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11. Filogranella Ben-Eliahu & Dafni, 1979 (Fig. 18) Type-species: Filogranella elatensis Ben-Eliahu & Dafni, 1979 Number of species: 1, maybe 3 Tubes white, opaque, circular in cross-section, with 5 longitudinal keels; peristomes absent. Granular overlay of the tube absent. Operculum and pseudoperculum absent (but see Remarks). Radioles arranged in semicircles, up to 9 per lobe. Inter-radiolar membrane and stylodes absent. Branchial eyes absent. Prominent prostomium with ocellar clusters. Mouth palps absent. 11–14 thoracic chaetigerous segments (Fig. 18A). Well-developed trilobed collar continuous with thoracic membranes ending in mid-thorax, around 5 th –6 th thoracic chaetiger. Tonguelets absent. Collar chaetae limbate (Fig. 18B). Apomatus chaetae present (Fig. 18C). Thoracic uncini saw- to rasp-shaped with about 16 teeth in profile and up to 4 teeth in a row above peg. Triangular depression absent. Abdominal chaetae flat sickle shaped with blunt teeth on the edge (Fig. 18E). Abdominal uncini rasp-shaped with > 20 teeth in profile, up to 7 teeth in a row; anterior peg blunt, slightly gouged underneath (Fig. 18D). Achaetous anterior abdominal zone short. Posterior capillary chaetae present. Posterior glandular pad absent. Remarks. Filogranella forms aggregations that resemble those of Filograna, but worms and their tubes are larger. However, Filograna differs from Filogranella in having fin-and-blade collar chaetae with proximal denticulate wing well separated from the distal limbate zone, a pair of well-developed thoracic membranes with apron, and rasp-shaped thoracic uncini (Ben-Eliahu & Dafni 1979). Filogranella aggregations have been mentioned from various locations around the world in diving guides (e.g., Allen & Steene 1994, 1996, Colin & Arneson 1995, Gosliner et al. 1996, Weinberg 1996, Debelius 1998, Fosså & Nilsen, 2000: Red Sea, Indonesia, Philippines, Caribbean). Some colonies have specimens with opercula with flat chitinous endplates, on a flat peduncle (more or less like that of Metavermilia) inserted as the second dorsal radiole. We suspect it is a complex of related species that needs to be sorted out. Vermiliopsis prampramiana Augener, 1918, regarded to be undeterminable by Zibrowius (1973b), might belong here by its general chaetation pattern and 9 thoracic chaetigers. Neovermilia aberrans Rullier & Amoureux, 1979, might rather belong in this genus. 1. ?Filogranella aberrans (Rullier & Amoureux, 1979), Brazil 2. Filogranella elatensis Ben-Eliahu & Dafni, 1979, Elat, Gulf of Aqaba; Indo-West Pacific, Central Atlantic, Caribbean; probably more than one species 3. ?Filogranella prampramiana (Augener, 1918), Gold Coast, Africa. 12. Filogranula Langerhans, 1884 (Fig. 19) Type-species: Filogranula gracilis Langerhans, 1884 Number of species: 6 Tube white, opaque, with elaborate peristomes; keel present. Granular overlay absent. Operculum with chitinous endplate, may have additional spines in the center. Peduncle cylindrical, smooth, without wings (however, see Remarks); inserted as second dorsal radiole on one side. Opercular constriction present (but for F. stellata, absent). Pseudoperculum absent. Radioles arranged in semi-circles, up to 7 per lobe. Inter-radiolar membrane and stylodes absent. Branchial eyes may be present. Mouth palps not observed. 7 thoracic chaetigerous segments. Collar generally non-lobed (may be trilobed) with entire edge, continuous with short thoracic membranes, ending at second thoracic chaetiger. Tonguelets absent. Collar chaetae fin-and-blade (Fig. 19A) and limbate. Apomatus chaetae present (Fig. 19B). Thoracic uncini saw- or saw-to-rasp-shaped 44 · Zootaxa 2036 © 2009 Magnolia Press TEN HOVE & KUPRIYANOVA
with 12–14 teeth in profile, up to 5 teeth in a row above anterior peg (P), blunt, gouged underneath (e.g., dental formula P:5:3:3:2:1:1:1:1:1:1:1:1:1, Fig. 19C). Triangular depression absent. Abdominal chaetae short, flat triangular with wide distal denticulate blade (Fig. 19E); abdominal uncini rasp-shaped (Fig. 19D). Achaetous anterior abdominal zone present. Long posterior capillary chaetae present. Posterior glandular pad absent. Remarks. Zibrowius (1983) and Kupriyanova (1993a) remark that the genera Chitinopoma and Filogranula are very similar in general structure of the operculum, chaetal structure, and length of the thoracic membranes. See also comments to Chitinopoma. Filogranula exilis is unusual as it shows a pinnulate radiole functioning as peduncle. 1. Filogranula annulata (O.G. Costa, 1861), Mediterranean, Portugal 2. Filogranula calyculata (O.G. Costa, 1861), Mediterranean, Lusitania, Mauretania 3. Filogranula exilis Imajima, 1979, Japan, Seychelles 4. Filogranula gracilis Langerhans, 1884, Eastern Atlantic, Hawaii? 5. Filogranula revizee Nogueira & Abbud, 2009, South Brazil 6. Filogranula stellata (Southward, 1963), Mediterranean Atlantic 13. Floriprotis Uchida, 1978 (Fig. 20) Type-species: Floriprotis sabiuraensis Uchida, 1978 Number of species: 1 Tube white, opaque, circular in cross-section, normally completely embedded into living corals; granular overlay absent. Operculum and pseudoperculum absent. Radioles arranged (semi-circular to) short pectinately, up to 20 per lobe. Inter-radiolar membrane present. Stylodes absent. Branchial eyes absent. Mouth palps absent. 7 thoracic chaetigerous segments. Collar trilobed with entire edge, tonguelets absent. Thoracic membranes long, apron present. A pair of pockets on the inner side of each thoracic membrane, between the second and third thoracic segments. Collar chaetae bayonet, with elongate rounded teeth at base of very short blade (Fig. 20A), and limbate. Apomatus chaetae absent. Thoracic uncini of Serpula type, with 4–5 teeth and pointed fang (Fig. 20B). Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge as in Serpula (Fig. 20D). Abdominal uncini similar to thoracic ones, saw-shaped with 4–5 teeth anteriorly, but rasp-to-saw-shaped (dental formula F:1:1:2:3:3:3:4:3) posteriorly (Fig. 20C), with 1 tooth proximally above fang (F) to 3 (exceptionally 4) teeth per row distally, 7–8 teeth in profile. Achaetous zone absent. Long posterior capillary chaetae present. Posterior glandular pad absent. Remarks. The species is an obligate symbiont of corals in the Indo-West Pacific (Uchida 1978, Bailey- Brock 1985, Nishi 1992b). The chaetation pattern of this species is very similar to that typical for Crucigera- Hydroides-Serpula-Spiraserpula group and, as it was suggested by its author, F. sabiuraensis is likely to be closer to that clade than to Protis. The unusual characteristic feature of this monotypic genus is the presence of pockets in the thoracic membranes, however, see Remarks following Serpula. Bailey-Brock (1985) noted that one Floriprotis specimen from Fiji did contain eggs in pockets with a few eggs under the overlapping flaps of thoracic membranes, but whether these eggs were fertilized remains unknown. Thus, there is no hard evidence that the pockets are used for incubation. Another special feature according to Uchida (1978, Plate V A) would be the presence of wart-like protuberances laterally in the collar segment. However, though never reported before, wart-like protuberances are found in some larger species of Hydroides and Serpula and in Protis hydrothermica (ten Hove & Zibrowius 1986, fig. 6q); the character had not been noted before in these taxa. TAXONOMY OF SERPULIDS: STATE OF AFFAIRS Zootaxa 2036 © 2009 Magnolia Press · 45
Page 1 and 2: ZOOTAXA 2036 Taxonomy of Serpulidae
Page 3 and 4: Zootaxa 2036: 1-126 (2009) www.mapr
Page 5 and 6: Introduction The family Serpulidae
Page 7 and 8: Morphology The tube Whereas tubes o
Page 9 and 10: FIGURE 2. Variability of serpulid t
Page 11 and 12: large crystals in the structure of
Page 13 and 14: 30A) most probably too are a series
Page 15 and 16: Throughout Spiraserpula, there is a
Page 17 and 18: FIGURE 3. Morphology of serpulid an
Page 19 and 20: FIGURE 5. General morphology of ser
Page 21 and 22: FIGURE 7. Serpulid morphology (cont
Page 23 and 24: occur incidentally in Pomatoceros,
Page 25 and 26: completely rasp-shaped (Knight-Jone
Page 27 and 28: Valid genera with diagnoses and lis
Page 29 and 30: 5. Apomatus globifer Théel, 1878,
Page 31 and 32: FIGURE 10. SEM micrographs of chaet
Page 33 and 34: tonguelets between ventral and late
Page 37 and 38: membrane. Radioles with ocellar clu
Page 39 and 40: Remarks. This relatively poorly kno
Page 41 and 42: 9. Ficopomatus Southern, 1921 (Fig.
Page 43: Filograna implexa M. Berkeley, 1835
Page 49 and 50: 14. Galeolaria Lamarck, 1818 (Fig.
Page 51 and 52: operculate H. cancerum Knight-Jones
Page 53 and 54: 14. Hydroides bulbosus ten Hove, 19
Page 55 and 56: 84. Hydroides trilobulus Chen & Wu,
Page 57 and 58: membrane and stylodes absent. Branc
Page 59 and 60: Schröder 1971), France (Fauvel 192
Page 61 and 62: most recent study (Kupriyanova et a
Page 65 and 66: FIGURE 29. Photos and SEMs of chaet
Page 69 and 70: absent. Thoracic uncini saw-to-rasp
Page 71 and 72: Tube white, opaque, circular in cro
Page 73 and 74: FIGURE 33. Photos (A-E) and a SEM m
Page 75 and 76: normal radiole and maximally coveri
Page 77 and 78: 31. Pomatoleios Pixell, 1913 (Fig.
Page 81 and 82: Jirkov (1997) further extended it t
Page 83 and 84: Paraprotula was based on the absenc
Page 85 and 86: limbate chaetae. Apomatus chaetae a
Page 91 and 92: 1. Salmacina amphidentata Jones, 19
Page 93 and 94: 7. Semivermilia torulosa (delle Chi
Page 95 and 96:
13. Serpula maorica (Benham, 1927),
Page 97 and 98:
FIGURE 47. SEM micrographs of chaet
Page 99 and 100:
limbate zone and proximal wing (Fig
Page 101 and 102:
FIGURE 49. SEM micrographs of chaet
Page 103 and 104:
46. Vitreotubus Zibrowius, 1979b (F
Page 105 and 106:
forming ventral apron. Collar chaet
Page 107 and 108:
** According to Ippolitov (2007: 26
Page 109 and 110:
38 (37) Thoracic uncini saw shaped
Page 111 and 112:
chaeta (pl. chaetae): chitinous bri
Page 113 and 114:
thoracic membranes: thin folds on b
Page 115 and 116:
Bush, K.J. (1907) Descriptions of t
Page 117 and 118:
Gravier, C. (1905 (1906?)) Sur deux
Page 119 and 120:
Hove, H.A. ten & Wolf, P.S. (1984)
Page 121 and 122:
Marsden, J.R. & Anderson, D.T. (198
Page 123 and 124:
Radl, E. (1912) Neue Lehre vom Zent
Page 125 and 126:
Verrill, A.E. (1873) Report upon th
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