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Hydroides albiceps a. AM W.46555, Darwin, Qld, b. AM W.21392 (previously as H. spiratubus holotype), Kimberley region, WA, operculum and coiling tube. Scale bar = 0.1 mm. 

Hydroides albiceps a. AM W.46555, Darwin, Qld, b. AM W.21392 (previously as H. spiratubus holotype), Kimberley region, WA, operculum and coiling tube. Scale bar = 0.1 mm. 

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Hydroides Gunnerus, 1768 is the largest and one of the economically most important genera of calcareous tubeworms (Serpulidae, Annelida) that includes a number of notorious fouling and bioinvading species. Although the representatives of the genus are typically found in shallow waters of tropical and subtropical areas worldwide, the species composi...

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Context 1
... OF AUSTRALIA absent, rarely present ( , mention only one species where it is present). Stylodes absent. Mouth palps absent. Prostomial eyes absent. Seven thoracic chaetigerous segments (checked in all Australian taxa), exceptionally more (nine in H. bisectus Imajima & ten Hove, 1989 andHydroides sp. 2 BastidaZavala & ten Hove, 2002;7-9 in H. bannerorum Bailey-Brock, 1991). Collar trilobed, tonguelets absent. Thoracic membranes long, forming ventral apron. Collar chaetae bayonet-type and limbate. All uncini saw-shaped with relatively few (up to seven) teeth; anterior fang simple pointed. Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge. Achaetous anterior abdominal zone absent. Posterior capillary chaetae present. Posterior glandular pad absent. Spines of verticil without external or lateral spinules, although tip of spines may be crescent shaped (Fig. 5) (Fig. 20) 21a. Dorsal spine narrow, with pointed tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21b. Dorsal spine more or less bulbous, thicker than other verticil spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22a. Dorsal spine with small subdistal internal spinule on the inner side, other verticil spines smooth . . . . . . . . . . H. recta (Fig. 23) 22b. Dorsal spine smooth, other verticil spines with basal internal spinule . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cf. recta (Fig. 24 (Fig. 28) 24b. Bulbous dorsal verticil spine slightly larger than other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Funnel radii with T-shaped tips, small tubercles on outer surface on bend of verticil spines absent . . . H. adamaformis (Fig. 1) 25b. Funnel radii with pointed tips, small tubercles on outer surface on bend of verticil spines present. . . . . H. tuberculata (Fig. 29) 26a. Dorsal spine with beak-like medial extension and bifid tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. malleolaspina (Fig. 15) 26b. Dorsal spine bulbous, without beak-like medial extension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. albiceps (Fig. 2) Hydroides adamaformis Pillai, 2009 ...
Context 2
... OF AUSTRALIA absent, rarely present ( , mention only one species where it is present). Stylodes absent. Mouth palps absent. Prostomial eyes absent. Seven thoracic chaetigerous segments (checked in all Australian taxa), exceptionally more (nine in H. bisectus Imajima & ten Hove, 1989 andHydroides sp. 2 BastidaZavala & ten Hove, 2002;7-9 in H. bannerorum Bailey-Brock, 1991). Collar trilobed, tonguelets absent. Thoracic membranes long, forming ventral apron. Collar chaetae bayonet-type and limbate. All uncini saw-shaped with relatively few (up to seven) teeth; anterior fang simple pointed. Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge. Achaetous anterior abdominal zone absent. Posterior capillary chaetae present. Posterior glandular pad absent. Spines of verticil without external or lateral spinules, although tip of spines may be crescent shaped (Fig. 5) (Fig. 20) 21a. Dorsal spine narrow, with pointed tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21b. Dorsal spine more or less bulbous, thicker than other verticil spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22a. Dorsal spine with small subdistal internal spinule on the inner side, other verticil spines smooth . . . . . . . . . . H. recta (Fig. 23) 22b. Dorsal spine smooth, other verticil spines with basal internal spinule . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cf. recta (Fig. 24 (Fig. 28) 24b. Bulbous dorsal verticil spine slightly larger than other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Funnel radii with T-shaped tips, small tubercles on outer surface on bend of verticil spines absent . . . H. adamaformis (Fig. 1) 25b. Funnel radii with pointed tips, small tubercles on outer surface on bend of verticil spines present. . . . . H. tuberculata (Fig. 29) 26a. Dorsal spine with beak-like medial extension and bifid tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. malleolaspina (Fig. 15) 26b. Dorsal spine bulbous, without beak-like medial extension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. albiceps (Fig. 2) Hydroides adamaformis Pillai, 2009 ...
Context 3
... OF AUSTRALIA absent, rarely present ( , mention only one species where it is present). Stylodes absent. Mouth palps absent. Prostomial eyes absent. Seven thoracic chaetigerous segments (checked in all Australian taxa), exceptionally more (nine in H. bisectus Imajima & ten Hove, 1989 andHydroides sp. 2 BastidaZavala & ten Hove, 2002;7-9 in H. bannerorum Bailey-Brock, 1991). Collar trilobed, tonguelets absent. Thoracic membranes long, forming ventral apron. Collar chaetae bayonet-type and limbate. All uncini saw-shaped with relatively few (up to seven) teeth; anterior fang simple pointed. Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge. Achaetous anterior abdominal zone absent. Posterior capillary chaetae present. Posterior glandular pad absent. Spines of verticil without external or lateral spinules, although tip of spines may be crescent shaped (Fig. 5) (Fig. 20) 21a. Dorsal spine narrow, with pointed tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21b. Dorsal spine more or less bulbous, thicker than other verticil spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22a. Dorsal spine with small subdistal internal spinule on the inner side, other verticil spines smooth . . . . . . . . . . H. recta (Fig. 23) 22b. Dorsal spine smooth, other verticil spines with basal internal spinule . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cf. recta (Fig. 24 (Fig. 28) 24b. Bulbous dorsal verticil spine slightly larger than other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Funnel radii with T-shaped tips, small tubercles on outer surface on bend of verticil spines absent . . . H. adamaformis (Fig. 1) 25b. Funnel radii with pointed tips, small tubercles on outer surface on bend of verticil spines present. . . . . H. tuberculata (Fig. 29) 26a. Dorsal spine with beak-like medial extension and bifid tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. malleolaspina (Fig. 15) 26b. Dorsal spine bulbous, without beak-like medial extension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. albiceps (Fig. 2) Hydroides adamaformis Pillai, 2009 ...
Context 4
... OF AUSTRALIA absent, rarely present ( , mention only one species where it is present). Stylodes absent. Mouth palps absent. Prostomial eyes absent. Seven thoracic chaetigerous segments (checked in all Australian taxa), exceptionally more (nine in H. bisectus Imajima & ten Hove, 1989 andHydroides sp. 2 BastidaZavala & ten Hove, 2002;7-9 in H. bannerorum Bailey-Brock, 1991). Collar trilobed, tonguelets absent. Thoracic membranes long, forming ventral apron. Collar chaetae bayonet-type and limbate. All uncini saw-shaped with relatively few (up to seven) teeth; anterior fang simple pointed. Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge. Achaetous anterior abdominal zone absent. Posterior capillary chaetae present. Posterior glandular pad absent. Spines of verticil without external or lateral spinules, although tip of spines may be crescent shaped (Fig. 5) (Fig. 20) 21a. Dorsal spine narrow, with pointed tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21b. Dorsal spine more or less bulbous, thicker than other verticil spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22a. Dorsal spine with small subdistal internal spinule on the inner side, other verticil spines smooth . . . . . . . . . . H. recta (Fig. 23) 22b. Dorsal spine smooth, other verticil spines with basal internal spinule . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cf. recta (Fig. 24 (Fig. 28) 24b. Bulbous dorsal verticil spine slightly larger than other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Funnel radii with T-shaped tips, small tubercles on outer surface on bend of verticil spines absent . . . H. adamaformis (Fig. 1) 25b. Funnel radii with pointed tips, small tubercles on outer surface on bend of verticil spines present. . . . . H. tuberculata (Fig. 29) 26a. Dorsal spine with beak-like medial extension and bifid tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. malleolaspina (Fig. 15) 26b. Dorsal spine bulbous, without beak-like medial extension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. albiceps (Fig. 2) Hydroides adamaformis Pillai, 2009 ...
Context 5
... OF AUSTRALIA absent, rarely present ( , mention only one species where it is present). Stylodes absent. Mouth palps absent. Prostomial eyes absent. Seven thoracic chaetigerous segments (checked in all Australian taxa), exceptionally more (nine in H. bisectus Imajima & ten Hove, 1989 andHydroides sp. 2 BastidaZavala & ten Hove, 2002;7-9 in H. bannerorum Bailey-Brock, 1991). Collar trilobed, tonguelets absent. Thoracic membranes long, forming ventral apron. Collar chaetae bayonet-type and limbate. All uncini saw-shaped with relatively few (up to seven) teeth; anterior fang simple pointed. Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge. Achaetous anterior abdominal zone absent. Posterior capillary chaetae present. Posterior glandular pad absent. Spines of verticil without external or lateral spinules, although tip of spines may be crescent shaped (Fig. 5) (Fig. 20) 21a. Dorsal spine narrow, with pointed tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21b. Dorsal spine more or less bulbous, thicker than other verticil spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22a. Dorsal spine with small subdistal internal spinule on the inner side, other verticil spines smooth . . . . . . . . . . H. recta (Fig. 23) 22b. Dorsal spine smooth, other verticil spines with basal internal spinule . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cf. recta (Fig. 24 (Fig. 28) 24b. Bulbous dorsal verticil spine slightly larger than other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Funnel radii with T-shaped tips, small tubercles on outer surface on bend of verticil spines absent . . . H. adamaformis (Fig. 1) 25b. Funnel radii with pointed tips, small tubercles on outer surface on bend of verticil spines present. . . . . H. tuberculata (Fig. 29) 26a. Dorsal spine with beak-like medial extension and bifid tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. malleolaspina (Fig. 15) 26b. Dorsal spine bulbous, without beak-like medial extension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. albiceps (Fig. 2) Hydroides adamaformis Pillai, 2009 ...
Context 6
... OF AUSTRALIA absent, rarely present ( , mention only one species where it is present). Stylodes absent. Mouth palps absent. Prostomial eyes absent. Seven thoracic chaetigerous segments (checked in all Australian taxa), exceptionally more (nine in H. bisectus Imajima & ten Hove, 1989 andHydroides sp. 2 BastidaZavala & ten Hove, 2002;7-9 in H. bannerorum Bailey-Brock, 1991). Collar trilobed, tonguelets absent. Thoracic membranes long, forming ventral apron. Collar chaetae bayonet-type and limbate. All uncini saw-shaped with relatively few (up to seven) teeth; anterior fang simple pointed. Triangular depression present. Abdominal chaetae flat trumpet-shaped with denticulate edge. Achaetous anterior abdominal zone absent. Posterior capillary chaetae present. Posterior glandular pad absent. Spines of verticil without external or lateral spinules, although tip of spines may be crescent shaped (Fig. 5) (Fig. 20) 21a. Dorsal spine narrow, with pointed tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21b. Dorsal spine more or less bulbous, thicker than other verticil spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22a. Dorsal spine with small subdistal internal spinule on the inner side, other verticil spines smooth . . . . . . . . . . H. recta (Fig. 23) 22b. Dorsal spine smooth, other verticil spines with basal internal spinule . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cf. recta (Fig. 24 (Fig. 28) 24b. Bulbous dorsal verticil spine slightly larger than other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Funnel radii with T-shaped tips, small tubercles on outer surface on bend of verticil spines absent . . . H. adamaformis (Fig. 1) 25b. Funnel radii with pointed tips, small tubercles on outer surface on bend of verticil spines present. . . . . H. tuberculata (Fig. 29) 26a. Dorsal spine with beak-like medial extension and bifid tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. malleolaspina (Fig. 15) 26b. Dorsal spine bulbous, without beak-like medial extension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. albiceps (Fig. 2) Hydroides adamaformis Pillai, 2009 ...
Context 7
... adamaformis is similar to H. tuberculata ( Fig. 29) in having thick bulbous verticil spines. According to , the difference between these two species lies in the number of verticil spines, which is six in H. adamaformis and five in H. tuberculata. However, in our opinion, the numerical difference might fall within the usual range of variability of the number of verticil spines in most species of Hydroides (e.g., 5-6 in H. tuberculata). Hydroides adamaformis differs from H. tuberculata in the shape of verticil spines and funnel radii. The verticil spines of H. adamaformis have an outpocket-shaped extension on each lateral side, ending in a tiny pointed knob (indicated by an arrow in Fig. 1), which is absent in H. tuberculata. Additionally, the tip of funnel radii is T-shaped in H. adamaformis, but pointed in H. ...
Context 8
... verticil small, with 7-11 spines (x=10.3, SD=0.6, n=4), straight, ending in pointed tips. AM W.44229 and AM W.45071 with all verticil spines smooth, similar in size and shape, spinules absent. SAM E3599 and ZMA with one dorsal verticil spine slightly larger than others with subterminal inner spinule, a tiny inner basal spinule present on each spine (Fig. 24). Funnel hardly chitinized, with 14-21 (x=19, SD=1.7, n=4) radii ending with pointed to swollen tips. Grooves separating radii extending 1/3 of length of wide part of funnel; funnel extending down to constriction for a length equalling ½ of that of the peduncle. Length of wide part of operculum about 0.8 mm, width 0.2 ...
Context 9
... with 90-95 (x=92, SD=2.5, n=3) chaetigers. Chaetae flat trumpet-shaped (Fig. 22e). Uncini saw- shaped anteriorly, with fang and five teeth; rasp-shaped posteriorly with 2-3 rows of teeth, five teeth in profile view (Fig. 22g). Capillaries present posteriorly (Fig. ...
Context 10
... with 90-95 (x=92, SD=2.5, n=3) chaetigers. Chaetae flat trumpet-shaped (Fig. 22e). Uncini saw- shaped anteriorly, with fang and five teeth; rasp-shaped posteriorly with 2-3 rows of teeth, five teeth in profile view (Fig. 22g). Capillaries present posteriorly (Fig. ...
Context 11
... with 90-95 (x=92, SD=2.5, n=3) chaetigers. Chaetae flat trumpet-shaped (Fig. 22e). Uncini saw- shaped anteriorly, with fang and five teeth; rasp-shaped posteriorly with 2-3 rows of teeth, five teeth in profile view (Fig. 22g). Capillaries present posteriorly (Fig. ...
Context 12
... collar chaetae bayonet with two heavy elongated conical teeth (Fig. 22b) and limbate. Uncini along entire thorax saw-shaped, with pointed fang and six curved teeth (Fig. ...
Context 13
... collar chaetae bayonet with two heavy elongated conical teeth (Fig. 22b) and limbate. Uncini along entire thorax saw-shaped, with pointed fang and six curved teeth (Fig. ...
Context 14
... unknown. BRANCHIAE: each lobe with seven radioles. PEDUNCLE: clear constriction below funnel absent. OPERCULUM: verticil with seven spines, dorsal spine swollen beak shaped, curved across the centre of verticil; other verticil spines similar in size, straight, conical, with a large subapical internal spinule each, meeting in the centre of the verticil (Fig. 26a, b). Funnel with 16 radii ending in dark, pointed tips. Grooves separating radii extending 2/5 of funnel length. Length of operculum 0.8 mm, width 0.4 ...
Context 15
... all Hydroides species described, only two species have all verticil spines similar in size and shape, with T-shaped to anchor-shaped tip: Hydroides qiui n. sp. and H. dirampha. Hydroides qiui n. sp. can be distinguished from H. dirampha by the following characters: (1) the funnel radii are T-shaped in H. qiui n. sp., but sharp in H. dirampha; (2) a smaller number of verticil spines in H. qiui n. sp. (8-9) vs. 11-15 in H. dirampha; (3) verticil spines in H. qiui n. sp. have a constriction under the anchor-shaped tips and are swollen at their base, while spines of H. dirampha are straight and flattened; (4) a basal internal spinule is absent in H. qiui n. sp., but present in H. dirampha. Combining all these differences, we conclude that H. qiui n. sp. is a new species. Other species that have all verticil spines equal in size and similarly shaped with dilated tip are H. bandaensis and H. microtis. However, while the tips of verticil spines in Hydroides qiui n. sp. (and H. dirampha) are anchor-shaped with sharp lateral points, they are globular and rounded laterally in H. bandaensis and H. microtis. Juveniles of the nominal taxa H. ancorispina, H. malleolaspina and H. novaepommeraniae Augener, 1925 may also show verticil spines with dilated tips, but the dorsalmost spine is larger, bulbous or even hammer-shaped. Hydroides sinensis Zibrowius, 1972, also with dilated verticil spines, can be immediately distinguished by its large subapical internal spinules. TUBE: white, about 1.7 (up to 2 mm) mm wide with lumen of about 1.1 (up to 1.3 mm) mm, sub-trapezoidal in cross section, with two to three longitudinal keels. BRANCHIAE: each lobe with 10-16 radioles (x=14, SD=1.9, n=10). PEDUNCLE: clear constriction present between peduncle and funnel. OPERCULUM: verticil small, with 6-10 short and slender spines (x=8, SD=1.3, n=12); dorsal verticil spine slightly larger, ending in a blunt, rectangular to dilated tip, with a small, internal, subterminal spinule. Other verticil spines ending in pointed tips, curved outwards (Fig. 23a, b). Central tooth absent. Funnel with 17-22 radii (x=19, SD=1.9, n=12), ending in bluntly rounded or swollen tips, usually thinly chitinized to and including constriction; base of funnel not chitinized. Grooves separating radii extending 1/3 of funnel length. Length of operculum about 1 (up to 1.1 mm) mm, width 1.1 (up to 1.2) ...
Context 16
... number of abdominal segments unknown. Uncini saw-shaped anteriorly, with pointed fang and 4-5 teeth; posterior chaetae unknown. SIZE: length unknown. Width of thorax 0.6 mm. COLOUR: verticil spines, funnel tips, and base of funnel dark brown. HABITAT: 20 m depth. OPERCULUM: with verticil inserted on a short stalk into proximal funnel. Verticil with 7-8 spines (x=7.4, SD=0.7, n=10). Each verticil spine sub-equal in size, curved outwards and ending with pointed tips, bearing a pair of outwardly curved lateral spinules at about half of their length, an inwardly curved radial spinule at the same level or slightly above, and a small basal radial spinule (Fig. 27). Distinct dorsal verticil spine absent. Funnel with 24-39 (x=31, SD=4.8, n=10) sharply pointed dark-brown chitinized radii, base of funnel not chitinized. Grooves separating radii extending 1/3 of funnel length. Length of operculum about 1.1 (up to 1.3) mm, width 0.8 (up to 1) ...
Context 17
... adamaformis is also similar to H. trilobula Chen & Wu, 1978 (as H. trilobulus, corrected herein) in rhomboidal spines. However, there are five verticil spines with one larger than others in H. adamaformis, but only three equal in size verticil spines in H. trilobula (according to the holotype, MBMCAS MBM79). BRANCHIAE: each lobe with up to 17 radioles, generally 11 radioles. PEDUNCLE: separate from funnel by a clear constriction. OPERCULUM: verticil with 6-13 (x=7.8, SD=1.2, n=56) spines, curved outward, with rectangular to clavate tips, without any accessory spinules. Dorsal verticil spine vesicular, consisting of a bulbous median part, and two latero-dorsal more or less triangular (seen from dorsal side) outpockets; the three parts vary considerably in relative size. Funnel with 14-31 (x=22.8, SD=3.1, n=56) clavate to bottle-shaped chitinized radii (Fig. 2a), bluntly rounded in juveniles, base of funnel not chitinized. Grooves separating radii extending 1/2 of funnel length. Length of operculum 1-2.1 mm, width 0.7-1.2 ...
Context 18
... (1906) described specimens from the Red Sea with an enlarged hook-like dorsal verticil spine bearing two lateral spinules under the name Hydroides monoceros. Fauvel (1953) considered both as separate but "closely allied" species based on his mistaken interpretation of the figures in Grube (1878 fig. 5a) and Gravier (1908 pl. 8 fig. 288) that the verticil should be symmetrical in H. minax, but asymmetrical in H. monoceros. Here we follow Hartman (1959) who synonymised H. monoceros with H. minax. Fig. 17 a-b; Map 15 Serpula (Hydroides) multispinosa Marenzeller, 1885: 216-217, pl. 4, fig. 2 [Enoshima Island, ...
Context 19
... (1906) described specimens from the Red Sea with an enlarged hook-like dorsal verticil spine bearing two lateral spinules under the name Hydroides monoceros. Fauvel (1953) considered both as separate but "closely allied" species based on his mistaken interpretation of the figures in Grube (1878 fig. 5a) and Gravier (1908 pl. 8 fig. 288) that the verticil should be symmetrical in H. minax, but asymmetrical in H. monoceros. Here we follow Hartman (1959) who synonymised H. monoceros with H. minax. Fig. 17 a-b; Map 15 Serpula (Hydroides) multispinosa Marenzeller, 1885: 216-217, pl. 4, fig. 2 [Enoshima Island, ...
Context 20
... each lobe with 14 radioles. PEDUNCLE: clear chitinized constriction below funnel present. OPERCULUM: verticil with 10 spines, nine of which similar in size and shape, curved outward, ending in outwardly curved pointed tips, without accessory spinules and an enlarged elongated dorsal spine somewhat swollen in middle, ending in a ventrally curved hook (Fig. 20). Funnel with 21 radii ending in swollen to T-shaped chitinized tips, base of funnel chitinized. Grooves separating radii extending 1/3 of funnel length. Length of operculum about 1.1 mm, width 0.8 mm. ...
Context 21
... each lobe with 7-11 radioles (x=8.6, SD=1.6, n=10). PEDUNCLE: clear chitinized constriction below funnel present. OPERCULUM: verticil inserted on a short stalk into proximal funnel. Verticil with 5-8 spines (x=7.1, SD=1, n=10), consisting of very large, dorsal spine made of a bulbous median part and two latero-dorsal extensions and 4-7 small outwardly curved spines positioned at the base of the dorsal spine, each ending in T-shaped tips, without accessory spinules (Fig. 28). Funnel with 18-25 (x=22, SD=3.3, n=10) chitinized radii tips, base of funnel half chinitized. Grooves separating radii extending 1/3 of funnel length. Length of operculum about 0.9 (up to 1.1) mm, width 0.4 (up to 0.6) ...
Context 22
... of H. albiceps show a high intraspecific variability in number, shape, and relative size of verticil spines. The number of verticil spines in Australian specimens is 6-13, while it is 8-10 in specimens from Southern China (Shin 1982;Sun et al. 2012b) and 10-14 in Japanese specimens ). The lateral extensions of the enlarged dorsal verticil spine can be situated at the tip only (Southern China specimens, ) or extend to the base of the verticil spine (specimens from Darwin, Fig. 2a). The enlarged dorsal verticil spine can be 1.5-4 times longer than smaller verticil spines in specimens collected in Darwin, which makes them similar to H. trivesiculosa . According to the descriptions of holotypes of these two species, H. albiceps differ from H. trivesiculosa in the size and shape of the dorsal verticil spine. Hydroides trivesiculosa has an exceptionally large dorsal vesicular verticil spine which is more than five times longer than the smaller verticil spines. The lateral outpockets of the dorsal verticil spine are straight in H. albiceps, while those of H. trivesiculosa are larger and curved ventrally. The enlarged dorsal verticil spine in some specimens of H. albiceps (e.g., AM W.3961) bears a stout conical medial process on the inner side of the verticil spine that extends to the base of dorsal verticil spine, which makes it even more similar to H. malleolaspina Straughan, 1967a. However, the dorsal verticil spine of H. malleolaspina is bifid-tipped, and the medial process is beak-like with a sharp tip ( Murray et al. 2010), while it is blunt and swollen in H. albiceps. compared H. albiceps, H. malleolaspina and H. trivesiculosa in detail, and showed an overlap in the number of spines and funnel radii, along with differences in the shapes of dorsal verticil spine and tubes. Hydroides albiceps is identical to H. spiratubus except for the roughly spirally coiled tube in the holotype of the latter (AM W.21443, Fig. 2b). However, spirally coiled tubes are very common in various species of Hydroides such as H. trivesiculosa (AM W.45423), H. tuberculata (AM W.45068), hence we synonymise H. spiratubus with H. ...
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... of H. albiceps show a high intraspecific variability in number, shape, and relative size of verticil spines. The number of verticil spines in Australian specimens is 6-13, while it is 8-10 in specimens from Southern China (Shin 1982;Sun et al. 2012b) and 10-14 in Japanese specimens ). The lateral extensions of the enlarged dorsal verticil spine can be situated at the tip only (Southern China specimens, ) or extend to the base of the verticil spine (specimens from Darwin, Fig. 2a). The enlarged dorsal verticil spine can be 1.5-4 times longer than smaller verticil spines in specimens collected in Darwin, which makes them similar to H. trivesiculosa . According to the descriptions of holotypes of these two species, H. albiceps differ from H. trivesiculosa in the size and shape of the dorsal verticil spine. Hydroides trivesiculosa has an exceptionally large dorsal vesicular verticil spine which is more than five times longer than the smaller verticil spines. The lateral outpockets of the dorsal verticil spine are straight in H. albiceps, while those of H. trivesiculosa are larger and curved ventrally. The enlarged dorsal verticil spine in some specimens of H. albiceps (e.g., AM W.3961) bears a stout conical medial process on the inner side of the verticil spine that extends to the base of dorsal verticil spine, which makes it even more similar to H. malleolaspina Straughan, 1967a. However, the dorsal verticil spine of H. malleolaspina is bifid-tipped, and the medial process is beak-like with a sharp tip ( Murray et al. 2010), while it is blunt and swollen in H. albiceps. compared H. albiceps, H. malleolaspina and H. trivesiculosa in detail, and showed an overlap in the number of spines and funnel radii, along with differences in the shapes of dorsal verticil spine and tubes. Hydroides albiceps is identical to H. spiratubus except for the roughly spirally coiled tube in the holotype of the latter (AM W.21443, Fig. 2b). However, spirally coiled tubes are very common in various species of Hydroides such as H. trivesiculosa (AM W.45423), H. tuberculata (AM W.45068), hence we synonymise H. spiratubus with H. ...

Citations

... Records of the nominal morphospecies Hydroides albiceps cover a wide geographic range from the Mediterranean, and Red Sea to the Indo-Pacific Ocean, South Japan and South Australia (reviewed in Sun et al., 2015). However, morphological criteria would not be enough for a taxonomic conclusion due to high intraspecific variability within the complex. ...
... Remarks. Hydroides elegans is by far the most important and successful invasive biofouling serpulid species, widely distributed in tropical, subtropical and even in temperate seas (ten Hove, 1970a;Sun et al., 2015;Schwan et al., 2016;Tovar-Hernández et al., 2016;Susick et al., 2020;Capa et al., 2021). The ability of settlement on various artificial surfaces and resistance to withstand environmental fluctuations (Mak and Huang, 1982;Qiu and Qian, 1997) are the main factors contributing to its success in ports around the world. ...
... Hydroides operculata had been reported from multiple localities of the Indo-Pacific as well as a Lessepsian migrant to the Mediterranean (Ben-Eliahu and ten Hove, 1992;Çinar, 2006;Sun et al., 2012Sun et al., , 2015. However, a partial revision of this nominal taxon by Sun et al. (2017) revealed a complex of morphologically similar and closely related regionally distributed species, including H. inornata (India and Hong Kong), H. basispinosa (Australia) and H. operculata (Gulf of Aden, Persian Gulf). ...
... In Australia, many of the earlier polychaete records were identified by European workers who largely reported them as European species (Hutchings and Glasby 1991). Recent reviews of these records have revealed greater polychaete diversity in Australian waters (Capa and Rouse 2007;Hutchings et al. 2012;Capa et al. 2013;Alvarez-Campos et al. 2015;Murray 2015a, 2015b;Sun et al. 2015). ...
Article
Sabellid fanworms (Sabellidae: Annelida) and particularly species of Branchiomma and Parasabella are difficult to discriminate due to a lack of unambiguous diagnostic characters and high intraspecific morphological plasticity. We studied specimens identified as Branchiomma and Parasabella from a marina in South Australia. Examination of morphological features alone failed to resolve the unambiguous identification of specimens to species. Therefore, two DNA markers, the nuclear internal transcribed spacer (ITS) and the mitochondrial cytochrome c oxidase I (cox1), were sequenced. Phylogenetic analyses and genetic distances were performed for each DNA marker, including Branchiomma and Parasabella sequences available in GenBank. Molecular analyses indicate that specimens belong to species that have previously been reported as cryptogenic and occurring in several worldwide harbour environments, but whose identity is still unsettled. A discussion about the relevance of several sources of information for species delineation, and the importance of taxonomy, is provided.
... Serpulidae from Hong Kong were most recently revised by Sun et al. [177], who provided illustrations, diagnoses and taxonomic keys. Tropical Australian sabellids belonging to 12 genera have been documented in a series of recent studies [68,69,[72][73][74]178]. Serpulids from Kimberley (Western Australia) were revised by Pillai [106] and those from Lizard Island (Queensland) by Kupriyanova et al. [179], whereas the revision of the genus Hydroides in Australia [180] includes both tropical and temperate species. The most comprehensive treatment of Australian sabellids and serpulids is still the interactive key by Wilson et al. [181], but it is outdated in the light of the recent studies. ...
... Sabellids from Australian temperate waters have been well documented in a series of recent papers [68,69,[72][73][74][200][201][202] along with records of temperate species. Most recent papers on temperate Serpulidae are Sun et al. [203] and Styan et al. [204], whereas the Australian Hydroides revision [180] also includes temperate species. ...
... suggest that this taxon is a complex of species. Two nominal Hydroides species, H. brachyacantha Rioja, 1941 and H. operculata (Treadwell, 1929), are examples of complexes of morphologically similar species [180,251]. Similarly, the invasive status attributed to serpulids Spirobranchus kraussii (Baird, 1865) and S. tetraceros (Schmarda, 1861) [365,439] is unjustified, as both are members of species complexes [84,193,252]. ...
Article
Full-text available
Sabellida Levinsen, 1883 is a large morphologically uniform group of sedentary annelids commonly known as fanworms. These annelids live in tubes made either of calcareous carbonate or mucus with agglutinated sediment. They share the presence of an anterior crown consisting of radioles and the division of the body into thorax and abdomen marked by a chaetal and fecal groove inversion. This study synthesises the current state of knowledge about the diversity of fanworms in the broad sense (morphological, ecological, species richness), the species occurrences in the different biogeographic regions, highlights latest surveys, provides guidelines for identification of members of each group, and describe novel methodologies for species delimitation. As some members of this group are well-known introduced pests, we address information about these species and their current invasive status. In addition, an overview of the current evolutionary hypothesis and history of the classification of members of Sabellida is presented. The main aim of this review is to highlight the knowledge gaps to stimulate research in those directions.
... The genus is one of the most ecologically and economically important groups of marine invertebrates (Lewis et al., 2006;Streftaris and Zenetos, 2006;Link et al., 2009) because it includes notorious biofoulers and bioinvaders that have been transported around the world on ship hulls, leading to high economic impact due to the costs associated with removal of the tubes from submerged structures (Hadfield, 1998). Their ecological importance has driven studies on taxonomy (e.g., Pillai, 1972;ten Hove, 2002, 2003;Sun et al., 2015), phylogeny (Sun et al., 2012, developmental, larval ecology and biofouling (e.g., Hadfield, 1998Hadfield, , 2011Qiu and Qian, 1998;Qian, 1999), ecotoxicology (e.g., Gopalakrishnan et al., 2007;Vijayaragavan and Vivek Raja, 2018), and climate change (e.g., Chan et al., 2012;Lane et al., 2013). Investigations of mitochondrial markers such as partial sequences of cytochrome c oxidase I (cox1) and cytochrome b (cob) genes have shown a significant variability of these sequences within Hydroides (Sun et al., 2012, indicating fast evolution of mitochondrial genes in this genus, yet our knowledge on the mitochondrial genomes of this group is very limited. ...
Article
Mitochondrial genomes are frequently applied in phylogenetic and evolutionary studies across metazoans, yet they are still poorly represented in many groups of invertebrates, including annelids. Here, we report ten mitochondrial genomes from the annelid genus Hydroides (Serpulidae) and compare them with all available annelid mitogenomes. We detected all 13 protein coding genes in Hydroides spp., including the atp8 which was reported as a missing gene in the Christmas Tree worm Spirobranchus giganteus, another annelid of the family Serpulidae. All available mitochondrial genomes of Hydroides show a highly positive GC skew combined with a highly negative AT skew – a feature consistent with that found only in the mitogenome of S. giganteus. In addition, amino acid sequences of the 13 protein-coding genes showed a high genetic distance between the Hydroides clade and S. giganteus, suggesting a fast rate of mitochondrial sequence evolution in Serpulidae. The gene order of protein-coding genes within Hydroides exhibited extensive rearrangements at species level, and were different from the arrangement patterns of other annelids, including S. giganteus. Phylogenetic analyses based on protein-coding genes recovered Hydroides as a monophyletic group sister to Spirobranchus with a long branch, and sister to the fan worm Sabellidae. Yet the Serpulidae+Sabellidae clade was unexpectedly grouped with Sipuncula, suggesting that mitochondrial genomes alone are insufficient to resolve the phylogenetic position of Serpulidae within Annelida due to its high base substitution rates. Overall, our study revealed a high variability in the gene order arrangement of mitochondrial genomes within Serpulidae, provided evidence to question the conserved pattern of the mitochondrial gene order in Annelida and called for caution when applying mitochondrial genes to infer their phylogenetic relationships.
... Hydroides elegans populations grow at high densities, changing the ecosystems they colonize (Bastida-Zavala, 2009) and competing for food and space with native species. Although H. elegans has originally been described from Australia (Haswell, 1883), and some authors speculate that it is native from there (Zibrowius, 1992;Ben-Eliahu and ten Hove, 2011;Sun et al., 2015), there is no clear evidence of its Australian origin because this species was also found in the Hawaiian coast at natural and artificial substrates (Long, 1974;Bailey-Brock, 1976;Sun et al., 2015), which suggests a broader Indo-Western Pacific origin. The true origin of its geographical distribution is unknown, so it is impossible to classify it as a native species of a territory. ...
... Hydroides elegans populations grow at high densities, changing the ecosystems they colonize (Bastida-Zavala, 2009) and competing for food and space with native species. Although H. elegans has originally been described from Australia (Haswell, 1883), and some authors speculate that it is native from there (Zibrowius, 1992;Ben-Eliahu and ten Hove, 2011;Sun et al., 2015), there is no clear evidence of its Australian origin because this species was also found in the Hawaiian coast at natural and artificial substrates (Long, 1974;Bailey-Brock, 1976;Sun et al., 2015), which suggests a broader Indo-Western Pacific origin. The true origin of its geographical distribution is unknown, so it is impossible to classify it as a native species of a territory. ...
Article
Full-text available
For decades, inputs of nutrients and organic matter into the Mar Menor coastal lagoon have favored the change from an original oligotrophic to a eutrophic state. The lagoon reached a stage of severe eutrophication and “environmental collapse” during the spring of 2016. This paper describes the massive growth of Serpulidae (Annelida, Polychaeta) forming large aggregations and reef structures after the environmental collapse caused by the eutrophic crisis. Four species belonging to the genera Hydroides and Serpula were identified; the identified species are Hydroides elegans, Hydroides dianthus, Serpula concharum, and Serpula vermicularis. The presence of bi-operculate and tri-operculate specimens is also documented in this study.
... For example, in the case of identifying nonnative species, failure to accurately identify the species may lead to costly consequences (Hutchings 2018). For many polychaetes only one or so species within a genus is invasive (Sun et al. 2015;lavesque et al. 2020 a, b) so it is critical to distinguish between native and non-native species in the genus. ...
Article
Full-text available
Researchers are continuing to identify polychaetes using inappropriate references and failing to appreciate that many if not most species have restricted distributions. using Marphysa sanguinea (Montagu, 1813) as a case example, we discuss the loss of valuable data by misidentifying a species. We suggest ways in which this problem can be addressed by both taxonomists, ecologists and other researchers. Furthermore, this situation is not unique to polychaetes but applies to many other groups of marine invertebrates.
... One of these aggregations was removed and the tubeworms identified in the laboratory as Hydroides ezoensis. The features required to identify this species are described by Sun et al. (2015). The walls were also colonised by another non-native species, the barnacle Austrominius modestus, which was found in all quadrat and wall scrape samples. ...
Article
The non-native bivalve Theora lubrica was recorded for the first time for the U.K. in April 2018, from Lake Lothing (Oulton Broad), Lowestoft, Suffolk. This represents the most northerly European record to date for the species. Several other non-native or cryptogenic species were recorded from the same location; in particular, the most northerly European locality of the tubeworm Hydroides ezoensis was confirmed. Theora lubrica has also been found in the vicinity of Southampton Water and the earliest known U.K. records (May 2011) of the non-native mussel Arcuatula senhousia, from the same area, are documented here.
... elegans (Haswell 1883) are commonly found on ship hulls and are subsequently introduced to novel ecosystems via boat traffic (Yan and Huang 1993). These species are of particular interest because both organisms adhere to and accumulate on economically significant artificial substrates (Sun et al. 2012(Sun et al. , 2015. Large, persistent populations of tubeworms can incur substantial financial costs as their calcareous tubes degrade artificial structures (Sun et al. 2012). ...
Article
Full-text available
Anthropogenic activities have accelerated the movement of non-indigenous species throughout the world. One approach to predict the spread of non-indigenous species is to employ bioclimatic envelope models which often assume niche conservation among sympatric, closely related species. Here, we test this assumption by comparing early developmental progress of two non-indigenous calcareous biofouling tubeworms Hydroides elegans and H. dirampha. In the subtropical Hong Kong monsoon climate, H. dirampha and H. elegans experience dramatic seasonal fluctuations in temperature (from 17 to 30 °C) and salinity (from 15 to 34 psu). Hydroides elegans was previously shown to be sensitive to lower salinity and warmer temperature while H. dirampha persisted in the field under these seasonal conditions. Thus, we hypothesize that the observed shift in abundance is due to the resilience of early stages of H. dirampha to the interactive stress of warming and lower salinity. Larval survival, growth, clearance, and settlement rate of H. dirampha were quantified in a 2 × 3 factorial experiment (24 and 28 °C; 20, 26, 32 psu). Stage-dependent tolerance was observed: cleavage was hindered by low salinity. However, larval growth and clearance did not follow this trend and instead peaked at 26 psu. Similarly, settlement rate did not decrease with freshening; rather, it peaked at 26 psu under warming. The salinity tolerance of H. dirampha is compared with that determined by Qiu and Qian (Mar Ecol Prog Ser 168: 127–134, 1998) for H. elegans. Differences in larval physiological tolerances could shape abundance and distribution of a single species as well as broader community structure.
... Lineages of each species were recovered as a single monophyletic group, with genetic distance of COI among lineages ranging from 9.8% to 30.9% in H exaltata, 15.9% to 16.9% in H. longispinosa, and 10% to 19.4% in the H. operculata-complex. Hydroides operculata, which has a long and convoluted taxonomic history and has been reported nearly world-wide (Ben-Eliahu, 1991; Ben-Eliahu and ten Hove, 1992;Çinar, 2006;Kubal et al., 2012;Sun et al., 2012Sun et al., , 2015, has been recently revised as a species complex consisting of at least three morphologically similar species: H. inornata, H. basispinosa and H. operculata, with different distribution ranges. (Fig. 7C; details see Sun et al., 2017). ...
... Hydroides trivesiculosa, which is morphologically similar to H. albiceps, was nested within the two main clades of the latter (Fig. 2, clade a, b), though with low nodal support. Although previous taxonomic studies of H. albiceps indicated variations in number, shape and relative size of the spines among specimens from different locations (Imajima, 1976;Fiege and Sun, 1999;Sun et al., 2015), the overlap of these morphological features made it difficult to distinguish specimens from different locations. Our phylogenetic results, however, indicate that there are at least four highly divergent lineages of H. albiceps morphotypes that potentially constitute separate species. ...
Article
Hydroides is a large and diverse group of calcareous tubeworms (Serpulidae, Annelida) recognised by a distinctive but variable two-tiered operculum. Despite considerable research using several species of Hydroides as models in ecological and biofouling studies, phylogenetic and biogeographic relationships within the genus are still poorly understood. Using combined mitochondrial (COI, cytochrome b) and nuclear (18S, 28S and ITS) gene markers for 284 individuals of 45 morphospecies of Hydroides, we investigated the global phylogenetic and biogeographic relationships within the genus. Phylogenetic topologies were well supported and indicated high genetic diversity within Hydroides, revealed potential cryptic species. Present results also include the first COI barcoding data enabling rapid and effective species identification of Hydroides on a global scale. Phylogenetic relationships within Hydroides were more concordant with geographical distributions than morphological similarity of their opercula. Molecular divergence estimates suggested the origin and subsequent diversification in the western Tethys Sea followed by a shift of the historical centre of diversity from the Indo-Mediterranean region to the central Indo-Pacific during the last 50 million years. Further studies on population genetics of species consisting of multiple lineages would provide a better understanding on the status of potential cryptic species. Furthermore, paleogeographic studies based on fossil Hydroides tubes would provide evidence to test this biogeographic hypothesis.
... Hydroides también tiene varios representantes invasores en puertos de todo el mundo (Link et al. 2009;Otani y Yamanishi, 2010). Sus tubos también forman densas agregaciones en Biología y Sociedad, febrero 2018 55 estructuras sumergidas como tuberías, pilotes, embarcaciones, boyas, cabos, cajas de siembra, etc., por lo que causan afectaciones en la acuicultura, en la navegación y en las plantas de generación de energía (Sun et al. 2015). La especie más estudiada es H. elegans, una especie de origen australiano (Sun et al. 2015), o criptogénica (de origen desconocido) para otros autores (Bastida-Zavala et al. 2017). ...
... Sus tubos también forman densas agregaciones en Biología y Sociedad, febrero 2018 55 estructuras sumergidas como tuberías, pilotes, embarcaciones, boyas, cabos, cajas de siembra, etc., por lo que causan afectaciones en la acuicultura, en la navegación y en las plantas de generación de energía (Sun et al. 2015). La especie más estudiada es H. elegans, una especie de origen australiano (Sun et al. 2015), o criptogénica (de origen desconocido) para otros autores (Bastida-Zavala et al. 2017). Es uno de los organismos incrustantes más problemáticos en las aguas tropicales y templadas del mundo (ten Hove y Weerdenburg, 1974). ...
Article
Full-text available
En esta contribución contribución se enfatiza la importancia de los gusanos poliquetos perforadores de moluscos de interés comercial en México y del estudio de las especies exóticas invasoras. Asimismo, se presenta el estado del conocimiento de ambos rubros en México. Es necesario sensibilizar a los tomadores de decisiones sobre la necesidad de apoyar proyectos de investigación y contrataciones en los dos temas. Recomendamos modificar la sección de poliquetos de la Lista de especies exóticas invasoras en México, publicada en el Diario Oficial de la Federación en 2016.