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We report on a detailed study of longitudinal strength in the nucleon resonance region, presenting new results from inclusive electron-proton cross sections measured at Jefferson Lab Hall C in the four-momentum transfer range 0.2

Ophryotrocha bohnorum sp. nov. urn:lsid:zoobank.org:act: 181A86F6-7F8D-401C-B252-21FF39F6C5C0 Fig. 9 Etymology Ophryotrocha bohnorum sp. nov. is named for Jeffrey and Brenda Bohn and their family in recognition of their enduring support of deep-sea research. Material examined Holotype TONGA • 1 spec. (fixed in ethanol and a midbody piece used for DNA extraction); Lau Back-Arc Basin, Southern Valu Fa Ridge, Hine Hina Vent Field, active hydrothermal vents; 22.539° S, 176.718° W; depth 1845–1906 m; 22–23 May 2005; Greg Rouse, Fredrik Pleijel and Robert Vrijenhoek leg.; collecting event: ROV Jason II dive 146; GenBank: OP311742 (COI), OP304895 (16S), OP311649 (H3); SIO-BIC A14092. Paratypes TONGA • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14094 • 4 specs (fixed in formalin); same collection data as for holotype; SIO-BIC A14095 • 1 spec. (fixed in ethanol and a midbody piece used for DNA extraction); same collection data as for holotype; GenBank: OP311744 (COI); SIO-BIC A14165 • 1 spec. (fixed in ethanol and a midbody piece used for DNA extraction); same collection data as for holotype; GenBank: OP311743 (COI); SIO-BIC A14166 • 1 spec. (fixed in ethanol); same locality as for holotype; 22.532° S, 176.719° W; depth 1818–1907 m; 21–22 May 2005; Greg Rouse, Fredrik Pleijel and Robert Vrijenhoek leg.; collecting event: ROV Jason II dive 145; SIO-BIC A14088 • 1 spec. (fixed in ethanol); same collection data as for preceding; SIO-BIC A14089 • 1 spec. (fixed in ethanol); same collection data as for preceding; SIO-BIC A14090 • 3 specs (fixed in formalin); same collection data as for preceding; SIO-BIC A14091 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14093. Description In life, white, yellow gut, with white eggs mid-body (Fig. 9A), opaque white after preservation. Body 12 mm long with 60+ segments, tapering slightly along body.Prostomium rounded, wider than long,with a slightly rise posteriorly in the middle. Paired antennae long cirriform, inserted dorsally, tapering distally. Paired palps cirriform, slightly shorter than antennae, inserted dorsal-laterally (Fig. 9A). Peristomium two equal rings, similar in length to first chaetiger, slightly longer than following chaetigers, with two brown spots located dorsal-laterally between the two rings (Fig. 9A–B). Maxillae P-type, maxillary carriers comb-like, with large main fang, 8 pairs of free denticles. Posterior 4 denticles (D1–D4) heavily sclerotized, with large main fang and sharp teeth; anterior 4 denticles (D5–D8) translucent, with a small main fang and tiny teeth, D5–D6 overlap with D3–D4 (Fig. 9C). Mandibles transparent, rod-like shafts, cutting plates triangular, with blunt teeth on the anterior edge (Fig. 9D). Parapodia uniramous, acicular lobe rounded with a small tip in the middle of the distal margin, dorsal cirri short, rounded lobes, ventral cirri long extending from distal margin of acicular lobe (Fig. 9E). Supra-acicular chaetae simple, distally serrated, with a small hook on the tip (Fig. 9F). Sub-acicular chaetae all compound, shafts bifid on the top, serrated blades with a small hook similar as supra-acicular chaetae on the tip (Fig. 9G). Pygidium with two long digitiform cirri (Fig. 9A). Distribution Only known from vents at the Lau Back-Arc Basin, southwest Pacific Ocean at depths of 1845–1907 m. Remarks The phylogenetic results (Fig. 1) show that Ophryotrocha bohnorum sp. nov. is most closely related to an undescribed species complex of Ophryotrocha (O. Seep4) from eastern Pacific methane seeps (Thornhill et al. 2012). There is no morphological information available for these specimens. A clade comprising other eastern Pacific species, Ophryotrocha globopalpata from hydrothermal vents, O. flabella from a whale fall and O. Seep3 (Thornhill et al. 2012), is then the well supported sister group to this clade. Based on this topology Ophryotrocha bohnorum appears to have independently colonized hydrothermal vents from O. globopalpata (Fig. 1). Ophryotrocha bohnorum has hooked tips of the supra- and sub-acicular chaetae and red-brown spots located dorsal-laterally between two peristomial segments, features not seen in other species from hydrothermal vents. Only four species of Ophryotrocha, O. atlantica Hilbig & Blake, 1991, O. mediterranea Martin Abello & Cartes, 1991, O. pachysoma Hilbig & Blake, 1991, and O. socialis Ockelmann & Åkesson, 1990, have been described with chaetae with hooked tips. Ophryotrocha bohnorum can be easily distinguished from these species by its transparent mandibles with serrated anterior edge., Published as part of Zhang, Dongsheng, Zhou, Yadong, Yen, Nicole, Hiley, Avery S. & Rouse, Greg W., 2023, Ophryotrocha (Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species, pp. 167-194 in European Journal of Taxonomy 864 on pages 187-189, DOI: 10.5852/ejt.2023.864.2101, http://zenodo.org/record/7867628, {"references":["Thornhill D. J., Struck T. H., Ebbe B., Lee R. W., Mendoza G. F., Levin L. A. & Halanych K. M. 2012. Adaptive radiation in extremophilic Dorvilleidae (Annelida): diversification of a single colonizer or multiple independent lineages? Ecology and Evolution 2: 1958 - 1970. https: // doi. org / 10.1002 / ece 3.314"]}

Ophryotrocha kailae sp. nov. urn:lsid:zoobank.org:act: FC8AD84C-5C47-49D2-BEC4-55FC04F3DFBC Fig. 5 Etymology Named for Kaila Pearson, an expert on another group of vent and seep-associated polychaetes, phyllodocids belonging to Galapagomystides Blake, 1985. Material examined Holotype EAST PACIFIC OCEAN • 1 spec. (anterior fixed in formalin, posterior fixed in ethanol and used for DNA extraction); Southern East Pacific Rise, active hydrothermal vents; 31.151° S, 111.932° W; depth 2237 m; 29 Mar. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4094; GenBank: OP311745 (COI); SIO-BIC A14100. Paratypes EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, Saguaro Vent Field, active hydrothermal vents; 31.865° S, 112.044° W; depth 2235 m; 28 Mar. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4093; GenBank: OP311748 (COI); SIO-BIC A14099 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, German Flats Vent Field, active hydrothermal vents; 37.793° S, 110.916° W; depth 2216 m; 22 Mar. 2005; Greg Rouse, Nerida Wilson, Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4088; GenBank: OP311746 (COI), OP304894 (16S), OP311650 (H3); SIO-BIC A14101 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311747 (COI); SIO-BIC A14102 • 1 spec. (fixed in formalin); same collection data as for holotype; SIO-BIC A14103 • 1 spec. (anterior fixed in formalin, posterior fixed in ethanol); same collection data as for holotype; SIO-BIC A14104. Description In life, translucent with light yellow gut and white eggs mid-body, body opaque white after preservation. Body length ~ 4.5 mm with more than 30 segments, similar width through the body, slightly tapering posteriorly (Fig. 5A). Eyes not visible. Prostomium rounded, wider than long, with paired short digitiform antennae inserted dorsally, paired digitiform palps inserted ventral-laterally, similar in length with antennae. Peristomium two rings, similar in length to following segments (Fig. 5A). Mandibles heavily sclerotized, shafts rod-like, cutting plates curved, with single blunt peak, lateral wings weakly sclerotized (Fig. 5B). Maxillae P-type, forceps comb-like, with large main large fang. Two rows of 7 free denticles, posterior most free denticles (D1) comb-like, D2–D7 shovel-shaped (Fig. 5C). Parapodia uniramous, acicular lobe with rounded distal margin, dorsal cirri subtriangular, barely extending beyond distal margin of acicular lobe, ventral cirri stubby (Fig. 5C). Supra-acicular chaetae simple, distally serrated, tapering into a fang (Fig. 5D–E). Sub-acicular chaetae compound, with serrated blades (Fig. 5D, F). Sub-acicular chaetal lobe with one simple chaeta (Fig. 5D, G). Pygidium with two anal cirri inserted laterally (Fig. 5A inset). Distribution Known only from vents at 2216–2237 m along the southern East Pacific Rise (Pacific Antarctic Ridge). Remarks While the DNA data suggests Ophryotrocha kailae sp. nov. is most closely related to O. pruittae sp. nov. (Fig. 1), morphologically it resembles Ophryotrocha akessoni in having similar mandibles with curving cutting plates, which are otherwise not seen in the vent clade that also includes O. charlottae sp. nov., O. jiaolongi, O. marinae sp. nov. and O. pruittae. Ophryotrocha kailae differs from O. akessoni in the form of its head appendages and possibly pygidial cirri (Table 3).

Ophryotrocha pruittae sp. nov. urn:lsid:zoobank.org:act: BE33115E-1B4C-405A-A425-AF518EE5E093 Fig. 8 Ophryotrocha cf. akessoni sp. 2 – Goffredi et al. 2017: supplemental, table 1. Etymology Named for Jessica Pruitt, an aficionada and expert on deep-sea Ophryotrocha. Material examined Holotype MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Gulf of California, Alarcón Rise, active hydrothermal vents; 23.377° N, 108.531° W; depth 2309 m; 22 Apr. 2015; Greg Rouse leg.; collecting event: ROV Doc Ricketts dive 754; GenBank: OP311761 (COI); ICML-EMU-13288, (ex SIO-BIC A13689). Paratypes MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: KY701728 (16S), OP311652 (H3); SIO-BIC A6322 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14123 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14124 • 1 spec. (fixed in ethanol and most tissue used for DNA extraction); same collection data as for holotype; SIO-BIC A14125 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14126 • at least 4 specs (1 fixed in formalin, 3 individuals and additional fragments fixed in ethanol); same collection data as for holotype; SIO-BIC A14127 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14128 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14129 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14130 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14131 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14132. Description In life, golden color (Fig. 8A), opaque white after preservation. Body 10.5 mm long, 50+ segments of similar width through the body. Eyes not visible. Prostomium rounded, wider than long, with paired digitiform antennae inserted dorsally, paired digitiform palps inserted ventral-laterally, similar in length with antennae. Peristomium two equal rings, similar size to the following segments (Fig. 8A). Maxillae P-type, forceps comb-like, with large main large fang, fused together basally. Four rows of seven free denticles, the posterior most free denticles (D1) comb-like, similar to the forceps, other free denticles shovel-shaped with fine teeth, D2–D4 smaller than D5–D7 (Fig. 8B). Mandibles heavily sclerotized, shafts rod-like, cutting plates L-shape, anterior edge flat with pointed lateral peaks weakly sclerotized, lateral wings weakly sclerotized (Fig. 8C). Parapodia uniramous, acicular lobe triangular with a blunt point distally, dorsal cirri cirriform, long, extend beyond distal margin of acicular lobe, ventral cirri short and stubby (Fig. 8D). Supra-acicular chaetae simple distally serrated, tapering into a large main fang, 2–5 per fascicle (Fig. 8D–E). Five to eight compound sub-acicular chaetae, with serrated blades (Fig. 8D, F). Sub-acicular chaetal lobe holds one or two simple chaetae (Fig. 8D, G). Pygidium with two short conical cirri inserted laterally (Fig. 8A). Distribution Only known from the Alarcón Rise vents in the southern Gulf of California at 2309 m depth. Remarks Ophryotrocha pruittae sp. nov. was initially reported in Goffredi et al. (2017) as Ophryotrocha cf. akessoni sp. 2 with a partial DNA sequence for mitochondrial 16S rRNA lodged on GenBank (KY701727). Ophryotrocha pruittae has a rounded prostomium, two equal segments of peristomium, digitiform antennae and palps and P-type maxillae, which are all features found in the vent-clade of Clade B. It differs from these species by having the mandibles with the L-shape cutting plates and two conical anal cirri, while O. marinae sp. nov. has triangular cutting plates and two digitiform anal cirri, O. akessoni / O. cf. akessoni has curved cutting plates, O. charlottae sp. nov. has subtriangular cutting plates and two lateral cirri and one median anal cirrus, O. kailae sp. nov. has curved cutting plates and two digitiform anal cirri (Table 3). Ophryotrocha pruittae showed four rows of maxillae. As discussed above for O. charlottae and O. marinae it is possible that the outermost pair of rows represent molted jaws as has been observed in other Ophryotrocha by Paxton (2004)., Published as part of Zhang, Dongsheng, Zhou, Yadong, Yen, Nicole, Hiley, Avery S. & Rouse, Greg W., 2023, Ophryotrocha (Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species, pp. 167-194 in European Journal of Taxonomy 864 on pages 185-187, DOI: 10.5852/ejt.2023.864.2101, http://zenodo.org/record/7867628, {"references":["Goffredi S. K., Johnson S., Tunnicliffe V., Caress D., Clague D., Escobar E., Lundsten L., Paduan J. B., Rouse G., Salcedo D. L., Soto L. A., Spelz-Madero R., Zierenberg R. & Vrijenhoek R. 2017. Hydrothermal vent fields discovered in the southern Gulf of California clarify role of habitat in augmenting regional diversity. Proceedings. Biological Sciences / The Royal Society 284: 20170817. https: // doi. org / 10.1098 / rspb. 2017.0817","Paxton H. 2004. Jaw growth and replacement in Ophryotrocha labronica (Polychaeta, Dorvilleidae). Zoomorphology 123: 147 - 154. https: // doi. org / 10.1007 / s 00435 - 004 - 0097 - 4"]}

Ophryotrocha charlottae sp. nov. urn:lsid:zoobank.org:act: 2AC55385-EA0C-41E5-8EAF-4D6F32818D94 Fig. 4 Etymology Named in honor of Charlotte Seid, collection manager of the Benthic Invertebrate Collection at Scripps Oceanography, for her dedication to facilitating biodiversity research. Material examined Holotype EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, northwest of Easter Island, active hydrothermal vents; 23.823° S, 115.456° W; depth 2649 m; 2 Apr. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4097; GenBank: OP311739 (COI), OP304893 (16S), OP311648 (H3); SIO-BIC A14096. Paratypes EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and midsection tissue used for DNA extraction); same collection data as for holotype; GenBank: OP311741 (COI); SIO-BIC A14163 • 1 spec. (fixed in ethanol and midsection tissue used for DNA extraction); same collection data as for holotype; GenBank: OP311740 (COI); SIO-BIC A14164. Other material EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and entirely used for DNA extraction); same collection data as for holotype; GenBank: OP311738 (COI); SIO-BIC A14097 • 7 or more specs (7 fixed in glutaraldehyde, additional material fixed in formalin and entirely used for slides of parapodia, additional tissue fixed in ethanol); same collection data as for holotype; SIO-BIC A14098 • 1 spec. (fixed in ethanol and midsection tissue used for DNA extraction); same collection data as for holotype; SIO-BIC A14187 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14188. Description In life light brown (Fig. 4A), opaque white after preservation. Body ~ 3 mm long, with more than 35 segments of similar width, slightly tapering posteriorly (Fig. 4A). Prostomium rounded, wider than long, with paired digitiform antennae inserted dorsally, paired digitiform palps inserted ventral-laterally, similar in length with antennae. Peristomium two equal rings, each similar in size to the following segments (Fig. 4A). Eyes not visible. Maxillae P-type, forceps comb-like, with large main fang, fused together basally. Four rows of seven free denticles, the posterior-most free denticles (D1) comb-like, like the forceps, other free denticles (D2–D7) shovel-shaped with fine teeth (Fig. 4B). Mandibles heavily sclerotized, shafts rod-like, cutting plates sub-triangular, lateral wings weakly sclerotized (Fig. 4C). Parapodia uniramous, acicular lobe with rounded distal margin, dorsal cirri enlarged fusiform, similar in size with acicular lobe, reaching distal margin of acicular lobe, ventral cirri short and stubby (Fig. 4D). Supra-acicular chaetae simple, distally serrated, tapering into a main fang (Fig. 4D–E). Sub-acicular chaetae compound, hooked with serrated blades (Fig. 4D, F). Sub-acicular chaetal lobe also with one or two simple chaetae (Fig. 4D, G). Pygidium with two digitiform cirri inserted laterally, a small median papilla posteriorly placed (Fig. 4A). Distribution Known only from vents at 2649 m depth near Easter Island (Rapa Nui) at the southern end of the East Pacific Rise (Pacific Antarctic Ridge). Remarks Ophryotrocha charlottae sp. nov. resembles O. cf. akessoni, which also occurs on the Southeast Pacific Ridge. They are similar in the shape of the prostomium, peristomium, antennae, palps, jaws, and chaetae. Ophryotrocha charlottae has distinctive parapodia with enlarged fusiform dorsal cirri, which easily distinguishes it from O. akessoni / O. cf. akessoni, O. jiaolongi and O. marinae sp. nov. Also, O. charlottae has two long digitiform anal cirri and a media papilla while O. cf. akessoni only has two short nub-like anal cirri (Fig. 3A). Ophryotrocha kailae sp. nov. is another species from southern end of the East Pacific Rise (Pacific Antarctic Ridge). It differs from O. charlottae in the form of antennae, palps, mandibles, dorsal cirri and pygidium (Fig. 5). Ophryotrocha charlottae can also be easily distinguished from other species of the “ akessoni ” clade, based on mandibles and parapodia (Table 3). The four rows of maxillae found in O. charlottae differ from that in close relatives such as O. akessoni, O. cf. akessoni (Fig. 3C) and O. jiaolongi that show only two rows (Blake 1985; Zhang et al. 2017), but four rows were also observed in O. marinae and O. pruittae sp. nov. (see below). It is possible that the outermost pair of rows represent molted jaws as has been observed in other Ophryotrocha by Paxton (2004), so this should not be interpreted as diagnostic without further study.

Ophryotrocha marinae sp. nov. urn:lsid:zoobank.org:act: 5B66661D-DDF9-425D-860F-412872058C27 Figs 6–7 Ophryotrocha cf. akessoni sp. 1 – Goffredi et al. 2017: supplemental, table 1. Ophryotrocha cf. akessoni – Salcedo et al. 2019: 6, table 1. Etymology Named in honor of Marina McCowin for her dedication in the study of the fauna associated with seeps and vents. Marina has studied Siboglinidae and Ophryotrocha marinae sp. nov. was notable for being associated in large numbers with the tubes of Oasisia alvinae Jones, 1985 (Fig. 6A). Material examined Holotype MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Gulf of California, Pescadero Basin, active hydrothermal vents; 23.960° N, 108.863° W; depth 3676–3756 m; 18 Apr. 2015; Greg Rouse leg.; collecting event: ROV Doc Ricketts dive 750 (specimens associated with tubes of Oasisia alvinae); GenBank: OP311750 (COI);, ICML-EMU-13289, (ex SIO-BIC A14109). Paratypes MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311749 (COI), KY701727 (16S), OP311651 (H3); SIO-BIC A6308 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311755 (COI); SIO-BIC A14108 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311751 (COI); SIO-BIC A14110 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14111 • ca 35 specs (10 fixed in formalin, ca 25 fixed in ethanol); same collection data as for holotype; SIO-BIC A14112. • 11 specs (5 fixed in paraformaldehyde / glutaraldehyde, 6 fixed in ethanol); Gulf of California, Pescadero Basin, Auka Vent Field, region of P Vent and Z Vent; 23.96° N, 108.86° W; depth 3648–3671 m; 3 Nov. 2021; Greg Rouse leg.; collecting event: ROV SuBastian dive 475; GenBank: OP561817 (COI); SIO-BIC A14031. Other material MEXICO • 7 specs (3 fixed in formalin, 4 fixed in ethanol); Gulf of California, Pescadero Basin, Auka Vent Field, Matterhorn area, active hydrothermal vents; 23.95° N, 108.86° W; depth 3650 m; 14 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 193, sample S0193-S4 (suction sampler chamber 4, specimens associated with tubes of Oasisia alvinae); SIO-BIC A9974 • 7 specs (2 fixed in formalin, 5 fixed in ethanol); same collection data as for preceding; SIO-BIC A9975 • 4 specs (fixed in ethanol); same locality as for preceding; 23.95369° N, 108.86231° W; depth 3668 m; 29 Oct. 2021; Greg Rouse leg.; collecting event: ROV SuBastia n dive 470, sample S0470-S3 (suction sampler chamber 3, among tube worms); SIO-BIC A13987 • 2 specs (1 anterior fragment fixed in ethanol, 1 anterior fixed in formalin and posterior fixed in ethanol and used for DNA extraction); Gulf of California, Pescadero Basin, Auka Vent Field, Z Mound area, active hydrothermal vents; 23.95616° N, 108.86191° W; depth 3688 m; 17 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 196, sample S0196-PC1 (push core 1 at microbial mat with venting); GenBank: OP311752 (COI); SIO-BIC A9987 • at least 15 specs (3 fixed in formalin, 12 fixed in RNAlater, additional material fixed in ethanol); same locality as for preceding; 23.9561° N, 108.8619° W; depth 3688 m; 21 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 200, samples S0200-S1 and S0200-S2 (suction sampler chambers 1 and 2, microbial mat with venting); SIO-BIC A10029 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311753 (COI); SIO-BIC A14114 • 4 specs (3 fixed in ethanol, 1 anterior fixed in formalin and posterior fixed in ethanol); Gulf of California, Pescadero Basin, Auka Vent Field, oily mat site southeast of Z Vent; 23.95° N, 108.86° W; depth 3650–3662 m; 5 Nov. 2021; Greg Rouse leg.; collecting event: ROV SuBastian dive 477; SIO-BIC A14045 • 7 specs (6 fixed in paraformaldehyde, 1 fixed in ethanol); Gulf of California, Pescadero Basin, midway between Auka and JaichMaa ‘ja’ag Vent Fields; 23.95369° N, 108.86231° W; depth 3663–3687 m; 1 Nov. 2021; Greg Rouse leg.; collecting event: ROV SuBastian dive 473; SIO-BIC A14007 • 11 specs (2 fixed in formalin, 9 fixed in ethanol); Gulf of California, Pescadero Basin, JaichMaa ‘ja’ag Vent Field, Cavern Tay Ujaa (Big Cave), active hydrothermal vents; 23.94157° N, 108.85570° W; depth 3675 m; 18 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 197, sample S0197-S6 (suction sampler chamber 6, specimens associated with small tubes of Oasisia alvinae); SIO-BIC A10002 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311754 (COI); SIO-BIC A14113 • ca 50 specs (fixed in ethanol); Gulf of California, Guaymas Basin, sedimented hydrothermal vents; 27.016° N, 111.410° W; depth 2012 m; 13 Nov. 2009; Anna-Louise Reysenbach leg.; collecting event: HOV Alvin dive 4558; SIO-BIC A14115 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311756 (COI); SIO-BIC A14116 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311757 (COI); SIO-BIC A14117 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; SIO-BIC A14118 • ca 15 specs (fixed in ethanol); Gulf of California, Guaymas Basin, sedimented hydrothermal vents; 27.015° N, 111.410° W; depth 2010 m; 15 Nov. 2009; Anna-Louise Reysenbach leg.; collecting event: HOV Alvin dive 4560; SIO-BIC A14119 • 1 speci. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311758 (COI); SIO-BIC A14120 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311759 (COI); SIO-BIC A14121 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311760 (COI); SIO-BIC A14122. Description In life, golden color with white eggs mid-body (Fig. 7A–B), opaque white after preservation. Body length 10.5 mm, with 50+ segments, slightly dorso-ventrally compressed, widest anteriorly, gradually tapering posteriorly (Fig. 7A–B). Prostomium rounded, wider than long, posterior medial area slightly raised, with paired digitiform antennae, tapering distally, inserted dorsally, paired digitiform palps similar in length with antennae, inserted ventral-laterally (Fig. 7A). Peristomium with two rings, subequal in length to the following segments (Fig. 7A). Eyes not visible. Mandibles heavily sclerotized, with rod-like shafts, sub-triangular shape cutting plates, with single blunt peak anteriorly, lateral wings weakly sclerotized (Fig. 7C). Maxillae P-type, forceps comb-like, with large main fang. Four rows of free denticles, the posterior-most free denticles (D1) comb-like, like the forceps, other free denticles shovel-shaped with fine teeth, D2–D4 smaller than D5–D7 (Fig. 7D). Parapodia uniramous, acicular lobe with rounded distal margin, dorsal cirri long digitiform, extending beyond distal margin of acicular lobe, ventral cirri short and stubby (Fig. 7E). Supra-acicular chaetae simple, distally serrated, tapering into a main fang (Fig. 7F). Sub-acicular chaetae compound, with serrated blades (Fig. 7G). Sub-acicular chaetal lobe usually with one simple chaeta (Fig. 7H). Pygidium with two digitiform cirri inserted laterally (Fig. 7A). Distribution Known from Gulf of California hydrothermal vents of the Pescadero Basin at over 3500 m and the Guaymas Basin sedimented vents at ~ 2000 m. Found in huge numbers on tubes of Oasisia (Fig. 6A), or on microbial mats near active flow (Fig. 6B). Remarks This species was initially reported in Goffredi et al. (2017) as Ophryotrocha cf. akessoni sp. 1 with a partial DNA sequence for mitochondrial 16S rRNA lodged on GenBank (KY701727). Ophryotrocha marinae sp. nov. most closely resembles Ophryotrocha jiaolongi described from hydrothermal vents of the Indian Ocean, including sharing distinctive mandibles that distinguish these two taxa from all other Ophryotrocha. Morphologically, O. marinae differs from O. jiaolongi based on body color, the form of antennae and palps, and in lacking a median pygidial papilla (Table 3). In life, O. marinae is golden while O. jiaolongi is white and translucent. Ophryotrocha marinae also has antennae and palps that are distally tapering and longer than the length of prostomium, while in O. jiaolongi antennae and palps are shorter than the length of prostomium. The minimum COI uncorrected distance obtained between O. marinae and O. jiaolongi specimens was relatively small at 3.73% (Table 2). Other relatively small distances that are currently known are up to 4.8% between O. notoglandulata Pfannenstiel, 1972 and O. japonicus Paxton and Åkesson, 2010 (both Japanese taxa) and 6.1% between O. flabella and O. globopalpata (both from deep waters of the eastern Pacific). Given the morphological differences between O. marinae and O. jiaolongi, the reciprocal monophyly based on numerous COI sequences and vast geographic separation, we regard them here as separate species. One notable difference between O. jiaolongi and O. marinae was that the latter species showed four rows of maxillae compared to the two rows in O. jiaolongi found by Zhang et al. (2017). As discussed above for O. charlottae sp. nov. it is possible that the outermost pair of rows represent molted jaws as has been observed in other Ophryotrocha by Paxton (2004)., Published as part of Zhang, Dongsheng, Zhou, Yadong, Yen, Nicole, Hiley, Avery S. & Rouse, Greg W., 2023, Ophryotrocha (Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species, pp. 167-194 in European Journal of Taxonomy 864 on pages 181-185, DOI: 10.5852/ejt.2023.864.2101, http://zenodo.org/record/7867628, {"references":["Goffredi S. K., Johnson S., Tunnicliffe V., Caress D., Clague D., Escobar E., Lundsten L., Paduan J. B., Rouse G., Salcedo D. L., Soto L. A., Spelz-Madero R., Zierenberg R. & Vrijenhoek R. 2017. Hydrothermal vent fields discovered in the southern Gulf of California clarify role of habitat in augmenting regional diversity. Proceedings. Biological Sciences / The Royal Society 284: 20170817. https: // doi. org / 10.1098 / rspb. 2017.0817","Salcedo D. L., Soto L. A. & Paduan J. B. 2019. Trophic structure of the macrofauna associated to deepvents of the southern Gulf of California: Pescadero Basin and Pescadero Transform Fault. PloS One 14: e 0224698. https: // doi. org / 10.1371 / journal. pone. 0224698","Zhang D., Zhou Y., Wang C. & Rouse G. W. 2017. A new species of Ophryotrocha (Annelida, Eunicida, Dorvilleidae) from hydrothermal vents on the Southwest Indian Ridge. ZooKeys 686: 1 - 9. https: // doi. org / 10.3897 / zookeys. 687.13046","Paxton H. 2004. Jaw growth and replacement in Ophryotrocha labronica (Polychaeta, Dorvilleidae). Zoomorphology 123: 147 - 154. https: // doi. org / 10.1007 / s 00435 - 004 - 0097 - 4"]}

Ophryotrocha charlottae sp. nov. urn:lsid:zoobank.org:act: 2AC55385-EA0C-41E5-8EAF-4D6F32818D94 Fig. 4 Etymology Named in honor of Charlotte Seid, collection manager of the Benthic Invertebrate Collection at Scripps Oceanography, for her dedication to facilitating biodiversity research. Material examined Holotype EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, northwest of Easter Island, active hydrothermal vents; 23.823° S, 115.456° W; depth 2649 m; 2 Apr. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4097; GenBank: OP311739 (COI), OP304893 (16S), OP311648 (H3); SIO-BIC A14096. Paratypes EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and midsection tissue used for DNA extraction); same collection data as for holotype; GenBank: OP311741 (COI); SIO-BIC A14163 • 1 spec. (fixed in ethanol and midsection tissue used for DNA extraction); same collection data as for holotype; GenBank: OP311740 (COI); SIO-BIC A14164. Other material EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and entirely used for DNA extraction); same collection data as for holotype; GenBank: OP311738 (COI); SIO-BIC A14097 • 7 or more specs (7 fixed in glutaraldehyde, additional material fixed in formalin and entirely used for slides of parapodia, additional tissue fixed in ethanol); same collection data as for holotype; SIO-BIC A14098 • 1 spec. (fixed in ethanol and midsection tissue used for DNA extraction); same collection data as for holotype; SIO-BIC A14187 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14188. Description In life light brown (Fig. 4A), opaque white after preservation. Body ~ 3 mm long, with more than 35 segments of similar width, slightly tapering posteriorly (Fig. 4A). Prostomium rounded, wider than long, with paired digitiform antennae inserted dorsally, paired digitiform palps inserted ventral-laterally, similar in length with antennae. Peristomium two equal rings, each similar in size to the following segments (Fig. 4A). Eyes not visible. Maxillae P-type, forceps comb-like, with large main fang, fused together basally. Four rows of seven free denticles, the posterior-most free denticles (D1) comb-like, like the forceps, other free denticles (D2–D7) shovel-shaped with fine teeth (Fig. 4B). Mandibles heavily sclerotized, shafts rod-like, cutting plates sub-triangular, lateral wings weakly sclerotized (Fig. 4C). Parapodia uniramous, acicular lobe with rounded distal margin, dorsal cirri enlarged fusiform, similar in size with acicular lobe, reaching distal margin of acicular lobe, ventral cirri short and stubby (Fig. 4D). Supra-acicular chaetae simple, distally serrated, tapering into a main fang (Fig. 4D–E). Sub-acicular chaetae compound, hooked with serrated blades (Fig. 4D, F). Sub-acicular chaetal lobe also with one or two simple chaetae (Fig. 4D, G). Pygidium with two digitiform cirri inserted laterally, a small median papilla posteriorly placed (Fig. 4A). Distribution Known only from vents at 2649 m depth near Easter Island (Rapa Nui) at the southern end of the East Pacific Rise (Pacific Antarctic Ridge). Remarks Ophryotrocha charlottae sp. nov. resembles O. cf. akessoni, which also occurs on the Southeast Pacific Ridge. They are similar in the shape of the prostomium, peristomium, antennae, palps, jaws, and chaetae. Ophryotrocha charlottae has distinctive parapodia with enlarged fusiform dorsal cirri, which easily distinguishes it from O. akessoni / O. cf. akessoni, O. jiaolongi and O. marinae sp. nov. Also, O. charlottae has two long digitiform anal cirri and a media papilla while O. cf. akessoni only has two short nub-like anal cirri (Fig. 3A). Ophryotrocha kailae sp. nov. is another species from southern end of the East Pacific Rise (Pacific Antarctic Ridge). It differs from O. charlottae in the form of antennae, palps, mandibles, dorsal cirri and pygidium (Fig. 5). Ophryotrocha charlottae can also be easily distinguished from other species of the “ akessoni ” clade, based on mandibles and parapodia (Table 3). The four rows of maxillae found in O. charlottae differ from that in close relatives such as O. akessoni, O. cf. akessoni (Fig. 3C) and O. jiaolongi that show only two rows (Blake 1985; Zhang et al. 2017), but four rows were also observed in O. marinae and O. pruittae sp. nov. (see below). It is possible that the outermost pair of rows represent molted jaws as has been observed in other Ophryotrocha by Paxton (2004), so this should not be interpreted as diagnostic without further study., Published as part of Zhang, Dongsheng, Zhou, Yadong, Yen, Nicole, Hiley, Avery S. & Rouse, Greg W., 2023, Ophryotrocha (Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species, pp. 167-194 in European Journal of Taxonomy 864 on pages 177-179, DOI: 10.5852/ejt.2023.864.2101, http://zenodo.org/record/7867628, {"references":["Blake J. A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific. I. Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae, and Maldanidae. Bulletin of the Biological Society of Washington 6: 67 - 101.","Zhang D., Zhou Y., Wang C. & Rouse G. W. 2017. A new species of Ophryotrocha (Annelida, Eunicida, Dorvilleidae) from hydrothermal vents on the Southwest Indian Ridge. ZooKeys 686: 1 - 9. https: // doi. org / 10.3897 / zookeys. 687.13046","Paxton H. 2004. Jaw growth and replacement in Ophryotrocha labronica (Polychaeta, Dorvilleidae). Zoomorphology 123: 147 - 154. https: // doi. org / 10.1007 / s 00435 - 004 - 0097 - 4"]}

Ophryotrocha kailae sp. nov. urn:lsid:zoobank.org:act: FC8AD84C-5C47-49D2-BEC4-55FC04F3DFBC Fig. 5 Etymology Named for Kaila Pearson, an expert on another group of vent and seep-associated polychaetes, phyllodocids belonging to Galapagomystides Blake, 1985. Material examined Holotype EAST PACIFIC OCEAN • 1 spec. (anterior fixed in formalin, posterior fixed in ethanol and used for DNA extraction); Southern East Pacific Rise, active hydrothermal vents; 31.151° S, 111.932° W; depth 2237 m; 29 Mar. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4094; GenBank: OP311745 (COI); SIO-BIC A14100. Paratypes EAST PACIFIC OCEAN • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, Saguaro Vent Field, active hydrothermal vents; 31.865° S, 112.044° W; depth 2235 m; 28 Mar. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4093; GenBank: OP311748 (COI); SIO-BIC A14099 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, German Flats Vent Field, active hydrothermal vents; 37.793° S, 110.916° W; depth 2216 m; 22 Mar. 2005; Greg Rouse, Nerida Wilson, Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4088; GenBank: OP311746 (COI), OP304894 (16S), OP311650 (H3); SIO-BIC A14101 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311747 (COI); SIO-BIC A14102 • 1 spec. (fixed in formalin); same collection data as for holotype; SIO-BIC A14103 • 1 spec. (anterior fixed in formalin, posterior fixed in ethanol); same collection data as for holotype; SIO-BIC A14104. Description In life, translucent with light yellow gut and white eggs mid-body, body opaque white after preservation. Body length ~ 4.5 mm with more than 30 segments, similar width through the body, slightly tapering posteriorly (Fig. 5A). Eyes not visible. Prostomium rounded, wider than long, with paired short digitiform antennae inserted dorsally, paired digitiform palps inserted ventral-laterally, similar in length with antennae. Peristomium two rings, similar in length to following segments (Fig. 5A). Mandibles heavily sclerotized, shafts rod-like, cutting plates curved, with single blunt peak, lateral wings weakly sclerotized (Fig. 5B). Maxillae P-type, forceps comb-like, with large main large fang. Two rows of 7 free denticles, posterior most free denticles (D1) comb-like, D2–D7 shovel-shaped (Fig. 5C). Parapodia uniramous, acicular lobe with rounded distal margin, dorsal cirri subtriangular, barely extending beyond distal margin of acicular lobe, ventral cirri stubby (Fig. 5C). Supra-acicular chaetae simple, distally serrated, tapering into a fang (Fig. 5D–E). Sub-acicular chaetae compound, with serrated blades (Fig. 5D, F). Sub-acicular chaetal lobe with one simple chaeta (Fig. 5D, G). Pygidium with two anal cirri inserted laterally (Fig. 5A inset). Distribution Known only from vents at 2216–2237 m along the southern East Pacific Rise (Pacific Antarctic Ridge). Remarks While the DNA data suggests Ophryotrocha kailae sp. nov. is most closely related to O. pruittae sp. nov. (Fig. 1), morphologically it resembles Ophryotrocha akessoni in having similar mandibles with curving cutting plates, which are otherwise not seen in the vent clade that also includes O. charlottae sp. nov., O. jiaolongi, O. marinae sp. nov. and O. pruittae. Ophryotrocha kailae differs from O. akessoni in the form of its head appendages and possibly pygidial cirri (Table 3)., Published as part of Zhang, Dongsheng, Zhou, Yadong, Yen, Nicole, Hiley, Avery S. & Rouse, Greg W., 2023, Ophryotrocha (Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species, pp. 167-194 in European Journal of Taxonomy 864 on pages 179-181, DOI: 10.5852/ejt.2023.864.2101, http://zenodo.org/record/7867628, {"references":["Blake J. A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific. I. Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae, and Maldanidae. Bulletin of the Biological Society of Washington 6: 67 - 101."]}

Ophryotrocha cf. akessoni Blake, 1985 Fig. 3 Material examined EAST PACIFIC OCEAN • 6 specs (5 fixed in formalin, 1 fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, German Flats Vent Field, active hydrothermal vents; 37.793° S, 110.916° W; depth 2216 m; 22 Mar. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4088; GenBank: OP311735 (COI); SIO-BIC A14105 • 2 specs (1 fixed in formalin, 1 fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311736 (COI); SIO-BIC A14106 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Southern East Pacific Rise, active hydrothermal vents; 31.151° S, 111.932° W; depth 2237 m; 29 Mar. 2005; Greg Rouse, Nerida Wilson and Robert Vrijenhoek leg.; collecting event: HOV Alvin dive 4094; GenBank: OP311737 (COI), OP304892 (16S), OP311647 (H3); SIO-BIC A14107. Description Body about 7.5 mm in length and ~ 30 segments (Fig. 3A). Prostomium rounded, wider than long, slightly rise medio-posteriorly, with paired digitiform antennae inserted dorsally, paired digitiform palps similar in length with antennae, inserted ventral-laterally (Fig. 3A). Peristomium with two rings, subequal in length to the following segments, with ciliary band on each ring (Fig. 3A). Mandibles heavily sclerotized, with rod-like shafts, curved shape cutting plates, with single blunt peak, lateral wings weakly sclerotized (Fig. 3B). Maxillae P-type, forceps comb-like, with large main fang. Two rows of 7 free denticles, posterior most denticles (D1) comb-like, with smaller main fang. The other six denticles (D2–D7) small, shovel-shaped, with fine teeth (Fig. 3C). Parapodia uniramous, with long dorsal cirri and short stubby ventral cirri, acicular lobes sub-triangular distally (Fig. 3D). Supra-acicular chaetae simple, distally serrated, tapering into a main fang (Fig. 3D–E). Sub-acicular chaetae compound, with serrated blades (Fig. 3F–G). Sub-acicular chaetal lobe holds one simple chaeta (Fig. 3D, G). Pygidium with two short nub-like papillae laterally (Fig. 3A). Remarks The specimens were collected from two sites of hydrothermal vents on the southern East Pacific Rise, also known as the Pacific Antarctic Ridge. Though collected well south of the type locality, they agree with Ophryotrocha akessoni described from Galapagos Rift vents in most characters, though there are differences in jaw structure. Blake (1985) described a replacement of mandibles in O. akessoni from serrated cutting plates in juveniles to curved cutting plates in adults. In our specimens (Fig. 3B), mandibles with curved cutting plates resemble the adult mandibles in O. akessoni. Paxton (2004) and Macnaughton et al. (2010) inferred that mandibles of a range of species of Ophryotrocha show basically adult size and shape of cutting plates since the larval stage, they only lengthen and enlarge their proximal shafts with maturity. It is possible that the juvenile mandibles in Blake’s (1985) original description of O. akessoni may belong to another Ophryotrocha. Furthermore, O. akessoni was described with a P-type tending to K-type maxillae, with lateral teeth on the forceps (Blake 1985). However, the forceps of K-type maxillae are unidentate or bidentate with the lateral dentition completely reduced (Paxton 2004; Macnaughton et al. 2010). Thus, according to the original description, it may be that O. akessoni has P-type maxillae as seen in our specimens (Fig. 3C). Blake (1985) described the presence of two anal cirri but provided no further details or drawings. Our specimens had two short papillae which could correspond to those of Blake’s specimens, but examination of the types is needed. Based on the variation in morphological characters (Table 3) and the lack of any DNA sequences for O. akessoni from the type locality, we consider it prudent to report our specimens as O. cf. akessoni., Published as part of Zhang, Dongsheng, Zhou, Yadong, Yen, Nicole, Hiley, Avery S. & Rouse, Greg W., 2023, Ophryotrocha (Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species, pp. 167-194 in European Journal of Taxonomy 864 on pages 175-176, DOI: 10.5852/ejt.2023.864.2101, http://zenodo.org/record/7867628, {"references":["Blake J. A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific. I. Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae, and Maldanidae. Bulletin of the Biological Society of Washington 6: 67 - 101.","Paxton H. 2004. Jaw growth and replacement in Ophryotrocha labronica (Polychaeta, Dorvilleidae). Zoomorphology 123: 147 - 154. https: // doi. org / 10.1007 / s 00435 - 004 - 0097 - 4","Macnaughton M. O., Worsaae K. & Eibye-Jacobsen D. 2010. Jaw morphology and ontogeny in five species of Ophryotrocha. Journal of Morphology 271: 324 - 339. https: // doi. org / 10.1002 / jmor. 10800"]}

Dorvilleids belonging to Ophryotrocha Claparède & Mecznikow, 1869 are known from deep-sea hydrothermal vents in the Pacific, Atlantic, Indian and Southern Oceans. However, how they colonized and diversified in these ecosystems has not been assessed in detail. Here, a collection of Pacific hydrothermal vent Ophryotrocha was examined using morphology and DNA markers (COI, 16S and H3). Five new species were revealed, largely expanding the diversity of the group at this habitat type. They are Ophryotrocha charlottae sp. nov., O. kailae sp. nov., O. marinae sp. nov., O. pruittae sp. nov. from eastern Pacific, and O. bohnorum sp. nov. from the western Pacific. Phylogenetic analyses based on the concatenated alignments of all three genes suggest vent habitants have been colonized several times independently within Ophryotrocha. One clade of six vent species was recovered, indicative of diversification following a colonization of hydrothermal vents, likely in the eastern Pacific. An Indian Ocean species, O. jiaolongi, was nested inside this clade and was closely related to one of the new species from the Gulf of California, diverging from it by less than 4% on COI.

Ophryotrocha bohnorum sp. nov. urn:lsid:zoobank.org:act: 181A86F6-7F8D-401C-B252-21FF39F6C5C0 Fig. 9 Etymology Ophryotrocha bohnorum sp. nov. is named for Jeffrey and Brenda Bohn and their family in recognition of their enduring support of deep-sea research. Material examined Holotype TONGA • 1 spec. (fixed in ethanol and a midbody piece used for DNA extraction); Lau Back-Arc Basin, Southern Valu Fa Ridge, Hine Hina Vent Field, active hydrothermal vents; 22.539° S, 176.718° W; depth 1845–1906 m; 22–23 May 2005; Greg Rouse, Fredrik Pleijel and Robert Vrijenhoek leg.; collecting event: ROV Jason II dive 146; GenBank: OP311742 (COI), OP304895 (16S), OP311649 (H3); SIO-BIC A14092. Paratypes TONGA • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14094 • 4 specs (fixed in formalin); same collection data as for holotype; SIO-BIC A14095 • 1 spec. (fixed in ethanol and a midbody piece used for DNA extraction); same collection data as for holotype; GenBank: OP311744 (COI); SIO-BIC A14165 • 1 spec. (fixed in ethanol and a midbody piece used for DNA extraction); same collection data as for holotype; GenBank: OP311743 (COI); SIO-BIC A14166 • 1 spec. (fixed in ethanol); same locality as for holotype; 22.532° S, 176.719° W; depth 1818–1907 m; 21–22 May 2005; Greg Rouse, Fredrik Pleijel and Robert Vrijenhoek leg.; collecting event: ROV Jason II dive 145; SIO-BIC A14088 • 1 spec. (fixed in ethanol); same collection data as for preceding; SIO-BIC A14089 • 1 spec. (fixed in ethanol); same collection data as for preceding; SIO-BIC A14090 • 3 specs (fixed in formalin); same collection data as for preceding; SIO-BIC A14091 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14093. Description In life, white, yellow gut, with white eggs mid-body (Fig. 9A), opaque white after preservation. Body 12 mm long with 60+ segments, tapering slightly along body.Prostomium rounded, wider than long,with a slightly rise posteriorly in the middle. Paired antennae long cirriform, inserted dorsally, tapering distally. Paired palps cirriform, slightly shorter than antennae, inserted dorsal-laterally (Fig. 9A). Peristomium two equal rings, similar in length to first chaetiger, slightly longer than following chaetigers, with two brown spots located dorsal-laterally between the two rings (Fig. 9A–B). Maxillae P-type, maxillary carriers comb-like, with large main fang, 8 pairs of free denticles. Posterior 4 denticles (D1–D4) heavily sclerotized, with large main fang and sharp teeth; anterior 4 denticles (D5–D8) translucent, with a small main fang and tiny teeth, D5–D6 overlap with D3–D4 (Fig. 9C). Mandibles transparent, rod-like shafts, cutting plates triangular, with blunt teeth on the anterior edge (Fig. 9D). Parapodia uniramous, acicular lobe rounded with a small tip in the middle of the distal margin, dorsal cirri short, rounded lobes, ventral cirri long extending from distal margin of acicular lobe (Fig. 9E). Supra-acicular chaetae simple, distally serrated, with a small hook on the tip (Fig. 9F). Sub-acicular chaetae all compound, shafts bifid on the top, serrated blades with a small hook similar as supra-acicular chaetae on the tip (Fig. 9G). Pygidium with two long digitiform cirri (Fig. 9A). Distribution Only known from vents at the Lau Back-Arc Basin, southwest Pacific Ocean at depths of 1845–1907 m. Remarks The phylogenetic results (Fig. 1) show that Ophryotrocha bohnorum sp. nov. is most closely related to an undescribed species complex of Ophryotrocha (O. Seep4) from eastern Pacific methane seeps (Thornhill et al. 2012). There is no morphological information available for these specimens. A clade comprising other eastern Pacific species, Ophryotrocha globopalpata from hydrothermal vents, O. flabella from a whale fall and O. Seep3 (Thornhill et al. 2012), is then the well supported sister group to this clade. Based on this topology Ophryotrocha bohnorum appears to have independently colonized hydrothermal vents from O. globopalpata (Fig. 1). Ophryotrocha bohnorum has hooked tips of the supra- and sub-acicular chaetae and red-brown spots located dorsal-laterally between two peristomial segments, features not seen in other species from hydrothermal vents. Only four species of Ophryotrocha, O. atlantica Hilbig & Blake, 1991, O. mediterranea Martin Abello & Cartes, 1991, O. pachysoma Hilbig & Blake, 1991, and O. socialis Ockelmann & Åkesson, 1990, have been described with chaetae with hooked tips. Ophryotrocha bohnorum can be easily distinguished from these species by its transparent mandibles with serrated anterior edge.

Ophryotrocha pruittae sp. nov. urn:lsid:zoobank.org:act: BE33115E-1B4C-405A-A425-AF518EE5E093 Fig. 8 Ophryotrocha cf. akessoni sp. 2 – Goffredi et al. 2017: supplemental, table 1. Etymology Named for Jessica Pruitt, an aficionada and expert on deep-sea Ophryotrocha. Material examined Holotype MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Gulf of California, Alarcón Rise, active hydrothermal vents; 23.377° N, 108.531° W; depth 2309 m; 22 Apr. 2015; Greg Rouse leg.; collecting event: ROV Doc Ricketts dive 754; GenBank: OP311761 (COI); ICML-EMU-13288, (ex SIO-BIC A13689). Paratypes MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: KY701728 (16S), OP311652 (H3); SIO-BIC A6322 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14123 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14124 • 1 spec. (fixed in ethanol and most tissue used for DNA extraction); same collection data as for holotype; SIO-BIC A14125 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14126 • at least 4 specs (1 fixed in formalin, 3 individuals and additional fragments fixed in ethanol); same collection data as for holotype; SIO-BIC A14127 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14128 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14129 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14130 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14131 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14132. Description In life, golden color (Fig. 8A), opaque white after preservation. Body 10.5 mm long, 50+ segments of similar width through the body. Eyes not visible. Prostomium rounded, wider than long, with paired digitiform antennae inserted dorsally, paired digitiform palps inserted ventral-laterally, similar in length with antennae. Peristomium two equal rings, similar size to the following segments (Fig. 8A). Maxillae P-type, forceps comb-like, with large main large fang, fused together basally. Four rows of seven free denticles, the posterior most free denticles (D1) comb-like, similar to the forceps, other free denticles shovel-shaped with fine teeth, D2–D4 smaller than D5–D7 (Fig. 8B). Mandibles heavily sclerotized, shafts rod-like, cutting plates L-shape, anterior edge flat with pointed lateral peaks weakly sclerotized, lateral wings weakly sclerotized (Fig. 8C). Parapodia uniramous, acicular lobe triangular with a blunt point distally, dorsal cirri cirriform, long, extend beyond distal margin of acicular lobe, ventral cirri short and stubby (Fig. 8D). Supra-acicular chaetae simple distally serrated, tapering into a large main fang, 2–5 per fascicle (Fig. 8D–E). Five to eight compound sub-acicular chaetae, with serrated blades (Fig. 8D, F). Sub-acicular chaetal lobe holds one or two simple chaetae (Fig. 8D, G). Pygidium with two short conical cirri inserted laterally (Fig. 8A). Distribution Only known from the Alarcón Rise vents in the southern Gulf of California at 2309 m depth. Remarks Ophryotrocha pruittae sp. nov. was initially reported in Goffredi et al. (2017) as Ophryotrocha cf. akessoni sp. 2 with a partial DNA sequence for mitochondrial 16S rRNA lodged on GenBank (KY701727). Ophryotrocha pruittae has a rounded prostomium, two equal segments of peristomium, digitiform antennae and palps and P-type maxillae, which are all features found in the vent-clade of Clade B. It differs from these species by having the mandibles with the L-shape cutting plates and two conical anal cirri, while O. marinae sp. nov. has triangular cutting plates and two digitiform anal cirri, O. akessoni / O. cf. akessoni has curved cutting plates, O. charlottae sp. nov. has subtriangular cutting plates and two lateral cirri and one median anal cirrus, O. kailae sp. nov. has curved cutting plates and two digitiform anal cirri (Table 3). Ophryotrocha pruittae showed four rows of maxillae. As discussed above for O. charlottae and O. marinae it is possible that the outermost pair of rows represent molted jaws as has been observed in other Ophryotrocha by Paxton (2004).

Ophryotrocha marinae sp. nov. urn:lsid:zoobank.org:act: 5B66661D-DDF9-425D-860F-412872058C27 Figs 6–7 Ophryotrocha cf. akessoni sp. 1 – Goffredi et al. 2017: supplemental, table 1. Ophryotrocha cf. akessoni – Salcedo et al. 2019: 6, table 1. Etymology Named in honor of Marina McCowin for her dedication in the study of the fauna associated with seeps and vents. Marina has studied Siboglinidae and Ophryotrocha marinae sp. nov. was notable for being associated in large numbers with the tubes of Oasisia alvinae Jones, 1985 (Fig. 6A). Material examined Holotype MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); Gulf of California, Pescadero Basin, active hydrothermal vents; 23.960° N, 108.863° W; depth 3676–3756 m; 18 Apr. 2015; Greg Rouse leg.; collecting event: ROV Doc Ricketts dive 750 (specimens associated with tubes of Oasisia alvinae); GenBank: OP311750 (COI);, ICML-EMU-13289, (ex SIO-BIC A14109). Paratypes MEXICO • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311749 (COI), KY701727 (16S), OP311651 (H3); SIO-BIC A6308 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311755 (COI); SIO-BIC A14108 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for holotype; GenBank: OP311751 (COI); SIO-BIC A14110 • 1 spec. (fixed in ethanol); same collection data as for holotype; SIO-BIC A14111 • ca 35 specs (10 fixed in formalin, ca 25 fixed in ethanol); same collection data as for holotype; SIO-BIC A14112. • 11 specs (5 fixed in paraformaldehyde / glutaraldehyde, 6 fixed in ethanol); Gulf of California, Pescadero Basin, Auka Vent Field, region of P Vent and Z Vent; 23.96° N, 108.86° W; depth 3648–3671 m; 3 Nov. 2021; Greg Rouse leg.; collecting event: ROV SuBastian dive 475; GenBank: OP561817 (COI); SIO-BIC A14031. Other material MEXICO • 7 specs (3 fixed in formalin, 4 fixed in ethanol); Gulf of California, Pescadero Basin, Auka Vent Field, Matterhorn area, active hydrothermal vents; 23.95° N, 108.86° W; depth 3650 m; 14 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 193, sample S0193-S4 (suction sampler chamber 4, specimens associated with tubes of Oasisia alvinae); SIO-BIC A9974 • 7 specs (2 fixed in formalin, 5 fixed in ethanol); same collection data as for preceding; SIO-BIC A9975 • 4 specs (fixed in ethanol); same locality as for preceding; 23.95369° N, 108.86231° W; depth 3668 m; 29 Oct. 2021; Greg Rouse leg.; collecting event: ROV SuBastia n dive 470, sample S0470-S3 (suction sampler chamber 3, among tube worms); SIO-BIC A13987 • 2 specs (1 anterior fragment fixed in ethanol, 1 anterior fixed in formalin and posterior fixed in ethanol and used for DNA extraction); Gulf of California, Pescadero Basin, Auka Vent Field, Z Mound area, active hydrothermal vents; 23.95616° N, 108.86191° W; depth 3688 m; 17 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 196, sample S0196-PC1 (push core 1 at microbial mat with venting); GenBank: OP311752 (COI); SIO-BIC A9987 • at least 15 specs (3 fixed in formalin, 12 fixed in RNAlater, additional material fixed in ethanol); same locality as for preceding; 23.9561° N, 108.8619° W; depth 3688 m; 21 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 200, samples S0200-S1 and S0200-S2 (suction sampler chambers 1 and 2, microbial mat with venting); SIO-BIC A10029 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311753 (COI); SIO-BIC A14114 • 4 specs (3 fixed in ethanol, 1 anterior fixed in formalin and posterior fixed in ethanol); Gulf of California, Pescadero Basin, Auka Vent Field, oily mat site southeast of Z Vent; 23.95° N, 108.86° W; depth 3650–3662 m; 5 Nov. 2021; Greg Rouse leg.; collecting event: ROV SuBastian dive 477; SIO-BIC A14045 • 7 specs (6 fixed in paraformaldehyde, 1 fixed in ethanol); Gulf of California, Pescadero Basin, midway between Auka and JaichMaa ‘ja’ag Vent Fields; 23.95369° N, 108.86231° W; depth 3663–3687 m; 1 Nov. 2021; Greg Rouse leg.; collecting event: ROV SuBastian dive 473; SIO-BIC A14007 • 11 specs (2 fixed in formalin, 9 fixed in ethanol); Gulf of California, Pescadero Basin, JaichMaa ‘ja’ag Vent Field, Cavern Tay Ujaa (Big Cave), active hydrothermal vents; 23.94157° N, 108.85570° W; depth 3675 m; 18 Nov. 2018; Greg Rouse and Ekin Tilic leg.; collecting event: ROV SuBastian dive 197, sample S0197-S6 (suction sampler chamber 6, specimens associated with small tubes of Oasisia alvinae); SIO-BIC A10002 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311754 (COI); SIO-BIC A14113 • ca 50 specs (fixed in ethanol); Gulf of California, Guaymas Basin, sedimented hydrothermal vents; 27.016° N, 111.410° W; depth 2012 m; 13 Nov. 2009; Anna-Louise Reysenbach leg.; collecting event: HOV Alvin dive 4558; SIO-BIC A14115 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311756 (COI); SIO-BIC A14116 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311757 (COI); SIO-BIC A14117 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; SIO-BIC A14118 • ca 15 specs (fixed in ethanol); Gulf of California, Guaymas Basin, sedimented hydrothermal vents; 27.015° N, 111.410° W; depth 2010 m; 15 Nov. 2009; Anna-Louise Reysenbach leg.; collecting event: HOV Alvin dive 4560; SIO-BIC A14119 • 1 speci. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311758 (COI); SIO-BIC A14120 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311759 (COI); SIO-BIC A14121 • 1 spec. (fixed in ethanol and the posterior end used for DNA extraction); same collection data as for preceding; GenBank: OP311760 (COI); SIO-BIC A14122. Description In life, golden color with white eggs mid-body (Fig. 7A–B), opaque white after preservation. Body length 10.5 mm, with 50+ segments, slightly dorso-ventrally compressed, widest anteriorly, gradually tapering posteriorly (Fig. 7A–B). Prostomium rounded, wider than long, posterior medial area slightly raised, with paired digitiform antennae, tapering distally, inserted dorsally, paired digitiform palps similar in length with antennae, inserted ventral-laterally (Fig. 7A). Peristomium with two rings, subequal in length to the following segments (Fig. 7A). Eyes not visible. Mandibles heavily sclerotized, with rod-like shafts, sub-triangular shape cutting plates, with single blunt peak anteriorly, lateral wings weakly sclerotized (Fig. 7C). Maxillae P-type, forceps comb-like, with large main fang. Four rows of free denticles, the posterior-most free denticles (D1) comb-like, like the forceps, other free denticles shovel-shaped with fine teeth, D2–D4 smaller than D5–D7 (Fig. 7D). Parapodia uniramous, acicular lobe with rounded distal margin, dorsal cirri long digitiform, extending beyond distal margin of acicular lobe, ventral cirri short and stubby (Fig. 7E). Supra-acicular chaetae simple, distally serrated, tapering into a main fang (Fig. 7F). Sub-acicular chaetae compound, with serrated blades (Fig. 7G). Sub-acicular chaetal lobe usually with one simple chaeta (Fig. 7H). Pygidium with two digitiform cirri inserted laterally (Fig. 7A). Distribution Known from Gulf of California hydrothermal vents of the Pescadero Basin at over 3500 m and the Guaymas Basin sedimented vents at ~ 2000 m. Found in huge numbers on tubes of Oasisia (Fig. 6A), or on microbial mats near active flow (Fig. 6B). Remarks This species was initially reported in Goffredi et al. (2017) as Ophryotrocha cf. akessoni sp. 1 with a partial DNA sequence for mitochondrial 16S rRNA lodged on GenBank (KY701727). Ophryotrocha marinae sp. nov. most closely resembles Ophryotrocha jiaolongi described from hydrothermal vents of the Indian Ocean, including sharing distinctive mandibles that distinguish these two taxa from all other Ophryotrocha. Morphologically, O. marinae differs from O. jiaolongi based on body color, the form of antennae and palps, and in lacking a median pygidial papilla (Table 3). In life, O. marinae is golden while O. jiaolongi is white and translucent. Ophryotrocha marinae also has antennae and palps that are distally tapering and longer than the length of prostomium, while in O. jiaolongi antennae and palps are shorter than the length of prostomium. The minimum COI uncorrected distance obtained between O. marinae and O. jiaolongi specimens was relatively small at 3.73% (Table 2). Other relatively small distances that are currently known are up to 4.8% between O. notoglandulata Pfannenstiel, 1972 and O. japonicus Paxton and Åkesson, 2010 (both Japanese taxa) and 6.1% between O. flabella and O. globopalpata (both from deep waters of the eastern Pacific). Given the morphological differences between O. marinae and O. jiaolongi, the reciprocal monophyly based on numerous COI sequences and vast geographic separation, we regard them here as separate species. One notable difference between O. jiaolongi and O. marinae was that the latter species showed four rows of maxillae compared to the two rows in O. jiaolongi found by Zhang et al. (2017). As discussed above for O. charlottae sp. nov. it is possible that the outermost pair of rows represent molted jaws as has been observed in other Ophryotrocha by Paxton (2004).

Preclinical models like cancer cell lines and patient-derived xenografts (PDXs) are vital for studying disease mechanisms and evaluating treatment options. It is essential that they accurately recapitulate the disease state of interest to generate results that will translate in the clinic. Prior studies have demonstrated that preclinical models do not recapitulate all biological aspects of human tissues, particularly with respect to the tissue of origin gene expression signatures. Therefore, it is critical to assess how well preclinical model gene expression profiles correlate with human cancer tissues to inform preclinical model selection and data analysis decisions. Here we evaluated how well preclinical models recapitulate human cancer and non-diseased tissue gene expression patterns with respect to the most variable genes, tumor purity, and tissue specificity by using publicly available gene expression profiles across multiple sources. We found that using the full gene set improves correlations between preclinical model and tissue global gene expression profiles, confirmed that GBM PDX global gene expression correlation to GBM tumor global gene expression outperforms GBM cell line to GBM tumor global gene expression correlations, and demonstrated that preclinical models in our study often failed to reproduce tissue-specific expression. While including additional genes for global gene expression comparison between cell lines and tissues decreases the overall correlation, it improves the relative rank between a cell line and its tissue of origin compared to other tissues. Our findings underscore the importance of using the full gene expression set measured when comparing preclinical models and tissues and confirm that tissue-specific patterns are better preserved in GBM PDX models than in GBM cell lines. Future studies can build on these findings to determine the specific pathways and gene sets recapitulated by particular preclinical models to facilitate model selection for a given study design or goal.

Tamoxifen is widely used in the adjuvant treatment of estrogen receptor–positive (ER+) breast cancer and is an important drug for pre-menopausal women and post-menopausal patients who cannot tolerate aromatase inhibitors. Despite the clear clinical benefit in improving relapse-free and overall survival in these patients, an adverse effect of tamoxifen is a 2- to 7-fold increased risk of uterine cancer (UC) after 2-5 years of treatment. To date, the mechanism of tamoxifen-driven tumorigenesis is not well understood, and preventive approaches are lacking. Here, to molecularly characterize tamoxifen-associated uterine cancers (TA-UCs) and gain insights into their unique evolution, we performed whole-exome sequencing of 21 TA-UCs (discovery cohort) and droplet digital PCR (ddPCR) of an additional 40 TA-UCs (validation cohort) obtained from the ‘Tamoxifen Associated Malignancies: Aspects of Risk’ (TAMARISK) study. In addition, we used in vivo mouse models to: (i) further investigate tamoxifen-activated molecular pathways that may be involved in TA-UC tumorigenesis; and (ii) offer mechanistic insights. Overall, we discovered that TA-UCs were genomically similar to non–TA-UCs from The Cancer Genome Atlas (TCGA) project, with one profound exception: TA-UCs are characterized by a lower-than-expected frequency of mutations in two highly prevalent UC driver genes in the PI3K pathway: PIK3CA (14% [3/21] vs 48% [265/554] in non–TA-UC; P=0.003, Fisher’s exact test; Q=0.02, Benjamini-Hochberg FDR) and PIK3R1 (0%, [0/21] vs 31% [174/554]; P=0.001; Q=0.01). We used ddPCR in the independent TA-UC validation cohort and confirmed the low frequency of mutations in PIK3CA (7.5% [3/40] vs 21% [144/685] in control UCs from the Dana-Farber contribution to the AACR GENIE project; P=0.04). We next performed mouse in vivo studies and demonstrated that tamoxifen activated the PI3K pathway and increased cell proliferation in normal mouse uterine tissue through paracrine and autocrine effects, both of which were abrogated by the PI3K inhibitor alpelisib. Taken together, we describe a distinct and novel pathway of carcinogenesis in which tamoxifen acts as a driver event in the uterus and promotes tumor development in a mutation-independent manner. Indeed, tamoxifen may increase the risk of UC by activating the PI3K pathway, which can substitute for the early acquisition of oncogenic PIK3CA or PIK3R1 mutations observed in non–TA-UC tumors. Furthermore, the ability of a PI3K inhibitor to reduce cell proliferation in our mouse model raises the possibility that downregulating the PI3K pathway may prevent or significantly reduce TA-UC development, offering a potential future therapeutic and prevention strategy for specific high-risk patients undergoing tamoxifen therapy. Citation Format: Kirsten Kubler, Agostina Nardone, Shankara Anand, Daniel Gorvich, Marjolein Droog, Francisco Hermida-Prado, Tara Akshi, Avery S Feit, Gabriella Cohen, Gwen Dackus, Matthew Pun, Yanan Kuang, Justin Cha, Mendy Miller, William J Gibson, Cloud P Paweletz, Eliezer M Van Allen, Flora E van Leeuwen, Petra Nederlof, Harry Hollema, Quang-Dé Nguyen, Marian JE Mourits, Ignaty Leshchiner, Chip Stewart, Ursula A Matulonis, Wilbert Zwart, Yosef E Maruvka, Gad Getz, Rinath Jeselsohn. Tamoxifen instigates uterine cancer development by activating PI3K signaling and supersedes PIK3CA driver mutations [abstract]. In: Proceedings of the 2021 San Antonio Breast Cancer Symposium; 2021 Dec 7-10; San Antonio, TX. Philadelphia (PA): AACR; Cancer Res 2022;82(4 Suppl):Abstract nr GS2-09.

Chromatin immunoprecipitation sequencing (ChIP-seq) and the Assay for Transposase-Accessible Chromatin with high-throughput sequencing (ATAC-seq) have become essential technologies to effectively measure protein-DNA interactions and chromatin accessibility. However, there is a need for a scalable and reproducible pipeline that incorporates proper normalization between samples, correction of copy number variations, and integration of new downstream analysis tools. Here we present Containerized Bioinformatics workflow for Reproducible ChIP/ATAC-seq Analysis (CoBRA), a modularized computational workflow which quantifies ChIP-seq and ATAC-seq peak regions and performs unsupervised and supervised analyses. CoBRA provides a comprehensive state-of-the-art ChIP-seq and ATAC-seq analysis pipeline that can be used by scientists with limited computational experience. This enables researchers to gain rapid insight into protein-DNA interactions and chromatin accessibility through sample clustering, differential peak calling, motif enrichment, comparison of sites to a reference database, and pathway analysis. CoBRA is publicly available online at https://bitbucket.org/cfce/cobra.

Here, we present the first mitochondrial genome of

Echinoids are key components of modern marine ecosystems. Despite a remarkable fossil record, the emergence of their crown group is documented by few specimens of unclear affinities, rendering their early history uncertain. The origin of sand dollars, one of its most distinctive clades, is also unclear due to an unstable phylogenetic context. We employ 18 novel genomes and transcriptomes to build a phylogenomic dataset with a near-complete sampling of major lineages. With it, we revise the phylogeny and divergence times of echinoids, and place their history within the broader context of echinoderm evolution. We also introduce the concept of a chronospace – a multidimensional representation of node ages – and use it to explore methodological decisions involved in time calibrating phylogenies. We find the choice of clock model to have the strongest impact on divergence times, while the use of site-heterogeneous models and alternative node prior distributions show minimal effects. The choice of loci has an intermediate impact, affecting mostly deep Paleozoic nodes, for which clock-like genes recover dates more congruent with fossil evidence. Our results reveal that crown group echinoids originated in the Permian and diversified rapidly in the Triassic, despite the relative lack of fossil evidence for this early diversification. We also clarify the relationships between sand dollars and their close relatives and confidently date their origins to the Cretaceous, implying ghost ranges spanning approximately 50 million years, a remarkable discrepancy with their rich fossil record.

Differentiation of cultured skeletal myoblasts is induced by extrinsic signals that include reduction in ambient mitogen concentration and increased cell density. Using an established murine myoblast cell line (C2C12), we have found that experimental reduction of the nucleoporin p62 (Nup62) content of myoblasts enhances differentiation in high-mitogen medium, while forced expression of Nup62 inhibits density-induced differentiation. In contrast, differentiation of myoblasts induced by low-mitogen medium was unaffected by ectopic Nup62 expression. Further analyses suggested that Nup62 content affects density-induced myoblast differentiation through a mechanism involving activation of p38 MAP kinase. Nuclear pore complex (NPC) composition, in particular changes in NUP62 content, may be altered during viral infection, differentiation, and in neoplastic growth. The results support a functional role for changes in Nup62 composition in NPCs and density-induced myogenic differentiation, and suggest a link between loss of Nup62 content and induction of an intracellular stress signaling pathways.

In tropical regions, widespread loss of native forest and savanna vegetation is increasing extreme heat, particularly in agricultural regions. Using the case of rising extreme heat from lost forest and savanna vegetation in Brazilian Amazon and Cerrado regions, we modeled losses to soy production, the region’s principal economic activity. We assessed two types of extreme-heat regulation values: the value of avoided extreme-heat exposure of soy from the conservation of neighboring ecosystems and the value of lost revenue due to increased extreme heat exposure from increased ecosystem conversion. Our modeling combines empirical estimates of (1) the influence of ecosystem conversion on extreme heat over neighboring cropland, (2) the impacts of extreme heat on agricultural yields, and (3) native vegetation area, agricultural area, and crop prices. We examine lost soy value from land conversion over the period 1985 to 2012, potential losses from further conversion under plausible land and climate change scenarios (2020–2050), and the future value of conservation of the region’s remaining ecosystem area near soy. Soy revenue lost due to extreme heat from native vegetation loss (1985–2012) totaled 99 (2005USD) ha−1 for 2012-2013 growing season. By 2050, agricultural growth, ecosystem conversion, and climate change could boost extreme-heat regulation values by 25% to 95%. Future values were strongly sensitive to changes in agricultural density, rates of native vegetation loss, and climate. Extreme-heat regulation values were largest in the Cerrado biome and the southeastern Amazon. Relative to land values, the value of extreme heat regulation was largest relative to the carbon value of biomass in the Cerrado. By regulating the exposure of agriculture to extreme heat, ecosystem conservation can create considerable value for the soy sector.

Epilepsy is one of the most common diseases of the central nervous system, impacting nearly 50 million people around the world. Heterogeneous in nature, epilepsy presents in children and adults alike. Currently, surgery is one treatment approach that can completely cure epilepsy. However, not all individuals are eligible for surgical procedures or have successful outcomes. In addition to surgical approaches, antiepileptic drugs (AEDs) have also allowed individuals with epilepsy to achieve freedom from seizures. Others have found treatment through nonpharmacologic approaches such as vagus nerve stimulation, or responsive neurostimulation. Difficulty in accessing samples of human brain tissue along with advances in sequencing technology have driven researchers to investigate sampling liquid biopsies in blood, serum, plasma, and cerebrospinal fluid within the context of epilepsy. Liquid biopsies provide minimal or non-invasive sample collection approaches and can be assayed relatively easily across multiple time points, unlike tissue-based sampling. Various efforts have investigated circulating nucleic acids from these samples including microRNAs, cell-free DNA, transfer RNAs, and long non-coding RNAs. Here, we review nucleic acid-based liquid biopsies in epilepsy to improve understanding of etiology, diagnosis, prediction, and therapeutic monitoring.

Alzheimer’s disease is the most common neurodegenerative disease and affects persons of all races, ethnic groups, and sexes. The disease is characterized by neuronal loss leading to cognitive decline and memory loss. There is no cure and the effectiveness of existing treatments is limited and depends on the time of diagnosis. The long prodromal period, during which patients’ ability to live a normal life is not affected despite neuronal loss, often leads to a delayed diagnosis because it can be mistaken for normal aging of the brain. In order to make a substantial impact on AD patients, early diagnosis may provide a greater therapeutic window for future therapies to slow AD-associated neurodegeneration. Current gold standards for disease detection include magnetic resonance imaging and positron emission tomography scans, which visualize amyloid β and phosphorylated tau depositions and aggregates. Liquid biopsies, already an active field of research in precision oncology, are hypothesized to provide early disease detection through minimally or non-invasive sample collection techniques. Liquid biopsies in Alzheimer’s disease have been studied in cerebrospinal fluid, blood, ocular, oral, and olfactory fluids. However, most of the focus has been on blood and cerebrospinal fluid due to biomarker specificity and sensitivity attributed to the effects of the blood-brain barrier and inter-laboratory variation during sample collection. Many studies have identified amyloid β and phosphorylated tau levels as putative biomarkers, however, advances in next-generation sequencing-based liquid biopsy methods have led to significant interest in identifying nucleic acids species associated with Alzheimer’s disease from liquid tissues. Differences in cell-free RNAs and DNAs have been described as potential biomarkers for AD and hold the potential to affect disease diagnosis, treatment, and future research avenues.

Reward benefits to memory formation have been robustly linked to dopaminergic activity. Despite established characterization of dopaminergic mechanisms as operating at multiple timescales, potentially supporting distinct functional outcomes, the temporal dynamics by which reward might modulate memory encoding are just beginning to be investigated. In the present study, we leveraged a mixed block/event experimental design to disentangle transient and sustained reward influences on task engagement and subsequent recognition memory in an adapted monetary-incentive encoding (MIE) paradigm. Across three behavioral experiments, transient and sustained reward modulation of item and context memory was probed, at both 24-hour and ~10-minute retention intervals, to investigate the importance of overnight consolidation. In general, we observed that transient reward was associated with enhanced item memory encoding, while sustained reward modulated task engagement at encoding (specifically, increasing response speed) but did not confer a benefit to subsequent memory performance. Notably, reward benefits to item memory performance were somewhat inconsistent across the three experiments, and in contrast to predictions, we did not observe reward modulation of context memory performance or amplification of reward effects by overnight consolidation. Taken together, the observed pattern of behavior suggests potentially distinct roles for transient and sustained reward in memory encoding and cognitive performance and suggests that further investigation of the temporal dynamics of dopaminergic contributions to memory formation will advance understanding of motivated memory.

Echinoids are key components of modern marine ecosystems. Despite a remarkable fossil record, the emergence of their crown group is documented by few specimens of unclear affinities, rendering much of their early history uncertain. The origin of sand dollars, one of its most distinctive clades, is also unclear due to an unstable phylogenetic context and discrepancies between molecular divergence times and fossil evidence. We employ seventeen novel genomes and transcriptomes to build a phylogenomic dataset with a near-complete sampling of major lineages. With it, we revise the phylogeny and divergence times of echinoids, and place their history within the broader context of echinoderm evolution. We also introduce the concept of a chronospace—a multidimensional representation of node ages—and use it to explore the effects of using alternative gene samples, models of molecular evolution, and clock priors. We find the choice of clock model to have the strongest impact on divergence times, while the use of site-heterogeneous models shows little effects. The choice of loci shows an intermediate impact, affecting mostly deep Paleozoic nodes, for which clock-like genes recover dates more congruent with fossil evidence. Our results reveal that crown group echinoids originated in the Permian and diversified rapidly in the Triassic, despite the relative lack of fossil evidence for this early diversification. We also clarify the relationships among sand dollars and their close relatives, showing that the genus Apatopygus represents a relict lineage with a deep Jurassic origin. Surprisingly, the origin of sand dollars is confidently dated to the Cretaceous, implying ghost ranges spanning approximately 50 million years, a remarkable discrepancy with their rich fossil record.

Large-scale agriculture in the state of Mato Grosso, Brazil is a major contributor to global food supplies, but its continued productivity is vulnerable to contracting wet seasons and increased exposure to extreme temperatures. Sowing dates serve as an effective adaptation strategy to these climate perturbations. By controlling the weather experienced by crops and influencing the number of successive crops that can be grown in a year, sowing dates can impact both individual crop yields and cropping intensities. Unfortunately, the spatiotemporally resolved crop phenology data necessary to understand sowing dates and their relationship to crop yield are only available over limited years and regions. To fill this data gap, we produce a 500 m rainfed soy (Glycine max) sowing and harvest date dataset for Mato Grosso from 2004 to 2014 using a novel time series analysis method for Moderate Resolution Imaging Spectroradiometer (MODIS) satellite imagery, adapted for implementation in Google Earth Engine (GEE). Our estimates reveal that soy sowing and harvest dates varied widely (about 2 months) from field to field, confirming the need for spatially resolved crop timing information. An interannual trend toward earlier sowing dates occurred independently of variations in wet season onset, and may be attributed to an improvement in logistic or economic constraints that previously hampered early sowing. As anticipated, double cropped fields in which two crops are grown in succession are planted earlier than single cropped fields. This difference shrank, however, as sowing of single cropped fields occurred closer to the wet season onset in more recent years. The analysis offers insights about sowing behavior in response to historical climate variations which could be extended to understand sowing response under climate change in Mato Grosso., Soy cultivation; Sowing date; Mato Grosso; Climate change; Remote sensing; Time series analysis.

Alzheimer's disease (AD) is the most common neurodegenerative disease and affects persons of all races, ethnic groups, and sexes. The disease is characterized by neuronal loss leading to cognitive decline and memory loss. There is no cure and the effectiveness of existing treatments is limited and depends on the time of diagnosis. The long prodromal period, during which patients' ability to live a normal life is not affected despite neuronal loss, often leads to a delayed diagnosis because it can be mistaken for normal aging of the brain. In order to make a substantial impact on AD patient survival, early diagnosis may provide a greater therapeutic window for future therapies to slow AD-associated neurodegeneration. Current gold standards for disease detection include magnetic resonance imaging and positron emission tomography scans, which visualize amyloid β and phosphorylated tau depositions and aggregates. Liquid biopsies, already an active field of research in precision oncology, are hypothesized to provide early disease detection through minimally or non-invasive sample collection techniques. Liquid biopsies in AD have been studied in cerebrospinal fluid, blood, ocular, oral, and olfactory fluids. However, most of the focus has been on blood and cerebrospinal fluid due to biomarker specificity and sensitivity attributed to the effects of the blood-brain barrier and inter-laboratory variation during sample collection. Many studies have identified amyloid β and phosphorylated tau levels as putative biomarkers, however, advances in next-generation sequencing-based liquid biopsy methods have led to significant interest in identifying nucleic acid species associated with AD from liquid tissues. Differences in cell-free RNAs and DNAs have been described as potential biomarkers for AD and hold the potential to affect disease diagnosis, treatment, and future research avenues.

c-MYC (MYC) is a major driver of prostate cancer tumorigenesis and progression. Although MYC is overexpressed in both early and metastatic disease and associated with poor survival, its impact on prostate transcriptional reprogramming remains elusive. We demonstrate that MYC overexpression significantly diminishes the androgen receptor (AR) transcriptional program (the set of genes directly targeted by the AR protein) in luminal prostate cells without altering AR expression. Importantly, analyses of clinical specimens revealed that concurrent low AR and high MYC transcriptional programs accelerate prostate cancer progression toward a metastatic, castration-resistant disease. Data integration of single-cell transcriptomics together with ChIP-seq revealed an increased RNA polymerase II (Pol II) promoter-proximal pausing at AR-dependent genes following MYC overexpression without an accompanying deactivation of AR-bound enhancers. Altogether, our findings suggest that MYC overexpression antagonizes the canonical AR transcriptional program and contributes to prostate tumor initiation and progression by disrupting transcriptional pause release at AR-regulated genes.STATEMENT OF SIGNIFICANCEAR and MYC are key to prostate cancer etiology but our current understanding of their interplay is scarce. Here we show that the oncogenic transcription factor MYC can pause the transcriptional program of the master transcription factor in prostate cancer, AR, while turning on its own, even more lethal program.

Deoxyguanosine kinase (dGK) is reported responsible for the phosphorylation of deoxyadenosine (dA) and deoxyguanosine (dG) in the mitochondrial purine salvage pathway. Antiviral nucleoside analogs known as nucleoside reverse transcriptase inhibitors (NRTIs) must be phosphorylated by host enzymes for the analog to become active. We address the possibility that NRTI purine analogs may be competitive inhibitors of dGK. From a group of such analogs, we demonstrate that entecavir (ETV) competitively inhibited the phosphorylation of dG and dA in rat mitochondria. Mitochondria from the brain, heart, kidney, and liver showed a marked preference for phosphorylation of dG over dA (10-30-fold) and ETV over dA (2.5-4-fold). We found that ETV inhibited the phosphorylation of dG with an IC

ChIP-seq and ATAC-seq have become essential technologies used as effective methods of measuring protein-DNA interactions and chromatin accessibility. However, there is a need for a scalable and reproducible pipeline that incorporates correct normalization between samples, adjustment of copy number variations, and integration of new downstream analysis tools. Here we present CoBRA, a modularized computational workflow which quantifies ChIP and ATAC-seq peak regions and performs unsupervised and supervised analysis. CoBRA provides a comprehensive state-of-the-art ChIP and ATAC-seq analysis pipeline that is usable by scientists with limited computational experience. This enables researchers to gain rapid insight into protein-DNA interactions and chromatin accessibility through sample clustering, differential peak calling, motif enrichment, comparison of sites to a reference DB and pathway analysis.Code availability: https://bitbucket.org/cfce/cobra

Protocol for uploading mitochondrial COI sequences to GenBank

SummaryDrugs that block the activity of the methyltransferase EZH2 are in clinical development for the treatment of non-Hodgkin lymphomas harboring gain-of-function EZH2 mutations that enhance its polycomb repressive function. In contrast, in castration-resistant prostate cancer (CRPC) we have previously reported that EZH2 plays a non-canonical role as a transcriptional activator. In this setting, we now show that EZH2 inhibitors can also block the non-canonical activity of EZH2 and inhibit the growth of CRPC cells. Gene expression and epigenomic profiling of cells treated with EZH2 inhibitors demonstrated that rather than de-repressing tumor suppressor genes silenced by PRC2, EZH2 inhibitors downregulate a set of DNA repair genes that are directly regulated by EZH2. In addition, genome-wide CRISPR/Cas9-mediated loss-of-function screens in the presence of EZH2 inhibitors identified these DNA repair genes to underlie the growth-inhibitory function of these compounds. Interrogation of public data from diverse solid tumor types expressing wild-type EZH2 showed that expression of DNA damage repair genes is significantly correlated with cellular sensitivity to EZH2 inhibitors. Consistent with these findings, treatment of CRPC cells with EZH2 inhibitors dramatically enhanced their sensitivity to genotoxic stress. These studies reveal a previously unappreciated mechanism of action of EZH2 inhibitors and provide a mechanistic basis for potential new combination cancer therapies.

This protocol provides thorough instructions for how to assemble and annotate full mitochondrial genomes from NGS genome skimming data (specifically paired-end, FASTQ, gzipped reads). The steps are categorized in the following sections, which include detailed explanations throughout (e.g. why specific steps are executed in the manner instructed): I. Download Programs and Dependencies II. Choose Working Directory III. Simple Stats with SeqKit IV. Clean and Trim Reads with Trimmomatic V. Downsample Reads with MITObim VI. Reformat Reads to Non-Interleaved with BBMap VII. Mitogenome Assembly and Annotation: MitoFinder with MetaSPAdes VIII. Mitogenome Assembly with NOVOPlasty Furthermore, there are supplementary tutorial videos for each of the aforementioned protocol sections, which show all of the corresponding steps being executed. These videos may be used for extra guidance while running the protocol, and are especially helpful to see how long the commands take to execute and successfully terminate in real-time with real NGS genome skimming data.

Anxiety disorders are highly prevalent and disabling but seem particularly tractable to investigation with translational neuroscience methodologies. Neuroimaging has informed our understanding of the neurobiology of anxiety disorders, but research has been limited by small sample sizes and low statistical power, as well as heterogenous imaging methodology. The ENIGMA‐Anxiety Working Group has brought together researchers from around the world, in a harmonized and coordinated effort to address these challenges and generate more robust and reproducible findings. This paper elaborates on the concepts and methods informing the work of the working group to date, and describes the initial approach of the four subgroups studying generalized anxiety disorder, panic disorder, social anxiety disorder, and specific phobia. At present, the ENIGMA‐Anxiety database contains information about more than 100 unique samples, from 16 countries and 59 institutes. Future directions include examining additional imaging modalities, integrating imaging and genetic data, and collaborating with other ENIGMA working groups. The ENIGMA consortium creates synergy at the intersection of global mental health and clinical neuroscience, and the ENIGMA‐Anxiety Working Group extends the promise of this approach to neuroimaging research on anxiety disorders., Anxiety disorders are highly prevalent and disabling but seem particularly tractable to investigation with translational neuroscience methodologies. Neuroimaging has informed our understanding of the neurobiology of anxiety disorders, but research has been limited by small sample sizes and low statistical power, as well as heterogenous imaging methodology. The ENIGMA‐Anxiety Working Group has brought together researchers from around the world, in a harmonized and coordinated effort to address these challenges and generate more robust and reproducible findings. The ENIGMA consortium creates synergy at the intersection of global mental health and clinical neuroscience, and the ENIGMA‐Anxiety Working Group extends the promise of this approach to neuroimaging research on anxiety disorders.

Polynoidae Kinberg, 1856 has five branchiate genera: Branchipolynoe Pettibone, 1984, Branchinotogluma Pettibone, 1985, Branchiplicatus Pettibone, 1985, Peinaleopolynoe Desbruyères & Laubier, 1988, and Thermopolynoe Miura, 1994, all native to deep-sea, chemosynthetic-based habitats. Of these, Peinaleopolynoe has two accepted species; Peinaleopolynoe sillardi Desbruyères & Laubier, 1988 (Atlantic Ocean) and Peinaleopolynoe santacatalina Pettibone, 1993 (East Pacific Ocean). The goal of this study was to assess the phylogenetic position of Peinaleopolynoe, utilizing DNA sequences from a broad sampling of deep-sea polynoids. Representatives from all five branchiate genera were included, several species of which were sampled from near the type localities; Branchinotogluma sandersi Pettibone, 1985 from the Galápagos Rift (E/V “Nautilus”); Peinaleopolynoe sillardi from organic remains in the Atlantic Ocean; Peinaleopolynoe santacatalina from a whalefall off southern California (R/V “Western Flyer”) and Thermopolynoe branchiata Miura, 1994 from Lau Back-Arc Basin in the western Pacific (R/V “Melville”). Phylogenetic analyses were conducted using mitochondrial (COI, 16S rRNA, and CytB) and nuclear (18S rRNA, 28S rRNA, and H3) genes. The analyses revealed four new Peinaleopolynoe species from the Pacific Ocean that are formally described here: Peinaleopolynoe orphanae Hatch & Rouse, sp. nov., type locality Pescadero Basin in the Gulf of California, Mexico (R/V “Western Flyer”); Peinaleopolynoe elvisi Hatch & Rouse, sp. nov. and Peinaleopolynoe goffrediae Hatch & Rouse, sp. nov., both with a type locality in Monterey Canyon off California (R/V “Western Flyer”) and Peinaleopolynoe mineoi Hatch & Rouse, sp. nov. from Costa Rica methane seeps (R/V “Falkor”). In addition to DNA sequence data, the monophyly of Peinaleopolynoe is supported by the presence of ventral papillae on segments 12–15. The results also demonstrated the paraphyly of Branchinotogluma and Lepidonotopodium Pettibone, 1983 and taxonomic revision of these genera is required. We apply the subfamily name Lepidonotopodinae Pettibone 1983, for the clade comprised of Branchipolynoe, Branchinotogluma, Bathykurila, Branchiplicatus, Lepidonotopodium, Levensteiniella Pettibone, 1985, Thermopolynoe, and Peinaleopolynoe.

This study presents strategies to render triple-negative breast cancers sensitive to CDK4/6 inhibitors., Inhibitors of cyclin-dependent kinases CDK4 and CDK6 have been approved for treatment of hormone receptor–positive breast cancers. In contrast, triple-negative breast cancers (TNBCs) are resistant to CDK4/6 inhibition. Here, we demonstrate that a subset of TNBC critically requires CDK4/6 for proliferation, and yet, these TNBC are resistant to CDK4/6 inhibition due to sequestration of CDK4/6 inhibitors into tumor cell lysosomes. This sequestration is caused by enhanced lysosomal biogenesis and increased lysosomal numbers in TNBC cells. We developed new CDK4/6 inhibitor compounds that evade the lysosomal sequestration and are efficacious against resistant TNBC. We also show that coadministration of lysosomotropic or lysosome-destabilizing compounds (an antibiotic azithromycin, an antidepressant siramesine, an antimalaria compound chloroquine) renders resistant tumor cells sensitive to currently used CDK4/6 inhibitors. Lastly, coinhibition of CDK2 arrested proliferation of CDK4/6 inhibitor-resistant cells. These observations may extend the use of CDK4/6 inhibitors to TNBCs that are refractory to current anti-CDK4/6 therapies.

Rates of deforestation reported by Brazil's official deforestation monitoring system have declined dramatically in the Brazilian Amazon. Much of Brazil's success in its fight against deforestation has been credited to a series of policy changes put into place between 2004 and 2008. In this research, we posit that one of these policies, the decision to use the country's official system for monitoring forest loss in the Amazon as a policing tool, has incentivized landowners to deforest in ways and places that evade Brazil's official monitoring and enforcement system. As a consequence, we a) show or b) provide several pieces of suggestive evidence that recent successes in protecting monitored forests in the Brazilian Amazon may be doing less to protect the region's forests than previously assumed.