Wiklund H et al. (2019), Abyssal fauna of polymetallic nodule exploratio...

ECB-ART-49890

Zookeys 2019 Oct 28;883:1-82. doi: 10.3897/zookeys.883.36193.

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Abyssal fauna of polymetallic nodule exploration areas, eastern Clarion-Clipperton Zone, central Pacific Ocean: Annelida: Capitellidae, Opheliidae, Scalibregmatidae, and Travisiidae.

Wiklund H , Neal L , Glover AG , Drennan R , Muriel Rabone , Dahlgren TG .

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We present DNA taxonomy of abyssal polychaete worms from the eastern Clarion-Clipperton Zone (CCZ), central Pacific Ocean, using material collected as part of the Abyssal Baseline (ABYSSLINE) environmental survey cruises 'AB01' and 'AB02' to the UK Seabed Resources Ltd (UKSRL) polymetallic nodule exploration contract area 'UK-1', the Ocean Mineral Singapore exploration contract area 'OMS-1' and an Area of Particular Environmental Interest, 'APEI-6'. This is the fourth paper in a series to provide regional taxonomic data with previous papers reporting on Cnidaria, Echinodermata and Mollusca. Taxonomic data are presented for 23 species from 85 records within four polychaete families: Capitellidae, Opheliidae, Scalibregmatidae and Travisiidae, identified by a combination of morphological and genetic data, including molecular phylogenetic analyses. Two taxa (genetically separated from one another) morphologically matched the same known cosmopolitan species, Ophelina abranchiata that has a type locality in a different ocean basin and depth from where no genetic data was available. These two species were assigned the open nomenclature 'cf.' as a precautionary approach in taxon assignments to avoid over-estimating species ranges. Twelve (12) taxa are here described as new species, Ammotrypanella keenani sp. nov., Ammotrypanella kersteni sp. nov., Ophelina curli sp. nov., Ophelina ganae sp. nov., Ophelina juhazi sp. nov., Ophelina martinezarbizui sp. nov., Ophelina meyerae sp. nov., Ophelina nunnallyi sp. nov., Oligobregma brasierae sp. nov., Oligobregma tani sp. nov., Oligobregma whaleyi sp. nov. and Travisia zieglerae sp. nov. For the remaining nine taxa, we have determined them to be potentially new species, for which we make the raw data, imagery and vouchers available for future taxonomic study. The CCZ is a region undergoing intense exploration for potential deep-sea mineral extraction from polymetallic nodules. We present these data to facilitate future taxonomic and environmental impact study by making both data and voucher materials available through curated and accessible biological collections.

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	Figure 1. Map over sampling sites. A UK-1 Stratum-A B UK-1 Stratum-B study areas, both within the UK Seabed Resources UK-1 exploration contract area C OMS Stratum-A study area, in the Ocean Mineral Singapore (OMS) polymetallic nodule exploration contract area D Area of Particular Interest APEI-6. Inset map showing location of Clarion-Clipperton Fracture Zone. Bathymetric survey and sample localities from the AB01 2013 RV Melville survey cruise and AB02 2015 Thomas G. Thompson survey cruise, data courtesy Craig R. Smith (University of Hawaii), UK Seabed Resources Ltd and Seafloor Investigations, LLC.
	Figure 2. Capitellidae sp. NHM_1486 (specimen NHM_1486). A Lab image, whole specimen B Live image, anterior C Lab image, anterior, highlighting first three chaetigers (faded stain, pr = prostomium, pb = proboscis) D Lab image, thoracic capillary chaetae E Lab image, transitionary chaetigers between thorax and abdomen (faded stain) (t = thorax, h = hooks, a = abdomen) F Lab image, hooks G Lab image, whole specimen (methyl green stain). Scale bars: 1 mm (A, C); 25 μm (D, F).
	Figure 3. Notomastus sp. NHM_162. A Lab image, whole specimen (specimen NHM_162) B Live image, whole specimen (specimen NHM_162) C Lab image, biannulated chaetigers (specimen NHM_162) D Lab image, prostomium (specimen NHM_162, pr = prostomium) E Lab image, whole specimen, dorsal (specimen NHM_915B, no = notopodial lobe, nu = neuropodial lobe) F Lab image, whole specimen, ventral (specimen NHM_915B, nu = neuropodial lobe) G Lab image, thoracic chaetae, (specimen NHM_162) H Lab image, thoracic hooks (specimen NHM_162) I Lab image, whole specimen (specimen NHM_162, methyl green stain). Scale bars: 1 mm (A, E, F, I); 50 μm (G, H).
	Figure 4. Phylogenetic analysis of Capitellidae. 50% majority rule tree from the Bayesian analyses using 18S and 16S, with posterior probability values on nodes. Twenty-four taxa from GenBank were included, and Echiura was chosen as outgroup following the annelid phylogeny of Weigert and Bleidorn (2016).
	Figure 5. Ammotrypanella arctica holotype (BMNH 1921.12392). A Whole specimen (fragmented) B Anterior-most fragment C Posterior-most fragment. Scale bars: 1 mm (A, B, C).
	Figure 6. Ammotrypanella keenani sp. nov. A Lab images, whole specimens (holotype [specimen NHM_1250], post-staining, faded stain [left], paratype NHM_1871, unstained [right]) B Live images, whole specimens (holotype [left], paratype NHM_1871 [right]) C Lab image, prostomium (paratype NHM_1871, damaged, no = everted nuchal organ) D Lab image, prostomium (holotype, p = palpode) E Lab image, detail of palpode (holotype) F Lab image, posterior branchiae (paratype NHM_1871, br = branchiae) G Lab image, posterior branchiae (holotype, br = branchiae) H Live image, anterior (paratype NHM_1871, pp = parapodia, no = nuchal organ) I Lab image, detail of capillary chaetae (holotype) J Lab image, detail of anal funnel, ventro-distal (left) and latero-distal (right) views (holotype, stained, cp = cushioned pad, vc = ventral cirrus). Morphological features in plates C–H, J have been outlined with a fine white line to improve clarity of those features. Scale bars: 1 mm (A, F,G); 100 μm (E, I).
	Figure 7. Ammotrypanella kersteni sp. nov. holotype (specimen NHM_254) live images. A Live image, whole specimen B Live image, detail of anterior (upper) and posterior (lower), with branchiae outlined in a fine black line (e = eggs, br = branchiae).
	Figure 8. Ammotrypanella kersteni sp. nov. holotype (specimen NHM_254). A Lab images, whole specimens, dorsal (upper) and ventral (lower) B Lab image, anterior C Lab image, detail of palpode D Lab image, mid-body parapodia (p = parapodium) E Lab image, detail of parapodium F Lab image, posterior and anal funnel. Morphological features in plates B, D, F have been outlined with a fine white or black line to improve clarity of those features. Scale bars: 2 mm (A); 1 mm (B, D, F); 100 μm (C, E).
	Figure 9. Ammotrypanella sp. NHM_1653 (specimen NHM_1653). A Lab image, whole specimen (faded stain B Live image, whole specimen C Lab image, anterior (faded stain) D Lab image, detail of palpode E Lab image, posterior (faded stain, br = branchiae) F Lab image, anterior-midbody (faded stain, pp = parapodia) G Lab image, detail of capillary chaetae H Lab image, detail of posterior and anal tube. Morphological features in plates C, E, F, G have been outlined with a very fine white to improve clarity of those features. Scale bars: 1 mm (A); 100 μm (D, G, H).
	Figure 10. Ammotrypanella sp. NHM_2114 (specimen NHM_2114). A Live images, whole specimen with detail of anterior B Lab images, fragmented whole specimen, post-staining, (faded stain, anterior fragment [left], posterior fragment [right],) C Lab image, anterior fragment, post-staining, faded stain D Lab image, detail of anterior and palpode E Lab image, detail of capillary chaetae. Scale bars: 1 mm (B); 250 μm (C) 50 μm (D); 25 μm (E).
	Figure 11. Ophelina curli sp. nov. holotype (specimen NHM_2112). A Lab image, whole specimen (post-staining, very faded stain) B Live image, with detail of anterior C Lab image, anterior and proboscis, lateral view, stained D Lab image, detail of palpode E Lab image, anterior and proboscis, ventral view (stained, pb = proboscis) F Lab image, mid-body parapodium (stained, pp = parapodia) G Lab image, detail of capillary chaeta H Lab image, anal funnel. Scale bars: 1 mm (A, E); 100 μm (D, G)
	Figure 12. Ophelina ganae sp. nov. A Lab image, whole specimen (holotype [specimen NHM_1137] B Live images, whole specimens (holotype [bottom], paratype NHM_598 [middle], paratype NHM_1309 [top]) C Lab image, detail of palpode (holotype) D SEM image, second chaetiger with lateral organ (paratype NHM_1309, lo = lateral organ) E SEM images, lateral organs, (a) pre-chaetigerous segment, (b) chaetiger 1, (c) chaetiger 2, (d) chaetiger 17 (last chaetiger) (paratype NHM_1309) F Lab image, detail of capillary chaetae (holotype) G Live image, with detail of potential anal funnel (specimen NHM_473, vc = potential ventral cirrus). H Lab image, detail of posterior (holotype) I Lab image, whole specimen (paratype NHM_598, stained). Morphological features in plates G, H have been outlined with a fine white or black line to improve clarity of those features. Scale bars: 1 mm (A); 100 μm (C); 20 μm (D); 5 μm (E); 50 μm (F); 100 μm (H); 1 mm (I).
	Figure 13. Ophelina juhazi sp. nov. holotype (specimen NHM_1073). A Live images, whole specimen (center) with detail of anterior (left) and posterior (right) B Lab image, whole specimen C Lab image, anterior and prostomium D Lab image, detail of palpode E Lab image, anterior and parapodia (stained, pp = parapodia) F Lab image, detail of capillary chaetae G Lab image, posterior and anal tube (stained). Morphological features in plates C, E, G have been outlined with a very fine white line to improve clarity of those features. Scale bars: 1 mm (B); 100 μm (D); 50 μm (F); 1 mm (E).
	Figure 14. Ophelina martinezarbizui sp. nov. A Live images, whole specimen (center) with detail of anterior (left) and anal funnel (right) (holotype [specimen NHM_681], e = eggs, vc = enlarged ventral cirrus, c = cirri) B Lab image, whole specimen (holotype). C Lab image, detail of palpode (paratype NHM_2116) D Live image, prostomium, “royal crown” palpode (specimen NHM_2092) E Lab image, ventral posterior and branchiae (paratype NHM_1766, br = branchiae) F Lab image, lateral posterior and branchiae (paratype NHM_1766, br = branchiae) G Lab image, detail of anal funnel (paratype NHM_1766, vc = enlarged ventral cirri [folded over], c = cirri, br = branchiae H Lab image, detail of capillary chaetae (holotype). Morphological features in plates A, E, G have been outlined with a fine white or black line to improve clarity of those features. Scale bars: 1 mm (B, E, F); 100 μm (C, H).
	Figure 15. Ophelina martinezarbizui sp. nov. (specimen NHM_2088). A Live image, whole specimen B SEM image, dorsal anterior C SEM image, lateral anterior D SEM image, dorsal posterior, br = branchiae) E SEM image, detail of mid-body lateral organ (lo = lateral organ). Scale bars: 300 μm (B, C, D); 40 μm (E).
	Figure 16. Ophelina meyerae sp. nov. holotype (specimen NHM_1241). A Live images, whole specimen with detail of anterior (p = palpode) B Lab image, whole specimen, lateral view (faded stain) C Lab image, anterior and branchiae (faded stain) (br = branchiae) D Lab image, mid-body branchiae (faded stain, br = branchiae) E Lab image, anterior parapodia (faded stain, pp = parapodia) F Lab image, detail of capillary chaetae (br = branchiae) G Lab image, detail of capillary chaetae (br = branchiae). Morphological features in plates C–F have been outlined with a fine white line to improve clarity of those features. Scale bars: 1 mm (B–E); 100 μm (F, G).
	Figure 17. Ophelina nunnallyi sp. nov. A Lab image, whole specimen (paratype NHM_700) B Live images, whole specimens (holotype [specimen NHM_683] [left], paratype NHM_700 [right]) C SEM image, anterior and palpode (specimen NHM_1309A, m = mouth) D Lab image, detail of palpode, (paratype NHM_1273) E Lab image, anterior and branchiae (paratype NHM_1273, br = branchiae) F Lab image, parapodia (holotype, pp = parapodia) G Lab image, capillary chaeta (holotype) H Live image, anal funnel (holotype, c = cirri) I Lab image, detail of anal funnel (holotype, c = cirri). Morphological features in plates E, F, H, I have been outlined with a fine white line to improve clarity of those features. Scale bars: 1 mm (A, E, F); 200 μm (C, D); 100 μm (G, I).
	Figure 18. Ophelina cf. abranchiata sp. NHM_1769 (specimen NHM_1769). A Live image, whole specimen B Lab image, whole specimen (very faded post-stain,) C Lab image, detail of prostomium and palpode D Lab image, detail of chaetae E Lab image, whole specimen (stained). Scale bars: 1 mm (B, E); 50 μm (C, D).
	Figure 19. Ophelina cf. abranchiata sp. NHM_2017 (specimen NHM_2017). A Live image, whole specimen B Lab image, whole specimen (stained,) C Lab image, whole specimen (faded stain) D Lab image, anterior (stained, arrow highlighting dark banding) E Lab image, detail of anterior F Lab image, detail of capillary chaetae. Scale bars: 1 mm (B, C); 100 μm (E, F).
	Figure 20. Ophelina sp. NHM_689 (specimen NHM_689). A Lab image, whole specimen, lateral view (pre-stain) B Lab image, whole specimen, ventral view (faded stain, vg = ventral groove) C Live image, whole specimen D Lab image, damaged anterior (stained, br = branchiae, pr = prostomium) E Lab image, detail of anterior branchiae (br = branchiae) F Lab image, posterior branchiae (stained, br = branchiae) G Lab image, mid-body parapodia (stained, pp = parapodia) H Lab image, detail of capillary chaetae. Morphological features in plates B, D–G have been outlined with a fine white line to improve clarity of those features. Scale bars: 1 mm (A, B); 0.5 mm (D); 50 μm (E); 0.25 (F, H).
	Figure 21. Ophelina sp. NHM_1068. A Lab image, whole specimen (specimen NHM_1068,) B Lab image, whole specimen (specimen NHM_1874, faded stain) C Live image, whole specimen (specimen NHM_1874) D Live image, anterior, with branchiae outlined in a fine black line (specimen NHM_1068, br = branchiae) E Lab image, detail of palpode (specimen NHM_1068) F Lab image, detail of anterior parapodia and branchiae (specimen NHM_1874, stained) G Lab image, posterior (specimen NHM_1068) H Lab image, detail of anal funnel (specimen NHM_1068). Scale bars: 1 mm (A, B); 100 μm (E).
	Figure 22. Ophelina sp. NHM_1331 (specimen NHM_1331). A Lab image, whole specimen, dorsal view (faded stain) B Lab image, whole specimen, lateral view (faded stain) C Live image, whole specimen D Lab image, anterior (faded stain, br = branchiae, no = nuchal organ) E Lab image, detail of anterior (br = branchiae, no = nuchal organ) F Lab image, posterior, anal funnel (br = branchiae) G Lab image, detail of posterior and anal funnel (br = branchiae). Morphological features in plates B, D–G have been outlined with a fine white line to improve clarity of those features. Scale bars: 1 mm(A, B); 100 μm (E, G).
	Figure 23. Phylogenetic analysis of Opheliidae. 50% majority rule tree from the Bayesian analyses using 18S and 16S, with posterior probability values on nodes. Twenty-nine taxa from GenBank were included, and Capitellidae and Echiura was chosen as outgroup following the annelid phylogeny of Weigert and Bleidorn (2016).
	Figure 24. Oligobregma brasierae sp. nov. A Lab image, whole specimen (holotype [specimen NHM_823], pre-staining) B Lab image, dorsal anterior (holotype, faded stain, h = prostomial horns, pe = peristomial ring) C Lab image, dorsal segments, quadriannulate chaetigers (holotype, faded stain) D Lab image, ventral segments (holotype, faded stain, vm = ventral midline) E Lab image, ventral anterior, (holotype, pre-staining) F Lab image, mid-body parapodia (holotype, shirlastained, dc = dorsal cirrus, vc = ventral cirrus, ip = interramal papilla) G Lab image, detail of dorsal cirrus (paratype NHM_684, ig = internal gland) H Lab image, detail of hirstute notopodial spines on chaetiger 1 (paratype NHM_684, ig = internal gland) I Lab image, detail of capillary and lyrate chaetae (paratype NHM_684) J Lab images, posterior (paratype NHM_684, [bottom-left panel], ac = anal cirri, ig = internal gland). Morphological features in plates B–D, F, G, H have been outlined with a fine white or black line to improve clarity of those features. Scale bars: 1 mm (A, E); 50 μm (G); 25 μm (H, I); 100 μm (J).
	Figure 25. Oligobregma brasierae sp. nov. A Live image, whole specimen, ventral view (holotype [specimen NHM_823]) B Live image, whole specimen, dorsal view (paratype NHM_684) C Live image, dorsal anterior, with prostomial features outlined in a fine white line(paratype NHM_684, h = prostomial horns, pe = peristomial ring).
	Figure 26. Oligobregma tani sp. nov. holotype (specimen NHM_1454). A Lab image, whole specimen (pre-stain,) B Live image, whole specimen C Lab image, dorsal anterior, (stained, h = prostomial horns, pe = peristomial ring) D Lab image, lateral anterior, (stained, h = prostomial horns, pe = peristomial ring) E Lab image, ventral anterior, (stained, vm = ventral midline) F Lab image, mid-body parapodia (faded stain, dc = dorsal cirrus, vc = ventral cirrus, ip = interramal papilla) G Lab image, detail of dorsal cirrus, with no internal gland visible (dc = dorsal cirrus) H Lab image, detail of hirsute spines on notopodia on chaetiger 1 ([left panel]) I Lab image, detail of capillary and lyrate chaetae on chaetiger 12 J Lab image, detail lyrate chaetae. Morphological features in plates F, G have been outlined with a fine white line to improve clarity of those features. Scale bars: 0.5 mm (A, C, E); 50 μm (G–I).
	Figure 27. Oligobregma whaleyi sp. nov. holotype (specimen NHM_822). A Lab (upper) and live (lower) images, whole specimens [lab image] B Lab image, mid-body segments and annulation C Lab image, ventral midbody (vm = ventral midline) D Live (left) and lab (right) images of prostomium (pe = peristomial ring, h = prostomial horns) E Lab images, midbody and posterior parapodia, chaetigers 15 (left) and 24 (right) (dc = dorsal cirrus, vc = ventral cirrus, ip = interramal papilla) F Lab image, detail of dorsal cirrus and internal gland (dc = dorsal cirrus, ig = internal gland) G Lab image, detail of notopodial spines on chaetiger 1 H Lab image, detail of capillary chaetae (lc = lyrate chaetae) I Lab image, detail of lyrate chaetae. Morphological features in plates B–D, F, G, H have been outlined with a fine white or black line to improve clarity of those features. Scale bars: 1 mm (A); 100 μm (F–H); 50 μm (I).
	Figure 28. Scalibregmatidae sp. NHM_2308 (specimen NHM_2308). A Lab image, whole specimen, dorsal view (pre-stain) B Lab image, dorsal anterior (stained) C Lab image, ventral anterior, (shirla stained, vm = ventral midline) D Lab image, detail of hirsute spines on chaetigers 1 and 2 E Lab image, detail of hirsute spine tips F Lab image, detail of lyrate chaeta. Scale bars: 1 mm (A–C); 100 μm (D); 50 μm (E); 10 μm (F).
	Figure 29. Travisia zieglerae sp. nov. A Lab image, whole specimen, pre-stain (holotype [specimen NHM_1431]) B Live images, whole specimens (specimen NHM_1911 [left], specimen NHM_188 [right]) C Lab image, lateral anterior, (holotype, stained, pr = prostomium, pp = parapodial lappets) D Lab image, distal anterior, (holotype, stained, m = mouth) E Lab image, lateral posterior, (holotype, stained, pp = parapodia, io = interramal organs) F Detail of capillary chaeta (paratype NHM_140) G Lab image, pygidium, distal view (left) and lateral view (right), with pygidial features outlined in a fine white line (holotype, stained, vl = ventral lobe). Scale bar: 1 mm (A).
	Figure 30. Travisia sp. NHM_1244. A Lab images, whole specimens (specimen NHM_1244, unstained [top], specimen NHM_1863, faded stain [bottom]) B Live image, whole specimen (specimen NHM_1863) C Lab image, dorsal anterior (specimen NHM_1863, stained, pr = prostomium) D Lab image, ventral anterior, (stained) (specimen NHM_1863, m = mouth) E Lab image, lateral posterior (specimen NHM_1863, stained, pp = parapodia, io = interramal organs) F Lab image, detail of capillary chaetae (specimen NHM_1244) G Lab image, pygidium, distal view (lower left) and lateral view (upper right), with pygidial features outlined in a fine white line (specimen NHM_1863, stained, vl = ventral lobe). Scale bars: 1 mm (A); 50 μm (F).
	Figure 31. Comparison between Travisia sp. NHM_1244 and Travisia zieglerae sp. nov., and holotypes of Travisia glandulosa (BMNH 1921.5.1.2431) and Travisia gravieri (BMNH 1921.5.1.2429). A Lab images, whole specimens (Travisia sp. NHM_1244 specimen NHM_1863 [left], Travisia zieglerae sp. nov. holotype [specimen NHM_1431] [right], stained,) B Lab images, comparison of prostomia (Travisia sp. NHM_1244 specimen NHM_1863 [left], Travisia zieglerae sp. nov. holotype [right], stained) C Lab images, comparison of pygidia (Travisia sp. NHM_1244, specimen NHM_1863 [left], Travisia zieglerae sp. nov. holotype [right], stained) DTravisia glandulosa holotype, with detail of pygidium ETravisia gravieri holotype, with detail of pygidium. Morphological features in plates C–E have been outlined with a very fine white line to improve clarity of those features. Scale bars: 1 mm (A, D, E).
	Figure 32. Phylogenetic analysis of Scalibregmatidae and Travisiidae. 50% majority rule tree from the Bayesian analyses using 18S and 16S, with posterior probability values on nodes. Twenty taxa from GenBank were included, and Opheliidae was chosen as outgroup following the annelid phylogeny of Weigert and Bleidorn (2016).

References [+] :

Amon, Megafauna of the UKSRL exploration contract area and eastern Clarion-Clipperton Zone in the Pacific Ocean: Annelida, Arthropoda, Bryozoa, Chordata, Ctenophora, Mollusca. 2017, Pubmed