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First described in 2004 off California, Osedax worms are now known from many of the world's oceans, ranging from 10 to over 4000 m in depth. Currently, little is known about species ranges, since most descriptions are from single localities. In this study, we used new sampling in the north-eastern Pacific and available GenBank data from off Japan and Brazil to report expanded ranges for five species: Osedax frankpressi, O. knutei, O. packardorum, O. roseus and O. talkovici. We also provided additional DNA sequences from previously reported localities for two species: Osedax priapus and O. randyi. To assess the distribution of each species, we used cytochrome c oxidase subunit I (COI) sequences to generate haplotype networks and assess connectivity amongst localities where sampling permitted. Osedax frankpressi, O. packardorum, O. priapus, O. roseus and O. talkovici all had one or more dominant COI haplotypes shared by individuals at multiple localities, suggesting high connectivity throughout some or all of their ranges. Low ΦST values amongst populations for O. packardorum, O. roseus and O. talkovici confirmed high levels of gene flow throughout their known ranges. High ΦST values for O. frankpressi between the eastern Pacific and the Brazilian Atlantic showed little gene flow, reflected by the haplotype network, which had distinct Pacific and Atlantic haplotype clusters. This study greatly expands the ranges and provides insights into the phylogeography for these nine species.

Osedax ventana n. sp. Figure 4D Osedax ‘nude-palp-B’ (Jones et al. 2008; Rouse et al. 2011; Rouse et al. 2009; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A1643 palps only (GenBank COI sequence EU235218), fixed in formalin preserved in ethanol, collected from cow bones deployed at 2898 meters depth in Monterey Submarine Canyon, California (36°36.606’N; 122°26.122’W) ROV Tiburon dive number 1069, Oct. 1, 2007. Diagnosis and description. Holotype represented by fragments of four palps (Fig. 4D), still within a gelatinous tube. Oviduct not discerned. In life, palps reddish with two blood vessels in each, lacking obvious pinnules (Fig. 4D). Distribution. Known from Monterey Bay, California from 2898 meters depth (Table 2). It has only been found in experimentally deployed cow bones. Etymology. This species is named (noun in apposition) for the ROV Ventana, which was used to collect many Osedax specimens. Remarks. Osedax ventana n. sp. is known from only a single partial specimen, sequenced by Jones et al. (2008). It was found on a deployed cow bone and was not discovered during any subsequent sampling at the same site. Osedax ventana n. sp. is part of Osedax Clade II and closest relative (with low support) to Osedax ryderi n.sp., from which it shows an uncorrected divergence of 12% (Table 4). Given the holotype is a fragment, there is little, except for its unique COI sequence, to distinguish Osedax ventana n. sp. from other Clade II species. Three other Osedax species: O. crouchi, O. nordenskjoeldi and O. rogersi, were similarly erected based on fragmented specimens and DNA data (Amon et al. 2014).

Osedax sigridae n. sp. Figure 2 ���green palp��� (Katz et al. 2010; Katz et al. 2011; Katz & Rouse 2013; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A7809 female (GenBank COI sequence FJ347641), collected from an experimentally deployed whale carcass (Eschrichtius robustus) deployed at 1820 meters depth in Monterey Submarine Canyon, California (36��42.496���N; 122��6.316���W) ROV Tiburon dive number 1163, Dec. 20, 2007. Fixed in formalin preserved in ethanol. Paratypes: All females. Fixed in formalin preserved in ethanol; SIO-BIC A7811 same locality and date as holotype (GenBank COI sequence FJ347642); SIO-BIC A1650 (GenBank COI sequence FJ347639) and SIO-BIC A7810 (GenBank COI sequence FJ347640), collected on cow bones deployed at 1820 meters depth in Monterey Submarine Canyon, California (36��42.496���N; 122��6.316���W) ROV Tiburon dive number 1119, Aug. 16, 2007. Diagnosis and description. Holotype female (Fig. 2D); trunk 1 mm long, 0.5 mm wide; crown of pinnulated palps contracted, 2 mm long. Tube gelatinous, up to 10 mm long, 5 mm across. Oviduct shorter than palps (Fig. 2C). In life, palps green-yellow (Figs 2A, C, D). Pinnules of all four palps oriented dorsally (Fig. 2D). No obvious pigmentation on trunk, or clear demarcation into upper and lower trunk. Ovisac spheroidal with roots as simple lobes (Figs 2B, D). Males not found. Distribution. Known from Monterey Bay, California from 1820 meters depth (Table 2). It has been found in whale and cow bones. Etymology. This species is named (noun in the genitive case) for Sigrid Katz, whose Ph.D. project involved detailed study of the anatomy of this species (Katz et al. 2010; Katz et al. 2011). Remarks. Osedax sigridae n. sp. is part of Osedax clade I and closest relative to the undescribed Japanese taxon Osedax ���Sagami-5��� (Fig. 1). All four available COI sequences for Osedax sigridae n. sp. (Table 3) show less than 1% uncorrected sequence divergence, while the minimum uncorrected distance to Osedax ���Sagami-5��� is 11% (Table 4). Careful searching for males in some of the tubes of the Osedax sigridae n. sp. females shown in Figure 2A failed, though Katz & Rouse (2013) did find sperm in the ovisac of females. The most distinguishing features of the Osedax sigridae n. sp. specimens collected to date are the distinctive green/yellow palps and the lobate roots (Fig. 2)., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 459, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Katz, S., Klepal, W. & Bright, M. (2010) The skin of Osedax (Siboglinidae, Annelida): An ultrastructural investigation of its epidermis. Journal of Morphology, 271, 1272 - 1280. https: // doi. org / 10.1002 / jmor. 10873","Katz, S., Klepal, W. & Bright, M. (2011) The Osedax trophosome: organization and ultrastructure. Biological Bulletin, 220, 128 - 139. https: // doi. org / 10.1086 / BBLv 220 n 2 p 128","Katz, S. & Rouse, G. W. (2013) The reproductive system of Osedax (Annelida, Siboglinidae): sperm ultrastructure, ovary structure and fertilization. Invertebrate Biology, 132, 368 - 385. https: // doi. org / 10.1111 / ivb. 12037","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z"]}

Osedax westernflyer n. sp. Figure 5 ���nude-palp-D��� (Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009) ���Sagami 8��� (GenBank COI sequence FM998110; Pradillon et al. unpublished) Material examined. Holotype: SIO-BIC A 1645, Female (GenBank COI sequence MG262303), fixed in formalin preserved in ethanol, collected from a whale carcass (Eschrichtius robustus) deployed at 1820 meters depth in Monterey Submarine Canyon, California (36��42.496���N; 122��6.316���W), ROV Doc Ricketts dive number 12, March 13, 2009. Paratypes: All females, fixed in formalin preserved in ethanol, SIO-BIC A7802, A7803 (GenBank COI sequence MG262302, MG262304), same locality and date as holotype. Diagnosis and description. Holotype female (Fig. 5C); in life trunk 2.8 mm long, 0.5 mm wide; crown of four palps, 3.5 mm long. Tube gelatinous, thin, fitting about trunk and base of crown, 3 mm long (Fig. 5C). Oviduct among palps 2.3 mm long (Figs 5 A-C). In life, palps with white patches with no visible blood vessels, without obvious pinnules (Figs 5 A-C). Trunk with demarcation into upper and lower trunk (Fig. 5C), white pigment patch mid-ventrally on anterior tip of trunk (Figs 5B, C). Ovisac 1.5 mm by 1 mm with lobate greenish roots extending outwards (Figs 5B, C). Males not found. Distribution. Known from Monterey Bay, California from 1820 meters depth and Sagami, Bay Japan (Table 2). It has been found in cow and whale bones. Etymology. This species is named (noun in apposition) for the Research Vessel Western Flyer, which has been critical to Osedax studies. Remarks. Osedax westernflyer n. sp. is part of Osedax Clade II and is closest relative to O. knutei n. sp., with a minimum divergence of 16 % for COI, though it shows a smaller distance, 14% to O. docricketts n. sp. (Table 4). The four available COI sequences for Osedax westernflyer n. sp. from California (Table 3) are less that 1% divergent. The holotype and two paratypes all showed distinct white pigment on the palps, though these were all collected from one patch of bone and the color may have been caused by a bacterial coating. A single COI sequence on GenBank (FM998110), referred to Osedax sp. Sagami-8 (Pradillon et al., unpublished), is less than 1% divergent from the four California COI sequences (Table 4), leading us to propose that Osedax westernflyer n. sp. is also found in Japan. There is little to distinguish Osedax docricketts n. sp. from other Clade II species, apart from the white pigmentation at the anterior end of the trunk and on the pigmentation on palps., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 465, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Rouse, G. W., Goffredi, S. K., Johnson, S. B. & Vrijenhoek, R. C. (2011) Not whale-fall specialists, Osedax worms also consume fishbones. Biology Letters, 7, 736 - 739. https: // doi. org / 10.1098 / rsbl. 2011.0202","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z"]}

Osedax talkovici n. sp. Figure 3 ‘yellow patch’ (Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A 7821 female (GenBank COI sequence MG262313), fixed in glutaraldehyde-preserved in ethanol, collected from turkey (Meleagris gallopavo) bone deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W) ROV Doc Ricketts dive number 233, June 2, 2011. Paratypes: Females, all fixed in formalin-preserved in ethanol, SIO-BIC A1649 (GenBank COI sequence FJ347621), SIO-BIC A7822, A7823, A7824, A7825, A7826 (GenBank COI sequences FJ347620, 431196, 431197, 431201, 431205), whale (Eschrichtius robustus) deployed at 633 meters depth in Monterey Submarine Canyon, California (36°48.178’N; 121°59.677’W) ROV Tiburon dive number 1160, Dec. 18, 2007; Female SIO-BIC A7829 (GenBank COI sequence MG262311) and dwarf males (allotypes) SIO-BIC A7830 whale fall at 633 meters depth in Monterey Submarine Canyon, California (36°48.178’N; 121°59.677’W) ROV Doc Ricketts dive number 205, October 26, 2010; Female SIO-BIC A7827 (GenBank COI sequence MG262310) and dwarf males (allotypes) SIO-BIC A7828, whale (Eschrichtius robustus) deployed at 633 meters depth in Monterey Submarine Canyon, California (36°48.178’N; 121°59.677’W) ROV Doc Ricketts dive number 233, June 2, 2011; Female SIO-BIC A7831 (GenBank COI sequence MG262312) on green turtle bone (Chelonia mydas), deployed at same locality and date as holotype. Diagnosis and description. Holotype female (Figs 3B, C); in life trunk 1.8 mm long, 0.3 mm wide; crown of palps contracted, 1.5 mm long. Tube gelatinous. Oviduct 0.75 mm (Figs 3C, D). In life, palps reddish with blood (Figs 3A–E). Pinnules of all four palps oriented dorsally (Figs 3A, D, E). Trunk with bright yellow patch on midventral anterior edge (Figs 3A, B, D). Clear demarcation into upper and lower trunk (Fig. 3D). Ovisac spheroidal in holotype (Figs 3B, C), extending laterally, or as simple lobes in other specimens. Roots of holotype longer that the remaining body, extending posteriorly as a single lobate mass (Figs 3B, C). In other specimens (Figs 3A, D, E), roots as simple lobes or extending laterally in two lobes. Males dwarfs, resembling trochophore larvae, with spermatids and sperm (Figs 3F, G). Distribution. Known from Monterey Bay, California from 633 to 1018 meters depth (Table 2). It has been found in whale, elephant seal, cow, turkey, turtle and teleost bones. Etymology. This species is named (noun in the genitive case) for Mark Talkovic, Senior ROV pilot for MBARI, who collected many bones containing Osedax over the years. Remarks. Osedax talkovici n. sp. is part of Osedax Clade I and forms a well-supported clade with the undescribed species O. ‘mediterranea’ and a clade comprised of Osedax sigridae n. sp. and the undescribed species O. Sagami-5 (Fig. 1). It has a minimum uncorrected distance, for COI, of at least 16% from each of these (Table 4). The 15 available COI sequences for Osedax talkovici n. sp. (Table 3) show less that 1% sequence divergence. The most distinguishing feature of Osedax talkovici n. sp. is the bright yellow patch ventrally on the anterior trunk (Fig. 3).

Osedax packardorum n. sp. Figure 16 ‘orange collar’ (Braby et al. 2007; Rouse et al. 2009; Rouse et al. 2015; Tresguerres et al. 2013; Vrijenhoek et al. 2009) ‘sp.4 SBJ-2006’ (GenBank COI sequence DQ996639, DQ996641-2) Material examined. Holotype: Female, fixed in formalin preserved in ethanol, SIO-BIC A 1641 (GenBank COI sequence EU223341), collected from whale carcass (Balaenoptera musculus) deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W), ROV Tiburon dive number 1049, Oct. 25, 2006. Paratypes: Female and dwarf males (allotypes), fixed in formalin preserved in ethanol SIO-BIC A7840, A7842 (female GenBank COI sequences EU223339, EU223343), same locality and date as holotype; Female, fixed in formalin preserved in ethanol, SIO-BIC A7841(GenBank COI sequence EU223340), same locality and date as holotype; Female, sectioned for histology, same locality and date as holotype, SIO-BIC A7843 (GenBank COI sequence EU223342); Female, fixed in formalin preserved in ethanol, SIO-BIC A7844 (GenBank COI sequence FJ431200), collected from whale fall (Eschrichtius robustus) at 663 meters depth in Monterey Submarine Canyon, California (36°48.178’N; 121°59.677’W) ROV Tiburon dive number 1160, Dec. 18, 2007. Diagnosis and description. Holotype female, in life trunk 11 mm long, 0.6 mm wide; crown of palps contracted, curled, 2 mm long (image not shown as the trunk is damaged). Palps up to 12 mm long when extended in other specimens (Figs 16A, B). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (Figs 16B, D, E). Oviduct extends from trunk into crown for 3 mm (Fig. 16B). In life, palps reddish with two major blood vessels in each (Figs 16A, B). Tube a gelatinous loose mass around trunk (Figs 16A, B). Trunk with pale to dark orange ring around anterior margin; ring broken mid-dorsally by oviduct and midventrally by and oval unpigmented patch (Figs 16A, B, E). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (Fig. 16G). Ovisac in holotype an ellipsoidal mass (not shown); extending laterally as lobes in other specimens (Fig. 16C). Roots of holotype extend from ovisac on one side as one mass; in other specimens, roots as long branching masses. Males dwarfs, with spermatids and sperm (Figs 16F, G), chaetal-bearing segments not inflated (Fig. 16H); found in tube lumen of females. Distribution. Known from Monterey Bay, California at 349, 633, and 1018 meters depths (Table 2). Found in whale and cow bones. Etymology. This species is named (noun in the genitive case) in honor of the Packard family whose foundation supports MBARI and enabled the discovery of all of the Osedax species in California. Remarks. Osedax packardorum n. sp. is part of Osedax Clade IV and closest relative to Osedax lehmani n. sp., (Fig. 1), from which it has a minimum uncorrected distance for COI of 8-9% (Table 4). As pointed out above, this is one of the smaller interspecific distances among Osedax species and the two species are morphologically very similar. All 27 available COI sequences for Osedax packardorum n. sp. (Table 3) comprise less than 1% sequence divergence. The most distinguishing feature of Osedax packardorum n. sp. is the orange ring around the anterior part of the trunk. (Fig. 16 A-C, E).

Osedax ventana n. sp. Figure 4D Osedax ���nude-palp-B��� (Jones et al. 2008; Rouse et al. 2011; Rouse et al. 2009; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A1643 palps only (GenBank COI sequence EU235218), fixed in formalin preserved in ethanol, collected from cow bones deployed at 2898 meters depth in Monterey Submarine Canyon, California (36��36.606���N; 122��26.122���W) ROV Tiburon dive number 1069, Oct. 1, 2007. Diagnosis and description. Holotype represented by fragments of four palps (Fig. 4D), still within a gelatinous tube. Oviduct not discerned. In life, palps reddish with two blood vessels in each, lacking obvious pinnules (Fig. 4D). Distribution. Known from Monterey Bay, California from 2898 meters depth (Table 2). It has only been found in experimentally deployed cow bones. Etymology. This species is named (noun in apposition) for the ROV Ventana, which was used to collect many Osedax specimens. Remarks. Osedax ventana n. sp. is known from only a single partial specimen, sequenced by Jones et al. (2008). It was found on a deployed cow bone and was not discovered during any subsequent sampling at the same site. Osedax ventana n. sp. is part of Osedax Clade II and closest relative (with low support) to Osedax ryderi n.sp., from which it shows an uncorrected divergence of 12% (Table 4). Given the holotype is a fragment, there is little, except for its unique COI sequence, to distinguish Osedax ventana n. sp. from other Clade II species. Three other Osedax species: O. crouchi, O. nordenskjoeldi and O. rogersi, were similarly erected based on fragmented specimens and DNA data (Amon et al. 2014)., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 464, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Jones, W. J., Johnson, S. B., Rouse, G. W. & Vrijenhoek, R. C. (2008) Marine worms (genus Osedax) colonize cow bones. Proceedings of the Royal Society B, 275, 387 - 391. https: // doi. org / 10.1098 / rspb. 2007.1437","Rouse, G. W., Goffredi, S. K., Johnson, S. B. & Vrijenhoek, R. C. (2011) Not whale-fall specialists, Osedax worms also consume fishbones. Biology Letters, 7, 736 - 739. https: // doi. org / 10.1098 / rsbl. 2011.0202","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Amon, D. J., Wiklund, H., Dahlgren, T. G., Copley, J. T., Smith, C. R., Jamieson, A. J. & Glover, A. G. (2014) Molecular taxonomy of Osedax (Annelida: Siboglinidae) in the Southern Ocean. Zoologica Scripta, 43, 405 - 417. https: // doi. org / 10.1111 / zsc. 12057"]}

Osedax bryani n. sp. Figure 18 ���MB17��� (Rouse et al. 2015; Salath�� & Vrijenhoek 2012) Material examined. Holotype: SIO-BIC 4619, Female (GenBank COI sequence JX280609), fixed in formalin preserved in ethanol, collected from a whale carcass deployed at 1820 meters depth in Monterey Submarine Canyon, California (36��42.496���N; 122��6.316���W), ROV Doc Ricketts dive number 12, March 13, 2009. Diagnosis and description. Holotype and only available specimen female (Figs18A, B); in life trunk 4 mm long, 1.8 mm wide; crown of palps somewhat contracted, curled, 4.5 mm long. Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps. Oviduct extends from trunk into crown for 5 mm (Figs 18A, B). In life, pinnules of palps reddish, otherwise no obvious pigment. Trunk with white ring around anterior margin; ring broken mid-dorsally by oviduct and mid-ventrally by and oval unpigmented patch. Clear demarcation of upper and lower trunk. Ovisac in holotype, damaged, a large ellipsoidal mass (Figs 18A, B). Green root tissue visible but torn. Possible male found in tube of holotype, with chaetal-bearing segments inflated; found in tube lumen of females, lost following microscope slide preparation (Fig. 18C). Distribution. Known from Monterey Bay, California from 1820 meters depth in whale bone. Etymology. This species is named (noun in the genitive case) in honor of Bryan Touryan-Schaefer, ROV Pilot/ Technician for MBARI, who collected many bones with Osedax over the years. Remarks. Osedax bryani n. sp. is part of Osedax Clade V and is sister taxon to the O. rubiplumus and O. roseus clade (Fig. 1). It shows a minimum uncorrected distance, for COI, of ~15% to O. rubiplumus (Table 4). Osedax bryani n. sp. shares distinguishing feature of a white ring around the anterior part of the trunk with O. randyi n. sp. and O. frankpressi and as with Clade II of Osedax, these species likely need to be distinguished with DNA data. The documentation of dwarf males in Osedax bryani n. sp. is the first for this species, which was listed as unknown in Rouse et al. (2015)., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 483, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Salathe, R. M. & Vrijenhoek, R. C. (2012) Temporal variation and lack of host specificity among bacterial endosymbionts of Osedax bone worms (Polychaeta: Siboglinidae). BMC Evolutionary Biology, 12, 189. https: // doi. org / 10.1186 / 1471 - 2148 - 12 - 189"]}

Osedax packardorum n. sp. Figure 16 ���orange collar��� (Braby et al. 2007; Rouse et al. 2009; Rouse et al. 2015; Tresguerres et al. 2013; Vrijenhoek et al. 2009) ���sp.4 SBJ-2006��� (GenBank COI sequence DQ996639, DQ996641-2) Material examined. Holotype: Female, fixed in formalin preserved in ethanol, SIO-BIC A 1641 (GenBank COI sequence EU223341), collected from whale carcass (Balaenoptera musculus) deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W), ROV Tiburon dive number 1049, Oct. 25, 2006. Paratypes: Female and dwarf males (allotypes), fixed in formalin preserved in ethanol SIO-BIC A7840, A7842 (female GenBank COI sequences EU223339, EU223343), same locality and date as holotype; Female, fixed in formalin preserved in ethanol, SIO-BIC A7841(GenBank COI sequence EU223340), same locality and date as holotype; Female, sectioned for histology, same locality and date as holotype, SIO-BIC A7843 (GenBank COI sequence EU223342); Female, fixed in formalin preserved in ethanol, SIO-BIC A7844 (GenBank COI sequence FJ431200), collected from whale fall (Eschrichtius robustus) at 663 meters depth in Monterey Submarine Canyon, California (36��48.178���N; 121��59.677���W) ROV Tiburon dive number 1160, Dec. 18, 2007. Diagnosis and description. Holotype female, in life trunk 11 mm long, 0.6 mm wide; crown of palps contracted, curled, 2 mm long (image not shown as the trunk is damaged). Palps up to 12 mm long when extended in other specimens (Figs 16A, B). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (Figs 16B, D, E). Oviduct extends from trunk into crown for 3 mm (Fig. 16B). In life, palps reddish with two major blood vessels in each (Figs 16A, B). Tube a gelatinous loose mass around trunk (Figs 16A, B). Trunk with pale to dark orange ring around anterior margin; ring broken mid-dorsally by oviduct and midventrally by and oval unpigmented patch (Figs 16A, B, E). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (Fig. 16G). Ovisac in holotype an ellipsoidal mass (not shown); extending laterally as lobes in other specimens (Fig. 16C). Roots of holotype extend from ovisac on one side as one mass; in other specimens, roots as long branching masses. Males dwarfs, with spermatids and sperm (Figs 16F, G), chaetal-bearing segments not inflated (Fig. 16H); found in tube lumen of females. Distribution. Known from Monterey Bay, California at 349, 633, and 1018 meters depths (Table 2). Found in whale and cow bones. Etymology. This species is named (noun in the genitive case) in honor of the Packard family whose foundation supports MBARI and enabled the discovery of all of the Osedax species in California. Remarks. Osedax packardorum n. sp. is part of Osedax Clade IV and closest relative to Osedax lehmani n. sp., (Fig. 1), from which it has a minimum uncorrected distance for COI of 8-9% (Table 4). As pointed out above, this is one of the smaller interspecific distances among Osedax species and the two species are morphologically very similar. All 27 available COI sequences for Osedax packardorum n. sp. (Table 3) comprise less than 1% sequence divergence. The most distinguishing feature of Osedax packardorum n. sp. is the orange ring around the anterior part of the trunk. (Fig. 16 A-C, E)., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on pages 479-481, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Braby, C. E., Rouse, G. W., Johnson, S. B., Jones, W. J. & Vrijenhoek, R. C. (2007) Bathymetric and temporal variation among Osedax boneworms and associated megafauna on whale-falls in Monterey Bay, California. Deep-Sea Research Part I, 54, 1773 - 1791. https: // doi. org / 10.1016 / j. dsr. 2007.05.014","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Tresguerres, M., Katz, S. & Rouse, G. W. (2013) How to get into bones: proton pump and carbonic anhydrase in Osedax boneworms. Proceedings of the Royal Society of London Series B-Biological Sciences, 280, 20130625. https: // doi. org / 10.1098 / rspb. 2013.0625"]}

Osedax tiburon n. sp. Figure 4A, B, C ���nude-palp-A��� (Jones et al. 2008; Rouse et al. 2011; Rouse et al. 2009; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A7817 female (GenBank COI sequence FJ347622), fixed in formalin preserved in ethanol, collected from whale carcass (Eschrichtius robustus) deployed at 1820 meters depth in Monterey Submarine Canyon, California (36��42.496���N; 122��6.316���W) ROV Tiburon dive number 1163, Dec. 20, 2007. Paratypes: All females, fixed in formalin preserved in ethanol, SIO-BIC A1642 (GenBank COI sequence EU223359), SIO-BIC A7820 (GenBank COI sequence EU223356, EU223357), same locality as holotype, ROV Tiburon dive number 1119, Aug. 16, 2007; SIO-BIC A7818, A7819 (GenBank COI sequences FJ347623, FJ347624, respectively) same locality and date as holotype. Diagnosis and description. Holotype female (Fig. 4B); preserved trunk 0.75 mm long, 0.2 mm wide; crown of four palps, 3 mm long. Tube gelatinous 6 mm long (Fig. 4A). Oviduct not discerned. In life, palps reddish with blood (Fig. 4A). Four palps without obvious pinnules (Fig. 4B, C). Trunk with no obvious demarcation into upper and lower trunk (Fig. 4B, C). Ovisac mostly missing in holotype, some traces of roots present (Fig. 4A). In paratype ovisac and root complex extends laterally on either side of trunk and posteriorly (Fig. 4C). Males not seen. Distribution. Known from Monterey Bay, California from 1820 meters depth (Table 2). Etymology. This species is named (noun in apposition) for the ROV Tiburon, which was used to collect the first Osedax specimens. Remarks. Osedax tiburon n. sp. is part of Osedax Clade II and is closest relative (though with low support) to Osedax nordenskjoeldi from which it shows a divergence of 15%, though it shows a smaller pairwise distance (14%) to O. ryderi n. sp. (Table 4). The seven available COI sequences for Osedax tiburon n. sp. (Table 3) show less than 1% sequence divergence. It has been found in whale bones and experimentally deployed cow bones. There is little to distinguish Osedax tiburon n. sp. from other 'nude palp' species in Clade II, which share a similar morphology. The species status of Osedax tiburon n. sp. is supported largely on its COI divergence from other Osedax., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 462, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Jones, W. J., Johnson, S. B., Rouse, G. W. & Vrijenhoek, R. C. (2008) Marine worms (genus Osedax) colonize cow bones. Proceedings of the Royal Society B, 275, 387 - 391. https: // doi. org / 10.1098 / rspb. 2007.1437","Rouse, G. W., Goffredi, S. K., Johnson, S. B. & Vrijenhoek, R. C. (2011) Not whale-fall specialists, Osedax worms also consume fishbones. Biology Letters, 7, 736 - 739. https: // doi. org / 10.1098 / rsbl. 2011.0202","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032"]}

Osedax docricketts n. sp. Figure 4E ‘nude-palp-C’ (Higgs et al. 2014; Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009) ‘Sagami-6’ (GenBank COI sequence series FM998088-FM998107; Pradillon et al. unpublished) Holotype: SIO-BIC A 1644, Female (GenBank COI sequence FJ347626), fixed in formalin preserved in ethanol, collected from whale fall deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W), ROV Tiburon dive number 1049, Oct. 25, 2006. Diagnosis and description. Holotype incomplete female; preserved trunk 0. 5 mm long, 0.2 mm wide; crown of four palps, 2 mm long (Fig. 4E). Tube gelatinous cylinder around trunk and base of palps. Oviduct not discerned. In life, palps white with some reddish patches. Palps without obvious pinnules (Fig. 4E). Trunk with no obvious demarcation into upper and lower trunk, or any pigmentation. Ovisac mostly missing in holotype, some traces of roots present. Males not seen. Distribution. Known from Monterey Bay, California from 1820 meters depth, and Sagami Bay, Japan (Table 2). It has been found living on cow and whale bones. Etymology. This species is named (noun in apposition) for the ROV Doc Ricketts, which was used to collect many Osedax specimens. Remarks. Osedax docricketts n. sp. is part of Osedax Clade II, which all show apinnulate palps, and is closest relative to the clade comprised of Osedax westernflye r n. sp. and O. knutei n. sp. (minimum divergence of 15%) though it shows smaller distance (14%) to other taxa such as O. lonnyi n. sp. (Table 4). The four available COI sequences for Osedax docricketts n. sp. from California (Table 3) show less that 1% sequence divergence. The only retained specimen is the holotype as the others were destroyed for sequencing and the holotype itself is only a partial specimen. Twenty additional COI sequences were referred to Osedax sp. Sagami-6 (Pradillon et al., GenBank, unpublished). Together, the four California and 11 of the Japanese sequences (FM998088, FM998089, FM998091, FM998093, FM998094, FM998098, FM998099, FM998100, FM998101, FM998103, FM998107) encompass less than 1% sequence divergence; so, we propose that Osedax docricketts n. sp. is also found in Japan. The nine remaining Japanese specimens with>2.4% uncorrected pairwise distance from the 11 Japanese sequences (GenBank sequences: FM998090, FM998092, FM998095, FM998096, FM998097, FM998102, FM998104, FM998105, FM998106) require further investigation before their assignment to Osedax docricketts n. sp. can be justified. See Table 4 for comparison of California Osedax docricketts n. sp. versus Japanese Sagami-6 representative sequences. There is little to distinguish Osedax docricketts n. sp. from other Clade II species.

Osedax randyi n. sp. Figure 17 ���white collar��� (Rouse et al. 2015; Vrijenhoek et al. 2009) ���Sagami-7��� (GenBank COI sequence FM998108-9; Pradillon et al. unpublished) Material examined. Holotype: Female, fixed in formalin preserved in ethanol, SIO-BIC A 7845 (GenBank COI sequence FJ347615), collected from a whale carcass (Balaenoptera musculus) deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W), ROV Tiburon dive number 1049, Oct. 25, 2006. Paratypes: Female, fixed in formalin preserved in ethanol SIO-BIC A1648, SIO-BIC A7846 (GenBank COI sequence FJ347612, FJ347613), same locality and date as holotype; Female and dwarf males (allotypes), fixed in formalin, female sectioned, males preserved in ethanol SIO-BIC A7847 (female GenBank COI sequence FJ347614), same locality and date as holotype. Diagnosis and description. Holotype female, in life trunk 5 mm long, 0.8 mm wide; crown of palps somewhat contracted, 4.5 mm long (Fig. 17A). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (Figs 17A, B, D). Oviduct extends from trunk into crown for 1-2 mm (Figs 17A, B). In life, palps reddish with pigmented white outer margins (Fig. 17A). Trunk with white ring around anterior margin; ring broken mid-dorsally by oviduct and mid-ventrally by and oval unpigmented patch (Figs 17A, B). Clear demarcation of upper and lower trunk (Fig. 17A). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (Fig. 17E). Ovisac in holotype, damaged, an ellipsoidal mass; similar shape in paratypes. Roots incomplete, simple lobes where present in all specimens. Males dwarfs, with spermatids and sperm, chaetal-bearing segments inflated (Fig. 17C), found in tube lumen of females. Distribution. Known from Monterey Bay, California from 1018 meters depth, and from Sagami Bay, Japan (Table 2). It has been found in whale and cow bones. Etymology. This species is named (noun in the genitive case) in honor of Randy Prickett, Senior ROV pilot for MBARI, who collected many bones and Osedax over the years. Remarks. Osedax randyi n. sp. is part of Osedax Clade IV and closest relative to the undescribed Osedax species from California known as O. ���MB16��� (Fig. 1) with a minimum uncorrected COI- distance of ~6-7% (Table 4). As previously noted, this is among the smaller interspecific distances among Osedax species. Two COI sequences, referred to Osedax sp. Sagami-7 (Pradillon et al., GenBank unpublished, Table 3) are much less than 1% divergent from the six California Osedax randyi n. sp. sequences (example distance in Table 4). Thus, it is reasonable to propose that Osedax randyi n. sp. is also found in Japan. The most distinguishing features of Osedax randyi n. sp. are the white ring around the anterior part of the trunk and white pigment on the palps of females (Fig. 17A, B), though a similar whitish ring is found on females of O. bryani n. sp. (Fig. 18A, B) and O. frankpressi (Rouse et al. 2004), with the latter species also showing white pigmentation on the palps. DNA sequencing is most likely the best way to distinguish Osedax randyi n. sp. from these species., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on pages 481-483, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z","Rouse, G. W., Goffredi, S. K. & Vrijenhoek, R. C. (2004) Osedax: Bone-eating marine worms with dwarf males. Science, 305, 668 - 671. https: // doi. org / 10.1126 / science. 1098650"]}

Osedax bryani n. sp. Figure 18 ‘MB17’ (Rouse et al. 2015; Salathé & Vrijenhoek 2012) Material examined. Holotype: SIO-BIC 4619, Female (GenBank COI sequence JX280609), fixed in formalin preserved in ethanol, collected from a whale carcass deployed at 1820 meters depth in Monterey Submarine Canyon, California (36°42.496’N; 122°6.316’W), ROV Doc Ricketts dive number 12, March 13, 2009. Diagnosis and description. Holotype and only available specimen female (Figs18A, B); in life trunk 4 mm long, 1.8 mm wide; crown of palps somewhat contracted, curled, 4.5 mm long. Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps. Oviduct extends from trunk into crown for 5 mm (Figs 18A, B). In life, pinnules of palps reddish, otherwise no obvious pigment. Trunk with white ring around anterior margin; ring broken mid-dorsally by oviduct and mid-ventrally by and oval unpigmented patch. Clear demarcation of upper and lower trunk. Ovisac in holotype, damaged, a large ellipsoidal mass (Figs 18A, B). Green root tissue visible but torn. Possible male found in tube of holotype, with chaetal-bearing segments inflated; found in tube lumen of females, lost following microscope slide preparation (Fig. 18C). Distribution. Known from Monterey Bay, California from 1820 meters depth in whale bone. Etymology. This species is named (noun in the genitive case) in honor of Bryan Touryan-Schaefer, ROV Pilot/ Technician for MBARI, who collected many bones with Osedax over the years. Remarks. Osedax bryani n. sp. is part of Osedax Clade V and is sister taxon to the O. rubiplumus and O. roseus clade (Fig. 1). It shows a minimum uncorrected distance, for COI, of ~15% to O. rubiplumus (Table 4). Osedax bryani n. sp. shares distinguishing feature of a white ring around the anterior part of the trunk with O. randyi n. sp. and O. frankpressi and as with Clade II of Osedax, these species likely need to be distinguished with DNA data. The documentation of dwarf males in Osedax bryani n. sp. is the first for this species, which was listed as unknown in Rouse et al. (2015).

Osedax knutei n. sp. Figure 6 ���nude-palp-E��� (Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009; Worsaae et al. 2016) Material examined. Holotype: SIO-BIC A 7812, Female (GenBank COI sequence FJ347635), fixed in formalin preserved in ethanol, collected from whale fall (Balaenoptera musculus) deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W), ROV Tiburon dive number 1049, Oct. 25, 2006; Allotype, dwarf male from tube of holotype SIO-BIC A7813. Paratypes: All females, fixed in formalin preserved in ethanol; SIO-BIC A1646 (GenBank COI sequence FJ347634), SIO-BIC A7814, (GenBank COI sequence FJ347632), same locality and date as holotype; SIO-BIC A7815, (GenBank COI sequence JF509952), collected from fish bones deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W), ROV Doc Ricketts dive number 209, Oct. 29, 2010; SIO-BIC A7816, (GenBank COI sequences MG262306), collected from turkey (Meleagris gallopavo) bone deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W), ROV Doc Ricketts dive number 233, June 2, 2011. Diagnosis and description. Holotype female preserved partially dissected from whale bone (Fig. 6A); trunk contracted 1.6 mm long, 0.3 mm wide; crown of apinnulate palps contracted, 2.3 mm long. Tube not kept for holotype, but forms a tight sheath in other specimens, from which palps emerge. Oviduct not visible in holotype (Fig. 6A), but is slightly shorter (1.5 mm) than contracted palps in paratype SIO-BIC A7816, and much shorter than extended palps in live specimen spawning eggs (Fig. 6D). In life, palps reddish with two blood vessels each (Fig. 6E). Trunk with no obvious pigmentation or demarcation into upper and lower trunk (Figs 6A, D). Ovisac represented as lobes on either side of trunk in holotype, extending laterally, or as long lobes in other specimens. Roots of holotype longer that the remaining body, extending in several directions (Fig. 6A). In other specimens, roots also extend in various directions (Fig. 6D). Males dwarfs, 450 ��m long by 100 ��m wide, with anterior prototroch and sperm, two posterior expanded segments bearing hooked chaetae (Figs 6B, C). Distribution. Known from Monterey Bay, California from 1018 to 2898 meters depth (Table 2). It has been found in whale, cow, turkey and teleost bones. Etymology. This species is named (noun in the genitive case) for Knute Brekke, Chief ROV pilot for MBARI, who expertly collected many bones and Osedax over the years. Remarks. Osedax knutei n. sp. is part of Osedax Clade II and closest relative to Osedax westernflyer n. sp, (Fig. 1) with a minimum uncorrected distance for COI of 16% (Table 4). Twelve COI sequences for Osedax knutei n. sp. (Table 3) generally showed less that 1.2% sequence divergence from each other. An exceptional sequence (JF509952) from paratype SIO-BIC A7815 dissected from fishbone (Rouse et al. 2011), diverged by 3.2% to 4.5% from the other Osedax knutei n. sp. specimens (see example distance in Table 4). The tentative assignment of paratype SIO-BIC A7815 to Osedax knutei n. sp. requires further investigation. Two new COI sequences for Osedax knutei n. sp. were also deposited on GenBank for which no voucher specimens were retained; MG262305 was recovered from a fishbone deployed at 2898 meters and MG262307 was from a turkey deployed at 1018 meters, Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on pages 465-468, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Rouse, G. W., Goffredi, S. K., Johnson, S. B. & Vrijenhoek, R. C. (2011) Not whale-fall specialists, Osedax worms also consume fishbones. Biology Letters, 7, 736 - 739. https: // doi. org / 10.1098 / rsbl. 2011.0202","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z","Worsaae, K., Rimskaya-Korsakova, N. N. & Rouse, G. W. (2016) Neural reconstruction of bone-eating Osedax spp. (Annelida) and evolution of the siboglinid nervous system. BMC Evolutionary Biology, 16, 83. https: // doi. org / 10.1186 / s 12862 - 016 - 0639 - 7"]}

Osedax lehmani n. sp. Figures 14, 15 ���yellow collar���, (Braby et al. 2007; Huusgaard et al. 2012; Katz & Rouse 2013; Rouse et al. 2009; Rouse et al. 2015; Tresguerres et al. 2013; Vrijenhoek et al. 2009; Worsaae et al. 2016) ���sp.3 SBJ-2006��� (GenBank COI sequences DQ996629- DQ99638) ���sp.4 SBJ-2006��� (GenBank COI sequences DQ996640, DQ99643) ���orange collar��� (GenBank COI sequence EU267672) Material examined. Holotype: SIO-BIC A7804, Female (GenBank COI sequence DQ996629), fixed in glutaraldehyde, preserved in ethanol, collected from a whale carcass (Eschrichtius robustus) deployed at 385 meters depth in Monterey Submarine Canyon, California (36��47.401���N; 122�� 53.235���W), ROV Tiburon dive number 933, Jan. 6, 2006. Paratypes: Females, fixed in formalin or glutaraldehyde, preserved in ethanol, same locality and date as holotype, SIO-BIC A1640 (GenBank COI sequence DQ996630), same locality and date as holotype, SIO-BIC A7806 (GenBank COI sequence DQ996631); SIO-BIC A7807 and A7808 (GenBank COI sequence EU223332 and EU223337, respectively), same locality as holotype, ROV Tiburon dive number 1070, Jan. 11, 2007; dwarf males (Allotypes) SIO-BIC A7805 (no GenBank COI sequence), several in tubes of various females, same locality and date as holotype. Diagnosis and description. Holotype female, in life trunk 8 mm long, 0.6 mm wide; crown of palps contracted, curled, 1.5 mm long (Fig. 14C). Palps up to 10 mm long when extended in other specimens (Figs 14A, D). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (Figs 14B, D, E, 15A). Oviduct extends from trunk into crown for 1.75 mm (Figs 14 B-D). In life, palps reddish with two major blood vessels in each (Figs 14A, D). Tube a gelatinous loose mass around trunk (Fig. 14D). Trunk with pale yellow ring around anterior margin; ring broken mid-dorsally by oviduct (Figs 14B, D). and mid-ventrally by an oval unpigmented patch (Fig. 14C). Clear demarcation of upper and lower trunk with oviduct emerging onto outer surface of upper trunk (Fig. 14B). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (Fig. 15B). Ovisac in holotype extending laterally as two masses; simple lobes in other specimens (Fig. 14C). Roots of holotype extend from ovisac lobes (Fig. 14C); in other specimens, roots as simple lobes or extending laterally in two lobe or lobes (Fig. 14B). Males dwarfs, with spermatids and sperm, chaetalbearing segments not inflated; found in tube lumen of females (Fig. 15 C-D). Distribution. Known from Monterey Bay, California only from 349 meters depth (Table 2). It has been found in whale and cow bones. Etymology. This species is named (noun in the genitive case) in memory of Alan George Lehman, father of Ellen Lehman, in recognition of her long and continued support of the Scripps Oceanographic Collections. Remarks. Osedax lehmani n. sp. is part of Osedax Clade IV and is closest relative to Osedax packardorum n. sp., (Fig. 1), with a minimum uncorrected distance for COI of 8-9% (Table 4). This is one of the smaller intraspecific distances among Osedax species, with others being between O. frankpressi and the O. '1336_61_2' specimen from the Atlantic at ~10%; and O. randyi n. sp. and O. ���MB16 at ~ 6-7% (Table 4). The majority of the 31 available COI sequences for Osedax lehmani n. sp. (Table 3) show less than 1% divergence, though a few sequences e.g. DQ996640 are around 2% divergent from the majority and DQ996631 shows the greatest distance, at ~3% (Table 4), so the possibility of there being a cryptic species complex for O. lehmani n. sp. needs further investigation. The most distinguishing feature of Osedax lehmani n. sp. is the yellow ring around the anterior part of the trunk. (Fig. 14 A-D). This is the only Osedax species from Monterey Bay that was not found deeper than 389 meters depth., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 479, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Braby, C. E., Rouse, G. W., Johnson, S. B., Jones, W. J. & Vrijenhoek, R. C. (2007) Bathymetric and temporal variation among Osedax boneworms and associated megafauna on whale-falls in Monterey Bay, California. Deep-Sea Research Part I, 54, 1773 - 1791. https: // doi. org / 10.1016 / j. dsr. 2007.05.014","Huusgaard, R. S., Vismann, B., Kuhl, M., Macnaugton, M., Colmander, V., Rouse, G. W., Glover, A. G., Dahlgren, T. & Worsaae, K. (2012) The potent respiratory system of Osedax mucofloris (Siboglinidae, Annelida) - a prerequisite for the origin of bone-eating Osedax? PLoS ONE, 7, e 35975. https: // doi. org / 10.1371 / journal. pone. 0035975","Katz, S. & Rouse, G. W. (2013) The reproductive system of Osedax (Annelida, Siboglinidae): sperm ultrastructure, ovary structure and fertilization. Invertebrate Biology, 132, 368 - 385. https: // doi. org / 10.1111 / ivb. 12037","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Tresguerres, M., Katz, S. & Rouse, G. W. (2013) How to get into bones: proton pump and carbonic anhydrase in Osedax boneworms. Proceedings of the Royal Society of London Series B-Biological Sciences, 280, 20130625. https: // doi. org / 10.1098 / rspb. 2013.0625","Worsaae, K., Rimskaya-Korsakova, N. N. & Rouse, G. W. (2016) Neural reconstruction of bone-eating Osedax spp. (Annelida) and evolution of the siboglinid nervous system. BMC Evolutionary Biology, 16, 83. https: // doi. org / 10.1186 / s 12862 - 016 - 0639 - 7"]}

Osedax ryderi n. sp. Fig. 8 ‘nude-palp-G’, ‘nude palp #20’ (Higgs et al. 2014; Rouse et al. 2015) Material examined. Holotype: Female, SIO-BIC 4617 (GenBank COI sequence KP119563), fixed in glutaraldehyde-preserved in ethanol, collected from a natural whale fall (Eschrichtius robustus) found at 2898 meters depth in Monterey Submarine Canyon, California (36°36.606’N; 122°26.122’W) ROV Doc Ricketts dive number 234, June 3, 2011. Allotype: Male found in tube of holotype, SIO-BIC A4618. Diagnosis and description. Holotype female preserved partially dissected from bone; trunk contacted 0.6 mm long, 0.3 mm wide; crown of apinnulate palps contracted, 0.7 mm long (Fig. 8B). Tube not kept for holotype, but forms a tapering sheath in life, from which palps were not seen emerging (Fig. 8A). Oviduct slightly longer than palps visible in holotype (Fig. 8B). In life, palps reddish with two blood vessels each (Fig. 8A). Trunk with no obvious pigmentation or demarcation into upper and lower trunk. Ovisac represented as lobes on either side of trunk in holotype, extending laterally, or as long lobes in other specimens. Roots of holotype longer that the remaining body, extending in several directions as a single lobate mass (Fig. 8B). Male in a tube of holotype a dwarf, 210 µm long by 50 µm wide, with anterior prototroch and sperm squeezed out of head, two posterior expanded segments bearing hooked chaetae (Figs 8C, D). Another possible male, also in tube, resembled newlyrecruited Osedax larva (Fig. 8E). Distribution. Known from Monterey Bay, California from 2898 meters depth (Table 2). It has been found in whale and turtle bones. Etymology. This species is named (noun in the genitive case) for Ryder Williams, for his generosity in sharing his mom for our research efforts. Remarks. Osedax ryderi n. sp. is part of Osedax Clade II and is the well-supported sistergroup to Osedax ventana n. sp., another Monterey Bay species with an uncorrected divergence of 12%. Two other COI sequences obtained for specimens from the type locality, for which no voucher material was obtained, differed from the type specimen by less than 1% (MG262308 from whale bone and MG262309 from turtle bone). There is little to distinguish Osedax ryderi n. sp. from other Clade II species, apart from genetic differences.

Osedax sigridae n. sp. Figure 2 ‘green palp’ (Katz et al. 2010; Katz et al. 2011; Katz & Rouse 2013; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A7809 female (GenBank COI sequence FJ347641), collected from an experimentally deployed whale carcass (Eschrichtius robustus) deployed at 1820 meters depth in Monterey Submarine Canyon, California (36°42.496’N; 122°6.316’W) ROV Tiburon dive number 1163, Dec. 20, 2007. Fixed in formalin preserved in ethanol. Paratypes: All females. Fixed in formalin preserved in ethanol; SIO-BIC A7811 same locality and date as holotype (GenBank COI sequence FJ347642); SIO-BIC A1650 (GenBank COI sequence FJ347639) and SIO-BIC A7810 (GenBank COI sequence FJ347640), collected on cow bones deployed at 1820 meters depth in Monterey Submarine Canyon, California (36°42.496’N; 122°6.316’W) ROV Tiburon dive number 1119, Aug. 16, 2007. Diagnosis and description. Holotype female (Fig. 2D); trunk 1 mm long, 0.5 mm wide; crown of pinnulated palps contracted, 2 mm long. Tube gelatinous, up to 10 mm long, 5 mm across. Oviduct shorter than palps (Fig. 2C). In life, palps green-yellow (Figs 2A, C, D). Pinnules of all four palps oriented dorsally (Fig. 2D). No obvious pigmentation on trunk, or clear demarcation into upper and lower trunk. Ovisac spheroidal with roots as simple lobes (Figs 2B, D). Males not found. Distribution. Known from Monterey Bay, California from 1820 meters depth (Table 2). It has been found in whale and cow bones. Etymology. This species is named (noun in the genitive case) for Sigrid Katz, whose Ph.D. project involved detailed study of the anatomy of this species (Katz et al. 2010; Katz et al. 2011). Remarks. Osedax sigridae n. sp. is part of Osedax clade I and closest relative to the undescribed Japanese taxon Osedax ‘Sagami-5’ (Fig. 1). All four available COI sequences for Osedax sigridae n. sp. (Table 3) show less than 1% uncorrected sequence divergence, while the minimum uncorrected distance to Osedax ‘Sagami-5’ is 11% (Table 4). Careful searching for males in some of the tubes of the Osedax sigridae n. sp. females shown in Figure 2A failed, though Katz & Rouse (2013) did find sperm in the ovisac of females. The most distinguishing features of the Osedax sigridae n. sp. specimens collected to date are the distinctive green/yellow palps and the lobate roots (Fig. 2).

Osedax ryderi n. sp. Fig. 8 ���nude-palp-G���, ���nude palp #20��� (Higgs et al. 2014; Rouse et al. 2015) Material examined. Holotype: Female, SIO-BIC 4617 (GenBank COI sequence KP119563), fixed in glutaraldehyde-preserved in ethanol, collected from a natural whale fall (Eschrichtius robustus) found at 2898 meters depth in Monterey Submarine Canyon, California (36��36.606���N; 122��26.122���W) ROV Doc Ricketts dive number 234, June 3, 2011. Allotype: Male found in tube of holotype, SIO-BIC A4618. Diagnosis and description. Holotype female preserved partially dissected from bone; trunk contacted 0.6 mm long, 0.3 mm wide; crown of apinnulate palps contracted, 0.7 mm long (Fig. 8B). Tube not kept for holotype, but forms a tapering sheath in life, from which palps were not seen emerging (Fig. 8A). Oviduct slightly longer than palps visible in holotype (Fig. 8B). In life, palps reddish with two blood vessels each (Fig. 8A). Trunk with no obvious pigmentation or demarcation into upper and lower trunk. Ovisac represented as lobes on either side of trunk in holotype, extending laterally, or as long lobes in other specimens. Roots of holotype longer that the remaining body, extending in several directions as a single lobate mass (Fig. 8B). Male in a tube of holotype a dwarf, 210 ��m long by 50 ��m wide, with anterior prototroch and sperm squeezed out of head, two posterior expanded segments bearing hooked chaetae (Figs 8C, D). Another possible male, also in tube, resembled newlyrecruited Osedax larva (Fig. 8E). Distribution. Known from Monterey Bay, California from 2898 meters depth (Table 2). It has been found in whale and turtle bones. Etymology. This species is named (noun in the genitive case) for Ryder Williams, for his generosity in sharing his mom for our research efforts. Remarks. Osedax ryderi n. sp. is part of Osedax Clade II and is the well-supported sistergroup to Osedax ventana n. sp., another Monterey Bay species with an uncorrected divergence of 12%. Two other COI sequences obtained for specimens from the type locality, for which no voucher material was obtained, differed from the type specimen by less than 1% (MG262308 from whale bone and MG262309 from turtle bone). There is little to distinguish Osedax ryderi n. sp. from other Clade II species, apart from genetic differences., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on pages 469-471, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Higgs, N. D., Glover, A. G., Dahlgren, T. G., Smith, C. R., Fujiwara, Y., Pradillon, F., Johnson, S. B., Vrijenhoek, R. C. & Little, C. T. S. (2014) The morphological diversity of Osedax worm borings (Annelida: Siboglinidae). Journal of the Marine Biological Association of the UK, 94, 1429 - 1439. https: // doi. org / 10.1017 / S 0025315414000770","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032"]}

Osedax lonnyi n. sp. Figure 7 ‘nude-palp-F’ (Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: Female, SIO-BIC A1647 (GenBank COI sequence FJ347643), fixed in glutaraldehyde-preserved in ethanol, collected from natural whale fall (Eschrichtius robustus) found at 2898 meters depth in Monterey Submarine Canyon, California (36°36.606’N; 122°26.122’W) ROV Tiburon dive number 1162, Dec. 19, 2007. Diagnosis and description. Holotype female; in life trunk 1 mm long, 0.5 mm wide (Fig. 7A), crown of four palps, 2.5 cm long (Fig. 7A), likely contracted even at this length. Tube gelatinous, thin, fitting about trunk and base of crown, 4 cm long (Fig. 7A). In life, palps with visible blood vessels and greenish tips (Fig. 7A, inset). Palps without obvious pinnules, oviduct extends slightly beyond palps (Fig. 7A inset). Trunk with pigmented ring demarking upper and lower trunk (Fig. 7B). Ovisac 2 mm by 1 mm with damaged lobate greenish roots extending outwards (Fig. 7B). Males not found. Distribution. Known from Monterey Bay, California from 2898 meters depth (Table 2). It has only been found in a whale bone fragment, adjacent to the main skeleton. Etymology. This species is named (noun in the genitive case) for Lonny Lundsten, Senior Research Technician at MBARI, for his enthusiasm and assistance on many Osedax expeditions. Remarks. Osedax lonnyi n. sp. is part of Osedax Clade II and is the well-supported sistergroup to Osedax rogersi, an Antarctic species with an uncorrected divergence of 12%. Osedax lonnyi n. sp. is remarkable for the very long palps (2.5 cm) that it shows relative to the very short trunk (1.5 mm). The species is known from only a single specimen (holotype) collected in 2007 and no further specimens were found during any subsequent sampling at the same site. This is somewhat surprising as it is one of the larger species of Clade II and the whale fall was sampled a number of times.

Siboglinidae Caullery, 1914 Osedax Rouse, Goffredi & Vrijenhoek 2004 Diagnosis. Siboglinids with females having a contractile trunk, bulbous ovisac, and branching ���roots���. A crown of palps usually present, with or without pinnules. Trunk lies within transparent tube emergent from bone surface. Mostly males are dwarfs resembling larvae, exceptionally having a crown, contractile trunk, bulbous testis sac, and branching ���roots���, as in O. priapus. Crown in females, when present, comprised of cylindrical oviduct plus four palps. Osedax priapus, the only species known to produce adult bone-eating males, has crown with only two palps. No mouth or obvious gut. Cylindrical trunk comprised mostly of longitudinal muscles and glands, with large dorsal and ventral blood vessels present. Oviduct or sperm duct runs dorsally along trunk surface into posterior ovisac, or testis sac. Ovisac or testis sac enclosed by epidermis and trophosome with bacteriocytes, which also extends outwards as vascularized ���roots���. No chaetae or segmentation apparent in females or bone-eating males. In most species, paedomorphic dwarf males cluster around oviduct in gelatinous tube surrounding trunk of female. The dwarf males possess anterior prototroch and posterior hooked chaetae arranged on two segments. Hooks, lacking rostrum, comprise capitium with curved teeth over subrostral process. Internally, males contain spermatids and sperm anteriorly. Remarks. This diagnosis is revised from that in Rouse et al. (2004) to accommodate the diversity of Osedax forms. These include Osedax jabba n. sp. where the females lack palps entirely and Osedax priapus where the males are not paedomorphic dwarfs, but also consume bone and have similar anatomy to females. Many Osedax species have palps that lack obvious pinnules. The dorsal placement of the oviduct reflects the reorientation of Osedax as reported in Huusgaard et al. (2012) and Worsaae et al. (2016)., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 459, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Caullery, M. (1914) Sur les Siboglinidae, type nouveau d'invertebres receuillis par l'expedition du Siboga. Bulletin de la Societe Zoologique de France (Evolution et Zoologie), 39, 350 - 353.","Rouse, G. W., Goffredi, S. K. & Vrijenhoek, R. C. (2004) Osedax: Bone-eating marine worms with dwarf males. Science, 305, 668 - 671. https: // doi. org / 10.1126 / science. 1098650","Huusgaard, R. S., Vismann, B., Kuhl, M., Macnaugton, M., Colmander, V., Rouse, G. W., Glover, A. G., Dahlgren, T. & Worsaae, K. (2012) The potent respiratory system of Osedax mucofloris (Siboglinidae, Annelida) - a prerequisite for the origin of bone-eating Osedax? PLoS ONE, 7, e 35975. https: // doi. org / 10.1371 / journal. pone. 0035975","Worsaae, K., Rimskaya-Korsakova, N. N. & Rouse, G. W. (2016) Neural reconstruction of bone-eating Osedax spp. (Annelida) and evolution of the siboglinid nervous system. BMC Evolutionary Biology, 16, 83. https: // doi. org / 10.1186 / s 12862 - 016 - 0639 - 7"]}

Osedax knutei n. sp. Figure 6 ‘nude-palp-E’ (Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009; Worsaae et al. 2016) Material examined. Holotype: SIO-BIC A 7812, Female (GenBank COI sequence FJ347635), fixed in formalin preserved in ethanol, collected from whale fall (Balaenoptera musculus) deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W), ROV Tiburon dive number 1049, Oct. 25, 2006; Allotype, dwarf male from tube of holotype SIO-BIC A7813. Paratypes: All females, fixed in formalin preserved in ethanol; SIO-BIC A1646 (GenBank COI sequence FJ347634), SIO-BIC A7814, (GenBank COI sequence FJ347632), same locality and date as holotype; SIO-BIC A7815, (GenBank COI sequence JF509952), collected from fish bones deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W), ROV Doc Ricketts dive number 209, Oct. 29, 2010; SIO-BIC A7816, (GenBank COI sequences MG262306), collected from turkey (Meleagris gallopavo) bone deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W), ROV Doc Ricketts dive number 233, June 2, 2011. Diagnosis and description. Holotype female preserved partially dissected from whale bone (Fig. 6A); trunk contracted 1.6 mm long, 0.3 mm wide; crown of apinnulate palps contracted, 2.3 mm long. Tube not kept for holotype, but forms a tight sheath in other specimens, from which palps emerge. Oviduct not visible in holotype (Fig. 6A), but is slightly shorter (1.5 mm) than contracted palps in paratype SIO-BIC A7816, and much shorter than extended palps in live specimen spawning eggs (Fig. 6D). In life, palps reddish with two blood vessels each (Fig. 6E). Trunk with no obvious pigmentation or demarcation into upper and lower trunk (Figs 6A, D). Ovisac represented as lobes on either side of trunk in holotype, extending laterally, or as long lobes in other specimens. Roots of holotype longer that the remaining body, extending in several directions (Fig. 6A). In other specimens, roots also extend in various directions (Fig. 6D). Males dwarfs, 450 µm long by 100 µm wide, with anterior prototroch and sperm, two posterior expanded segments bearing hooked chaetae (Figs 6B, C). Distribution. Known from Monterey Bay, California from 1018 to 2898 meters depth (Table 2). It has been found in whale, cow, turkey and teleost bones. Etymology. This species is named (noun in the genitive case) for Knute Brekke, Chief ROV pilot for MBARI, who expertly collected many bones and Osedax over the years. Remarks. Osedax knutei n. sp. is part of Osedax Clade II and closest relative to Osedax westernflyer n. sp, (Fig. 1) with a minimum uncorrected distance for COI of 16% (Table 4). Twelve COI sequences for Osedax knutei n. sp. (Table 3) generally showed less that 1.2% sequence divergence from each other. An exceptional sequence (JF509952) from paratype SIO-BIC A7815 dissected from fishbone (Rouse et al. 2011), diverged by 3.2% to 4.5% from the other Osedax knutei n. sp. specimens (see example distance in Table 4). The tentative assignment of paratype SIO-BIC A7815 to Osedax knutei n. sp. requires further investigation. Two new COI sequences for Osedax knutei n. sp. were also deposited on GenBank for which no voucher specimens were retained; MG262305 was recovered from a fishbone deployed at 2898 meters and MG262307 was from a turkey deployed at 1018 meters

Osedax lehmani n. sp. Figures 14, 15 ‘yellow collar’, (Braby et al. 2007; Huusgaard et al. 2012; Katz & Rouse 2013; Rouse et al. 2009; Rouse et al. 2015; Tresguerres et al. 2013; Vrijenhoek et al. 2009; Worsaae et al. 2016) ‘sp.3 SBJ-2006’ (GenBank COI sequences DQ996629- DQ99638) ‘sp.4 SBJ-2006’ (GenBank COI sequences DQ996640, DQ99643) ‘orange collar’ (GenBank COI sequence EU267672) Material examined. Holotype: SIO-BIC A7804, Female (GenBank COI sequence DQ996629), fixed in glutaraldehyde, preserved in ethanol, collected from a whale carcass (Eschrichtius robustus) deployed at 385 meters depth in Monterey Submarine Canyon, California (36°47.401’N; 122° 53.235’W), ROV Tiburon dive number 933, Jan. 6, 2006. Paratypes: Females, fixed in formalin or glutaraldehyde, preserved in ethanol, same locality and date as holotype, SIO-BIC A1640 (GenBank COI sequence DQ996630), same locality and date as holotype, SIO-BIC A7806 (GenBank COI sequence DQ996631); SIO-BIC A7807 and A7808 (GenBank COI sequence EU223332 and EU223337, respectively), same locality as holotype, ROV Tiburon dive number 1070, Jan. 11, 2007; dwarf males (Allotypes) SIO-BIC A7805 (no GenBank COI sequence), several in tubes of various females, same locality and date as holotype. Diagnosis and description. Holotype female, in life trunk 8 mm long, 0.6 mm wide; crown of palps contracted, curled, 1.5 mm long (Fig. 14C). Palps up to 10 mm long when extended in other specimens (Figs 14A, D). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (Figs 14B, D, E, 15A). Oviduct extends from trunk into crown for 1.75 mm (Figs 14 B-D). In life, palps reddish with two major blood vessels in each (Figs 14A, D). Tube a gelatinous loose mass around trunk (Fig. 14D). Trunk with pale yellow ring around anterior margin; ring broken mid-dorsally by oviduct (Figs 14B, D). and mid-ventrally by an oval unpigmented patch (Fig. 14C). Clear demarcation of upper and lower trunk with oviduct emerging onto outer surface of upper trunk (Fig. 14B). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (Fig. 15B). Ovisac in holotype extending laterally as two masses; simple lobes in other specimens (Fig. 14C). Roots of holotype extend from ovisac lobes (Fig. 14C); in other specimens, roots as simple lobes or extending laterally in two lobe or lobes (Fig. 14B). Males dwarfs, with spermatids and sperm, chaetalbearing segments not inflated; found in tube lumen of females (Fig. 15 C-D). Distribution. Known from Monterey Bay, California only from 349 meters depth (Table 2). It has been found in whale and cow bones. Etymology. This species is named (noun in the genitive case) in memory of Alan George Lehman, father of Ellen Lehman, in recognition of her long and continued support of the Scripps Oceanographic Collections. Remarks. Osedax lehmani n. sp. is part of Osedax Clade IV and is closest relative to Osedax packardorum n. sp., (Fig. 1), with a minimum uncorrected distance for COI of 8-9% (Table 4). This is one of the smaller intraspecific distances among Osedax species, with others being between O. frankpressi and the O. '1336_61_2' specimen from the Atlantic at ~10%; and O. randyi n. sp. and O. ‘MB16 at ~ 6-7% (Table 4). The majority of the 31 available COI sequences for Osedax lehmani n. sp. (Table 3) show less than 1% divergence, though a few sequences e.g. DQ996640 are around 2% divergent from the majority and DQ996631 shows the greatest distance, at ~3% (Table 4), so the possibility of there being a cryptic species complex for O. lehmani n. sp. needs further investigation. The most distinguishing feature of Osedax lehmani n. sp. is the yellow ring around the anterior part of the trunk. (Fig. 14 A-D). This is the only Osedax species from Monterey Bay that was not found deeper than 389 meters depth.

Osedax lonnyi n. sp. Figure 7 ���nude-palp-F��� (Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: Female, SIO-BIC A1647 (GenBank COI sequence FJ347643), fixed in glutaraldehyde-preserved in ethanol, collected from natural whale fall (Eschrichtius robustus) found at 2898 meters depth in Monterey Submarine Canyon, California (36��36.606���N; 122��26.122���W) ROV Tiburon dive number 1162, Dec. 19, 2007. Diagnosis and description. Holotype female; in life trunk 1 mm long, 0.5 mm wide (Fig. 7A), crown of four palps, 2.5 cm long (Fig. 7A), likely contracted even at this length. Tube gelatinous, thin, fitting about trunk and base of crown, 4 cm long (Fig. 7A). In life, palps with visible blood vessels and greenish tips (Fig. 7A, inset). Palps without obvious pinnules, oviduct extends slightly beyond palps (Fig. 7A inset). Trunk with pigmented ring demarking upper and lower trunk (Fig. 7B). Ovisac 2 mm by 1 mm with damaged lobate greenish roots extending outwards (Fig. 7B). Males not found. Distribution. Known from Monterey Bay, California from 2898 meters depth (Table 2). It has only been found in a whale bone fragment, adjacent to the main skeleton. Etymology. This species is named (noun in the genitive case) for Lonny Lundsten, Senior Research Technician at MBARI, for his enthusiasm and assistance on many Osedax expeditions. Remarks. Osedax lonnyi n. sp. is part of Osedax Clade II and is the well-supported sistergroup to Osedax rogersi, an Antarctic species with an uncorrected divergence of 12%. Osedax lonnyi n. sp. is remarkable for the very long palps (2.5 cm) that it shows relative to the very short trunk (1.5 mm). The species is known from only a single specimen (holotype) collected in 2007 and no further specimens were found during any subsequent sampling at the same site. This is somewhat surprising as it is one of the larger species of Clade II and the whale fall was sampled a number of times., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 468, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z"]}

Osedax westernflyer n. sp. Figure 5 ‘nude-palp-D’ (Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009) ‘Sagami 8’ (GenBank COI sequence FM998110; Pradillon et al. unpublished) Material examined. Holotype: SIO-BIC A 1645, Female (GenBank COI sequence MG262303), fixed in formalin preserved in ethanol, collected from a whale carcass (Eschrichtius robustus) deployed at 1820 meters depth in Monterey Submarine Canyon, California (36°42.496’N; 122°6.316’W), ROV Doc Ricketts dive number 12, March 13, 2009. Paratypes: All females, fixed in formalin preserved in ethanol, SIO-BIC A7802, A7803 (GenBank COI sequence MG262302, MG262304), same locality and date as holotype. Diagnosis and description. Holotype female (Fig. 5C); in life trunk 2.8 mm long, 0.5 mm wide; crown of four palps, 3.5 mm long. Tube gelatinous, thin, fitting about trunk and base of crown, 3 mm long (Fig. 5C). Oviduct among palps 2.3 mm long (Figs 5 A-C). In life, palps with white patches with no visible blood vessels, without obvious pinnules (Figs 5 A-C). Trunk with demarcation into upper and lower trunk (Fig. 5C), white pigment patch mid-ventrally on anterior tip of trunk (Figs 5B, C). Ovisac 1.5 mm by 1 mm with lobate greenish roots extending outwards (Figs 5B, C). Males not found. Distribution. Known from Monterey Bay, California from 1820 meters depth and Sagami, Bay Japan (Table 2). It has been found in cow and whale bones. Etymology. This species is named (noun in apposition) for the Research Vessel Western Flyer, which has been critical to Osedax studies. Remarks. Osedax westernflyer n. sp. is part of Osedax Clade II and is closest relative to O. knutei n. sp., with a minimum divergence of 16 % for COI, though it shows a smaller distance, 14% to O. docricketts n. sp. (Table 4). The four available COI sequences for Osedax westernflyer n. sp. from California (Table 3) are less that 1% divergent. The holotype and two paratypes all showed distinct white pigment on the palps, though these were all collected from one patch of bone and the color may have been caused by a bacterial coating. A single COI sequence on GenBank (FM998110), referred to Osedax sp. Sagami-8 (Pradillon et al., unpublished), is less than 1% divergent from the four California COI sequences (Table 4), leading us to propose that Osedax westernflyer n. sp. is also found in Japan. There is little to distinguish Osedax docricketts n. sp. from other Clade II species, apart from the white pigmentation at the anterior end of the trunk and on the pigmentation on palps.

Osedax randyi n. sp. Figure 17 ‘white collar’ (Rouse et al. 2015; Vrijenhoek et al. 2009) ‘Sagami-7’ (GenBank COI sequence FM998108-9; Pradillon et al. unpublished) Material examined. Holotype: Female, fixed in formalin preserved in ethanol, SIO-BIC A 7845 (GenBank COI sequence FJ347615), collected from a whale carcass (Balaenoptera musculus) deployed at 1018 meters depth in Monterey Submarine Canyon, California (36°46.308’N; 122° 4.981’W), ROV Tiburon dive number 1049, Oct. 25, 2006. Paratypes: Female, fixed in formalin preserved in ethanol SIO-BIC A1648, SIO-BIC A7846 (GenBank COI sequence FJ347612, FJ347613), same locality and date as holotype; Female and dwarf males (allotypes), fixed in formalin, female sectioned, males preserved in ethanol SIO-BIC A7847 (female GenBank COI sequence FJ347614), same locality and date as holotype. Diagnosis and description. Holotype female, in life trunk 5 mm long, 0.8 mm wide; crown of palps somewhat contracted, 4.5 mm long (Fig. 17A). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (Figs 17A, B, D). Oviduct extends from trunk into crown for 1-2 mm (Figs 17A, B). In life, palps reddish with pigmented white outer margins (Fig. 17A). Trunk with white ring around anterior margin; ring broken mid-dorsally by oviduct and mid-ventrally by and oval unpigmented patch (Figs 17A, B). Clear demarcation of upper and lower trunk (Fig. 17A). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (Fig. 17E). Ovisac in holotype, damaged, an ellipsoidal mass; similar shape in paratypes. Roots incomplete, simple lobes where present in all specimens. Males dwarfs, with spermatids and sperm, chaetal-bearing segments inflated (Fig. 17C), found in tube lumen of females. Distribution. Known from Monterey Bay, California from 1018 meters depth, and from Sagami Bay, Japan (Table 2). It has been found in whale and cow bones. Etymology. This species is named (noun in the genitive case) in honor of Randy Prickett, Senior ROV pilot for MBARI, who collected many bones and Osedax over the years. Remarks. Osedax randyi n. sp. is part of Osedax Clade IV and closest relative to the undescribed Osedax species from California known as O. ‘MB16’ (Fig. 1) with a minimum uncorrected COI- distance of ~6-7% (Table 4). As previously noted, this is among the smaller interspecific distances among Osedax species. Two COI sequences, referred to Osedax sp. Sagami-7 (Pradillon et al., GenBank unpublished, Table 3) are much less than 1% divergent from the six California Osedax randyi n. sp. sequences (example distance in Table 4). Thus, it is reasonable to propose that Osedax randyi n. sp. is also found in Japan. The most distinguishing features of Osedax randyi n. sp. are the white ring around the anterior part of the trunk and white pigment on the palps of females (Fig. 17A, B), though a similar whitish ring is found on females of O. bryani n. sp. (Fig. 18A, B) and O. frankpressi (Rouse et al. 2004), with the latter species also showing white pigmentation on the palps. DNA sequencing is most likely the best way to distinguish Osedax randyi n. sp. from these species.

Osedax talkovici n. sp. Figure 3 ���yellow patch��� (Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A 7821 female (GenBank COI sequence MG262313), fixed in glutaraldehyde-preserved in ethanol, collected from turkey (Meleagris gallopavo) bone deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W) ROV Doc Ricketts dive number 233, June 2, 2011. Paratypes: Females, all fixed in formalin-preserved in ethanol, SIO-BIC A1649 (GenBank COI sequence FJ347621), SIO-BIC A7822, A7823, A7824, A7825, A7826 (GenBank COI sequences FJ347620, 431196, 431197, 431201, 431205), whale (Eschrichtius robustus) deployed at 633 meters depth in Monterey Submarine Canyon, California (36��48.178���N; 121��59.677���W) ROV Tiburon dive number 1160, Dec. 18, 2007; Female SIO-BIC A7829 (GenBank COI sequence MG262311) and dwarf males (allotypes) SIO-BIC A7830 whale fall at 633 meters depth in Monterey Submarine Canyon, California (36��48.178���N; 121��59.677���W) ROV Doc Ricketts dive number 205, October 26, 2010; Female SIO-BIC A7827 (GenBank COI sequence MG262310) and dwarf males (allotypes) SIO-BIC A7828, whale (Eschrichtius robustus) deployed at 633 meters depth in Monterey Submarine Canyon, California (36��48.178���N; 121��59.677���W) ROV Doc Ricketts dive number 233, June 2, 2011; Female SIO-BIC A7831 (GenBank COI sequence MG262312) on green turtle bone (Chelonia mydas), deployed at same locality and date as holotype. Diagnosis and description. Holotype female (Figs 3B, C); in life trunk 1.8 mm long, 0.3 mm wide; crown of palps contracted, 1.5 mm long. Tube gelatinous. Oviduct 0.75 mm (Figs 3C, D). In life, palps reddish with blood (Figs 3A���E). Pinnules of all four palps oriented dorsally (Figs 3A, D, E). Trunk with bright yellow patch on midventral anterior edge (Figs 3A, B, D). Clear demarcation into upper and lower trunk (Fig. 3D). Ovisac spheroidal in holotype (Figs 3B, C), extending laterally, or as simple lobes in other specimens. Roots of holotype longer that the remaining body, extending posteriorly as a single lobate mass (Figs 3B, C). In other specimens (Figs 3A, D, E), roots as simple lobes or extending laterally in two lobes. Males dwarfs, resembling trochophore larvae, with spermatids and sperm (Figs 3F, G). Distribution. Known from Monterey Bay, California from 633 to 1018 meters depth (Table 2). It has been found in whale, elephant seal, cow, turkey, turtle and teleost bones. Etymology. This species is named (noun in the genitive case) for Mark Talkovic, Senior ROV pilot for MBARI, who collected many bones containing Osedax over the years. Remarks. Osedax talkovici n. sp. is part of Osedax Clade I and forms a well-supported clade with the undescribed species O. ���mediterranea��� and a clade comprised of Osedax sigridae n. sp. and the undescribed species O. Sagami-5 (Fig. 1). It has a minimum uncorrected distance, for COI, of at least 16% from each of these (Table 4). The 15 available COI sequences for Osedax talkovici n. sp. (Table 3) show less that 1% sequence divergence. The most distinguishing feature of Osedax talkovici n. sp. is the bright yellow patch ventrally on the anterior trunk (Fig. 3)., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on pages 460-462, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Rouse, G. W., Goffredi, S. K., Johnson, S. B. & Vrijenhoek, R. C. (2011) Not whale-fall specialists, Osedax worms also consume fishbones. Biology Letters, 7, 736 - 739. https: // doi. org / 10.1098 / rsbl. 2011.0202","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z"]}

Osedax tiburon n. sp. Figure 4A, B, C ‘nude-palp-A’ (Jones et al. 2008; Rouse et al. 2011; Rouse et al. 2009; Rouse et al. 2015; Vrijenhoek et al. 2009) Material examined. Holotype: SIO-BIC A7817 female (GenBank COI sequence FJ347622), fixed in formalin preserved in ethanol, collected from whale carcass (Eschrichtius robustus) deployed at 1820 meters depth in Monterey Submarine Canyon, California (36°42.496’N; 122°6.316’W) ROV Tiburon dive number 1163, Dec. 20, 2007. Paratypes: All females, fixed in formalin preserved in ethanol, SIO-BIC A1642 (GenBank COI sequence EU223359), SIO-BIC A7820 (GenBank COI sequence EU223356, EU223357), same locality as holotype, ROV Tiburon dive number 1119, Aug. 16, 2007; SIO-BIC A7818, A7819 (GenBank COI sequences FJ347623, FJ347624, respectively) same locality and date as holotype. Diagnosis and description. Holotype female (Fig. 4B); preserved trunk 0.75 mm long, 0.2 mm wide; crown of four palps, 3 mm long. Tube gelatinous 6 mm long (Fig. 4A). Oviduct not discerned. In life, palps reddish with blood (Fig. 4A). Four palps without obvious pinnules (Fig. 4B, C). Trunk with no obvious demarcation into upper and lower trunk (Fig. 4B, C). Ovisac mostly missing in holotype, some traces of roots present (Fig. 4A). In paratype ovisac and root complex extends laterally on either side of trunk and posteriorly (Fig. 4C). Males not seen. Distribution. Known from Monterey Bay, California from 1820 meters depth (Table 2). Etymology. This species is named (noun in apposition) for the ROV Tiburon, which was used to collect the first Osedax specimens. Remarks. Osedax tiburon n. sp. is part of Osedax Clade II and is closest relative (though with low support) to Osedax nordenskjoeldi from which it shows a divergence of 15%, though it shows a smaller pairwise distance (14%) to O. ryderi n. sp. (Table 4). The seven available COI sequences for Osedax tiburon n. sp. (Table 3) show less than 1% sequence divergence. It has been found in whale bones and experimentally deployed cow bones. There is little to distinguish Osedax tiburon n. sp. from other 'nude palp' species in Clade II, which share a similar morphology. The species status of Osedax tiburon n. sp. is supported largely on its COI divergence from other Osedax.

Osedax docricketts n. sp. Figure 4E ���nude-palp-C��� (Higgs et al. 2014; Rouse et al. 2011; Rouse et al. 2015; Vrijenhoek et al. 2009) ���Sagami-6��� (GenBank COI sequence series FM998088-FM998107; Pradillon et al. unpublished) Holotype: SIO-BIC A 1644, Female (GenBank COI sequence FJ347626), fixed in formalin preserved in ethanol, collected from whale fall deployed at 1018 meters depth in Monterey Submarine Canyon, California (36��46.308���N; 122�� 4.981���W), ROV Tiburon dive number 1049, Oct. 25, 2006. Diagnosis and description. Holotype incomplete female; preserved trunk 0. 5 mm long, 0.2 mm wide; crown of four palps, 2 mm long (Fig. 4E). Tube gelatinous cylinder around trunk and base of palps. Oviduct not discerned. In life, palps white with some reddish patches. Palps without obvious pinnules (Fig. 4E). Trunk with no obvious demarcation into upper and lower trunk, or any pigmentation. Ovisac mostly missing in holotype, some traces of roots present. Males not seen. Distribution. Known from Monterey Bay, California from 1820 meters depth, and Sagami Bay, Japan (Table 2). It has been found living on cow and whale bones. Etymology. This species is named (noun in apposition) for the ROV Doc Ricketts, which was used to collect many Osedax specimens. Remarks. Osedax docricketts n. sp. is part of Osedax Clade II, which all show apinnulate palps, and is closest relative to the clade comprised of Osedax westernflye r n. sp. and O. knutei n. sp. (minimum divergence of 15%) though it shows smaller distance (14%) to other taxa such as O. lonnyi n. sp. (Table 4). The four available COI sequences for Osedax docricketts n. sp. from California (Table 3) show less that 1% sequence divergence. The only retained specimen is the holotype as the others were destroyed for sequencing and the holotype itself is only a partial specimen. Twenty additional COI sequences were referred to Osedax sp. Sagami-6 (Pradillon et al., GenBank, unpublished). Together, the four California and 11 of the Japanese sequences (FM998088, FM998089, FM998091, FM998093, FM998094, FM998098, FM998099, FM998100, FM998101, FM998103, FM998107) encompass less than 1% sequence divergence; so, we propose that Osedax docricketts n. sp. is also found in Japan. The nine remaining Japanese specimens with>2.4% uncorrected pairwise distance from the 11 Japanese sequences (GenBank sequences: FM998090, FM998092, FM998095, FM998096, FM998097, FM998102, FM998104, FM998105, FM998106) require further investigation before their assignment to Osedax docricketts n. sp. can be justified. See Table 4 for comparison of California Osedax docricketts n. sp. versus Japanese Sagami-6 representative sequences. There is little to distinguish Osedax docricketts n. sp. from other Clade II species., Published as part of Rouse, Greg W., Goffredi, Shana K., Johnson, Shannon B. & Vrijenhoek, Robert C., 2018, An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California, pp. 451-489 in Zootaxa 4377 (4) on page 464, DOI: 10.11646/zootaxa.4377.4.1, http://zenodo.org/record/1165874, {"references":["Higgs, N. D., Glover, A. G., Dahlgren, T. G., Smith, C. R., Fujiwara, Y., Pradillon, F., Johnson, S. B., Vrijenhoek, R. C. & Little, C. T. S. (2014) The morphological diversity of Osedax worm borings (Annelida: Siboglinidae). Journal of the Marine Biological Association of the UK, 94, 1429 - 1439. https: // doi. org / 10.1017 / S 0025315414000770","Rouse, G. W., Goffredi, S. K., Johnson, S. B. & Vrijenhoek, R. C. (2011) Not whale-fall specialists, Osedax worms also consume fishbones. Biology Letters, 7, 736 - 739. https: // doi. org / 10.1098 / rsbl. 2011.0202","Rouse, G. W., Wilson, N. G., Worsaae, K. & Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032","Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. & Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z"]}