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Arthropoda

The Segmental 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 identi cal structures; yet if we compare the para podium of Tomopteris or of Phyllodoce with one of the foliaceous limbs of Branchipus or __; Apus, the correspond ences 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 II), 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. To and II, en'). 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 poisonjaws 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. to) 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 FIG. 6. - Diagram of the head of a Crustacean. Triprosthomerous.

FP, Frontal processes (observed in Cirrhiped nauplius-larvae) probably representing the prostomial tentacles of Chaetopods.

e, Eye.

Ant', First pair of antennae. Ant', Second pair of an- md, Mandible. [tennae. mx', First and second pairs of maxillae.

m, Mouth.

I, II, and III, The three prosthomeres.

IV, V, VI, The three somites following the mouth. P, Protocerebrum.

D, Deuterocerebrum.

T, Tritocerebrum.

(After Goodrich.) FIG. 8. - Diagram of the somite-appendage or parapodium of a Polychaet Chaetopod. The chaetae are omitted.

Ax, The axis.

nr.c, Neuropodial cirrhus.

nr.l', nr.l', Neuropodial lobes or endites. nt.c. Notopodial cirrhus.

nt.l', nt.l', Notopodial lobes or exites.

The parapodium is represented with its neural or ventral surface uppermost. (Original).

as, for instance, in the lobster's claw. Such chelate rami or limbbranchesare independently developed in Crustacea and inArachnida, 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 d z 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.

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.

A, A walking leg; p ' to p 4, Since the eye-stalks of Podophthalthe characteristic " pads "; f, mate Crustacea represent appendthe foot; cl', c1 2 , the two ages, we are forced to the conclusion claws. that the sessile eyes of other B, An oral papilla, one of Crustacea, and of other Arthropoda the second pair of post-oral generally, indicate the position of appendages. appendages which have atrophied.' C, One of the first post-oral From what has been said, it is pair of appendages or manapparent that we cannot, in attemptdibles; cl', el', the greatly ing to discover the affinities and enlarged claws. (Compare A.) divergences of the various forms of The appendages are repreArthropoda, attach a very high sented with the neural or phylogenetic value to the coincidence ventral surface uppermost. or divergence in form of the apOriginal. pendages belonging to the somites compared with one another.

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 8 limbs of lower After Lankester, Q. J. Mic. Sci. vol. xxi., 188e.

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.

I, 2; The two segments of the axis.

The gnathobase.

en' to en s , The five following " endites." f, The flabellum or anterior exite.

br, The bract or posterior exite.

' 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.

(4) Three of the rami of the primitive limb (endites 5 and 6, and exite I) 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 i)- 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 jawprocess 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, After Lankester, Q. J. Mic. Sci. vol. xxi., 1881.

FIG. II. - The first thoracic (fourth post-oral) appendage of Apus cancriformis (right side). Ax' to Ax el , the four segments of the axis with muscular bands.

En', Gnathobase.

En' to En', The elongated jointed endites (rami).

En s , 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.

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.

(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 Hexapods).

(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 FIG. 9. - Three somite-appendages or parapodia of Peripatus. 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).

e n / e n 2' (the eye-stalk and sessile lateral eyes of Arthropoda generally, exclusive of Peripatus).

(io) 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 nerveend-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.

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 bloodvessels - apparently formed from the same tissue elements as bloodvessels - 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 Pen 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))...,r,, f lens, cl, of crystalline body, cry, or of rhabdom, rhab; hy, hypodermis or epidermal cells; corn', laterallyplaced cells in the simpler stage, A, which like the nerve-end cells, vit' and ret', are corneagens or lens-producing; corn, specialized corneagen or lens-producing cells; vie, potential vitrella cells with cry', potential crystalline body now indistinguishable from retinula cells and rhabdomeres; vit, vitrella cell with cry, its contained cuticular product, the crystalline cone or body; ret', rhab', retinula cells and rhabdom of scorpion undifferentiated from adjacent cells, vit'; ret, retinula cell; rhab, rhabdom; nf, optic nerve-fibres. (Modified from Watase.) patus 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 closeset 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 subphylum 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.

Grade A. - Hyparthropoda. Hypothetical forms.

Grade B. - Protarthropoda.

(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 fanglike 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 Nereidophthalmous).

(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 (neuro mere) in each somite. (In this respect the Protarthropoda are at a lower stage than most of the existing Chaetopoda.) (1) The genital ducts are formed by the enlargement of the coelomoducts of the penultimate somite.

Class (Unica). - Onychophora.

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.

Gener

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 midventral 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.

(1) The genital ducts may be the coelomoducts of the penultimate or antepenultimate or adjacent somite, or of a somite placed near the middle of the series, or of a somite far forward in the series.

Class I (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.

Class 4.-Chilopoda.

Head triprosthomerous 1 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 pericardial blood-sinus.

Renal caecal tubes (Malpighian tubes) open into the proctodaeum. (See Centipede.) Class 5. - Hexapoda.

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, thhe 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 1 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.

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.

= Chaetopoda.

Grade' (of the Arthropoda). - M Onognatha, 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, eec.). 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.

= Diplopoda.

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.

= Arachnida.

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 pan 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, 18 94), p. 85; 2. Lankester, " Primitive Cell-layers of the Embryo," Annals and Mag. Nat. Hist. (1873), p. 33 6; 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. 142146; 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 Extremitat 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 he 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.)