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The Pauropoda

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The Pauropoda "* IX «- THE PAUROPODA THE members of this group are minute, elongate, soft-bodied arthropods of the myriapod type of structure (fig. 70 A, B), but because of their relatively few legs, usually nine pairs in the adult stage, they have been named pauropods (Lubbock, 1868 ). A pauro­ pod of average size is about a millimeter in length, but some species are only half as long, and others reach a length of nearly 2 mm. Probably owing to their small size, the pauropods have no circulatory system and no tracheae or other differentiated organs of respiration. They live in moist places under logs and stones, on the ground among decaying leaves, and in the soil to a depth of several inches. The feeding habits of the pauropods are not well known, but their food has been thought to be humus and decaying plant and animal tissue. Starling (1944) says that mold fungi were observed to be the usual food of Pauropus carolinensis and that a "correlation appears to exist between the optimum temperature for mold growth in gen­ eral and high incidence of pauropod population." He gives reasons for believing that pauropods, where abundant, regardless of their small size, play a Significantpart in soil formation. A typical adult pauropod (fig. 70 B) has a relatively small, conical head and an elongate body of 12 segments, counting as segments the first and the last body divisions, which are known respectively as the collum (Col) and the pygidium (Pyg). Statements by other writers as to the number of segments may vary, because some do not include the pygidium as a segment and some exclude both the collum and the pygidium, but such differences are merely a matter of definition for a "segment." 250 THE PAUROPODA The number of legs in an adult pauropod, except in one known species, is invariably nine pairs, the first pair being on the second body segment, the last on the tenth (fig. 70 B). Preceding the pygidium is a legless segment, and the pygidium itself never bears appendages. A ten-legged pauropod, Decapauropus (F), said to have 13 body segments, has been described by Remy (1931 ). In this form the usual eleventh segment appears to be partly subdivided ventrally into two parts, the anterior of which carries the extra pair of legs; but it might be suspected that the apparent division is super­ ficial and that Decapauropus differs from other genera only in hav­ ing legs on the eleventh segment. The body segmentation of most pauropods appears to be different on the dorsal and ventral surfaces (fig. 70 B, F). In the specimen shown at A all the segments between the collum and the pygidium appear to be fully and equally developed, but generally, as seen in the genus Pauropus (B), there are only six tergal plates on the dorsum, though there are ten corresponding segmental divisions of the venter. This condition suggests the diplopod type of segmentation, but in the extended pauropod the narrowed dorsal arcs of segments 4, 6, 8, and 10 are exposed between the tergal plates of the other seg­ ments. Hansen (1902 ) has noted that live specimens of Pauropodidae "are able to elongate their body or to shorten it by contraction in a very high degree." Each of the tergal plates except the first bears a pair of long, slender tactile setae. The first segment, or collum, of the pauropod body (fig. 70 B) is quite different from the collum of a diplopod; instead of being a large tergal plate, it forms but an inconspicuous fold dorsally behind the head and is much expanded below. It bears ventrally a pair of small papillae (H, g), which have commonly been regarded as vestigial legs, but according to Tiegs (1947 ) they neither develop in the manner of limbs nor have the structure of legs, and appear rather to be comparable to the exsertile vesicles of Symphyla and apterygote insects. The pauropod legs are all six-segmented (fig. 70 I), including the small pretarsus (Ptar ). Except in the legs of the first and the last pairs, the tarsus of the adult (Tar ) is divided into two subseg­ ments. Writers who do not regard the pretarsus as a segment, state that the leg is either "Six-segmented" or "five-segmented" according as the tarsus is divided or not. That the "segmentation" of the tarsus 251 ARTHROPOD ANATOMY is not a division into true segments is shown by the fact that no muscles are attached on the distal part, and none arises in the proximal part. The entire tarsus is traversed in the usual manner by the tendon of the flexormuscle of the pretarsus (flptar ), the branches of which arise in the tibia (Tb) and the femur (Fm). The muscula­ ture thus identifies the distal segments of the leg, and there is only one segment (Tr) in the trochanteral region between the coxa and the short femur; the leg, therefore, is Six-segmented in all cases, regardless of the subsegmentation of the tarsus. The structure of the pretarsus is variable; there is a median claw and usually a ventral lobe, or empodium, but one or two accessory lateral claws may be present (fig. 70 J), and the claws themselves may be padlike on their undersurfaces. The pauropod head has a unique feature in the presence of a pair of large, smooth, clear, oval areas on the sides (fig. 70 D, E, e) that have the appearance of eyes and were formerly supposed to be eyes. The sublying tissue, however, has nothing of the structure character­ istic of light receptors, and the organs are now given the meaning­ less name of "pseudoculi," their function being unknown. It is shown by Tiegs (1947) that the convex, cornealike cuticle of a pseudoculus is separated by an intervening space from a flat layer of large epi­ dermal cells beneath it, which are innervated from the protocephalon. Tiegs suggests that the organs might be responsive to vibrations, but that, since the subcuticular space is apparently filledwith liquid, the organs must be supposed to respond, by contact, to vibrations of solid objects rather than to air vibrations. Characteristic of the pauropods are the relatively large, branched antennae (fig. 70 D). Each antenna consists of a basal stalk, which is four-segmented in the adult stage and of two apical branches, one dorsal, the other ventral. The dorsal branch ends in a single slender multiarticulate flagellum; the ventral branch bears two shorter flagella and, arising between their bases, a club-shaped or globular appendage termed the globulus (f), supposed to be a special sense organ of some kind. The segments of the antennal stalk and the bases of the branches are individually musculated, but the flagella have no mus­ cles. The pauropod mouth parts include only a pair of mandibles (fig. 70 H, Md) and an underlip structure (Mx, mxS ), the latter, said to be composed of the appendages and sternum of the firstmaxillary 252 THE PAUROPODA Fig. 70. Pauropoda. (B, C, E from Tiegs, 1947; D, G, H, I from Silvestri, 1902. ) A, a pauropod (unidentified) with regular segmentation on the dorsum. B, Pauropus silvaticus Tiegs, fully extended adult. C, same, first instar. D, Stylo­ pauropus pubescens Hansen, head and right antenna, dorsal. E, Pauropus silva­ ticus Tiegs, underside of head, with right mandible in place and head apodemes. F, Decapauropus cuenoti Remy (from Remy, 1931 ). G, A1Zopauropus brevisetus Silv., mandible. H, same, head and collum segment, ventral. I, same, seg­ mentation and musculature of a leg. J, a pauropod leg with accessory pretarsal claw and empodium. Ant, antenna; Col, collum (first body segment ); Cx, coxa; d, ligament from mandible to head apodeme; e, "pseudoculus"; Ephy, epipharyngeal surface; f, "globulus" of antenna; f/ptar, flexor muscle of pretarsus; FIt, fultural sclerite; Fm, femur; g, ventral papillae of collum segment; hAp, head apodeme; Md, mandible; Mx, maxilla; mxS, maxillary sternum; Oe, oesophagus; Ptar, pretarsus; Pyg, pygidium; Tar, tarsus; Tb, tibia; Tr, trochanter. 253 ARTHROPOD ANATOMY segment, appears, therefore, to be comparable with the diplopod gnathochilarium. The pauropod mandibles, however, have no like­ ness to the jaws of a diplopod; the gnathal lobes are solid extensions from the elongate, tapering bases (G), which, instead of being fully exposed on the sides of the cranium, are deeply sunken into the head (E, Md), as are the mandibles of a chilopod. Insofar as the pauropod mandibles are solid, single-piece appendages they resem­ ble in particular the mandibles of a geophilid chilopod. In Pauropus, according to Tiegs (1947), the tapering basal part of each mandible is connected by a ligament (E, d) to the head apodeme (hAp) of the same side and by a lateral ligament (not shown in the figure) to the cranial wall just behind the pseudoculus. This lateral liga­ ment is suggestive of the slender rod connecting the end of the chilopod mandible with the cranial margin (fig. 59 B, mdr), which will be met with again in the Protura and Collembola. The ventrally concave distal ends of the pauropod mandibles are provided with combs of delicate curved blades (fig. 70 H), and together with a median elevation of the epipharyngeal surface, Tiegs says, they form the upper wall of a tubular food passage above the grooved Hoor of the preoral cavity. The mandibular musculature is described by Tiegs as consisting of protractor, retractor, depressor, arid adductor muscles, the last attached on the head apodemes. Associated with the mandibles is a suspensory apparatus of the same nature as the hypopharyngeal fulturae of the chilopods. It con­ sists of a pair of slender, transverse, premandibular sclerites in the ventral head wall attached laterally on the cranial margins (fig.
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