Revision of Pherecardia Horst 1886 (Annelida Amphinomidae)

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Pigmentation patterns: Pherecardia striata (Kinberg, 1857) Kwajalein, in situ, close-up of anterior ends of two specimens. Photo: S. Johnson. Zoosystema est indexé dans / Zoosystema is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Zoosystema est distribué en version électronique par / Zoosystema is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Zoosystema sont référencés par / Articles and nomenclatural novelties published in Zoosystema are referenced by: – ZooBank® (http://zoobank.org) Zoosystema est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris / Zoosystema is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Geodiversitas, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 691 ZOOSYSTEMA • 2025 • 47 (28) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:E708447E-3F83-4C84-914A-CF1255364654 Salazar-Vallejo S. I., Piotrowski C. N., Harris L. & Paulay G. 2025. — Revision of Pherecardia Horst, 1886 (Annelida, Amphinomidae). Zoosystema 47 (28): 691-720. https://doi.org/10.5252/zoosystema2025v47a28. http://zoosystema.com/47/28 ABSTRACT Amphinomid annelids are classified after the shape of caruncle, the presence and type of branchiae along body segments, and the type of chaetae present. Pherecardia Horst, 1886 was proposed to include species having a large caruncle with a heart-shaped median ridge and lateral lobes. However, generic delineation was not well understood at the time, and some species currently referred to the genus were described in Hermodice Kinberg, 1857 and Amphinome Bruguière, 1789. Pherecardia currently includes four shallow water species: P.maculata Imajima, 2003 from Japan; P.parva Monro, 1924 from the Macclesfield Bank; P.polylamellata de Silva, 1961 from Sri-Lanka; and P.striata ( Kinberg, 1857) from the French Polynesia. Two other species have also been proposed as congeneric: A. bruguieresi de Quatrefages, 1866; and A.formosa de Quatrefages, 1866. The genus is best known for its type species, Pherecardia striata (Kinberg, 1857), a common and widespread, large and readily recognized species characterized by numerous longitudinal dorsal pigmented stripes and recorded across the Indo-Pacific from East Africa to the East Pacific. This species has been regarded as relevant for coral reef ecology since it attacks injured predatory crown-of-thorns starfish (Acanthaster planci (Linnaeus, 1758)). Here we review the genus based on examination of type and additional material from 10museum collections and mitochondrial DNA comparisons of representative specimens from various localities. We recognize three species: P.striata, P.distincta (Hoagland, 1920) (new combination, including P.polylamellata as synonym), and P.maculata (retained pending the study of topotypical specimens). Pherecardia parva is transferred to Pherecardites Horst, 1912 and renamed as Pherecardites monroi because of homonymy with Pherecardites parva Horst, 1912. Amphinome bruguieresi is regarded as a junior synonym of Eurythoe indica (Schmarda, 1861); the status of A.formosa cannot be determined. Keys are included for species of Pherecardia and Pherecardites. Sergio I. SALAZAR-VALLEJO El Colegio de la Frontera Sur, Avenida Centenario km 5.5, Unidad Chetumal (México) [email protected] (corresponding author) Christina N. PIOTROWSKI Collections Manager of Invertebrate Zoology, California Academy of Sciences San Francisco, 55 Music Concourse Dr, California (United States) Leslie HARRIS Collection Manager, Allan Hancock Foundation Polychaete Collection Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California (United States) Gustav PAULAY University of Florida, Natural History Museum Gainesville, 3215 Hull Rd., Florida (United States) Submitted on 4 November 2024 | Accepted on 16 March 2025 | Published on 6 November 2025 Revision of Pherecardia Horst, 1886 (Annelida, Amphinomidae) KEY WORDS Caruncle, dorsal pigmentation, notopodia pigmentation, keys, new name, new combination. 692 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. RÉSUMÉ Révision de Pherecardia Horst, 1886 (Annelida, Amphinomidae). Les annélides polychètes amphinomidées sont classées en fonction de la forme de leur caroncule, de la présence et du type de branchies le long des segments du corps, et des types des soies présents. Pherecardia Horst, 1886 a été proposé pour inclure les espèces ayant une grande caroncule avec une bordure médiane en forme de cœur et des lobes latéraux. Cependant, la définition générique n’était pas bien comprise à cette époque, et certaines espèces actuellement référencées dans ce genre ont été décrites dans Hermodice Kinberg, 1857 ou dans Amphinome Bruguière, 1789. Pherecardia inclut actuellement quatre espèces d’eau peu profonde: P.maculata Imajima, 2003 du Japon, P.parva Monro, 1924 du banc Macclesfield, P.polylamellata de Silva, 1961 du Sri-Lanka, et P.striata (Kinberg, 1857) de Polynésie française. Deux autres espèces ont aussi été proposées comme congénères: A.bruguieresi de Quatrefages, 1866 et A.formosa de Quatrefages, 1866. Le genre Pherecardia est mieux connu pour son espèce type, P.striata (Kinberg, 1857), une espèce commune, largement répandue et aisément reconnaissable, caractérisée par les nombreuses rayures pigmentées dorso-longitudinales et recensée dans tout l’Indo-Pacifique, de l’Afrique de l’Est au Pacifique Est. Cette espèce a été considérée comme importante pour l’écologie des récifs de corail, car elle s’attaque aux étoiles de mer “couronne d’épines” (Acanthaster planci (Linnaeus, 1758), quand celles-ci sont blessées. Nous révisons ici le genre sur la base de l’examen du matériel type et de matériel supplémentaire provenant de dix collections muséales, ainsi que de comparaisons d’ADN mitochondrial de spécimens représentatifs provenant de diverses localités. Nous reconnaissons trois espèces: P.striata, P.distincta (Hoagland, 1920) (nouvelle combinaison, incluant P.polylamellata comme synonyme), et P.maculata (conservée en attendant l’étude des spécimens topotypiques). Pherecardia parva est transférée à Pherecardites Horst, 1912 et renommée Pherecardites monroi en raison de l’homonymie avec Pherecardites parva Horst, 1912. Amphinome bruguieresi est considéré comme un synonyme junior d’Eurythoe indica (Schmarda, 1861); le statut de A.formosa reste indéterminé. Des clefs sont incluses pour les espèces de Pherecardia et Pherecardites. MOTS CLÉS Caroncule, pigmentation dorsale, pigmentation dunotopodia, clefs, nom nouveau, combinaisons nouvelles. INTRODUCTION The taxonomy of amphinomid annelids has historically been based on the shape of the caruncle, combined with branchial and chaetal features. Horst (1886: 165) proposed Pherecardia Horst, 1886 with P.lobata Horst, 1886 as its only species, for a poorly preserved specimen of unknown provenance with distinctive caruncle with its median ridge being roughly heart-shaped, and provided the etymology in a foot note as derived from this feature after combining the Greek word phero, bear, carry, bring with the Latin cardio-, from Greek kardia, heart (Brown 1954: 184, 187). Pherecardia resembles Hermodice Kinberg, 1857, another amphinomid genus that includes species with a large caruncle. Several species now placed in Pherecardia, including the senior synonym of the type species, were originally placed in Hermodice, whereas others have been included in Amphinome Bruguière, 1789, another amphinomid genus that includes species of large size but with a rather short, simple caruncle. Pherecardia currently includes four recognized species (Read& Fauchald 2025): P.maculata Imajima, 2003 from the Sagami Sea, Japan; P.parva Monro, 1924 from the Macclesfield Bank, China Sea; P.polylamellata de Silva, 1961 from Sri Lanka, and P.striata (Kinberg, 1857) from Moorea, French Polynesia. Two other species have also been assigned to this genus in the past: Amphinome bruguieresi de Quatrefages, 1866 and A.formosa de Quatrefages, 1866 (Yáñez-Rivera& Salazar-Vallejo 2011). This transfer was tentative because we did not study their type material, and their status needs to be clarified. The most common and best-known species is Pherecardia striata, one of the largest amphinomids. It is commonly 1020cm long and reaches over 50cm in length (Fig.1E, Steve 2012), with large specimens common in Hawaii (Glasby & Bailey-Brock 2001). It is easily recognized by its distinctive color pattern of dorsal longitudinal stripes, interrupted in the intersegmental areas (Kinberg 1910, pl. 12, fig.8B). It has been observed attacking injured crown-of-spines starfishes in the Eastern Pacific (Glynn 1984). The species swarms and undergoes mass spawning in the water column, and large numbers of wasted bodies have been observed washed ashore (Bailey-Brock& Magalhães 2016). Two distinct pigmentation patterns have been observed in Pherecardia striata, suggesting that more than one species may be involved. The typical form has longitudinal stripes of similar length and width across each segment (Fig.1A, B), while stripes in the second form vary substantially in their thickness and length across each segment (Fig.1C-E). Cytochrome oxidaseI (COI) sequence data comparisons indicate that these color forms represent distinct genetic lineages. Acomplex pattern of striping is observed in the anterior region of some specimens (Fig.1E, F). To assess the morphological and genetic diversity within the genus we undertook a broad study of type and non-type specimens. Based on our investigations, three species are recognized in the genus. Pherecardia striata (Kinberg, 1857) is redescribed and restricted; P.distincta (Hoagland, 1920) is transferred from Hermodice to Pherecardia and reinstated, with P.polylamellata de Silva, 1961 included as a synonym; and P.maculata Imajima, 2003 is retained pending the study 693 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) of additional specimens. Pherecardia parva Monro, 1924 is transferred to Pherecardites Horst, 1912 and is redescribed and renamed as Pherecardites monroi n.nom. because it becomes a junior homonym of Pherecardites parva Horst, 1912, the type species of Pherecardites Horst, 1912. Study of the syntypes of A.bruguieresi revealed that it is a junior synonym of Eurythoe indica (Schmarda, 1861), and it is herein redescribed and illustrated. Amphinome formosa lacks type material: its status cannot be fixed. MATERIAL AND METHODS Total length and width of each specimen were measured, the latter at chaetiger 10 including chaetae, and the number of chaetigers were counted. Species are presented in chronological order of description. Species transferred to other genera are treated after Pherecardia and included to provide a placement for them, however the additional genera involved were not revised. Coordinates are A B FG D CE Fig. 1. – Pigmentation patterns of Pherecardia striata (Kinberg, 1857) (A, B, F, G) and P. distincta (Hoagland, 1920) (C, D, E); A, UF 5461; B, Kwajalein, in situ; C, UF 5343; D, Kwajalein, in situ; E, Hawaiian fisherman with a large specimen (and close-up of same); F, G, Kwajalein, in situ, close-up of anterior ends of two specimens (no scales available; photos: A, C: G. Paulay; B, D, F, G: S. Johnson; E, M. Tsukamoto, reproduced with permission). 694 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. included when available directly from collection labels. Photographs were made using a digital camera with an adapter for microscopes, illuminated with combinations of standard and LED microscope lamps. Specimens were temporarily stained with Methyl green or Shirlastain-A, thus some figures appear with a greenish or reddish hue in some plates. Images were stacked with HeliconFocus8 and plates were arranged with PaintShopPro9. Gustafson (1930: 317) noted that amphinomid chaetae include abundant, thin, usually smooth capillaries, and thicker variably ornamented chaetae or aciculars. These latter chaetae can be smooth, bifurcate, denticulate, or present some of these features variably combined; when they are denticulate in a single series, they are referred to as harpoon-chaetae. Aciculae among amphinomids have exposed tips, and these tips are usually subdistally swollen and can be blunt or sharp. In Pherecardia, aciculars can be harpoon-chaetae or have a short series of small distal denticles, tips are bent, often with a shallow constriction (Gustafson 1930: 317, fig.10). Acicular has been used in polychaete keys and glossaries (Fauchald 1977a: 156; Mikkelsen& Virnstein 1982: 2; Glasby etal. 2000: 403) for referring to thick, rod-like protruding chaetae. In Pherecardia, the lateral caruncular lobes show a pattern of branching, which could match blood irrigation, along each caruncular lobe. A central vein or vessel can be pinnately branched along the lobe, and this is herein referred to as venation. Further, in some specimens these areas can be variably swollen such that the caruncular lobe is not smooth, and when this feature was noted, it was referred to as rugose. As in many morphological features, some terms were taken from botany. Botanists refer to leaf venation in general, and they recognize different patterns of venation, being parallel venation and reticulate venation the most common ones (Sack& Scoffoni 2013). Then, for simplicity, this pinnate branching pattern will be referred to as venation whenever they are present and provided as details about caruncular lobes. There are two fleshy, longitudinal outgrowths or ridges running along the peristomium and often reaching the mouth; they have been referred to as palps when the functional palps were regarded as antennal palp or secondary antennae, or as lips. They are not palps (Orrhage 1990), and because they do not surround the mouth they could not be lips either. However, after their shape and proximity to the mouth, they are indicated as lips, and their proximity to the mouth is sometimes included in the descriptions below, especially if not reaching the mouth. An additional feature is the area between notopodia and parapodia, which can include some glands. When present, this feature is referred to as an interramal belt, for emphasizing its position, and if pigmented glands are distinct, it is referred to as interramal belt maculate, or non-maculate if there are no discrete spots in the interramal area. Tissues from some specimens were sequenced for mitochondrial cytochrome c oxidase I (COI) as described elsewhere (Piotrowski etal. 2024; Salazar-Vallejo etal. 2024) and combined with additional sequence data available in GenBank and BOLD (Table1). Sequences were analyzed with MEGA (Tamura etal. 2021) using Maximum Likelihood with 1000 bootstrap replicates, with TN93+I as the best fit model, and Pareurythoe borealis (M. Sars, 1862) as outgroup (Jimi etal. 2022). AbbreviAtions Institutions AM Australian Museum, Sydney; CAS California Academy of Sciences, San Francisco; LACM-AHF Los Angeles County Museum of Natural History, Allan Hancock Polychaete collection, Los Angeles; MAGNT Museum and Art Gallery, Northern Territory, Darwin; MNHN Muséum national d’Histoire naturelle, Paris; NHML Natural History Museum, London; RMNH Rijksmuseum voor Natuurlijke Historie (now NBC: Naturalis Biodiversity Center), Leiden; UF University of Florida, Natural History Museum, Gainesville; ZMA Zoological Museum, University of Amsterdam (this collection is now housed at NBC); ZMH Zoological Institute, University of Hamburg, Hamburg. Table 1. — COI sequences used in this study. Species Voucher Field Number GenBank BOLD Process ID Locality Source Pherecardia striata UF Annelida 5438 BKON-0952 MW278193 KANBI670-19 Oahu Kaneohe Bioblitz Pherecardia striata UF Annelida 5461 BKON-1328 MW278360 KANBI894-19 Oahu Kaneohe Bioblitz Pherecardia distincta UF Annelida 5343 BKON-0123 MW277733 KANBI074-19 Oahu Kaneohe Bioblitz Pherecardia distincta UF Annelida 5420 BKON-0638 MW278005 KANBI427-19 Oahu Kaneohe Bioblitz Pherecardia striata UF Annelida 2125 BMOO-14438 KC706793 GBAHO352-15 Moorea Moorea Biocode Pherecardia striata no voucher JF905665 GBAN5127-13 – Moorea Leray et al. 2013 Pherecardia cf. striata no voucher MZ568412 GBMNF499-22 – Japan? Jimi et al. 2022 Pherecardia striata UF Annelida 545 BMOO-00384 – MBMIE014-07 Moorea Moorea Biocode Pherecardia striata UF Annelida 523 BMOO-00702 – MBMIE033-07 Moorea Moorea Biocode Pherecardia striata UF Annelida 1269 BMOO-08306 – – Moorea Moorea Biocode Pherecardia striata UF Annelida 2312 BMOO-14426 – – Moorea Moorea Biocode Pherecardia striata no voucher BMOO-13917 – – Moorea Moorea Biocode Pherecardia striata CAS 187536 none – – Philippines This study Pherecardia distincta CAS 185402 none – – Philippines This study Pareurythoe borealis SIO BIC A2379 JN086550 GBAN6186-14 – Norway Borda et al. 2012 695 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) RESULTS Family AmphinomidAe Savigny in Lamarck, 1818 Subfamily AmphinominAe Savigny in Lamarck, 1818 Genus Pherecardia Horst, 1886 Pherecardia Horst, 1886: 165; 1909: 299 (syn.); 1911: 17-21 (syn.).— Fauchald 1977a: 102-103 (key and diagn.). Eucarunculata Malaquin & Dehorne, 1907: 358 (type species: E.grubei Malaquin and Dehorne, 1907, by monotypy). type species.— Pherecardia lobata Horst, 1886, by monotypy; junior synonym of Hermodice striata Kinberg, 1857. diAgnosis (modif. After horst 1886).— Amphinominae with median and lateral antennae and palps. Caruncle large, median ridge heart-shaped with several lateral digitate to foliose lobes. Branchiae present in all segments, bushy. Notochaetae capillaries and aciculars, smooth or serrated (harpoon-chaetae). Neurochaetae aciculars nonfurcate, often distally denticulate, hastate (tips bent). composition We recognize three species in Pherecardia below: P.striata, P.distincta, and P.maculata. The last is known only from a single small specimen and additional material is necessary to evaluate and better characterize it. The first two species were previously confused and regarded as synonyms but are readily differentiated by color pattern and DNA sequence data (Fig.8). Both range across most of the tropical Indo-Pacific to the American coasts. Pherecardia striata is evidently much more common. Perusing images in iNaturalist identified as Pherecardia, we note that only 18 pertain to P.distincta against 153 for P.striata, and one specimen from Timor-Leste was blue, probably after the presence of eggs (accessed 22.VII.2024). remArks Horst (1886: 165) diagnosed Pherecardia as having a caruncle with “a median, heart-shaped portion (= median ridge), which bears on each side several folded lobes” but he did not compare it against other genera. Pherecardia is similar to Hermodice but can be distinguished by caruncle shape. In Pherecardia the caruncle lateral lobes are separate from each other, often fused to the median ridge, whereas in Hermodice they are fused forming a single plate, and the median ridge is indistinct. Malaquin & Dehorne (1907: 360) differentiated Pherecardia (under the synonym Eucarunculata) from Hermodice Kinberg, 1857 by indicating that the caruncle lobes “are aligned such that they are convergent anteriorly. This is contrary to the caruncle arrangement in Hermodice, where the lateral lobes are convergent posteriorly along the median ridge.” Because Hermodice is restricted to the Atlantic and Mediterranean, whereas Pherecardia is found in the Indian and Pacific Oceans, there were no comparative illustrations until Hartman (1951: 24) included anterior ends of representatives of each genus. Malaquin& Dehorne (1907) also provided a detailed description of the caruncle and dorsal pigmentation and noted that anterior eyes were larger than posterior eyes. Amphinomids rarely evert their pharynx fully, as this organ does not need to be completely everted for capturing preyitems, but because of some specimens under stress evert a fairly large portion of the anterior gut, and even if the everted fraction constitutes the whole pharynx, it is rarely exposed for capturing food particles (Salazar-Vallejo 2023). In Pherecardia, the pharynx exhibits two morphological patterns, but the second may be a fixation artifact. Well-preserved specimens (Fig.2A, B) exhibit a roughly spoon-shaped pharynx with granular papillae present on the dorsal depressed region. The mouth opens in the center, with a distal furrow separating lateral cushions comprised of a series of crests running ventrally, which become slightly rugose mid-ventrally. The second pattern was observed in poorly preserved specimens, where the pharynx includes three rings: 1) a short outer yellowish oval ring with a thin mid-dorsal blade (Fig.2C, D); 2) a median pale ring with a granulose surface that is 3-4 times longer than the outer ring; and 3) an inner ring with a thinner wall that may be twice as long as the median one. There is one video showing a P.striata capturing a crab, but the pharynx is not shown (van Antwerp 2013). In Hermodice, the short cylindrical pharynx can be everted for capturing prey items offered by hand (Kosemen 2013), and in another video of a Chloeia Savigny in Lamarck, 1818, it eats a dying anchovy from the tail by sucking it up, and the pharynx is not exposed at all, although it can be expanded laterally for ingesting the head (MaverickDiving 2022). As is the case in some other amphinomids with indefinite growth, the number of segments in Pherecardia specimens is not diagnostic. The following morphological features are not useful for separating species of Pherecardia. The number of lateral lobes in the caruncle, which has been applied to sepakey to species of Pherecardia horst, 1886 (type localities after the species name) 1. Dorsum with numerous longitudinal stripes in each segment; dorsal cirri and branchiae colorful ................ 2 — Body without patterned pigmentation; dorsal cirri and branchiae pale ........ P.maculata Imajima, 2003 Japan 2. Longitudinal stripes more or less regular, of similar width and length; notopodia with anterior surface darker than posterior surface, without complete blackish ring; notochaetae very abundant and larger than dorsal cirri ..... .................................................................................... P.striata (Kinberg, 1857) restricted, French Polynesia — Longitudinal stripes irregular, of varied width and length; notopodia with distinct complete blackish ring; notochaetae moderately abundant, shorter than dorsal cirri ................................................................................ P. distincta (Hoagland, 1920) reinst., n. comb. Philippines (includes P.polylamellata de Silva, 1961 Sri Lanka). 696 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. rate species in the past, is problematic as it varies in number and complexity of caruncle lateral lobes due to growth and the caruncle is frequently altered by trauma (Fig.3). The fine denticulation of neurochaetae can vary within the same parapodium, however this character must be used with caution because denticles may be damaged or corroded by fixative, resulting in the reduction or loss of subdistal denticles (de Silva 1961: 171). Consequently, these two features should be avoided to diagnose taxa. Pherecardia striata (Kinberg, 1857), restricted (Figs1A, B, F, G; 2A, B; 3; 4; 6-8) Hermodice striata Kinberg, 1857: 13; 1910: 35 (diagn. repeated), pl. 12, fig.8.— Ehlers 1920: 13-14.— Augener 1927: 122 (partim); 1933: 188. Pherecardia lobata Horst, 1886: 165-166, pl.7, figs10-14; 1909: 300 (syn.); 1911: 17-21 (syn.); 1912: 32-33 (syn.)— Collin 1902: 741.— Fauvel 1919a: 336; 1919b: 349-350.— Day 1934: 27.— Bleeker & van der Spoel 1992: 152. Amphinome sericata Fischli, 1903: 95-98, pl.4, figs1, 2, pl.7, figs45-49, pl.8. figs79-80. Hermodice pennata Treadwell, 1906: 1165, fig.41 (junior syn. P.striata fide Hartman 1956: 251). Eucarunculata grubei Malaquin& Dehorne, 1907: 358-361, pl.51, fig.2, pl.53, figs12-15, 17-20.— Potts 1909: 365-366.— McIntosh 1925: 13-14, pl.1, figs3-4. Eucarunculata grubei var. gracilis Potts, 1909: 366-367. Eucarunculata grubei var. minuta Potts, 1909: 367. Pherecardia striata – Monro 1928: 77; 1933: 7.— Fauvel 1935: 292-293 (partim); 1936: 257.— Okuda 1937: 265-266, fig.5.— Monro 1939: 166-167 (epitokes).— Hartman 1948: 46; 1966: 181-182.— Day 1957: 67 (syn., partim); 1962: 636; 1967: 131, fig.3.2p-t.— Fauchald 1977b: 13.— Blake 1991: 84.— Imajima 2005: 86-88, fig.36A-G. — Cañete 2017: 199-201, Fig. 1. diAgnosis.— Pherecardia with dorsum bearing longitudinal stripes of similar width along body. Notopodia darker along anterior surface, posterior surface paler, not forming a complete blackish ring. Notochaetae abundant, longer than dorsal cirri along anterior chaetigers. type mAteriAl.— Holotype of Hermodice striata Kinberg, 1857. French Polynesia • 1specimen; Moorea; KSF Eugenies Expedition; Sta. 1245; coral gravel; SMNH 6080. Holotype of Pherecardia lobata Horst, 1886. Locality uncertain• 1specimen; RMNH VER.1295 (locality given as Ambon Bay, III.1901, R. Semon leg. on label; the holotype specimen matches the original description, the label indicates that it was collected in 1901, 15 years after the original description. Horst (1886) noted that the provenance of the specimen was uncertain. Thus, the locality (Ambon Bay) on the label is a later addition and the type locality must be considered uncertain). Holotype of Hermodice pennata Treadwell, 1906. Hawaiian Islands • 1specimen; USFS Albatross; Sta. 4162; Nihoa Island; 3844m; 8.VIII.1902; USNM 5209. AdditionAl mAteriAl.— Kenya • 1specimen; Mombasa; D.MacGregor leg., no further data; NHML 1961.8.32 (complete, longitudinal stripes well defined dorsally, ventrally along anterior body half; notopodia without black rings; caruncle with eight pairs of lateral lobes [last one tiny]; median antenna without tip, 1/3 as long as caruncle; pygidium with anus terminal, anal plate truncate, probably in regeneration; body 62mm long, 9mm wide, 62 chaetigers). Madagascar • 1specimen; Nossy Vorona; off ENE side; 13°25’27”S, 48°21’15”E; 4m depth; broken reef, extracted from large, dead Porites; 15.V.2008; A. Anker, G. Bakary, E. Boissin, H. Bruggemann, T.Horeau, F. Michonneau, G. Paulay& T. Werner leg.; UF 750 (complete; dorsal longitudinal stripes of similar width, often branching; caruncle, interramal areas and venter maculate; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, with seven pairs of lateral smooth lobes, with venation; notopodia with anterior surface darker, without distal black rings; pygidium with anus terminal, anal plate truncate, barely notched; 83mm long, 8mm wide, 64 chaetigers) • 1specimen; Nosy-Be; 7-26.V.2008; G. Paulay etal. leg.; UF 4230 (complete; dorsal longitudinal stripes of similar width, a few branching; caruncle interramal areas and venter maculate; anterior eyes three times as large as posterior ones, not protruded; caruncle with median ridge complete, with eight pairs of lateral smooth lobes, with venation; notopodia with anterior surface darker, without distal black rings; pygidium with anus terminal, anal plate truncate, barely notched; 108mm long, 10mm wide, 73 chaetigers). Mascarene Islands, La Réunion • 1specimen; Saint Leu, Maison Verte, Cimetière; 21°11’36”S, 55°16’56”E; 8-9m depth, fore reef, within dead digitate Acropora; 13.VIII.2007; H. Bruggemann, N.Hubert, F. Michonneau& G. Paulay leg.; UF 654 (anterior fragment; dorsal longitudinal stripes of similar width, a few branching; not maculate; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, with 7-8 pairs of lateral lobes, barely pinnate; notopodia with anterior surface darker, without distal black rings; 26mm long, 7mm wide, 26 chaetigers) •3specimens; Saint Leu, Sec Jaune; 21°9’11”S, 55°16’51”E; 6-15m depth; 17.VIII.2007; H. Bruggemann, N. Hubert, F. Michonneau& G.Paulay leg.; UF 664 (two complete, one without posterior region; dorsal longitudinal stripes of similar width, a few branching; not maculate; interramal belt and venter maculate; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, twisted, with seven pairs of lateral smooth lobes; notopodia with anterior surface darker, without distal black rings; pygidium with anus terminal, anal plate truncate, margin entire; 33-34mm long, 7-8mm wide, 50 chaetigers). Indonesia • 1specimen; Lesser Sunda Islands; RV Siboga Expedition; Sta. 60; Timor, Samau Island, Haingsisi; 23m depth, reef, Lithothamnion in 3m and less; 27-28.IV.1899; ZMA V.POL.1070.1 (complete, slightly twisted, breaking in two from chaetiger 17; some chaetae from chaetigers 10 and 25 removed for observation; dorsum with rather regular longitudinal stripes of similar width; caruncle with 6-8 lateral lobes with venation, not rugose; branchial filaments basally darker, paler distally; pygidium with anus terminal, anal plate truncate; body 53mm long, 7mm wide, 48 chaetigers) • 1specimen; Maluku; RV Siboga Expedition; Sta. 240; Banda anchorage; 9-45m depth; trawl, black sand and coral; 22.XI–1.XII.1899; ZMA V.POL.1070.2 (bent ventrally; juvenile; some parapodia previously removed [lost?], breaking in two parts; dorsum with longitudinal stripes of similar width; caruncle with 5-6 lateral lobes with venation, not rugose; branchial filaments basally darker, tips paler; pygidium with anus terminal, anal plate wider than long, medially notched; body 19mm long, 4mm wide, 35 chaetigers) •1specimen; Lesser Sunda Islands, RV Siboga Expedition; Sta. 300; 10°48.6’S,123°23.1’E; 918m; trawl, muddy bottom; 30.I.1900; ZMA V.POL.1070.4 (bent laterally, juvenile; dorsum with longitudinal stripes of similar width; caruncle with 5-6 lateral lobes with venation, not rugose; branchiae basally darker, tips paler; pygidium with anus terminal, anal plate slightly longer than wide, round; body 19mm long, 4mm wide, 35 chaetigers) • 1specimen; Ambon Island; 3.II.1902; V. Kamper leg.; RMNH VER.1296 (partially dried, with salt spherules over chaetae and branchiae; dorsum with rather regular longitudinal stripes; venter with paler longitudinal stripes, less regular; caruncle 697 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) in regeneration; eyes black, not protruded from prostomial margin; caruncle with lateral lobes with venation, not rugose; branchiae dark with tips pale; pygidium with anus terminal, anal plate truncate; body 93mm long, 13mm wide, 59 chaetigers) • 25 specimens; Ambon Island; 1887; H. Brock leg.; ZMH PE29 (17 complete; most grayish with purple to blackish longitudinal rather regular stripes, other brownish with more irregular streaking; caruncle lobes foliose, slightly indented, 6-10 pairs; eyes not protruded from prostomial margins; median antenna 1/4 to 2/3 as long as caruncle; pygidium with anus terminal, anal plate lobate, often slightly notched medially; complete with body 16-70mm long, 6-13mm wide, 3245 chaetigers • 1specimen; Ambon Island; 1887; H. Brock leg.; ZMH PE30 (soft, complete, unpigmented [this has been retained in P.striata with hesitation after it was found in the type locality of P.lobata Horst, 1912, which is a junior synonym of P.striata, but the fact that this was collected in the same locality as the type does not preclude it being something else]; left parapodia of chaetigers 24 and 25 previously removed [kept in container]; caruncle with eightpairs of lateral lobes; median antenna half as long as caruncle; pygidium with anus terminal, anal plate truncate; body 50mm long, 10mm wide, 52 chaetigers). East Timor • 1specimen; Timor, between Timor and Nusa Besi; anchorage; RV Siboga Expedition; Sta. 282; 08°25.2’S, 127°18.4’E; 2754m depth, trawl, sand and coral; 15-17.I.1900; ZMA V.POL.1010.3 A CD B Fig. 2. — Pharynx eversion in two species of Pherecardia Horst, 1886: A, B, Pherecardia striata (Kinberg, 1857), restricted, non-type specimen, UF 31: A, anterior region, right lateral view; B, frontal view after Shirlastain-A staining; C, D, Pherecardia distincta (Hoagland, 1920) reinstated, non-type specimen, NHML 1932.12.24.265–268, after Methyl green staining: C, anterior end, frontal view of pharynx; D, anterior region, left lateral view. Scale bars: A, C, 0.7 mm; B, D, 0.6 mm. 704 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. 65mm long, 7mm wide, and 90 segments. The caruncle was described as ‘very developed, reaching segment 4. Its shape is oval, with 7-8 foliose lobes fused medially by a ridge with a free margin.’ These features match Pherecardia; however, because there is no type specimen the status of the species cannot be determined (see below). Horst (1886) proposed Pherecardia, with P.lobata as its type species, based on a damaged specimen with most of its pigmentation faded away, but he noted that it was ‘marked on its dorsal side with faint longitudinal folds’ which may have corresponded with longitudinal stripes. Details of the prostomium were not provided because it was damaged. A D G IB IB * * HIJ E BC F Fig. 7. — Pherecardia striata (Kinberg, 1857) restricted, COI-sequenced non-type specimens: A, 6 mm wide specimen, UF 5438, anterior region, dorsal view; B, same, left lateral view; C, same, anterior end, dorsal view; D, 8 m wide specimen, UF 5461, anterior region, dorsal view; E, same, right lateral view; F, same, anterior end, dorsal view; G, same, chaetiger 8, left parapodium, anterior view (IB: interramal belt); H, same, after Methyl green staining (asterisks indicate dissected section; IB: interramal belt); I, cross section of interramal band, pigment concentrated in furrows; J, same, upper and lower neurochaetae. Scale bars: A, 1.5 mm; B, 1 mm; C, 0.6 mm; D, 2 mm; E, 0.9 mm; F, 0.7 mm, G, 0.5 mm; H, 0.4 mm; I, 0.1 mm; J, 30 µm. 705 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) Fischli (1903) described Amphinome sericata in detail with several illustrations, and although prostomial details were not given, the color plates portray the dorsum with rather regular longitudinal stripes of similar width, and this explains why Horst (1911) may have synonymized it with P.striata. Fischli lived in Linthal, Switzerland, but whether his type specimens are extent and if so where they are located, could not be clarified. Inquiries to nearby natural history museums about potential types were not answered. Treadwell (1906) proposed Hermodice pennata based on a 15-segment anterior fragment collected in Hawaii. He indicated that pigmentation included numerous longitudinal stripes extending across segments, and that the eyes were ‘prominent on dorsal view’, and his illustration shows them not protruded from the prostomial margin. The caruncle was described and illustrated as having seven smooth lobes per side. The holotype is in two fragments. The caruncle is as illustrated and while the pigmentation is now gone it is indicated in the original description. Based on examination of the type and Treadwell’s description we confirm that this species is a synonym of P.striata, as proposed by Augener (1927). Malaquin& Dehorne (1907) proposed Eucarunculata with E.grubei as its type species without citing any of the previous literature on the above species. They indicated that anterior eyes were slightly larger than posterior ones, and that the animal had 10-15 ‘dark longitudinal stripes running from segment to segment along the whole body.’ Their illustrations show the anterior end with longitudinal stripes of similar width. Horst (1909) indicated that E. grubei was a junior synonym of his P.lobata. We did not locate the type of E.grubei. Based on the description and illustration we agree with Horst that E.grubei is a synonym of P.stricta. Potts (1909) proposed two varieties for E.grubei based on two smaller specimens: E.g. var. gracilis and E.g. var. minuta, both with regular longitudinal stripes, but no type specimens were deposited. He noted in a footnote (page 365) that ‘the differences in form of caruncle and chaetae are probably due to varying age.’ His specimens have not been located, and it is likely that they were not deposited in a museum. The typical form in his treatment included a specimen 104mm long, 7mm wide, and with 73 segments; its caruncle had 6-7 lateral lobes, notochaetae included smooth capillaries and non-spurred harpoon chaetae, neurochaetae were distally falcate, either with subdistal denticles or smooth. His variety gracilis included one specimen 24mm long, and 4mm wide with 41 segments, with 5-6 lateral lobes in the caruncle, and the neurochaetae had ‘more distinct and numerous serrations’ than in the typical form. His variety minuta was based on a specimen 16mm long, 2mm wide, and with 34 segments. He noted that neurochaetae had distinct, variable serrations. Hartman (1959: 133) regarded both as junior synonyms of Pherecardia lobata. We agree with her conclusions. Differences between Pherecardia striata and P.distincta, are mostly in pigmentation pattern, as indicated in the key above. In P.striata the longitudinal stripes are more or less regular, having similar width and length, and the notopodia have a darker pigmentation along anterior surface than on posterior one, whereas in P.distincta the longitudinal stripes are irregular, of different width and length, and the notopodia have a blackish ring. Further, these two species are separated by 22.4-23.5 % K2P distance in COI (Fig.8). Intraspecific variation between the two sequenced species is 0.3-2% in P.striata, and 1.2% in P.distincta. The records by Amoureux (1977: 1095) and Rajasekaran & Fernando (2012: 3) could not be confirmed. Pherecardia distincta (Hoagland, 1920) reinst., n.comb. (Figs1C-E; 2C, D; 8-14) Hermodice distincta Hoagland, 1920: 612, pl.48, figs13-15. Hermodice pennata var. tutuilensis Treadwell, 1926: 2. Pherecardia striata – Fauvel 1935: 292 (partim). — Hartman 1940: 207 (partim).— Pillai 1965: 121, fig.4F-H (non (Kinberg, 1857)). Pherecardia polylamellata de Silva, 1961: 170, fig.4. diAgnosis.— Pherecardia with longitudinal stripes of variable width and length along dorsal surfaces of body. Notopodia with a black distal ring. Notochaetae moderately abundant, shorter than dorsal cirri along anterior chaetigers. t ype mAteriAl .— Holotype of Hermodice distincta Hoagland, 1920. Philippines • 1specimen; Mindanao Island, Misamis Oriental, Macajalar Bay, Suluan Point, Opol; USFS Albatross; unnumbered station; 2-4m depth; after dynamite explosion (original label indicates “EL”, and identification label indicates “Electric Light”), 4.VIII.1909; USNM 18953. Paratype of Pherecardia polylamellata de Silva, 1961. Sri Lanka, Gulf of Manaar • 1specimen; ; Ceylon Pearl Banks; no further data; NHML 1962.14.1. A dditionAl mAteriAl .— Madagascar • 1specimen; Nossy Vorona, off ENE side; 13°25’27”S, 48°21’15”E; 4m depth; broken reef, extracted from large, dead Porites; 15.V.2008; A. Anker, G.Bakary, E.Boissin, H.Bruggemann, T. Horeau, F. Michonneau, G.Paulay& T.Werner leg.; UF 749 (complete; dorsal longitudinal stripes irregular, often branching, thinner towards notopodia; some parapodial areas and branchiae maculate; anterior and posterior eyes of similar size, not protruded; caruncle with median ridge complete, with 6-7 pairs of lateral smooth lobes, with venation; notopodia with distal black rings; pygidium with anus prolapsed, anal plate semicircular, slightly notched; 150mm long, 20mm wide, 74 chaetigers) • 1specimen; Nossy Vorona, off ENE side; 13°25’27”S, 48°21’15”E; 4m depth; broken reef, extracted from large, dead Porites; 15.V.2008; A. Anker, G.Bakary, E. Boissin, H.Bruggemann, T. Horeau, F. Michonneau, G.Paulay& T.Werner leg.; UF 751 (complete; dorsal longitudinal stripes irregular, often branching, thinner towards notopodia; some parapodial areas and branchiae maculate; anterior and posterior eyes of similar size, not protruded; caruncle with median ridge complete, with sevenpairs of lateral smooth lobes, with venation; notopodia with distal black rings; pygidium slightly damaged, anus prolapsed, anal plate globose, margin entire; 90mm long, 14mm wide, 55 chaetigers). Mascarene Islands, La Réunion • 1specimen; Saint Leu, Maison Verte, Cimetière; 21°11’36”S, 55°16’56”E; 8-9m depth; fore reef, 706 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. within dead digitate Acropora; 13.VIII.2007; H. Bruggemann, N.Hubert, F.Michonneau& G. Paulay leg.; UF 655 (anterior fragment; dorsal longitudinal stripes irregular, dorsal ones wider, thinner towards notopodia; not maculate; anterior eyes three times as large as posterior ones, slightly protruded; caruncle with median ridge complete, with seven pairs of lateral pinnate lobes; notopodia with distal black rings; 21mm long, 5mm wide, 28 chaetigers). Gulf of Thailand • 1specimen; Thailand, Chantabun; no further data; MNHN A431 (complete; dorsal longitudinal stripes irregular, especially from chaetiger 12 where some stripes become wider than others; venter with midventral thin bluish line; caruncle with 10pairs of crenulated lobes; median antenna 1/3 as long as caruncle, slightly larger than lateral antennae and palps; eyes large, protruded from prostomial margin; anus terminal, anal plate medially notched; body 103mm long, 15mm wide, 82 chaetigers). Vietnam • 1specimen; Nha-trang; no further data; MNHN A398 (body twisted, anterior end heavily contracted; dorsal stripes fading, especially thinnest ones; midventral line faded; caruncle with 10 pairs of lateral crenulate lobes; median antenna lost, upper lips separate lateral antennae; eyes protruded from prostomial margins; anus terminal, anal plate rounded, not notched; body 62mm long, 15mm wide, 74 chaetigers). Indonesia • 1specimen; Java; “Outer Islands Archipelago”; no further data; ZMA V.POL. 1071 (epitoke, progressively wider medially and posteriorly; dorsal stripes irregular; eyes protruded from prostomial margins; caruncle with 10-11 lateral rugose (pinnate) lobes; median antenna 4/5 as long as caruncle; some median and posterior intersegmental areas broken, exposing inner organs; venter with a blackish, discontinuous midventral line; posterior region in regeneration, pygidium damaged; body 116mm long, 13mm wide, 70 chaetigers) • 4 specimens; Java Bay, near Batavia, Eil Edam; 19.IX.1930; Museum Buitenzorg, J. Versvey leg.; ZMH V12095 (soft, apparently spent epitokes based on the amount of foreign material accumulated on chaetae; longitudinal stripes irregular, with variable width and length, wider mid-dorsally, thinner marginally; stripes visible ventrally along a few chaetigers; eyes protruded from prostomial margins; caruncle with 8-10 pairs of lateral foliose, pinnate lobes; caruncle median ridge base and ridge areolated; median antenna 1/2-2/3 as long as caruncle, slightly longer to twice longer than lateral antennae and palps, both of similar shape and length; body 2850mm long, 8-12mm wide, 45-46 chaetigers • 3 specimens; Poeloe (Pulau) Weh; 1902; G.A.Y. van den Samde leg.; no further data; RMNH VER.1297 (twisted, two without posterior region; two females, ovigerous, one male epitoke; dorsum with purple to blackish longitudinal, rather regular stripes; eyes large, very close to each other on each side, not protruded from prostomial margins; notopodia with distal black rings; male epitoke with caruncle median ridge rugose, with 12-13 lateral foliose lobes; base of median ridge and supraocular areas between posterior eyes with low lobes; females with caruncle median ridge smooth, without additional lobes between posterior eyes; parapodia with many chaetae, capillaries not abundant; female epitoke without pygidium, with egg-sac extruded from last chaetiger; 75mm long, 17mm wide, 77 chaetigers; male epitoke; pygidium tapered, anus terminal, anal plate rounded; 108mm long, 13mm wide, 96 chaetigers). Philippines • 1specimen; Lubang Islands, Bahura Reef, N of Lubang Island; Sta. LB-003; 13.0°51.0’53.3”N, 120.0°11.0’31.6”E; 0-5m depth ; 6.IX.2011; B. Moore leg.; CASIZ 185402 (complete; dorsal longitudinal stripes irregular, often branching; notopodia with distal black ring; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, with 7-8 lateral pinnate lobes; pygidium with anus prolapsed, anal plate truncate, entire; 86mm long, 9mm wide, 71 chaetigers). Mariana Islands • 1specimen; Guam; right side of Cocos Island; no depth data; rubble; 23.IV.1999; G. Paulay leg.; UF 22 (complete, regenerating posterior segments; pharynx fully exposed; dorsal longitudinal stripes of differing width along segments, lateral stripes thinner; integument without maculae; notopodia with distal blackish rings, better defined along anterior segments; anterior eyes slightly larger than posterior eyes, barely protruded from prostomial margins; caruncle pale, with median ridge complete, with 11 pairs of lateral pinnate lobes; median antenna without tip, half as long as caruncle; pygidium with anus terminal, anal plate indistinct; 112mm long, 20mm wide, 74 chaetigers). Ph. striata MZ568412 Japan Ph. striata JF905665 Moorea Ph. striata UF523 Moorea Ph. striata UF2312 Moorea Ph. striata UF5483 Oahu Ph. striata UF545 Moorea Ph. striata UF2125 Moorea Ph. striata BMOO 13917 Moorea Ph. distincta CAS 185402 Philippines Ph. distinctaUF5343 Oahu Ph. distinctaUF5420 Oahu Pareurythoe borea/is JN086550 Norway 0.2 Ph. striata CAS 187536 Philippines Ph. striata UF1269 Moorea Ph. striata UF5461 Oahu Fig. 8. — Maximum likelihood tree of Pherecardia Horst, 1886 based on COI sequence data; both species have 100% bootstrap support. 707 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) Hawaii • 1specimen; Oahu, Kaneohe Bay, NW end of Ahu’olaki Island; the Sand Bar; 21°28’26”N, 157°49’11”W; 0-2m depth; 22.V.2017; Bioblitz team leg.; UF 5343 (without pygidium, regenerating posterior end, cut into two sections, left parapodium of chaetiger 8 removed for study; longitudinal brown stripes irregular, some wide, others narrow; notopodia with distal brown ring; anterior eyes larger than posterior eyes, protruded from prostomial margin; caruncle whitish, venation barely darker, with 12-14 pinnate lateral lobes, median ridge discontinuous; median antennae 2/3 as long as caruncle; anterior fragment 20mm long, 16mm wide, 13-14 chaetigers; posterior fragment 41mm long, 15mm wide, 28 chaetigers) • 1specimen; Oahu, Kaneohe Bay; 21°30’0”N, 157°48’0”W; 0-20m depth; 21.V.2017; Bioblitz team leg.; UF 5420 (anterior fragment, markedly contracted, bent ventrally; dorsal longitudinal stripes irregular, alternating wide and narrow, thinner stripes towards parapodia; notopodia with distal brown rings; anterior eyes larger, with lenses, not protruded from prostomial margin; caruncle whitish, without median ridge, with 7-8 lateral lobes per side; median antennae without tip, half as long as caruncle; fragment 32mm long, 12mm wide, 39 chaetigers). New Caledonia • 1specimen; Mathew Island, Northwest side; 22°20’29”S, 171°21’1”E; boulders and large rocks on black sand; 5m; 1.VIII.2017; S. Hannam, A. Reid, C. Bedford,& I. Middleton leg.; AM 51661 (juvenile, distorted due to label compression; longitudinal anastomosing stripes dorsally, ventrally visible along a few anterior chaetigers; caruncle grayish, with thin anastomosing stripes, 2/3 as wide as anterior end, with four lateral foliose, rugose lobes; median antenna 1/5 as long as caruncle; chaetae not removed to avoid further damage; body 13.5mm long, 1.5mm wide, 31chaetigers). French Polynesia • 3 specimens; Society Islands, Tahiti, Papeete Harbour; swimming at night; Crossland Pacific Expedition 192324; V.1923; B. Grey leg.; NHML 1941.4.4.146-152 (epitokes, twisted, without posterior region, gametes and debris adhered, stained pink the formalin solution (“formol in which they were preserved pink; under lens this seen to be eggs and sperm” as indicated by Monro 1939: 167); dark purple longitudinal stripes, becoming more irregular medially and posteriorly, visible ventrally; eyes not protruded from prostomial margin; caruncle grayish (pale green after Crossland), contracted, with 11-12 lateral foliose lobes; caruncle median ridge with base smooth; median antennae 1/4 AB CD EF Fig. 9. — Pherecardia distincta (Hoagland, 1919) n. comb., holotype, USNM 18953: A, anterior region, dorsal view (anterior end bent ventrally); B, anterior end, dorsal view, after Methyl green staining; C, chaetiger 20, left parapodium, anterior view; D, same, harpoon-notochaetae, neurochaetae, neuracicula; E, chaetigers 68-74, dorsum and left parapodia, seen from above, showing remaining pigmentation; F, posterior region, dorsal view. Scale bars: A, 1.8 mm; B, 0.5 mm; C, 0.6 mm; D, 80 µm; E, 1.9 mm. 708 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. as long as caruncle; eyes larger than in non-epitokous specimens, of similar size; one parapodium of each specimen removed for comparing cirri development; notopodial lobes blackish, dorsal cirrophores 3times wider and about as long as cirrostyles; body 54-165mm long, 13-20mm wide, 52-73 chaetigers) • 5 specimens; Society Islands, Tahiti, Crossland Pacific Expedition, 192324; C. Crossland leg.; no further data; NHML 1941.4.4.153-157 (3complete, 2 without posterior end; longitudinal brownish stripes along dorsum, less regular posteriorly or ventrally; notopodial lobes blackish; caruncle with fine stripes along ridges of lateral lobes, with 8 lateral foliose lobes; median antenna 3/4 to 4/5 as long as caruncle; posterior end tapered, anus terminal, anal plate short; body 56-76mm long, 3.5-6.0mm wide, 53-54 chaetigers). Panama • 8 specimens; Coiba National Park, Isla Coibita, North of Smithsonian beach house, small islet connected by causeway; 07°38.036’N, 81°41.847’W; subtidal, dead Pocillopora; 16.III.2005; L. Harris& I. Wehrtmann leg.; LACM-AHF 14779 (complete specimens, some regenerating anterior or posterior ends; irregular longitudinal stripes dorsally and black rings in notopodia; median antenna as long as caruncle in smallest and largest specimens; body 22-78mm long, 4-9mm wide, 34-61 chaetigers) • 2 specimens; Taboga Island, Taboga Channel; with an associated polycladid flatworm, 19.II.2007; A. Anker leg.; LACM-AHF 14780 (irregular longitudinal stripes dorsally and black rings in notopodia; largest one regenerating posterior region; body 30-57mm long, 7-12mm wide, 40-45 chaetigers) • 2 specimens; Secas Islands; RV Velero III; Sta. 454; 07°57’10”N, 82°00’45”W; shore, tide flats, coral; 6.II.1935; LACM-AHF 14781 (complete specimens; irregular longitudinal stripes and black rings in notopodia; median antenna 1/2-3/4 as long as caruncle; caruncle with eight pairs of lateral lobes; body 45-72mm long, 7-10mm wide, 47-52 chaetigers) • 1specimen; Gulf of Chiriqui, Coiba National Park; Isla Coibita, SE of Smithsonian beach house; causeway connecting third island; 7°41’38”N, 81°48’28”E; in dead Pocillopora; 16.III.2005; L. Harris& I. Wehrtmann leg.; LACM-AHF 14782 (fragmented in two pieces, anterior end in regeneration; irregular longitudinal stripes, notopodia with black ring; not measured) • 7 specimens; Secas Islands; Panamá; RV Velero III; Sta. 447; 07°57’10”N, 82°00’45”W; shallow water, coral; 4.II.1935; LACM-AHF 14783 (complete, some regeneration anterior or posterior regions, with little or no pigmentation, some specimens with asymmetrical caruncles; irregular longitudinal stripes dorsally and black rings around notopodia; body 26-92mm long, 5-11mm wide, 30-66 chaetigers) • 2 specimens; Gulf of Chiriqui; Uva Island; Canteras Islands; 3m depth; feeding on Acanthaster; 22.VI.1978; P.W. Glynn leg.; LACM-AHF 14784 (complete, heavily contracted, bent ventrally; irregular longitudinal stripes, and black ring around notopodia; body 45-54mm long, 10-12mm wide, 42-48 chaetigers) • 3 specimens; Taboga; DrTh.Mortensen’s Pacific Expedition 1914-1916; shore at low tide; T. Mortensen leg.; NHM 1928.9.13.15-17 (complete specimens, regenerating posterior region; smallest one smashed medially; median antenna 1/4 to 1/3 as long as caruncle (tips lost); caruncle with 6-7 to 7-8 pairs of lateral lobes; pygidium with anus terminal, anal plate round or truncate (barely bifid); body 68-105mm long, 8-10mm wide, 51-61 chaetigers) • 2 specimens; Taboga; from colony of Pocillopora; S.Y. St. George Expedition 1923-1924; C. Crossland leg.; NHML 1932.12.24.269-270 (complete specimens; largest bent laterally; longitudinal stripesbest preserved in largest specimen; median antenna 4/5 as long as caruncle; caruncle with 7-8 or 9-10 pairs of lateral lobes; pygidium with anus terminal, anal plate round; body 29-72mm long, 5-9mm wide, 37-55 chaetigers). Colombia • 4 specimens; Gorgona; S.Y. St. George Expedition 1923-1924; from coral; C. Crossland leg.; NHML 1932.12.24.265268 (soft, without posterior end, two with duplicate parapodia at chaetigers 10/11, one with pharynx exposed; dorsal longitudinal stripes better preserved in largest specimen; median antenna nearly as long as caruncle; caruncle with 7-8 or 9-10 pairs of lateral lobes; body 31-70mm long, 5-12mm wide, 35-43 chaetigers). other mAteriAl. — Maldives • https://www.inaturalist.org/ observations/150011379 southern Maldives (approximately 2°1’52”N, 73°31’13”E), (longitudinal stripes brownish, irregular, vary in width and shape along dorsal segments; blackish ring in notopodia distinct). description of type specimens Holotype of Hermodice distincta (USNM 18953) complete, anterior end damaged, with an anteroventral dissection, folded over in two posterior sections; right parapodia of chaetigers 6 and 14 previously removed (lost), left parapodium of chaetiger 20 removed for observation (retained in container). Dorsum with longitudinal stripes faded almost completely, wider stripes medially, thinner stripes lateral (Fig.9A). Dorsal and ventral cirri exposed, tapered, pale. Many chaetae broken. Venter with one thin, almost complete black line. Body 132mm long, 12mm wide, 87 chaetigers. Prostomium bent ventrally, ovoid, wider than long, anterior margin and longitudinal lips entire, ventrally restricted to chaetiger 1 (ventral dissection reaching its anterior margin). Lateral antennae and palps of similar length. Median antenna directed anteriorly, tip broken (Fig.9B). Eyes black, of similar size, protruded from prostomial margins. Mouth opening between chaetigers 2-3(4), damaged after ventral dissection. Caruncle pale, reaching chaetiger 3, 2/3 as wide as anterior end; median ridge cut along caruncle length, with 10 pairs of lateral foliose lobes, blades pinnate. Parapodia biramous; notopodia cylindrical with distal black ring, including chaetae and dorsal cirri (Fig.9C), barely visible dorsally (Fig.9E) in markedly bent portions along posterior regions. Dorsal cirrophores about as long as cirrostyles throughout body. Branchiae arborescent, each with 3-4 short main stems, with abundant filaments, progressively reduced in number posteriorly. Notochaetae including harpoon-chaetae (Fig.9D, left), and abundant smooth capillaries. Neuropodial lobes subcylindrical, neurochaetae arising directly from neurochaetal lobe. Ventral cirri with cirrophore 1/3 as long as cirrostyle in anterior and median chaetigers, 1/4 as long in posterior ones. Abundant capillary neurochaetae, acicular spines with slightly bent tips, some with tiny subdistal denticles (Fig.9D, center); neuraciculae subdistally swollen, tapered (Fig.9D, right). Posterior region tapered (Fig.9F); anus terminal, anal plate minute, rounded. Paratype of P.polylamellata (NHML 1962.14.1) complete, regenerating posterior region; body brownish with irregular, longitudinal stripes barely visible along anterior chaetigers (Fig.10C); palps and dorsal cirrostyles blackish; thin darker line midventrally along body; first right parapodium, notopodia of chaetiger 13 and 28, and neuropodia of chaetiger 15 and 28 previously removed; right parapodium of chaetiger 25 removed for observation; body 195mm long, 15mm wide, 91 chaetigers. Prostomium ovoid, wider than long; lateral antennae positioned on round projections, ceratostyles lost, ceratophores separated by dorsal longitudinal ‘lip’ projection reaching about half prostomial length (Fig.10B, restricted to chaetiger 1 ven- 709 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) trally. Palps biarticulate, positioned in a constriction between the projection of dorsal lips. Median antenna lost. Eyes black, globular, anterior eyes slightly smaller than posterior ones, both protruded from prostomial margins. Mouth opening between chaetigers 2 and 4. Caruncle pale, nearly reaching chaetiger 5, 1/3 as wide as anterior end; median ridge slightly damaged, extending throughout caruncle length (Fig.10A), with 10-11 lateral foliose lobes, corrugated (pinnate). Parapodia bearing arborescent branchiae with abundant filaments from chaetiger 1 to posterior segments, as long as dorsal cirrophores (Fig.10C). Notopodia cylindrical with dorsal cirrophores 4-5 times wider and slightly shorter than cirrostyles, notochaetae rise from a protruded cone, surrounding dorsal cirrophores with dorsal cirri at apex of the cone (Fig.10D); notopodia with a barely visible distal black ring (Fig.10E). Neuropodial lobes subcylindrical, neurochaetae arising from between these lobes. Ventral cirri with A CD E * MR B Fig. 10 . — Pherecardia distincta (Hoagland, 1920) n. comb., paratype of P. polylamellata de Silva, 1961, NHML 1962.14.1: A, anterior region, dorsal view; B, anterior end, after Shirlastain-A staining (MR: median ridge); C, chaetigers 13-15, dorsal view; D, chaetiger 25, right parapodium, posterior view, after Methyl green staining; E, same, after Shirlastain-A, enlarged (*: slightly darker distal ring in dorsal base of cirrophore). Scale bars: A, 1 mm; B, 0.4 mm; C, 1.4 mm; D, 1.1 mm; E, 0.9 mm. 710 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. cirrophores globose, about twice wider than and 1/4-1/5 as long as cirrostyle. Chaetae broken, damaged because of prolonged exposure to formalin (de Silva 1961: 171). Posterior end tapered; anus dorsal, extended along last three chaetigers; anal plate round. description of epitokes Three epitokes (NHML 1941.4.4.146–152), showing little modifications for swimming along median chaetigers (Fig.11); notopodia and neuropodia massive, with dorsal cirri slightly longer than ventral cirri, ceratophores and ceratostyles cylindrical; notopodia with black distal rings, visible once branchiae contract and expose the tips of notopodia (Fig.11A) or if branchiae are displaced (Fig.11B, C). vAriAtion Specimens with well-preserved color pattern have wider longitudinal stripes along mid-dorsal region and thinner stripes laterally, together with distinctive black rings in the notopodia. This pattern is not size-dependent; the same characters are observed in small (Fig.12A), medium (Fig.12C), and large (Fig.12E) specimens, and occur along the body, although longitudinal stripes become progressively thinner posteriorly, where they become of similar width along each segment; this pattern is shown in small (Fig.12B), medium (Fig.12D), and large specimens (Fig.12F). The caruncle has 8-10 foliose lobes, and small specimens exhibit distinctive venation along each lobe (Fig.13B), with pigmented areas transitioning into furrows such that the regular rugosity results in a pinnate appearance to each lobe (Fig.13B, E). Another variations are found in eye and caruncular development. A small 28mm long specimen (Fig.13A) has dorsal longitudinal stripes of differing length and width along anterior segments. The prostomium has black eyes protruded from the prostomial surface (Fig.13B), and despite its size the caruncle has nine pairs of lateral lobes. A larger, 50mm long specimen has the same pattern of irregular dorsal longitudinal stripes (Fig.13D), eyes more clearly protruded from the prostomial surface (Fig.13E), and the caruncle has 11-12 pairs of lateral lobes. The posterior region in both specimens (Fig.13C, F) share similar pigmentation patterns, with discontinuous longitudinal striping and thinner stripes near the parapodia, as shown in other specimens referred to above. The number of longitudinal dorsal stripes decreases with body size, and stripes are thinner in smaller specimens (Fig.14A), becoming wider and more irregular, and often stripes are narrower adjacent to parapodia (Fig.14D); however, the unpigmented spaces between stripes are obviously narrower near parapodia in all cases figured (Figs13D; 14D). The prostomium is more easily observed in the larger specimen with anterior eyes protruded from the prostomial surface. The caruncle has lateral pinnate lobes, but in the ABC Fig. 11. — Pherecardia distincta (Hoagland, 1920), reinstated, non-type, epitokes, NHML 1941.4.4.146–152: A, epitoke 54 mm long, 13 mm wide, chaetiger 25, right parapodium, posterior view; B, epitoke 130 mm long, 18 mm wide, chaetiger 32, left parapodium, posterior view; C, epitoke 165 mm long, 20 mm wide, chaetiger 27, left parapodium, posterior view. Scale bars: A, 0.5 mm; B, 0.8 mm; C, 1 mm. 711 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) two specimens figured in Figure 14 the median ridge is not fully developed and is even missing in the smaller specimen, such that counting the lobes is problematic; there are about eight lobes in the smaller specimen (Fig.14C), and about 14 in the larger one (Fig.14F), in which the posterior median ridge is visible but the middle section is missing. The interramal belt is indistinct (Fig.14B, E), with this area smoother than in P.striata. Branchiae are abundant and although they have paler tips, they are not as dark as in P.striata. Notopodia are protruded muscular cylinders, each with a distinct blackish distal ring (Fig.14B, E). Notochaetae are abundant but shorter and less numerous than in P.striata, surrounding the dorsal cirri upon notopodial contraction, with dorsal cirri having dorsal cirrophores as long as the cirrostyles or up to twice longer. The interramal belt is less plicate with no maculae (Fig.14H, I) as indiACE BDF Fig. 12. — Pherecardia distincta (Hoagland, 1920), reinstated, non-type complete specimens, NHML 1941.4.4.153–157: A, specimen 56 mm long, 3.5 mm wide, anterior region, dorsal view; B, same, posterior region; C, specimen 62 mm long, 7 mm wide, posterior region, dorsal view; D, same, posterior region, dorsal view; E, specimen 76 mm long, 6 mm wide, anterior region, dorsal view; F, same, posterior region, dorsal view. Scale bars: A, 0.9 mm; B, F, 0.5 mm; C, 1 mm; D, 0.7 mm; E, 1.2 mm. 712 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. cated above, no large glands are visible in the corresponding body wall. Neurochaetae include denticulate capillaries and aciculars with a small distal tooth. Subdistal denticles are barely visible, even in the upper chaetae (Fig.14J). remArks Pherecardia distincta can be distinguished from P.striata as indicated above. The main differences rely in the pigmentation pattern; in P.distincta the longitudinal stripes are irregular, varying in width and length, and notopodia have a blackish ring, whereas in P.striata the longitudinal stripes are rather regular, having similar width and length, and notopodia have a darker pigmentation along their anterior surfaces than along posterior ones. In addition, these two species are separated by 22.4-23.5% K2P distance in COI (Fig. 8). Intraspecific variation between the two sequenced P.distincta is 1.2%, while it ranges between 0.3-2% in P.striata. Pherecardia distincta (Hoagland, 1920) was described as having irregular blue-gray streaks and a dorsal chaetal “bundle borne on stout darkly pigmented lobe, also bearing dorsal cirrus.” Hoagland’s illustration of the anterior end shows eyes protruded from the prostomial margins, median antenna slightly longer than the laterals or palps, and the caruncle with 10 pairs of lateral, pinnate lamellae. Two of the labels with the holotype indicate the specimens were caught with electric light, suggesting that it was a swimming epitoke. Soon after its description, Augener (1927: 123) regarded P.distincta as a junior synonym of P.striata, because he found A DE F BC Fig. 13. — Pherecardia distincta (Hoagland, 1920), non-type specimens, ZMH V12095: A, smallest specimen (28 mm long), anterior region, dorsal view; B, same, close-up of prostomium and caruncle; C, same, posterior region, dorsal view; D, largest specimen (50 mm long), anterior region, dorsal view; E, same, close-up of prostomium and caruncle; F, same, posterior region. Scale bars: A, 1.3 mm; B, 0.3 mm; C, 1.2 mm; D, 1.5 mm; E, 0.4 mm, F, 1.1 mm. 713 Revision of Pherecardia Horst, 1886 ZOOSYSTEMA • 2025 • 47 (28) no differences in chaetae and regarded pigmentation pattern and stripes to be variable; Augener was following and expanding on Ehlers’ (1920: 13) synonymy. Hartman (1959: 134) regarded P.distincta as ‘questionably P.striata.’ Pherecardia polylamellata de Silva, 1961 was described (with specimen size not specified), as lacking pigmentation and having dorsal cirri with thicker cirrophores, and with dorsal cirri and branchiae having blackish tips. However, as indicated above, the type specimens exhibit remaining dorsal pigmentation as diffuse irregular longitudinal stripes, and notopodia have barely pigmented black distal rings. distribution From the Western Indian Ocean to the Eastern Pacific, including Hawaii and French Polynesia, in shallow water, coral or rocky bottoms. A DE BC F JIHG IB IB Fig. 14. — Pherecardia distincta (Hoagland, 1920) reinstated, n. comb., COI-sequenced non-type specimens: A, 8 mm wide specimen, UF 5420, anterior region, dorsal view; B, same, right lateral view; C, anterior end, dorsal view; D, 12 mm wide specimen, UF 5343, anterior region, dorsal view; E, same, right lateral view; F, anterior end, dorsal view; G, chaetiger 8, left parapodium, anterior view (IB: interramal belt); H, same, after Methyl green staining (IB: interramal belt); I, same, cross section of interramal area; J, same, tips of upper and lower neurochaetae. Scale bars: A, 2.1 mm; B, 1.2 mm; C, 0.6 mm; D, 2.2 mm; E, 1.6 mm; F, 0.9 mm; G, H, 0.8 mm; I, 0.3 mm; J, 30 µm. 720 ZOOSYSTEMA • 2025 • 47 (28) Salazar-Vallejo S. I. et al. prey revealed in the gut contents of two coral reef fish species. PLoS ONE8(4): e58076. https://doi.org/10.1371/journal. pone.0058076 mAlAQuin A. & dehorne A. 1907.— Les annélides polychètes de la Baie d’Amboine. Revue Suisse de Zoologie 15: 335-400. https:// www.biodiversitylibrary.org/page/10188313 mAverickdiving 2022.— Golden fireworm (Chloeia flava) consumes injured anchovy (Engraulis japonicus). https://www.youtube. com/watch?v=l3Nzj3j-WoM (seen 31 Jul. 2024). mcintosh W. c. 1925.— A second contribution to the marine polychaetes of South Africa. Union of South Africa, Fisheries and Marine Biological Survey Report 5(4): 1-93, pls 1-10. m ikkelsen p. s. & v irnstein r. W. 1982.— An illustrated glossary of polychaete terms. Harbor Branch Found, Technical Report 46: 1-92. http://fau.digital.flvc.org/islandora/object/fau:2692/datastream/OBJ/view/An_illusrated_glossary_of_polychaete_terms.pdf monro c. c. A. 1924.— On the Polychaeta collected by H.M.S. ‘Alert’, 1878-1882. Families Aphroditidae and Amphinomidae. Journal of the Linnean Society of London, Zoology 36(240): 65-77. https://doi.org/10.1111/j.1096-3642.1924.tb02208.x monro c. c. A. 1928.— Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16, 45. On the Polychaeta collected by Dr. Th. Mortensen off the coast of Panama. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i København 85: 75-103. monro c. c. A. 1933.— The Polychaeta Errantia collected by Dr.C.Crossland at Colón, in the Panama region, and the Galapagos Islands during the Expedition of the S.Y. ‘St. George.’ Proceedings of the General Meetings for Scientific Business of the Zoological Society of London 103(1): 1-96. https://doi. org/10.1111/j.1096-3642.1933.tb01578.x monro c. c. A. 1939.— On some tropical polychaetes in the British Museum, mostly collected by Dr. C. Crossland at Zanzibar, Tahiti and the Marquesas. Annals and Magazine of Natural History, series 11 4, 161-184. https://doi.org/10.1080/00222933908526982 okudA s. 1937.— Polychaetous annelids from the Palau Islands and adjacent waters, the South Sea Islands. Bulletin of the Biogeographical Society of Japan 7(12): 257-315. orrhAge l. 1990.— On the microanatomy of the supraoesophageal ganglion of some amphinomids (Polychaeta Errantia), with further discussion of the innervation and homologues of the polychaete palps. Acta Zoologica 71(1): 45-59. https://doi. org/10.1111/j.1463-6395.1990.tb01180.x pillAi t. g. 1965.— Annelida Polychaeta from the Philippines and Indonesia. Ceylon Journal of Science (Biological Science) 5(2): 110-177. piotroWski c. n , bolick h., hArris l., pAulAy g., cArrerApArrA l. f. & sAlAzAr-vAlleJo s. i. 2024.— Redescription of Iphione ovata Kinberg, 1856 and confirmation of its Indian Ocean-Eastern Pacific distribution (Annelida, Aphroditiformia, Iphionidae). Zoosystema 46 (1): 1-22. https://doi.org/10.5252/ zoosystema2024v46a1. http://zoosystema.com/46/1 potts f. A. 1909.— The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner, 20. Polychaeta of the Indian Ocean, 1. The Amphinomidae. Transactions of the Linnean Society of London, second series, Zoology 12(4): 355-371, pls 45-46. https://www.biodiversitylibrary.org/page/26710816 rAJAeskArAn r. & fernAndo o. J. 2012.— Polychaetes of Andaman and Nicobar Islands, in v enkAtArAmAn k., r AghunAthAn c. & sivAperumAn c. (eds), Ecology of Faunal Communities on the Andaman and Nicobar Islands. New York (NY), Springer: 1-22. reAd g. & fAuchAld k. (ed.) 2025.— World Polychaeta Database. Pherecardia Horst, 1886. [Accessed 15 Mar. 2025] https:// www.marinespecies.org/aphia.php?p=taxdetails&id=204530. sAck l. & scoffoni c. 2013.— Tansley Review: Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future. New Phytologist 198: 983-1000. https://doi.org/10.1111/nph.12253 sAlAzAr-vAlleJo s. i. 2020.— Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739: 1-114. https://doi.org/10.11646/zootaxa.4739.1.1 sAlAzAr-vAlleJo s. i. 2023.— Revision of Chloeia Savigny in Lamarck, 1818 (Annelida, Amphinomidae). Zootaxa 5328: 1-134. https://doi.org//10.11646/zootaxa.5238.1.1 sAlAzAr-vAlleJo s. i , piotroWski c. n. & pAulAy g. 2024.— Revision of Iphione Kinberg, 1856 (Annelida, Aphroditiformia, Iphionidae). Zootaxa 5548 (1), 1–84. https://doi.org/10.11646/ zootaxa.5548.1.1 s chmArdA l. k. 1861.— Neue Wirbellose Thiere beobachtet und gesammelt auf einer Reise um Die Erdr 1853 bis 1857. Erster Band: Turbellarien, Rotatorien und Anneliden, zweite halfte. Leipzig: Engelmann, 164 p., pls 16-37. https://www.biodiversitylibrary. org/page/50385225 s olís -W eiss v., b ertrAnd y., h elléouet m.-n. & p leiJel f. 2004.— Types of polychaetous annelids at the Muséum national d’Histoire naturelle, Paris. Zoosystema 26 (3): 377-384. https:// sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2004n3a2.pdf s teve l. 2012.— It came from beneath the sea (Honolulu, Hawaii, December 25, 2012). Large specimen caught with a fish hook. [Accessed 11 Sep. 2022] http://www.mikescatchreport.com/ new/122512hawaiiworm.html. tAmurA k., stecher g. & kumAr s. 2021.— MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution 38(7): 3022-3027. https://doi.org/10.1093/ molbev/msab120 treAdWell A. l. 1906.— Polychaetous annelids of the Hawaiian Islands collected by the steamer Albatross in 1902. Bulletin of the United States Fish Commission 23(3): 1145-1181. https://www. biodiversitylibrary.org/page/51691969 treAdWell A. l. 1926.— Polychaetous annelids from Fiji, Samoa, China and Japan. Proceedings of the United States National Museum 69(2641): 1-20. https://www.biodiversitylibrary.org/ page/15384893 vAn AntWerp n. 2013.— Lined fireworm eats crab: Komodo Indonesia. https://www.blueviews.net/Videos/marine-worms. html (seen on 31 Jul. 2024). yáñez-riverA b. & sAlAzAr-vAlleJo s. i. 2011.— Revision of Hermodice Kinberg, 1857 (Polychaeta: Amphinomidae). Scientia Marina 75: 251-262. https://doi.org/10.3989/ scimar.2011.75n2251 Submitted on4 November 2024; accepted on 16 March 2025; published on 6 November 2025.