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The Phreatoicoidea Family Phreatoicidae

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The Phreatoicoidea Family Phreatoicidae PAl'. & PHOC. ROY. Soc. TASMANIA, 194-:i (15TH DECEMBER, Hl4·L) 1 The Phreatoicoidea By G. E. NICHOLLS Professor of Biology, Un·i?Jersity of Western Australia (Read 16th November, 1948) PART II.-The PHREATOICIDAE Family Phreatoicidae Right mandible without lacinic~ mob1:lis. Body sub-cylindrical, pi eon compressed; head relatively long, generally with posterior process; cervical groove usually well developed; first peraeon segment normally not fused to the head; the tel son as a rule produced into terminal projection. Maxillula usually with few setospines on apex of proximal endite; coxae of peraeopods generally free from pleura of related segments; bases of hinder peraeopods moderately, or scarcely, expanded. As has already been noted (Part I, p. 25), the essential distinction between the Amphisopidae and the Phreatoicidae is found in the retention or the loss of the secondary cutting edge of the right mandible. Since a similar reduction in that appendage has also taken place independently in some Amphipoda as well as in several groups of the Isopocla, that moclification might be attributable to difference in dietary or mode of life, but such an explana­ tion will not avail for the Phreatoicidae, all of which seem to be humus feeders. ·within the sub-order, the degradation or complete disappearance of this lacinia could, of course, have occurred independently in more genera than one and, thus, forms which lack the right lncinin nwb-ais need not necessarily be near akin, but, on the other hand, its occurrence must have a phyletic significance, since its n~tention can only be interpreted as an inheritance from a common ancestor. Further, the absence of this cutting edge is generally associated with a more reduced condition of some of the other mouth parts; but, to that rule, the genus Ph1·eatoicus (s.s.) proves a notable excE~ption, the maxillula, at least, being as well, or even better, developed than it is in any Amphisopid form. Moreover, the retention of the more primitive condition of the mouth parts is usually accompanied by a greater development of the antennule, and the possession of large and prominent eyes, both features doubtless the attributes of the more aetive swimming mode of life. Part I, dealing with the An1phisopidae, was published in the 1942 volume of this Journal, and the literature referred to in Part II is given on pp. 4-5 of the 1942. volume. For explanation of lette-ring see Part l of this paper. Pap. wnd PToc. Rou. Soc. Tas., 1D42, p. 5. 2 THE PHREATOICOIDEA: PART II. Related to this, in the Amphisopidae, seems to be the condition of an apparently shortened head, on which the cervical groove can, in many species, no longer be traced, the process of cephalization having progressed so far that, in most members of that family, the first peraeon segment appears to have undergone considerable forward shifting, with the result that its appendage (the gnathopod) lies external to, instead of behind, the maxilliped. (1) In most of the Phreatoicidae, howevm·, the cervical groove persists and in many members of that family the forward shifting of the first peraeon segment seems to have been less marked, so that a visible gap may still exist between the attachments of maxilliped and gnathopod. But whether the head be short or long, a character which may perhaps be important is the relative length of the post-mandibular part of the head; associated with this is the forward production at its anterior angle into a 'posterior process'. In Pa·rwmphisopus palust1·is, although the head appears short, the post-mandibular region is relatively long and is produced into a well-marked process. Mesamphi­ sopus depressus, another species which retains many primitive features, also con­ forms to this type. All subterranean forms, whether Amphisopid or Phreatoicid, show, strongly marked, the opposite tendency (i.e., to head elongation), the lengthening being particularly noticeable in the region behind the mandible. Hyperoedesipus has no posterior process, but this is well developed in the species of Ph1·eatoicoides. In general, the head in Amphisopidae is short, and the reduc­ tion in length appears to have been effected mostly at the expense of the post­ mandibular region, while a posterior process is variable. It is well developed in !l1npkisopus spp., but absent in Phreatoicops'is. From these facts, one may perhaps draw the inferences that (1) the primitive condition was one in which the post-mandibular region was moderately long, this condition tending to become exaggerated in many subterranean species; (2) that a posterior process, moderately developed, was probably present in the ancestral form. Such a primitive condition of the head is characteristic of the majority of forms included in the second family of the sub-order-the Phnatoicidae. This includes some thirty-odd species and sub-species (more than twenty of which are new) and which are here referred to ten genera. In previous accounts, the eleven species known were all recorded under the name Phreatoicus. As noted in Part I, the Amphisopidae have a pan-Australian range and occur also in South Africa, whereas the members of the Phreatoicidae are restricted to New Zealand and the Bassian Tegion of Australia. 'l'hey fall, as will be seen from the key to genera given below, into three distinct groups, most readily separated by the form of the telson, and it is of interest that the species occurring on the periphery of the area (New Zealand and the northern fringe of the Bassian) probably most nearly resemble, in condition of telson and mouth parts, the typical Amphisopids from West Australia. A few species, found only in the Great Lake of Tasmania, retain what may well be the primitive condition of the telson, and, seemingly, a comparatively slight reduction of the condition recorded for the Carboniferous fossil, Acanthotelson. But the largest group of the species of the Phreatoicids cluster round cmstrnlis or joyneri, and these are strictly confined to the Australian Alps and Tasmania, being in the latter country, quite widespread and abundantly occurring forms. This probably represents the latest efflorescence of the sub-order, all its repre- ( 1 ) Part I, figs 1 and 2R G. E. NICHOLLS 3 sentatives being forms retaining· what must be assumed to be the docked stump only of a lJl'imarily elongate telson, reduced mouth parts, and uropods but slightly armed. This stump of the telson bears a te1·minal fringe of spines, in the great majority four in number. Such an armature characterizes the telsonic apex, not only of the species of Pantphreato·icus, but also of a very wide range of members of this sub-order, ·and it may well have been the number originally preserved on this tel sonic stump. A few, however (constituting the genus M etaphreato·icus), have six spines, an arrangement which might represent either an intermediate condition in the reduction of the telson, or be due to a secondary acquisition of a third pair of these spines developing upon the lateral border of the stump. In the present account it is regarded as secondary, (1) and the more general armature of four spines is considered as the earlier condition in this family. KEY TO GENERA A. Eyes large, prominent; tebon with terminal projection well developed, its pleura slig·htly produced; g·nathopod with spines on palm denticulate ; spine beneath insertion of rami of uropods stout, toothed. B. Telsonic projection cylindrical, backwardly directed. C. Telsonic projection relatively long, segments of peraeon deepe1· than long; bases of hinder peraeopods not expanded 11fesacnnthotelson Cl. Telsonic projection shortened; peraeon segments as long as deep; base3 of hinder peraeopods slig·htly expanded Colacanthotclson E 1 . Telsonic projection slig·htly flattened, sharply upturned; segments of peraeon deeper than long Onchotel::wn A 1. Eyes small or wanting·; teL'5onic projection reduced to sn1all upturned stump ; pleura of telson well developed; ~;rnathopod with spines on palm denticulate; spine beneath insertion of rami of uropod stout, toothed. B. Apex of telsonic projection arn1ed with two pairs of spines. C. Endopodite of first pleopod setose Paraphreatoirus C1 • Endopodite of first pleopod without setae Col1.dJotelson B 1 • Apex of telsonic projection with three pairs of spines MctaJJhrcatoiclu3 A'2. Eyes wanting; telson scarcely produced or upturned. its hinder dorsal border having an outline ranging from sub-triangular to a ·flattened convex; gnathopod with palm armed with stout sub-conical spines. B. Body vermiform, telson with flattened posterior surface; spine beneath insertion of rmni of uropods stout, shnple. C. Gnathopod practically chelate; bases of hinder peraeopods not expanded C1 • Gnathopod sub-chelate; bases of hinder peraeopods n1oderately expanded 1Veophreatoicus B 1 . Body sub-cylindrical, fusiform; bases of hinder peraeopods scarcely expanded. C. First pleopod with endopodite bearing plu1nose setae; penial stylet cylindrical with several terminal spines; spine beneath inHer- tion of rami toothed N otarn1Jhisopus C1 . First pleopod with endoporlite unarmed; penial stylet eurved. tapering, armeU with a stout spatulate spine; spine beneath insertion of rami of uropod toothed or absent Crwno~icu.R (1) There are six spines on the tnpering apex of Mesant']Jhisovus car;ens1:s {an undoubtedly prin1itlve species), but only four on the similar apex of Crenoicus mixtus; the condition of Paraphrt?atoicus 1·clictus, which retains the setosity of the first p]eopod endopodite, is, however, almost certainly primitive and in that speeies only four terminal spinr.:>s are present. 4 THE PHREATOICOIDEA: PART II. The family Phreatoicidae has been subdivided as follows:-- Sub-family VI.-PHREATOICINAE Genus-P hrect to,icus Genus-N eoplweatoicus Genus-Crenoicus Genus-N otarnphisopus Sub-family VII.-MESACANTHOTELSONINAE Genus-li!J esctcctnthotelson Genus-Onchotelson Genus-Colnccmthotelson Sub-family VIIL-PARAPHREATOICINAE Genus-Pctrctph~·eatoicus Genus-Colubotelson G en us-M etaphTeatoicus Sub-family VI.
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