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        <cito:cites>Grania ersei Coates, 1990</cito:cites>
        <cito:cites>Grania ersei Coates, 1990 p 17–20</cito:cites>
        <cito:cites>Coates and Stacey 1993 , p 406</cito:cites>
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        <dc:title>The diverse Grania fauna (Clitellata: Enchytraeidae) of the Esperance area, Western Australia, with descriptions of two new species</dc:title>
        <dc:creator>Rota, Emilia</dc:creator>
        <dc:creator>Wang, Hongzhu</dc:creator>
        <dc:creator>Erséus, Christer</dc:creator>
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        <bibo:journal>Journal of Natural History</bibo:journal>
        <dc:date>2007</dc:date>
        <bibo:pubDate>2010-07-29</bibo:pubDate>
        <bibo:volume>41</bibo:volume>
        <bibo:issue>17 - 20</bibo:issue>
        <bibo:pageStart>999</bibo:pageStart>
        <bibo:pageEnd>1023</bibo:pageEnd>
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        <dwc:authority>Rota and Erseus, 2000</dwc:authority>
        <dwc:authorityName>Rota and Erseus</dwc:authorityName>
        <dwc:authorityYear>2000</dwc:authorityYear>
        <dwc:box>[391,894,982,1006]</dwc:box>
        <dwc:class>Florideophyceae</dwc:class>
        <dwc:family>Acrochaetiaceae</dwc:family>
        <dwc:genus>Grania</dwc:genus>
        <dwc:kingdom>Plantae</dwc:kingdom>
        <dwc:order>Nemaliales</dwc:order>
        <dwc:pageId>9</dwc:pageId>
        <dwc:pageNumber>1006</dwc:pageNumber>
        <dwc:phylum>Rhodophyta</dwc:phylum>
        <dwc:rank>species</dwc:rank>
        <dwc:species>dolichura</dwc:species>
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        <spm:hasContent> ( Figure 4A–C)  Grania dolichura Rota and Erséus 2000, p 249–252, Figure 3A–H.  Material examined   WAM V 7308, one subadultfrom Stn  ES03-28  Band WAM V 7309, one puberal specimen (spermathecae not fully differentiated) from Stn  ES03-35B.  Description Body 9.7–10.5 mmlong, 0.17 mmwide at V, 0.20 mmat XII, comprising 84, 85 segments. Prostomium small, bluntly conical, 60–80 mmwide and 48–55 mmlong, epidermis 5.5– 6.0 mm thick at front ( Figure 4B). Chaetae ( Figure 4C) beginning in XVII–XVIII, in IV ventrally. Preclitellar chaetae up to 55 mmlong, postclitellar chaetae up to 57 mmlong. Epidermal gland cells conspicuous on anteriormost segments. A ‘‘copulatory gland’’ seen midventrally in XIV in puberal specimen. Spermathecal pores in lateral lines, slightly posterior to 4/5. Head organ containing two large ( 5–6 mm) inclusions in both specimens ( Figure 4B); other inclusions may be present but coelomocytes partly impede observation of the region. Anterior septa 7/8–8/9 thickened. Pharyngeal glands showing no ventral lobes in IV, nor any extra pairs in VII. Gut contents: brown organic matter and diatoms. Rectal ampulla spacious, bowl-shaped, filling most of pygidium, ending in a broad anus. Coelomocytes numerous, rather large ( 16–19 mmlong), filled with small, non-transparent, round granules ( Figure 4A). Dorsal blood vessel originating in L. Sperm sac reaching XXV. Sperm funnels three to fours times longer than wide (130–170 × 38–45 mm), cylindrical, with tall hyaline portion below collar. Sperm heads 20 mmlong, with evident helical structure. Vasa deferentia unmodified, showing conspicuous inner ciliation, 11 mmthick near funnel, then narrowing to about 6–8 mmthick. Penial apparati with bipartite aglandular sacs and trumpet-like, entally funnel-shaped stylets, latter 21–25 mmlong, 7 mmwide at junctions with vasa deferentia, terminating in a 1.5 mmwide tip. Stylets fully developed in puberal specimen. Each stylet housed in anterior portion of aglandular sac. Egg sac not developed. Spermathecal ampullae bulb- or pear-shaped, with small but evident inner cavity, empty (both specimens precopulatory). Ectal ducts narrow ( 18 mm), ectally bent, slightly expanded but devoid of glands at orifice.  Figure 4. (A–C)  Grania dolichura Rota and Erséus, 2000: (A) coelomocytes; (B) cephalic region, dorsolateral view; (C) ventral chaeta of segment LXVII. (D–G)  Grania ersei Coates, 1990: (D) tip of cephalic region in dorsolateral view, showing a shallow optical section of the prostomium and the typical bilobed outline of the head organ; (E) cephalic region of a different specimen. Deeper optical section through the head organ showing the location of inclusions inside the ‘‘empty’’ compartments; (F) coelomocytes; (G) terminal tract of the vas deferens and the enclosed penial stylet in an unusual loose state; the stylet length is more than twice the length of segment XII.  Remarks Neither of the Esperance specimens of  G. dolichurais fully mature but both are consistent with the original description ( Rota and Erséus 2000) from Tasmaniain diagnostic details of somatic (large size of head organ inclusions, long dorsal vessel, large chloragogen cells, thickening of anterior septa) and reproductive characters (clitellar pattern, penial bulb construction, stylet shape and size, length of sperm heads). From present observations on Tasmanian material (in E. R. collection), the two Australian populations are also consistent in the morphology of the rectum, a detail not noted previously. As concerns the pharyngeal glands, the Esperance specimens differ from the Tasmanian ones in lacking ventral lobes in IV and the extra glands in VII, while their sperm sacs extend more posteriorly. The new morphological information reported herein for  G. bykane(head organ with four small globular inclusions, entally constricted penial stylets, etc.; see above) does not strengthen the possibility of a close relationship between the two species advanced by Rota and Erséus (2000).  Distribution and habitat Originally reported from estuaries around Tasmania, where it is abundant and widespread (about 70 specimensfrom six stations) and covers a salinity range of 11–35psu,  G. dolichuranow appears to be a ‘‘southern Australian element’’ with a distribution possibly spanning across the Great Australian Bight and the South Australian Gulfs. Previously known as intertidal, in Esperance it is found in shallow subtidal ( 0.5–1.5 m) coarse sand colonized by  Posidoniaand in shelly heterogeneous sand.</spm:hasContent>
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        <spm:hasContent>   Grania ersei Coates, 1990 ( Figures 4D–G, 5A)    Grania ersei Coates, 1990 p 17–20, Figures 1A–D, 2;  Coates and Stacey 1993, p 406–408, Figure 10A–F.</spm:hasContent>
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        <spm:hasContent>  Material examined   WAM V 7310–7314, five specimensnumbered in this order: two from Stn  ES03-5 B, two from Stn  ES03-34 B, and one from Stn  ES03-39.  SMNHMain Coll. 90228-90236, nine specimens: one from Stn  ES03-4 B, two from Stn  ES03-5 B, one from Stn  ES03-21 C, and five from Stn  ES03-34  A. MCZR Oligochaeta0140–0145, six specimens: two from Stn  ES03-1 B, two from Stn  ES03-5 B, and two from Stn  ES03-15 C. All specimens fully mature, except those from Stns  ES03-1  Band ES03-39, and one from  ES03-15 C(puberal to precopulatory).  Figure 5. (A)  Grania ersei Coates, 1990, chaetae from different segments as indicated by Roman numerals. (B–D)  Grania quaerens sp. n.: (B) chaetae from different segments as indicated by Roman numerals; (C) spermatheca; (D) lateral view of the penial apparatus. (C) and (D) shown to the same scale. l, lateral chaeta; v, ventral chaeta.  Description  Body 6.1–8.1 mmlong ( n512), 0.15–0.24 mmwide at V, 0.17–0.29 mmat XII ( n516). Segments of adults 53–61 ( n512). Prostomium dome-shaped ( Figure 4D, E), 72–104 mmwide at 0/1, 56– 85 mmlong ( n57), epidermis not always reduced at front ( 5–9.5 mm). Lateral chaetae beginning in XIV– XV, more rarely in XVI–XIX. Chaetal shaft straight, stout, 3.5–6 mmthick at mid-point, proximally bent into an oblique, rather long, narrow foot with indistinct heel and low instep; tip of foot straight ( Figure 5A). Preclitellar chaetae 50–70 mmlong, with foot 13–19 mmlong; postclitellar chaetae 54–75 mmlong, with foot 16–23 mmlong. Chaetal index 3.65, n514, s50.303. Maximal length of shaft around XVIand in caudal segments ( 72–75 mm), thickness greatest in anterior body region. In the two specimensfrom ES03-34 B, chaetae smaller, shaft length by thickness ranging within 40–56 by 2.5–5 mm. Epidermal gland cells inconspicuous. Brain indented posteriorly. Head organ present, containing up to four equal- or unequalsized globules ( Figure 4D, E). Head organ not always detectable, partly because specimens on fixation had either everted their foregut or ejected refringent coelomocytes from head pore. Pharyngeal glands never extending into VII, always with ventral lobes in IV (one pair). Gut contents heterogeneous: both fine and coarse organic matter, undigested algal filaments, diatoms, fragmented sponge spicules. Rectal ampulla not or just enlarged at mid-length (olive-shaped). Nephridia not seen before clitellum. Coelomocytes numerous, narrow oval to drop-shaped, finely granular, refractile, up to 16 mmlong ( Figure 4F). Dorsal blood vessel arising in XXV. Sperm sac extending into XVI–XIX ( n516). Sperm heads 15–18 mmlong. Penial stylets ( Figure 4G) originating in posterior portion of penial apparatus, about 394– 500 mmlong ( n54), fully formed also in puberal specimens devoid of spermathecae. An adult specimen from ES03-5 Ahad two detached stylets at bottom of egg sac. Egg sac extending into XX–XXII ( n516). Ectal ducts of spermathecae very long, coiled, 13–14 mmthick, occasionally bearing a distinct, sessile ectal gland ( ES03-4 B, ES03-5 B). Sperm rings, 13– 18 mmin diameter, embedded in agranular walls of ental ampullar portion.  Remarks The Esperance material gives the opportunity to complete the original and subsequent accounts of this Western Australian taxon (Albany: Coates 1990; Rottnest Island: Coates and Stacey 1993) with respect to the occurrence of coelomocytes, the different texture of the walls in the ectal and ental portions of the spermathecal ampulla, and the presence of a head organ (although not always detectable). The latter observation rectifies what was stated previously ( Rota and Erséus 2000, p 249; based on material in C. E. collection). Apparently, there is considerable variation in chaetal size among populations from different geographic areas (Albany: 42–112mm; Rottnest: 40–68 mm; Esperance: 50– 75 mm), but the values of the chaetal index are likely to be typically low, due to the proportionally long ental foot (see Coates 1990, Figure 1B; Coates and Stacey 1993, Figure 10 A, B). As noted in other species, the penial stylets (extremely long in  G. ersei) appear fully developed before complete differentiation of the spermathecae has taken place.  Distribution and habitat   South( Albany, Esperance) and west ( Rottnest Island) coasts of Western Australia. Inintertidal and subtidal sands down to  26 m(new depth record), often among boulders and pebbles, with algal debris. This species can be locally very abundant: both previous accounts were based on rich collections (Albany material: 112 specimensfrom 26 stations; Rottnest material: 101 specimensfrom 25 stations).</spm:hasContent>
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