The Structure and Classification of the Arachnida

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The Structure and Classification of the Arachnida STRUCTURE AND CLASSIFICATION OF THE ARACHNIDA. 165 The Structure and Classification of the Arachnida. By E. Ray Lankestcr, M.A., L.L.D., F.R.S., Director of the Natural History Departments of the Brit,ish Museum. (Reprinted by kind permission of the proprietors from the tenth edition of the 'Encyclopedia Britannica.') ARACHNIDA is the name given in 1815 by Lamarck (Greek apa\vr), a spider) to a class which he instituted for the recep- tion of the spiders, scorpions, and mites previously classified by Linnaeus in the order Aptera of his great group Insecta. Lamarck at the same time founded the class Crustacea for the lobsters, crabs, and water-fleas, also until then included in the order Aptera of Linnams. Lamarck included the Tliysanura and the Myriapoda in his class Arachnida. The Insecta of Linnseus was a group exactly equivalent to the Arthropoda founded a hundred years later by Siebold and Stannius. It was thus reduced by Lamarck in area, and made to comprise only the six-legged, wing-bearing " In- secta." For these Lamarck proposed the name Hexapoda; but that name has been little used, and they have retained to this day the title of the much larger Linnsean group, viz. Insecta. The position of the Arachnida in the great sub- phylum Arthropoda, according to recent anatomical and embryological researches, is explained in another article (AKTHEOPODA). The Arachnida form a distinct class or line of descent in the grade Euarthropoda, diverging (perhaps in common at the start with the Crustacea) from primitive Buarthropods, which gave rise also to the separate lines of VOL. 48, PART 2. — NEW SERIES. 12 166 B. RAT LANKESTER. descent known as the classes Diplopoda, Crustacea, Ohilo- poda, and Hexapoda. TIG. 1.—Entosternum, entostcrnite or plastron of Limulus polyphemus, Linn. Dorsal surface. LAP, left anterior process; RAP, right anterior process ; PAN, pharyngeal notch; ALR, anterior lateral rod or tendon ; PLR, posterior lateral rod or tendon ; PLP, posterior lateral process. Natural size. (From Lankester, ' Q. J. Micr. Sci./ N.S., vol. xxiv, 1884..) FLP.- FIG. 2.—Ventral surface of the entosternum of Limulus poly- pliemus, Linn. Letters as in Fig. 1 with the addition of NF, neural fossa protecting the aggregated ganglia of the central nervous system; PVP, left posterior ventral process; PMP, pos- terior median process. Natural size. (From Lankester.) Limulus an Arachnid.—Modern views as to the classifi- cation and affinities of the Arachnida have been determined STRUCTURE AND CLASSIFICATION OF THE ARACHNLDA. 167 by the demonstration that Limulus and the extinct Eury- pterines (Pterygotus, etc.) are Arachnida; that is to say, are identical in the structure and relation of so many important parts with Scorpio, whilst differing in those respects from other Arthropoda that it is impossible to suppose that the identity is due to homoplasy or convergence, and the con- FIG. 3. FIG. 4. FIG. 3.—Entosternum of Scorpion (Palamnccus indus, De Geer); dorsal surface, asp, paired anterior process of the sub- neural arch; sap, sub-neural arch; ap, anterior lateral process (same as RAP and LAP in Fig. 1); Imp, lateral median process (same as ALB and PLR of Fig. 1); pp, posterior process (same as PLP in Fig. 1); pf, posterior flap or diaphragm of Newport; w1 and m-, perforations of tlie diaphragm for the passage of muscles; DR, the paired dorsal ridges; GC, gastric canal or foramen; AC, arterial canal or foramen. Magnified five times linear. (After Lankester, loc. cit.) FIG. 4.—Ventral surface of the same entosternum as that drawn in Fig. 3. Letters as in Fig. 3 with the addition of NC, neural canal or foramen. (After Lankester, loc. cit.) elusion must be accepted that the resemblances arise from close genetic relationship. The view that Limulus, the king- crab, is an Arachnid was maintained as long ago as 1829 by Straus-Durkheim (1), on the ground of its possession of an internal cartilaginous sternum—also possessed by the Arach- nida (see Figs. 1—6),—and of the similarity of the disposition of the six leg-like appendages around the mouth in the two 168 E. RAY LANKESTER. cases (see Figs. 45 and 63). The evidence of the exact equivalence of the segmentation and appendages of Limulus and Scorpio, and of a number of remarkable points of agree- ment in their structure, was furnished by Lankester in an article published in 1881 ("Limulus an Arachnid," 'Quart. Journ. Micr. Sci./ vol. xxi, N.S.), and in a series of subse- quent memoirs, in which the structure of the entosteruuin, of the coxal glands, of the eyes, of the veno-pericardiac muscles, FIG. 5. FIG. 6. FIG. 5.—Entosternum of one of the mygalomorphous spiders; ventral surface. Ph.N., pharyngeal notch. The three pairs of rod- like tendons correspond to the two similar pairs in Limulus, and the posterior median process with its repetition of triangular seg- ments closely resembles the same process in Limulus. Magnified five times linear. (From Lankester, loc. cit.) FIG. 6.—Dorsal surface of the same entosternum as that drawn in Fig. 5. Ph.N., pliaryngeal notch. (After Lankester, loc. cit.) of the respiratory lamellee, and of other parts, was for the first time described, and in which the new facts discovered were shown uniformly to support the hypothesis that Lirnulus is an Arachnid. A list of these memoirs is given at the close of this article (2, 3, 4, 5, and 13). The Burypterines (Gigan- tostraca) were included in the identification, although at that time they were supposed to possess only five pairs of anterior or prosomatic appendages. They have now been shown to possess six pairs (Fig. 47), as do Limulus and Scorpio. The various comparisons previously made between the r-4 W- •• PA. FIG. 7.—Diagram of the dorsal surface of Limulus poly- piiemus. oc, lateral compound eyes; oc', central monomeniscous eyes; PA, post-anal spine; I to VI, the six appendage-bearing somites of the prosoma; VIT, probably to be considered as the tcrgum of the genital somite; VII to XII, the six somites of the mesosoma; XIII to XVIII, the six somites of the metasoma, of which the first (marked XIII at the side and 7 on the tergum) is provided with a lateral spine, and is separated by ridges from the more completely fused five hinder somites lettered 8 to 12. [This is a new figure replacing the Pig. 7 given in the ' Encyclo- paedia. It is at present a matter for further investigation as to whether the praagenital somite is merely represented by the piece marked x at the hinder border of the prosoma, or whether the area marked VII is the tergum of the praegenital somite, and that marked VIII the tergum of the genital somite. The disposition of the muscles and of the entopophyses should, when carefully studied, be sufficient to settle this point.—E. R. L.j 170 E. BAY LANKESTER. FIG. 8. structure of Limulus and the Eurypterines on the one hand, and that of a typical Arachnid, such as Scorpio, on the other, had been vitiated by erroneous notions as to the origin of the nerves supplying the anterior appendages of Limulus (which were finally removed by Alphonse Milne- Edwards in his beautiful memoir [6] on the structure of that animal), and secondly by the erroneous identification of the double FIG. 9. •XV xvn FIG. S.—Diagram of the dorsal surface of a Scorpion to compare with Fig. 7. Letters and Roman numerals as in Fig. 7, excepting that VII is here certainly the terguni of the first somite of the mesosoma—the genital somite—and is not a survival of the embry- onic prsegeniul somite. (From Lankester, loc. cit.) The anus (not seen) is on the sternal surface. FIG. 9.—Ventral view of the posterior carapace or meso-mela- somatic (opisiliosomatic) fusion of Limulus polypliemus. The soft integument ami limbs of the mesosoma have been removed as well as all the viscera aud muscles, so that the inner surface of the terga of these somites with their entopophyses are seen.' The un- sef*mented dense chitinous, sternal plate of the metasoma (XIII to XVIII) is not removed. Letters as in Fig. 7. (After Lankester, loc. cit.) STEUCTUKJi AND CLASSIFICATION 01' THE AUACHNJDA. 171 sternal plates of Limulus, called " chilaria " by Owen, with a pair of appendages (7). Once the identity of the chilaria with the pentagonal sternal FIG. 10. plate of the scorpion is recognised — an identifica- tion first insisted on by Lan- kester—the whole series of segments and appendages in the two animals, Liniulus and Scorpio, are seen to corres- pond most closely, segment for segment, with one an- other (see Figs. 7 and 8). FIG. 11. ]?IG. 10.—Ventral view of a Scorpion, Palarnnseus indus, De Geer, to show the ariargemeut of the coxae of the limbs, the sternal elements, genital plate and pcctens. M, mouth behiud the oval median camerostome; I, the chelieerse; II, the chelse ; III to VI, the four pairs of walking legs; Yllffo, the genital somite or first somite of the mesosoma with the genital operculum (a fused pair of limbs); Vlllp, the pectiniferous somite; IXs^ to XUstg, the four pulmonary somites; met, the pentagonal metasternite of the prosoma behind all the coxse; x, the sternum of the pectinifer- ous somite; y, the broad first somite of the metasoma. FIG. 11.—Third leg of Limulus polyphemus, showing the division of the fourth segment of the leg by a groove S into two, thus giving seven segments to the leg as in Scorpion. (From a drawing by Mr. Pocock.) The structure of the prosomatic appendages or legs is also seen to present many significant points of agreement (see 172 E.
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