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PRO

Online Encyclopedia
Originally appearing in Volume V02, Page 299 of the 1911 Encyclopedia Britannica.
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PRO, Prosoma. dpm, Dorso-plastral muscle. art, Lateral artery. Ism', Tergo-sternal muscle (la-belled dv in fig. 31) of the second (pectiniferous) mesosomatic somite; this is the most anterior pair of the series of six, none are present in the genital somite. tsm', Tergo-sternal muscle of the fifth mesosomatic somite. tsms, Tergo-sternal muscle of the enlarged first metasomatic somite. Per, Pericardium. VPM' to VPM7, The series of seven pairs of veno-pericardiac muscles (labelled pv in fig. 31). There is some reason to admit the existence of another more anterior pair of these muscles in Scorpio; this would make the number exactly correspond with the number in Limulus. (After Lankester, Trans. Zool. Soc. vol. xi, i,83.) when by such an initial coincidence the two members have been particularized. The chances against these two selected members exhibiting another really independent homoplastic agreement are enormous: let us say io,000 to 1. The chances against yet another coincidence are a hundred million to one, and against yet one more " coincidence " they are the square of a hundred million to one. Homoplasy can only be assumed when the co-incidence is of a simple nature, and is such as may be reasonably supposed to have arisen by the action of like selective conditions upon like material in two separate lines of descent.' So, too, degeneration is not to be lightly assumed as the ex-planation of a simplicity of structure. There is a very definite criterion of the simplicity due to degeneration, which can in most cases be applied. Degenerative simplicity is never uniformly distributed over all the structures of the organism. It affects many or nearly all the structures of the body, but leaves some, it may be only one, at a high level of elaboration and complexity. Ancestral simplicity is more uniform, and does not co-exist with specialization and elaboration of a single organ. Further: degeneration cannot be inferred safely by the examination of an isolated case; usually we obtain a series of forms indicating the steps of a change in structure—and what we have to decide is whether the movement has been from the simple to the more complex, or from the more complex to the simple. The feathers of a peacock afford a convenient example of primitive and degenerative simplicity. The highest point of elaboration in colour, pattern and form is shown by the great eye-painted tail feathers. From these we can pass by gradual transitions in two directions, viz. either to the simple lateral tail feathers with a few rami only, developed only on one side of the shaft and of uniform metallic coloration—or to the simple contour feathers of small size, with the usual symmetrical series of numerous rami right and left of the shaft and no remarkable colouring. The one-sided specialization and the peculiar metallic colouring of the lateral tail feathers mark them as the extreme terms of a degenerative series, whilst the symmetry, likeness of constituent parts inter se, and absence of specialized pigment, as well as the fact that they differ little from any average feather of birds in general, mark the contour feather as primitively simple, and as the starting-point from which the highly elaborated eye-painted tail feather has gradually evolved. Applying these principles to the consideration of the Arachnida, we arrive at the conclusion that the smaller and simpler Arachnids are not the more primitive, but that the Acari or mites are, in fact, a degenerate group. This was maintained by Lankester in 1878 (19), again in 1881 (20); it was subsequently announced as a novelty by Claus in 1885 (21). Though the aquatic members of a class of animals are in some instances derived from terrestrial forms, the usual transition is from an aquatic ancestry to more recent land-living forms. There is no doubt, from a consideration of the facts of structure, that the aquatic water-breathing Arachnids, represented in the past by the Eurypterines and to-day by the sole survivor Limulus, have preceded the terrestrial air-breathing forms of that group. Hence we see at once that the better-known Arachnida form a series, leading from Limulus-like aquatic creatures through scorpions, spiders and harvest-men, to the degenerate Acari or mites. The spiders are specialized and reduced in apparent complexity, as compared with the scorpions, but they cannot be regarded as degenerate since the concentration of structure which occurs in them results in greater efficiency and power than are exhibited by the scorpion. The determination of the relative degree of perfection of organization attained by two animals ' A great deal of superfluous hypothesis has lately been put forward in the name of " the principle of convergence of characters " by a certain school of palaeontologists. The horse is supposed by these writers to have originated by separate lines of descent in the Old World and the New, from five-toed ancestors! And the important consequences following from the demonstration of the identity in structure of Limulus and Scorpio are evaded by arbitrary and even phantastic invocations of a mysterious transcendental force which brings about " convergence " irrespective of heredity and selection. Morphology becomes a farce when such assumptions are made. (E. R. L.)compared is difficult when we introduce, as seems inevitable, the question of efficiency and power, and do not confine the question to the perfection of morphological development. We have no measure of the degree of power manifested by various animals—though it would be possible to arrive at some conclusions as to how that "power" should be estimated. It is not possible here to discuss that matter further. We must be content to point out that it seems that the spiders, the pedipalps, and d, Chelicera. ch, Chela. cam, Camerostome. m, Mouth. ent, Entosternum. p, Pecten. stig', First pulmonary aperture. stig4, Fourth pulmonary aperture. dam, Muscle from carapace to a praeoral entosclerite. other large Arachnids have not been derived from the scorpions directly, but have independently developed from aquatic ancestors, and from one of these independent groups—probably through the harvest-men from the spiders—the Acari have finally resulted. After Benham, Trans. Zool. Soc. vol. xi., x883. Entap4, Fourth dorsal entapophysis of left side. tsm, Tergo-sternal muscles, six pairs as in Scorpio (labelled dv in fig. 31). VPM' to VPM8, The eight pairs of veno-pericardiac muscles (labelled pv in fig. 31). VPM" is probably represented in Scorpio, though not marked in figs. 30 and 31. Leaving that question for consideration in connexion with the systematic statement of the characters of the various groups of Arachnida which follows on p. 299, it is well now to consider the following question, viz., seeing that Limulus and Scorpio are such highly developed and specialized forms, and that they seem to constitute as it were the first and second steps in the series of recognized Arachnida—what do we know, or what are we led to suppose with regard to the more primitive Arachnida from which the Eurypterines and Limulus and Scorpio have sprung ? Do we know in the recent or fossil condition any such primitive Arachnids? Such a question is not only legitimate, but prompted by the analogy of at least one other great class of Arthropods. The great Arthropod class, the Crustacea, presents to the zoologist at the present day an immense range of forms, dam., en' P olv+ ~~ dv° stilt+ After Beck, Trans. Zool. Soc. VOL. xi., 1883. ad, Muscle from carapace tosternum. md, Muscle from tergite of genital somite to entosternum (same as dpm in fig. 30). dv' to dve, Dorso-ventral muscles (same as the series labelled tsm in hg. 30). pv1 to pv', The seven veno-pericardiac muscles of the right side (labelled VPM in fig. 30). to en- Sue, Suctorial pharynx. al, Alimentary canal. Ph, Pharynx. M, Mouth. Est, Entosternum. VS, Ventral venous sinus. chi, Chilaria. go, Genital operculum. br' to brs, Branchial append- ages. met, Unsegmented metasoma. comprising the primitive phyllopods, the minute copepods, the parasitic cirrhipedes and the powerful crabs and lobsters, and the highly elaborated sand-hoppers and slaters. It has been insisted, by those who accepted Lankester's original doctrine of the direct or genetic affinity of the Chaetopoda and Arthropoda, that Apus and Branchipus really come very near to the ancestral forms which connected those two great branches of Appendiculate (Parapodiate) animals.. On the other hand, the land crabs are at an immense distance from these simple forms. The record of the Crustacean family- __ps tree is, in fact, a fairly complete t\., I L pS one—the lower primitive members f t of the group are still represented f I r' ~ '.: c' by living forms in great abundance. Ut _ In the case of the Arachnida, if we 1 have to start their genealogical ' ' history with Limulus and Scorpio, 1't we are much in the same position as we should be in dealing with the Crustacea, were the whole of the Entomostraca and the whole of the Arthrostraca wiped out of existence and record. There is no possibility of doubt that the series of forms corresponding in the Arachnidan line of descent, to the forms dis- ro tinguished in the Crustacean line B of descent as the lower grade—the Entomostraca—have , ceased to exist, and not only so, but have left little evidence in the form of fossils as to their former existence and nature. It must, however, be admitted as probable that we should find some evidence, in ancient rocks or in the deep sea, of the early more primitive Arachnids. And it must be remembered that such forms must be expected to exhibit, when found, differences from Limulus and Scorpio as great as those which separate Apus and Cancer. The existing Arachnida, like the higher Crustacea, are " nomomeristic," that is to say, have a fixed typical number of somites to the body. Further, they are like the higher Crustacea, " somatotagmic," that is to say, they have this limited set of somites grouped in three (or more) " tagmata " or regions of a fixed number of similarly modified somites —each tagma differing in the modification of its fixed number of somites from that characterizing a neighbouring " tagma." The most primitive among the lower Crustacea, on the other hand, for example, the Phyllopoda, have not a fixed number of somites, some genera—even allied species—have more, some less, within wide limits; they are " anomomeristic." They also, as is generally the case with anomomeristic animals, do not exhibit any conformity to a fixed plan of " tagmatism " or division of the somites of the body into regions sharply marked off from one another; the head or prosomatic tagma is followed by a trunk consisting of somites which either graduate in character as we pass along the series or exhibit a large variety in different genera, families and orders, of grouping of the somites. They are anomotagmic, as well as anomomeristic. When it is admitted—as seems to be reasonable—that the primitive Arachnida would, like the primitive Crustacea, beanomomeristic and anomotagmic, we shall not demand of claimants for the rank of primitive Arachnids agreement with Limulus and Scorpio in respect of the exact number of their somites and the exact grouping of those somites; and when we see how diverse are the modifications of the branches of the appendages both in Arachnida and in other classes of Arthropoda (q.v.), we shall not over-estimate a difference in the form of this or that appendage exhibited by the claimant as compared with the higher Arachnids. With those considerations in mind, the claim of the extinct group of the trilobites to be considered as representatives of the lower and more primitive steps in the Arachnidan genealogy must, it seems, receive a favourable judgment. They differ from the Crustacea in that they have only a single pair of prae-oral appendages, the second pair being definitely developed as mandibles. This fact renders their association with the Crustacea impossible, if classification is to be the expression of genetic affinity inferred from structural coincidence. On the contrary, this particular point is one in which they agree with the higher Arachnida. But little is known of the structure of these extinct animals; we are therefore compelled to deal with such special points of resemblance and difference as their remains still exhibit. They had lateral eyes' which resemble no known eyes so closely as the lateral eyes of Limulus. The general form and structure of their prosomatic carapace are in many striking features identical with that of Limulus. The trilobation of the head and body—due to the expansion and flattening of the sides or "pleura" of the tegumentary skeleton—is so closely repeated in the young of Limulus that the latter has been called " the trilobite stage " of Limulus (fig. 42 compared with fig. 41). No Crustacean exhibits this trilobite form. But most important of the evidences presented by the trilobites of affinity with Limulus, and therefore with the Arachnida, is the tendency less marked in some, strongly carried out in others, to form a pygidial or telsonic shield—a fusion of the posterior somites of the body, which is precisely identical in character with the metasomatic carapace of Limulus. When to this is added the fact that a post-anal spine is developed to a large size in some trilobites (fig. 38), like that of Limulus and Scorpio, and that lateral spines on the pleura of the somites are frequent as in Limulus, and that neither metasomatic fusion of somites nor post-anal spine, nor lateral pleural spines are found in any Crustacean, nor all three together in any Arthropod besides the trilobites and Limulus—the claim of the trilobites to be considered as representing one order of a lower grade of Arachnida, comparable to the grade Entomostraca of the Crustacea, seems to be established. The fact that the single pair of prae-oral appendages of trilobites, known only as yet in one genus, is in that particular case a pair of uni-ramose antennae—does not render the association of trilobites and Arachnids improbable. Although the prae-oral pair of appendages in the higher Arachnida is usually chelate, it is not always so; in spiders it is not so; nor in many Acari. The bi-ramose structure of the post-oral limbs, demonstrated by Beecher in the trilobite Triarthrus, is no more inconsistent with its claim to be a primitive Arachnid than is the foliaceous modification of the limbs in Phyllopods inconsistent with their relationship to the Arthrostracous Crustaceans such as Gammarus and Oniscus. Thus, then, it seems that we have in the trilobites the representatives of the lower phases of the Arachnidan pedigree. The simple anomomeristic trilobite, with its equi-formal somites and equi-formal appendages, is one term of the series which ends in the even more simple but degenerate Acari. Between the two and at the highest point of the arc, so far as morphological differentiation is concerned, stands the scorpion; near to it in the trilobite's direction (that is, on the ascending side) are Limulus and the Eurypterines—with a long gap, due to obliteration of the record, separating them from the trilobite. On the ' A pair of round tubercles on the labram (camerostome or hypo-stoma) of several species of Trilobites has been described and held to be a pair of eyes (22). Sense-organs in a similar position were discovered in Limulus by Patten (42) in 1894. From Lankester, "Limulus an Arachnid." ps,-Muscular suctorial enlargement of the pharynx. sal, Prosomatic pair of gastric caeca in Scorpio, called salivary glands by some writers. c', and c2, The anterior two pairs of gastric caeca and ducts of the mesosomatic region. c', c4 and c6, Caeca and ducts of Scorpio not represented in Limulus. M, The Malpighian or renal caecal diverticula of Scorpio. pro, The proctodaeum or portion of gut leading to anus and formed embryo-logically by an inversion of the epiblast at that orifice. other side—tending downwards from the scorpion towards the Acari—are the Pedipalpi, the spiders, the book-scorpions, the harvest-men and the water-mites. The strange•nobody-crabs or Pycnogonids occupy a place on the ascending half of the arc below the Eurypterines and Limulus. They are strangely modified and degenerate, but seem to be (as explained in the systematic review) the remnant of an Arachnidan group holding the same relation to the scorpions which the Laemodipoda hold to the Podophthalmate Crustacea. We have now to offer a classification of the Arachnida and to pass in review the larger groups, with a brief statement of their structural characteristic#.. In the bibliography at the close of this article (referred to by leaded arabic numerals in brackets throughout these 'pages), the titles of works are given which contain detailed information as to the genera and species of each order or sub-order, their geographical distribution and their habits and economy so far as they have been ascertained. The limits of space do not permit of a fuller treatment of those matters here.
End of Article: PRO
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NIKOLAI MIKHAILOVICH PRJEVALSKY [PRZHEVALSKY] (1839...
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PROA (Malay, prau)

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