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| Peinaleopolynoe orphanae Hatch & Rouse
in Hatch, Liew, Hourdez & Rouse, 2020.
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Abstract
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.
Keywords: deep sea, molecular phylogeny, seeps, systematics, vents, whalefalls
Taxonomy
Polynoidae Kinberg, 1856
Lepidonotopodinae Pettibone, 1983
Peinaleopolynoe Desbruyères & Laubier, 1988, emended
Type species: Peinaleopolynoe sillardi Desbruyères & Laubier, 1988
Diagnosis (emended): Twenty-one segments. Elytra large, sub-reniform, overlapping, and covering dorsum. Elytra with or without papillae and/or posterior extensions. Chaetae extending beyond the edge of elytra. Nine or ten pairs elytra and elytrophores on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, or lacking on 19. Pharynx with either seven pairs of border papillae, six pairs of border papillae, or seven dorsal and six ventral border papillae. Bilobed prostomium with triangular anterior lobes bearing lateral antennae (= minute frontal filaments, sensu Pettibone 1993). Median antenna in anterior notch. Paired palps. Eyes lacking. Achaetous segment 1 not visible dorsally and contains dorsal and ventral pairs of smooth, tapering anterior cirri (= tentacular cirri, sensu Pettibone 1993). Parapodia biramous. Neuropodia ranging from ca. twice the length to almost as long as notopodia. Dorsal tubercles, in line with elytrophores, on non-elytrigerous segments possessing small groups of branchiae. Notochaetae bundles stout. Neurochaetae long, slender. Dorsal cirri present on non-elytrigerous segments. In specimens with nine pairs elytra, segment 19 modified, lacking dorsal cirri. Cylindrical cirrophores and long distal styles (extending far beyond length of chaetae) of dorsal cirri. Arborescent branchiae beginning on segment 2 or 3 and continuing to near end of body. Branchiae attached on bases of notopodia and on dorsal tubercles. Four pairs of ventral segmental papillae on segments 12–15. Pygidium with a pair of anal cirri.
Peinaleopolynoe santacatalina Pettibone, 1993
Peinaleopolynoe orphanae Hatch & Rouse, sp. nov.
Etymology: Peinaleopolynoe orphanae sp. nov. is named after Dr. Victoria J. Orphan, not only for her invaluable research on deep-sea microorganisms, but also for her exploration of deep-sea chemosynthetic ecosystems and her love of the animals that thrive there.
Ecology: Peinaleopolynoe orphanae sp. nov. is unusual among Peinaleopolynoe in that most specimens were associated with bacterial mats adjacent to hydrothermal vents in the Pescadero Basin at ~3700 m depth. One specimen (SIO-BIC A10926) was found at a cold seep with abundant vesicomyid clams suggesting that P. orphanae sp. nov. may be more of a habitat generalist than its close relatives.
Peinaleopolynoe orphanae sp. nov. displayed an interesting fighting behavior in situ (Fig. 6C, E), in which an individual used its everted pharynx to attack an opponent’s elytra; the two individuals attacked one another back and forth for several minutes (Suppl. material 2: movie). This may explain the damaged elytra with apparent bite marks on the posterior edges in the holotype (Fig. 7A) and in several other paratypes collected (Fig. 11A, B, E).
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| Figure 6. In situ photos of the new Peinaleopolynoe spp. A–E Peinaleopolynoe orphanae sp. nov. observed in the Pescadero Basin, Gulf of California, Mexico: C, E Peinaleopolynoe orphanae sp. nov. fighting behavior observed; the everted pharynx is used to bite off pieces of the opponent’s elytra |
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| Figure 6. In situ photos of the new Peinaleopolynoe spp. A–E Peinaleopolynoe orphanae sp. nov. observed in the Pescadero Basin, Gulf of California, Mexico: C, E Peinaleopolynoe orphanae sp. nov. fighting behavior observed; the everted pharynx is used to bite off pieces of the opponent’s elytra F Peinaleopolynoe elvisi sp. nov. holotype SIO-BIC A8488 observed and collected next to a whale bone in the Monterey Canyon, California G Peinaleopolynoe elvisi sp. nov. paratype SIO-BIC A9699 observed and collected on a pig bone deployment from Jaco Scar, Costa Rica H Peinaleopolynoe goffrediae sp. nov. holotype SIO-BIC A5485 observed and collected on a whalefall in the Monterey Canyon, California. |
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| Figure 11. Live dorsal views of Peinaleopolynoe orphanae sp. nov. Arrows indicate elytral bite marks from the fighting behavior A paratype SIO-BIC A10020 with blue elytra B paratype SIO-BIC A10024 with pink elytra C paratype SIO-BIC A6312 with white elytra D paratype SIO-BIC A6166 with black elytra E paratype SIO-BIC A10023 with red elytra.
Scale bars: 5 mm (A, B, D, E); 3 mm (C).
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| Figure 7. Live dorsal views of the new Peinaleopolynoe spp. A Peinaleopolynoe orphanae sp. nov. holotype SIO-BIC A6151 B Peinaleopolynoe elvisi sp. nov. holotype SIO-BIC A8488 C Peinaleopolynoe goffrediae sp. nov. holotype SIO-BIC A5485 D Peinaleopolynoe mineoi sp. nov. holotype SIO-BIC A10071. Scale bars: 6 mm (A); 8 mm (B, C); 1 mm (D). |
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| Figure 6. In situ photos of the new Peinaleopolynoe spp. F Peinaleopolynoe elvisi sp. nov. holotype SIO-BIC A8488 observed and collected next to a whale bone in the Monterey Canyon, California G Peinaleopolynoe elvisi sp. nov. paratype SIO-BIC A9699 observed and collected on a pig bone deployment from Jaco Scar, Costa Rica H Peinaleopolynoe goffrediae sp. nov. holotype SIO-BIC A5485 observed and collected on a whalefall in the Monterey Canyon, California. |
Peinaleopolynoe elvisi Hatch & Rouse, sp. nov.
Etymology: Peinaleopolynoe elvisi sp. nov. is named after the legendary King of Rock and Roll, Elvis Presley; the iridescent golden/pink elytra are reminiscent of the sparkly, sequined costumes he favored in his late career.
Ecology: All specimens of P. elvisi sp. nov. were found associated with vertebrate bones or wood (Table 5). Fig. 6F shows the holotype observed in situ on sediment next to a whalefall just before collection. Fig. 6G shows paratype SIO-BIC A9699 observed in situ on a deployed pig bone before collection.
Peinaleopolynoe goffrediae Hatch & Rouse, sp. nov.
Etymology: Peinaleopolynoe goffrediae sp. nov. is named after Dr. Shana K. Goffredi for her notable contribution to the exploration and research of deep-sea chemosynthetic ecosystems (especially whalefalls), focusing on symbiotic relationships between bacteria and marine invertebrates.
Ecology: Peinaleopolynoe goffrediae sp. nov. was only found associated with a whalefall (Table 5). Fig. 6H shows the holotype observed in situ on a whale carcass before collection.
Peinaleopolynoe mineoi Hatch & Rouse, sp. nov.
Etymology: Peinaleopolynoe mineoi sp. nov. is named after Ronald M. Mineo, MD, in recognition of support from the Mineo family, their interest in the deep sea, and support for our research.
Ecology: Peinaleopolynoe mineoi sp. nov. was found associated with bones and wood (Table 5). Like P. santacatalina and P. orphanae sp. nov., it may be more of a habitat generalist than other Peinaleopolynoe.
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| Figure 1. Maximum likelihood (ML) tree of the combined analysis from six genes (COI, 16S, 18S, 28S, H3, CytB) aligned with MAFFT and then concatenated. Numbers next to nodes are ML bootstrap percentages from RAxML, Bayesian inference (BI) posterior probability, and maximum parsimony (MP) jackknife support values, separated by slashes. Key: * indicates 95% bootstrap/jackknife or greater and 0.95 posterior probability or greater. – indicates the node was not found. Branchiae drawings at terminals indicate presence of arborescent or plicate branchiae; ~ indicates that on segments with two groups of branchiae, the position in Thermopolynoe branchiata is split into anterior and posterior groups, as opposed to upper and lower groups in remaining taxa. The seven paraphyletic groups of Branchinotogluma are highlighted in a yellow-orange gradient. |
Avery S. Hatch, Haebin Liew, Stéphane Hourdez and Greg W. Rouse. 2020. Hungry Scale Worms: Phylogenetics of Peinaleopolynoe (Polynoidae, Annelida), with Four New Species. ZooKeys. 932: 27-74. DOI: 10.3897/zookeys.932.48532
