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Originally appearing in Volume V02, Page 681 of the 1911 Encyclopedia Britannica.
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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 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.
End of Article: ARACHNIDA

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