier plates of chitin by grooves of delicate cuticle results in the hinging or jointing of the body and its appendages, and the consequent flexing and extending of the jointed pieces.
(b) Claws and fangs are developed on the branches or rami of the parapodia, not on the end of the axis or corm.
(c) The head is either deuterognathous, tritognathous, or tetartognathous.
(d) Rarely only one, and usually at least two, of the somites following the mandibular somite carry appendages modified as jaws (with exceptions of a secondary origin).
(e) The rest of the somites may all carry appendages, or only a limited number may carry appendages. In all cases the appendages primarily develop rami or branches which form the limbs, the primitive axis or corm being reduced and of insignificant size. In the most primitive stock all the post-oral appendages had gnathobasic outgrowths.
(f) The segmentation of the body is anomomeristic in the more archaic members of each class, nomomeristic in the higher members.
(g) The two eyes of Chaetopod structure have disappeared, and are replaced by the Euarthropod eyes.
(h) The muscles in all parts of the body consist of striped muscular fibre, never of unstriped muscular tissue.
(i) The coelomoducts are suppressed in most somites, and retained only as the single pair of genital ducts (very rarely more numerous) and in some also as the excretory glands (one or two pairs).
(j) The ventral nerve-cords approach one another in the mid-ventral line behind the mouth.
(k) The nerve-cells of the ventral nerve cords are segregated as paired ganglia in each somite, often united by meristic dislocation into composite ganglia.
(l) The genital ducts may be the coelomoducts of the penultimate or ante-penultimate or adjacent somite, or of a somite placed near the middle of the series, or of a somite far forward in the series.
Class 1 (of the Euarthropoda).—Diplopoda.
The head has but one prosthomere (monoprosthomerous), and is accordingly deuterognathous. This carries short-jointed antennae (in one case bi-ramose) and eyes, the structure and development of which require further elucidation. Only one somite following the first post-oral or mandibular segment has its appendages modified as jaws.
The somites of the body, except in Pauropus, either fuse after early development and form double somites with two pairs of appendages (Julus, &c.), or present legless and leg-bearing somites alternating.
Somites, anomomeristic, from 12 to 150 in the post-cephalic series.
The genital ducts open in the fourth, or between the fourth and fifth post-oral somite.
Terrestrial forms with small-jointed legs formed by adaptation of a single ramus of the appendage. Tracheae are present.
Note.—The Diplopoda include the Juliformia, the Symphyla (Scolopendrella), and Pauropoda (Pauropus). They were until recently classified with the Chilopoda (Centipedes), with which they have no close affinity, but only a superficial resemblance. (Compare the definition of the class Chilopoda.)
The movement of the legs in Diplopoda is like that of those of Peripatus, of the Phyllopod Crustacea, and of the parapodia of Chaetopoda, symmetrical and identical on the two sides of the body. The legs of Chilopoda move in alternating groups on the two sides of the body. This implies a very much higher development of nerves and muscles in the latter. (See Millipede.)
Class 2 (of the Euarthropoda).—Arachnida.
Head tritognathous and diprosthomerous—that is to say, with two prosthomeres, the first bearing typical eyes, the second a pair of appendages reduced to a single ramus, which is in more primitive forms antenniform, in higher forms chelate or retrovert. The ancestral stock was pantognathobasic—i.e. had a gnathobase or jaw process on every parapodium. As many as six pairs of appendages following the mouth may have an enlarged gnathobase actually functional as a jaw or hemignath, but a ramus is well developed on each of these appendages either as a simple walking leg, a palp or a chela. In the more primitive forms the appendage of every post-oral somite has a gnathobase and two rami; in higher specialized forms the gnathobases may be atrophied in every appendage, even in the first post-oral.
The more primitive forms are anomomeristic; the higher forms nomomeristic, showing typically three groups or tagmata of six somites each.
The genital apertures are placed on the first somite of the second tagma or mesosoma. Their position is unknown in the more primitive forms. The more primitive forms have branchial respiratory processes developed on a ramus of each of the post-oral appendages. In higher specialized forms these branchial processes become first of all limited to five segments of the mesosoma, then sunk beneath the surface as pulmonary organs, and finally atrophied, their place being taken by a well-developed tracheal system.
A character of great diagnostic value in the more primitive Arachnida is the tendency of the chitinous investment of the tergal surface of the telson to unite during growth with that of the free somites in front of it, so as to form a pygidial shield or posterior carapace, often comprising as many as fifteen somites (Trilobites, Limulus).
A pair of central monomeniscous diplostichous eyes is often present on the head. Lateral eyes also are often present which are monostichous with aggregated lenses (Limulus) or with isolated lenses (Scorpio), or are diplostichous with simple lens (Pedipalpi, Araneae, &c.).
Class 3 (of the Euarthropoda).—Crustacea.
Head tetartognathous and triprosthomerous—that is to say, with three prosthomeres; the first bearing typical eyes, the second a pair of antenniform appendages (often bi-ramose), the third a pair of appendages usually antenniform, sometimes claw-like. The ancestral stock was (as in the Arachnida) pantognathobasic, that is to say, had a gnathobase or jaw-process on the base of every post-oral appendage.
Besides the first post-oral or mandibular pair, at least two succeeding pairs of appendages are modified as jaws. These have small and insignificant rami, or none at all, a feature in which the Arachnida differ from them. The appendages of four or more additional following somites may be turned upwards towards the mouth and assist in the taking of food.
The more primitive forms (Entomostraca) are anomomeristic, presenting great variety as to number of somites, form of appendages, and tagmatic grouping; the higher forms (Malacostraca) are nomomeristic, showing in front of the telson twenty somites, of which the six hinder carry swimmerets and the five next in front ambulatory limbs. The genital apertures are neither far forward nor far backward in the series of somites, e.g. on the fourteenth post-oral in Apus, on the ninth post-oral in female Astacus and in Cyclops.
With rare exceptions, branchial plates are developed either by modification of a ramus of the limbs or as processes on a ramus, or upon the sides of the body. No tracheate Crustacea are known, but some terrestrial Isopoda develop pulmonary in-sinkings of the integument. A characteristic, comparable in value to that presented by the pygidial shield of Arachnida, is the frequent development of a pair of long appendages by the penultimate somite, which with the telson form a trifid, or, when that is small, a bifid termination to the body.
The lateral eyes of Crustacea are polymeniscous, with highly specialized retinulae like those of Hexapoda, and unlike the simpler compound lateral eyes of lower Arachnida. Monomeniscous eyes are rarely present, and when present, single, minute, and central in position.
Note.—The Crustacea exhibit a longer and more complete series of forms than any other class of Arthropoda, and may be regarded as preserving the most completely represented line of descent.
Head triprosthomerous and tetartognathous. The two somites following the mandibular or first post-oral or buccal somite carry appendages modified as maxillae. The fourth post-oral somite has its appendages converted into very large and powerful hemignaths, which are provided with poison-glands. The remaining somites carry single-clawed walking legs, a single pair to each somite. The body is anomomeristic, showing in different genera from 17 (inclusive of the anal and genital) to 175 somites behind that which bears the poison jaws. No tagmata are developed. The genital ducts open on the penultimate somite.
Tracheae are developed which are dendriform and with spiral thickening of their lining. Their trunks open at paired stigmata placed laterally in each somite of the trunk or in alternate somites. Usually the tracheae open by paired stigmata placed upon the sides of a greater or less number of the somites, but never quite regularly on alternating somites. At most they are present on all the pedigerous somites excepting the first and the last. In Scutigera there are seven unpaired dorsal stigmata, each leading into a sac whence a number of air-holding tubes project into the pencardial blood-sinus.
Renal caecal tubes (Malpighian tubes) open into the proctodaeum. (See Centipede.)
Head shown by its early development to be triprosthomerous and consequently tetartognathous. The first prosthomere has its appendages represented by the compound eyes and a protocerebrum, the second has the antennae for its appendages and a deutocerebral neuromere, the third has suffered suppression of its appendages (which corresponded to the second pair of antennae of Crustacea), but has a tritocerebrum and coelomic chamber. The mandibular somite bears a pair of gnathobasic hemignaths without rami or palps, and is followed by two jaw-bearing somites (maxillary and labial). This enumeration would give six somites in all to the head—three prosthomeres and three opisthomeres. Recent investigations (Folsom, 4) show the existence in the embryo of a prae-maxillary or supra-lingual somite which is suppressed during development. This gives seven somites to the Hexapod’s head, the tergites of which are fused to form a cephalic carapace or box. The number is significant, since it agrees with that found in Edriophthalmous Crustacea, and assigns the labium of the Hexapod to the same somite numerically as that which carries the labium-like maxillipedes of those Crustacea.
The somites following the head are strictly nomomeristic and nomotagmic. The first three form the thorax, the appendages of which are the walking legs, tipped with paired claws or ungues (compare the homoplastic claws of Scorpio and Peripatus). Eleven somites follow these, forming the abdominal “tagma,” giving thus twenty-one somites in all (as in the higher Crustacea). The somites of the abdomen all may carry rudimentary appendages in the embryo, and some of the hinder somites may retain their appendages in a modified form in adult life. Terminal telescoping of the abdominal somites and excalation may occur in the adult, reducing the obvious abdominal somites to as few as eight. The genital apertures are median and placed far back in the series of somites, viz. the female on the seventh abdominal (seventeenth of the whole series) and the male on the ninth or ante-penultimate abdominal (nineteenth of the whole series). The appendages of the eighth and tenth abdominal somites are modified as gonapophyses. The eleventh abdominal segment is the telson, usually small and soft; it carries the anus.
The Hexapoda are not only all confined to a very definite disposition of the somites, appendages and apertures, as thus indicated, but in other characters also they present the specialization of a narrowly-limited highly-developed order of such a class as the Crustacea rather than a range from lower more generalized to higher more specialized forms such as that group and also the Arachnida present. It seems to be a legitimate conclusion that the most primitive Hexapoda were provided with wings, and that the term Pterygota might be used as a synonym of Hexapoda. Many Hexapoda have lost either one pair or both pairs of wings; cases are common of wingless genera allied to ordinary Pterygote genera. Some Hexapods which are very primitive in other respects happen to be also Apterous, but this cannot be held to prove that the possession of wings is not a primitive character of Hexapods (compare the case of the Struthious Birds). The wings of Hexapoda are lateral expansions of the terga of the second and third thoracic somites. They appear to be serial equivalents (homogenous meromes) of the tracheal gills, which develop in a like position on the abdominal segments of some aquatic Hexapods.
The Hexapoda are all provided with a highly developed tracheal system, which presents considerable variation in regard to its stigmata or orifices of communication with the exterior. In some a serial arrangement of stigmata comparable to that observed in Chilopoda is found. In other cases (some larvae) stigmata are absent; in other cases again a single stigma is developed, as in the smaller Arachnida and Chilopoda, in the median dorsal line or other unexpected position. When the facile tendency of Arthropoda to develop tracheal air-tubes is admitted, it becomes probable that the tracheae of Hexapods do not all belong to one original system, but may be accounted for by new developments within the group. Whether the primitive tracheal system of Hexapoda was a closed one or open by serial stigmata in every somite remains at present doubtful, but the intimate relation of the system to the wings and tracheal gills cannot be overlooked.
The lateral eyes of Hexapoda, like those of Crustacea, belong to the most specialized type of “compound eye,” found only in these two classes. Simple monomeniscous eyes are also present in many Hexapods.
Renal excretory caeca (Malpighian tubes) are developed from the proctodaeum (not from mesenteron as in scorpion and Amphipoda).
Concluding Remarks on the Relationships to one another of the Classes of the Arthropoda.—Our general conclusion from a survey of the Arthropoda amounts to this, that whilst Peripatus, the Diplopoda, and the Arachnida represent terrestrial offshoots from successive lower grades of primitive aquatic Arthropoda which are extinct, the Crustacea alone present a fairly full series of representatives leading upwards from unspecialized forms. The latter were not very far removed from the aquatic ancestors (Trilobites) of the Arachnida, but differed essentially from them by the higher specialization of the head. We can gather no indication of the forefathers of the Hexapoda or of the Chilopoda less specialized than they are, whilst possessing the essential characteristics of these classes. Neither embryology nor palaeontology assists us in this direction. On the other hand, the facts that the Hexapoda and the Chilopoda have triprosthomerous heads, that the Hexapoda have the same total number of somites as the nomomeristic Crustacea, and the same number of opisthomeres in the head as the more terrestrial Crustacea, together with the same adaptation of the form of important appendages in corresponding somites, and that the compound eyes of both Crustacea and Hexapoda are extremely specialized and elaborate in structure and identical in that structure, all lead to the suggestion that the Hexapoda, and with them, at no distant point, the Chilopoda, have branched off from the Crustacean main stem as specialized terrestrial lines of descent. And it seems probable that in the case of the Hexapoda, at any rate, the point of departure was subsequent to the attainment of the nomomeristic character presented by the higher grade of Crustacea. It is on the whole desirable to recognize such affinities in our schemes of classification.
We may tabulate the facts as to head-structure in Chaetopoda and Arthropoda as follows:—
Grade x (below the Arthropoda).—Agnatha, Aprosthomera.
Without parapodial jaws; without the addition of originally post-oral somites to the prae-oral region, which is a simple prostomial lobe of the first somite; the first somite is perforated by the mouth and its parapodia are not modified as jaws.
Grade 1 (of the Arthropoda).—Monognatha, Monoprosthomera.
With a single pair of parapodial jaws carried by the somite which is perforated by the mouth; this is not the first somite, but the second. The first somite has become a prosthomere, and carries a pair of extensile antennae.
= Onychophora (Peripatus, &c.).
Grade 2 (of the Arthropoda).—Dignatha, Monoprosthomera.
The third somite as well as the second develops a pair of parapodial jaws; the first somite is a prosthomere carrying jointed antennae.
Grade 3 (of the Arthropoda).—Pantognatha, Diprosthomera.
A gnathobase is developed (in the primitive stock) on every pair of post-oral appendages; two prosthomeres present, the second somite as well as the first having passed in front of the mouth, but only the second has appendages.
Grade 4 (of the Arthropoda).—Pantognatha, Triprosthomera.
The original stock, like that of the last grade, has a gnathobase on every post-oral appendage, but three prosthomeres are now present, in consequence of the movement of the oral aperture from the third to the fourth somite. The later eyes are polymeniscous, with specialized vitrellae and retinulae of a definite type peculiar to this grade.
= Crustacea, Chilopoda, Hexapoda.
According to older views the increase of the number of somites in front of the mouth would have been regarded as a case of intercalation by new somite-budding of new prae-oral somites in the series. We are prohibited by a general consideration of metamerism in the Arthropoda from adopting the hypothesis of intercalation of somites. However strange it may seem, we have to suppose that one by one in the course of long historical evolution somites have passed forwards and the mouth has passed backwards. In fact, we have to suppose that the actual somite which in grades 1 and 2 bore the mandibles lost those mandibles, developed their rami as tactile organs, and came to occupy a position in front of the mouth, whilst its previous jaw-bearing function was taken up by the next somite in order, into which the oral aperture had passed. A similar history must have been slowly brought about when this second mandibulate somite in its turn became agnathous and passed in front of the mouth. The mandibular parapodia may be supposed during the successive stages of this history to have had, from the first, well-developed rami (one or two) of a palp-like form, so that the change required when the mouth passed away from them would merely consist in the suppression of the gnathobase. The solid palpless mandible such as we now see in some Arthropoda is, necessarily, a late specialization. Moreover, it appears probable that the first somite never had its parapodia modified as jaws, but became a prosthomere with tactile appendages before parapodial jaws were developed at all, or rather pari passu with their development on the second somite. It is worth while bearing in mind a second possibility as to the history of the prosthomeres, viz. that the buccal gnathobasic parapodia (the mandibles) were in each of the three grades of prosthomerism only developed after the recession of the mouth and the addition of one, of two, or of three post-oral somites to the prae-oral region had taken place. In fact, we may imagine that the characteristic adaptation of one or more pairs of post-oral parapodia to the purposes of the mouth as jaws did not occur until after ancestral forms with one, with two, and with three prosthomeres had come into existence. On the whole the facts seem to be against this supposition, though we need not suppose that the gnathobase was very large or the rami undeveloped in the buccal parapodia which were destined to lose their mandibular features and pass in front of the mouth.
References.—1. Bateson, Materials for the Study of Variation (Macmillan, 1894), p. 85; 2. Lankester, “Primitive Cell-layers of the Embryo.” Annals and Mag. Nat. Hist. (1873), p. 336; 3. Korschelt and Heider, Entwickelungsgeschichte (Jena, 1892), cap. xv. p. 389; 4. Folsom, “Development of the Mouth Parts of Anurida,” Bulletin Mus. Comp. Zool. Harvard College, vol. xxxvi. No. 5 (1900), pp. 142-146; 5. Lankester, “Observations and Reflections on the Appendages and Nervous System of Apus Cancriformis,” Quart. Journ. Micr. Sci. vol. xxi. (1881); 6. Hofer, “Ein Krebs mit einer Extremität statt eines Stielauges,” Verhandl. d. deutschen zool. Gesellsch. (1894); 7. Watase, “On the Morphology of the Compound Eyes of Arthropods,” Studies from the Biol. Lab. of the Johns Hopkins University, vol. iv. pp. 287-334; 8. Benham describes backward shifting of the oral aperture in certain Chaetopods, Proc. Zoolog. Soc. London (1900), No. lxiv. p. 976. N.B.—References to the early literature concerning the group Arthropoda will be found in Carus, Geschichte der Zoologie. The more important literature up to 1892 is given in the admirable treatise on Embryology by Professors Korschelt and Heider. Detailed references will be found under the articles on the separate groups of Arthropoda. (E. R. L.)
- The group Arthropoda itself, thus constituted, was precisely identical in its area with the Insecta of Linnaeus, the Entoma of Aristotle. But the word “Insect” had become limited since the days of Linnaeus to the Hexapod Pterygote forms, to the exclusion of his Aptera. Lamarck’s penetrating genius is chiefly responsible for the shrinkage of the word Insecta, since it was he who, forty years after Linnaeus’s death, set up and named the two great classes Crustacea and Arachnida (included by Linnaeus under Insecta as the order “Aptera”), assigning to them equal rank with the remaining Insecta of Linnaeus, for which he proposed the very appropriate class-name “Hexapoda.” Lamarck, however, appears not to have insisted on this name Hexapoda, and so the class of Pterygote Hexapods came to retain the group-name Insecta, which is, historically or etymologically, no more appropriate to them than it is to the classes Crustacea and Arachnida. The tendency to retain the original name of an old and comprehensive group for one of the fragments into which such group becomes divided by the advance of knowledge—instead of keeping the name for its logical use as a comprehensive term, including the new divisions, each duly provided with a new name—is most curiously illustrated in the history of the word physiology. Cicero says, “Physiologia naturae ratio,” and such was the meaning of the name Physiologus, given to a cyclopaedia of what was known and imagined about earth, sea, sky, birds, beasts and fishes, which for a thousand years was the authoritative source of information on these matters, and was translated into every European tongue. With the revival of learning, however, first one and then another special study became recognized—anatomy, botany, zoology, mineralogy, until at last the great comprehensive term physiology was bereft of all its once-included subject-matter, excepting the study of vital processes pursued by the more learned members of the medical profession. Professional tradition and an astute perception on their part of the omniscience suggested by the terms, have left the medical men in English-speaking lands in undisturbed but illogical possession of the words physiology, physic and physician.
- H. Milne-Edwards, who was followed by Huxley, long ago formulated the conclusion that the eye-stalks of Crustacea are modified appendages, basing his argument on a specimen of Palinurus (figured in Bateson’s book (1), in which the eye-stalk of one side is replaced by an antenniform palp. Hofer (6) in 1894 described a similar case in Astacus.
- Embryological evidence of this is still wanting. In the other classes of Arthropoda we have more or less complete embryological evidence on the subject. It appears from observation of the embryo that whilst the first prosthomere of Centipedes has its appendages reduced and represented only by eye-patches (as in Arachnida, Crustacea and Hexapoda), the second has a rudimentary antenna, which disappears, whilst the third carries the permanent antennae, which accordingly correspond to the second antennae of Crustacea, and are absent in Hexapoda.