taxonID	type	description	language	source
2D1987E4FFF07338FF23F888F11E660C.taxon	type_taxon	Type species: Dalhousia atlantica McIntosh, 1885, by monotypy.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF07338FF23F888F11E660C.taxon	diagnosis	Diagnosis: Hesioninae with two lateral antennae, and one median antenna on dorsal prostomial surface. Palps biarticulate, palpophores large, massive, palpostyles smaller, blunt. Eyes dark, black, brown or reddish, anterior ones larger than posterior ones, sometimes approaching each other in lateral view. Nuchal organs U-shaped. Peristomial dorsolateral and ventrolateral tubercles low, barely projected. Pharynx with upper jaw double, lower jaw transverse plate. Parapodia sesquiramous along chaetigers 1 – 3, biramous thereafter. Notochaetae from chaetiger 4, subdistally denticulate, delicate, sometimes abundant, usually very long, reaching neurochaetal tips. Neurochaetae compound falcigers, blades bidentate, guards approaching subdistal tooth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF07338FF23F888F11E660C.taxon	etymology	Etymology. McIntosh (1885: 186, footnote 2) indicated that the genus group name was named ‘ after the Earl of Dalhousie, K. T. ’ It was Fox Maule-Ramsay, 11 th Earl of Dalhousie (22 Apr. 1801 – 6 Jul. 1874), who under Queen Victoria was the Secretary of State for War (1855 – 1858) (Fryde et al. 1941). ‘ K. T. ’ stands for Knight of the Order of the Thistle, a Scotish order of chivalry. Gender. Feminine. Indicated by the declination of the nominative, and after the combination with the specific epithet, atlantica, used in its feminine acception to emphasize that the type specimen was found in the Atlantic Ocean.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF07338FF23F888F11E660C.taxon	discussion	Remarks. von Marenzeller (1904: 308), Chamberlin (1919: 190), Horst (1921: 80), and Pleijel (1998: 110) regarded Dalhousia as a junior synonym of Leocrates. It is herein regarded as distinct on the bases of the above diagnosis. It can be separated from other genera in the tribe by following the key above. Roule (1896 b: 454) rejected the independent status of Dalhousia McIntosh, 1885 because the morphological characters seemed insufficient, especially regarding the presence of the so-called frontal tubercle. However, McIntosh (1885: 187) included in the diagnosis the lack of median antenna. Further, Roule (1896: 454) regarded Dalhousia as a junior synonym of Fallacia de Quatrefages, 1866 probably because McIntosh (1885: 188) wrongly indicated that the pharynx was unarmed, but Fallacia is a junior synonym of Hesione Savigny in Lamarck, 1818, as indicated elsewhere (Salazar-Vallejo 2018). Roule (1906: 51) modified his perspective by clarifying that Dalhousia was proposed because it lacks median antenna, and pharyngeal jaws, and he regarded it as a junior synonym of Tyrrhena Claparède, 1868 based upon a damaged specimen and concluded that ‘ Ce genre, avec son unique espèce, doit probablement disparaitre de la nomenclature. ’ [This genus, with its only species, should probably disappear from nomenclature]. This conclusion was probably taken too literally, including by McIntosh himself, because he referred to Roule as the author for the species in his subsequent publications (McIntosh 1901: 227, 1908: 130). However, provided that both names refer to the same biological species, Dalhousia atlantica McIntosh, 1885 has priority over Tyrrhena atlantica Roule 1896. On the other hand, there are some differences worth mentioning based on the original descriptions. For example, McIntosh (1885: 187) indicated that eyes were reddish-brown, whereas Roule (1906: 54) reported them as purple and, in the same publication, he included a figure to show some features. This difference, however, might depend on the time spent in the ethanol before the study of specimens by these authors. Then, these differences were the high variation of the relative size of eyes (Roule 1906, Pl. 5, Fig. 37), their fusion, pigmentation of nuchal organs, and insertion of the median antenna: between anterior eyes in two cases, central in one (two if figure 36 is included), and between posterior eyes in the other. Regretfully, despite McIntosh (1885: 187) indicated the eyes were placed in a pigmented prostomial area, this pigmentation was apparently not taken into account by Roule (1906) for clarifying the relative size of eyes. Further, as indicated in the above key to Hesioninae genera, Dalhousia McIntosh, 1885 is very similar to Paradalhousia n. gen. by having palps biarticulate, jaws in the pharynx, their nuchal organs as two U-shaped lobes, and parapodia are sesquiramous anteriorly and biramous posteriorly. They differ in some features of the pharynx armature and in neurochaetal pigmentation. In Dalhousia the upper jaw is double, T-shaped, and the ventral one is a transverse plate, but there are no marginal denticles, and neurochaetae are often brownish, whereas in Paradalhousia upper and lower jaws are single, fang-shaped, with a marginal circle of denticles, and neurochaetae are pale.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF67334FF23FDBDF5B46311.taxon	description	Figures 4, 5 A – C, 9, 10	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF67334FF23FDBDF5B46311.taxon	materials_examined	Type material. Western Africa. Holotype (BMNH 1885: 12: 1: 139), HMS Challenger, Sta. 3, SW off Canary Islands (25 ° 45 ’ N, 20 ° 14 ’ W), 2745 m, Feb. 1873 (type lost; E. Sherlock in email, 2018). Additional material. Western Africa. Five specimens (MNHN A 71), R / V Talisman, combined from three stations by Louis Roule, but two are Dalhousiella carpenteri McIntosh, 1901. Field data after Vaillant (1888) as follows: 4 specimens, dredge 20 (33 ° 43 ’ N, 11 ° 22 ’ W), off Mazagan (El Yadida), Morocco, 1105 m, sand and sponges, 14 Jun. 1883; 1 specimen, dredge 46, between Lanzarote (Canary Islands) and Morocco, 1153 m, yellow sand, 26 Jun. 1883; and 2 specimens, dragage 49, off Lanzarote (Canary Islands), 865 – 927 m, yellow sand, date not indicated (used for variation as topotypes). One specimen (MNHN A 420), Albert 1 er de Monaco Expeditions, Sta. 198 (38 ° 26 ’ 25 ” N, 28 ° 38 ’ 55 ” W), off South of Fayal, 800 m, sand and mud, 25 Jul. 1888 [19 mm long, 2.3 mm wide]. One specimen (MOM 180771), Albert 1 er de Monaco Expeditions, Sta. 1116 (31 ° 43 ’ 30 ” N, 10 ° 46 ’ 45 ” W), off Esauria, Morocco, 2165 m, pink sand with globigerinans, 11 Jul. 1901 [19 mm long, 4 mm wide]. Cabo Verde Islands. One specimen (MOM 180817), Albert 1 er de Monaco Expeditions, Sta. 1203 (15 ° 54 ’ 00 ” N, 22 ° 54 ’ 45 ” W), 6.2 km off SW Boa-Vista Island, 91 m, rocky bottom, 18 Aug. 1901. Açores. 11 specimens (MOM 180228), Albert 1 er de Monaco Expeditions, Sta. 198 (38 ° 26 ’ 25 ” N, 28 ° 38 ’ 55 ” W), 800 m, sandy mud, 25 Jul. 1888 [16 – 30 mm long, 3 – 5 mm wide]. One specimen (MOM 180401), Albert 1 er de Monaco Expeditions, Sta. 578 (38 ° 26 ’ 00 ” N, 26 ° 30 ’ 45 ” W), 1165 m, sandy mud, 14 Jul. 1895 [26 mm long, 3.5 mm wide]. 11 specimens (MOM 181144 a), Albert 1 er de Monaco Expeditions, Sta. 584 (38 ° 30 ’ 30 ” N, 26 ° 50 ’ 15 ” W), 845 m, rock, 16 Jul. 1895 [13 – 33 mm long, 2 – 4 mm wide]. Two specimens (MOM 181144 b), Albert 1 er de Monaco Expeditions, Sta. 584 (38 ° 30 ’ 30 ” N, 26 ° 50 ’ 15 ” W), 845 m, rock, 16 Jul. 1895 [very poor condition]. Two specimens (MOM 180414), Albert 1 er de Monaco Expeditions, Sta. 587 (36 ° 36 ’ 40 ” N, 27 ° 17 ’ 15 ” W), 793 m, sand, 18 Jul. 1895 [12 – 22 mm long, 3 – 4 mm wide]. One specimen (MOM 180433), Albert 1 er de Monaco Expeditions, Sta. 597 (38 ° 27 ’ 00 ” N, 28 ° 03 ’ 25 ” W), close to Prainha de Pico, 523 m, rock, 23 Jul. 1895 [16 mm long, 2.5 mm wide]. One specimen (MOM 180445), Albert 1 er de Monaco Expeditions, Sta. 616 (38 ° 46 ’ 35 ” N, 28 ° 17 ’ 20 ” W), close to Rosales or São Jorge Point, 1022 m, gray sandy mud, 1 Aug. 1895 [14 mm long, 2 mm wide]. One specimen (MOM 180527), Albert 1 er de Monaco Expeditions, Sta. 837 (37 ° 55 ’ 00 ” N, 25 ° 24 ’ 15 ” W), 880 m, rock, 22 – 24 Jul. 1897 [17 mm long, 3 mm wide]. Two specimens (MOM 180888), Albert 1 er de Monaco Expeditions, Sta. 1344 (38 ° 45 ’ 30 ” N, 28 ° 07 ’ 45 ” W), 1095 m, volcanic sand, 18 Aug. 1901 [27 – 32 mm long, 4 – 5 mm wide]. One specimen (MOM 180995), Albert 1 er de Monaco Expeditions, Sta. 2214 (39 ° 26 ’ 10 ” N, 31 ° 21 ’ 30 ” W), drifting object with Amphinome pallasii de Quatrefages (probably mislabeled specimen), 1866, 2 Sep. 1904 [19.5 mm long, 2 mm wide]. Celtic Sea. Sixteen specimens (MNHN CENTOB Thalassa 1973 - 417), R / V Thalassa, Cruise 1973, Sta. Z 417 (48 ° 12 ’ 99 ” N, 09 ° 09 ’ 05 ” W), 865 m, corals, sand to muddy bottom, 21 – 29 Oct. 1973 [21 – 28 mm long, 2.5 – 4.0 mm wide]. Fifteen specimens (MNHN CENTOB Thalassa 1973 - 421), R / V Thalassa, Cruise 1973, Sta. Z 421 (48 ° 22 ’ 05 ” N, 09 ° 33 ’ 05 ” W), 950 m, calcareous green rocks, sand, 21 – 29 Oct. 1973 [12 – 38 mm long, 2 – 6 mm wide]. Sixteen specimens (MNHN CENTOB Thalassa 1973 - 452), R / V Thalassa, Cruise 1973, Sta. Z 452 (48 ° 41 ’ 05 ” N, 10 ° 53 ’ 00 ” W to 48 ° 39 ’ 00 ” N, 10 ° 52 ’ 02 ” W), 1420 – 1470 m, rocks, sand, corals, 21 – 29 Oct. 1973 [15 complete and one anterior fragment; 12 – 35 mm long, 2 – 5 mm wide]. Iberian Seas. One specimen (MNCN 16.01 / 121), Banco de Galicia (43 ° 07 ’ N, 12 ° 14 ’ W), no further field data [27 mm long, 3.5 mm wide]. One specimen (MNCN 16.01 / 10300), Campaña Fauna II, Sta. 171 A, E off Islas Sisargas, 89 – 96 m (strange depth, probably wrong), 27 Jun. 1991 [34 mm long, 3.5 mm wide]. One specimen (MNCN 16.01 / 10301), Campaña Fauna II, Sta. 172 A, Banco de Galicia (43 ° 07 ’ N, 12 ° 14 ’ W), 761 – 768 m, 28 Jun. 1991 [43 mm long, 7 mm wide]. 29 specimens (MNCN 16.01 / 10302), Campaña Fauna II, Sta. 173 A, Banco de Galicia (43 ° 07 ’ N, 12 ° 14 ’ W), 769 – 760 m, 28 Jun. 1991 [25 – 38 mm long, 2.5 – 4.0 mm wide]. 12 specimens (MNCN 16.01 / 10303), Campaña Fauna IV, Sta. 277 B 22, E side of Banco El Fidalgo, Islas Columbretas, Castellón, 25 m, 16 Jul. 1996 [29 – 36 mm long, 3 – 4 mm wide]. 10 specimens (MNCN 16.01 / 10304), Campaña Fauna II, Sta. 173 A, Banco de Galicia (43 ° 07 ’ N, 12 ° 14 ’ W), 769 – 760 m, 28 Jun. 1991 [16 – 27 mm long, 2.5 – 4.0 mm wide]. 18 specimens (MNCN 16.01 / 10305), Campaña Fauna II, Sta. 173 A, Banco de Galicia (43 ° 07 ’ N, 12 ° 14 ’ W), 769 – 760 m, 28 Jun. 1991 [29 – 37 mm long, 3 – 4 mm wide]. Faroe Islands. One specimen (NHMD 109402), BIOFAR, SW off Faroe Islands, Sta. 493 (60 ° 49.38 ’ N, 09 ° 53.27 ’ W), 800 m, mud, 24 Jul. 1989 [24 mm long, 3 mm wide]. Eight specimens (NHMD 109404), BIOFAR, SW off Faroe Islands, Sta. 515 (60 ° 41.8 ’ N, 11 ° 46.5 ’ W), 700 m, sand, mud, 26 Jul. 1989 [one polynoid without elytrae and posterior end; other specimens 16 – 31 mm long, 2 – 4 mm wide]. One specimen (NHMD 237478), Shamrock Canyon, Sta. 1883 / 1 (47 ° 48.9 ’ N, 08 ° 08.87 ’ W), 1350 m, 19 Apr. 1977, Shackelton, coll. [26 mm long, 4 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF67334FF23FDBDF5B46311.taxon	description	Description. Largest non-type specimen (MNHN CENTOB Thalassa 1973 - 417), complete, tapered, slightly damaged, tentacular and dorsal cirri broken, most notochaetal bundles broken, bent laterally and ventrally. Body colorless, stomach darker in some specimens (Fig. 9 A), 29 mm long, 5 mm wide, 16 chaetigers; right parapodium of chaetiger 8 removed for observing parapodial features. Eyes blackish, posterior prostomial projections blackish; feebly pigmented areas include areas around eyes, projected anteriorly into palpophores and along lateral ciliated bands (Fig. 10 B), darker in other specimens (Fig. 9 D, E). Prostomium as long as wide, wider anteriorly. Lateral antennae longer than prostomium, slightly longer than palps; palpophores slightly longer than palpostyles. Median antenna missing, scar inserted centrally between eyes (other large specimens with median antenna surpassing anterior prostomial margin). Eyes blackish, anterior ones emarginate, twice larger and slightly more separated than posterior reniform eyes; in lateral view, eyes distinct. Nuchal organs lobes blackish, slightly longer than wide, pigment spot medially indented, lobes subrectangular, slightly divergent; lateral ciliated areas visible dorsally (Fig. 9 B, E). Lateral cushions swollen, entire anteriorly, bifid posteriorly, longitudinal striae visible. Peristomial dorsolateral tubercles low, oval, barely projected, ventrolateral tubercles indistinct. Pharynx evert- ed, anterior margin ciliated, lateral vesicles not swollen, left one slightly more projected; basal ring smooth. Jaws brownish, upper jaw double, lower jaw transverse with exposed region semicircular, almost transparent (Figs 5 A – C, 9 C, F). Longest tentacular cirri without tips, reaching chaetiger 6. Dorsal cirri broken, shorter than body width. Chaetigers 1 – 3 without notochaetae, notochaetae present along chaetigers 4 – 16, mostly broken (abundant, about 50 per bundle in another large specimen), subdistally denticulate, denticles fine. Notacicular lobes triangular, tapered (Fig. 9 G), or blunt (Fig. 10 G); neuracicular lobes as long as wide, blunt. Neurochaetae brownish, handle and basal portion of blades pigmented, about 50 per bundle, blades denticulate, decreasing in size ventrally, 3 – 20 times longer than wide (Fig. 10 H, I), tips of longest blades difficult to be seen, guards approaching subdistal tooth. Posterior region tapered; prepygidial segment with dorsal cirri 3 times longer than ventral ones; pygidium with anus terminal, anal cirri missing (other large specimens with broken cirri, reaching chaetiger 13). Gonads visible inside parapodia; oocytes not seen. Variation. Five presumptive topotype specimens (MNHN A 71) with body grayish, twisted, most cirri missing, many chaetae broken, eyes and nuchal organs brownish to blackish (Fig. 10 A, E). Body obconic, 20 – 32 mm long, 3 – 4 mm wide, 16 chaetigers. Lateral antennae shorter than prostomium or palps, broken in several specimens; palpophores slightly longer than palpostyles, bent ventrally (Fig 10 B, F). Median antenna short, not reaching prosto- mial anterior margin, inserted between eyes. Nuchal organs lobes partially covered by anterior margin of tentacular belt, parallel, blackish; lateral ciliated bands wide, visible dorsally. Pharynx exposed; jaws with exposed portions hyaline, transverse to pharynx; upper jaw double, separated, lower jaw double, fused as a transverse plate (Fig. 10 C). Dorsal cirri broken, shorter than body width (excluding parapodia). Notochaetae present along chaetigers 4 – 16; notacicular and neuracicular lobes conical, three times longer than wide, tapered (Fig. 10 D, G). Notochaetae about 30 per bundle, subdistally denticulate, denticles fine. Neurochaetae golden, 30 – 40 per bundle, blades bidentate, 8 – 18 times longer than wide, guards approaching subdistal tooth. Posterior region tapered into a blunt cone; prepygidial segment without cirri, pygidium with anus terminal, anal cirri missing. Gonads with spermatids; ovaries seen in parapodia by transparency, oocytes about 100 µm in diameter (Fig. 10 G).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF67334FF23FDBDF5B46311.taxon	discussion	Remarks. The holotype of Dalhousia atlantica McIntosh, 1885 is lost (E. Sherlock, 2018 in email). Pettibone (1970) redescribed and illustrated the type specimen, although she included another species from Indonesia, D. indica (Horst, 1921) n. comb., as a junior synonym. As indicated in the key above and in the corresponding redescription, they are different. The above description, as well as the variation, are based upon topotype and non-type specimens.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFF67334FF23FDBDF5B46311.taxon	distribution	Distribution. Originally described from off the Canary Islands, in deep water sediments (2745 m), it can be found from the Faroe Islands to Western Africa, in 760 – 1470 m depth. There is a surface record, in a drifting object by Fauvel (1914); the specimen (MOM 180995) matches the species and it is herein regarded as a mislabeled specimen. Other records for shallow water in Western Africa by Fauvel (1950: 349, 1953 a: 18) cannot be confirmed because his specimens were not available; additional records correspond with Paralamprophaea greeffiana (Augener, 1918) n. comb. (see below).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFB7337FF23F962F06F6335.taxon	description	Figs 11, 12	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFB7337FF23F962F06F6335.taxon	materials_examined	Type material. Indonesia. Holotype (ZMA V. Pol. 2481), Sulawesi, Banda Sea, R / V Siboga Exped., Sta. 221 (06 ° 24 ’ S, 124 ° 39 ’ E), 2798 m, deep sea trawl, muddy bottom, 4 Nov. 1899. Additional material. Wallis and Futuna. Two specimens (MNHN Musorstom 7 - 631), Sta. 631 (11 ° 54.0 ’ S, 179 ° 31.6 ’ W), 600 m, 29 May 1992 [21 – 22 mm long, 3 mm wide]. New Caledonia. One specimen (MNHN Halipro 2 - 04), R / V Tangaroa, Sta. 4 (23 ° 30.90 ’ S, 169 ° 33.93 ’ E to 23 ° 30.74 ’ S, 169 ° 33.66 ’ E), 790 – 845 m, 6 Nov. 1996 [8 mm long, 1 mm wide]. Two specimens (MNHN Halipro 2 - 25), R / V Tangaroa, Sta. 25 (25 ° 16.64 ’ S, 170 ° 23.69 ’ E to 25 ° 17.45 ’ S, 170 ° 23.93 ’ E), 1100 – 1380 m, 11 Nov. 1996 [30 – 33 mm long, 3 – 5 mm wide]. One specimen (MNHN Halipro 2 - 60), R / V Tangaroa, Sta. 60 (24 ° 52.06 ’ S, 168 ° 42.31 ’ E to 24 ° 52.11 ’ S, 168 ° 43.68 ’ E), 1133 – 1280 m, 18 Nov. 1996 [27 mm long, 3 mm wide]. One specimen (MNHN Halipro 2 - 83), R / V Tangaroa, Sta. 83 (23 ° 38.95 ’ S, 168 ° 15.79 ’ E to 23 ° 38.54 ’ S, 168 ° 15.80 ’ E), 820 – 1060 m, 22 Nov. 1996 [27 mm long, 4 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFB7337FF23F962F06F6335.taxon	description	Description. Holotype (ZMA V. Pol. 2481) complete, straight, integument, damaged (Fig. 11 A), without pygidium; posterior chaetigers almost broken off, damaged after several previous manipulations and dissections. Two pin perforations middorsally by chaetigers 6 and 7 / 8; left parapodia of chaetigers 1, 2, 6, 7, 13, and right parapodium of chaetiger 6 previously removed; two irregular ventral dissections along chaetigers 1 – 6 / 7 (Fig. 11 C). Body 17 mm long, 2 mm wide, 16 chaetigers; dorsum colorless, venter with brownish margins along midventral depression, interrupted segmentally. Prostomium as long as wide, slightly wider anteriorly and posteriorly (Fig. 11 B). Lateral antennae with ceratophores distinct, slightly shorter than prostomium, longer than palps; palpophores twice longer than palpostyles. Eyes brownish, barely defined. Median antenna broken, inserted centrally between eyes. Eyes brownish, anterior left eye reniform, right one round, larger and more distant to each other than posterior round ones (illustrated as round by Pettibone). Nuchal organs lobes as long as wide, parallel, slightly expanded posteriorly, well separated, barely pigmented in holotype, darker in non-type specimens. Tentacular cirri missing. Lateral cushions low, barely projected, entire, longitudinal striae visible. Peristomial dorsolateral and ventrolateral tubercles low, barely projected. Pharynx not exposed, inner features observed by previous dissection (Fig. 11 C). Anterior margin smooth. Upper jaw double, delicate, small; lower jaw not visible, probably damaged by dissection. Chaetigers 1 – 3 without notochaetae; following parapodia biramous, notochaetae present along chaetigers 4 – 16, about 40 per bundle, reaching neurochaetal tips, delicate, tips curled, subdistally denticulate, denticles fine. Notacicular lobes tapered, blunt in anterior (Fig. 11 D) and posterior chaetigers (Fig. 11 F). Neuracicular lobes round, blunt, barely projected in anterior chatigers (Fig. 11 D), blunt, conical in posterior chaetigers (Fig. 11 F). Neurochaetae about 20 per bundle (30 in posterior segments), blades bidentate, 6 – 50 times longer than wide (Fig. 11 E, G), in middle chaetigers blades slightly narrower than handle width, or 1 / 3 – 1 / 5 as wide as handle’s width, longest thin blades subdistally expanded, denticles tiny, guards approaching subdistal tooth. Posterior region tapered. Prepygidial segment and pygidum missing. Oocytes not seen. Variation. Body size 8 – 33 mm long; neurochaetal blade length smaller in larger specimens. Body dorsum, nuchal organs and neurochaetal pigmentation variable; usually darker in larger specimens; dorsal cirrophores and neuropodia paler. Best preserved specimen (Fig. 12 A) with brownish pigmentation dorsally, progressively paler medially and posteriorly. Nuchal organs depressed, medially darker, expanded laterally (Fig. 12 B). Peristomial middorsal and dorsolateral tubercles darker, ventrolateral tubercles paler (Fig. 12 C). Notacicular and neuracicular lobes blunt, tapered (Fig. 12 D); neurochaetae golden to brownish, blades 10 - 20 times longer than wide (Fig. 12 E, insets).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFB7337FF23F962F06F6335.taxon	discussion	Remarks. Dalhousia indica (Horst, 1921) was included as a junior synonym of D. atlantica McIntosh, 1885 by Pettibone (1970) and Pleijel (1998: 160). They are different species because D. indica has nuchal organs lobes as long as wide and its eyes are brownish, anterior ones twice larger than posterior ones, whereas in D. atlantica nuchal organs lobes are slightly longer than wide, and eyes are blackish, anterior ones 2 – 3 times larger than posterior ones. Consequently, D. indica is reinstated. It has been newly combined into Dalhousia because of the development of the nuchal organs, being posteriorly expanded, and by the large size of notochaetae which reach neurochaetal tips in the same parapodia. The brownish pigmentation of neurochaetae has faded off in the type specimen, which is a damaged juvenile, but in more recently collected specimens, neurochaetal handles and at least the basal to middle part of blades are brownish, and even notochaetal bases are brownish as well, as indicated in the original description.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFB7337FF23F962F06F6335.taxon	distribution	Distribution. Originally described from the Banda Sea, Indonesia in deep water (2798 m), it has been found in Wallis and Futuna, and New Caledonia, in sediments at depths of 600 – 1380 m.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFF7332FF23FA20F09C63A1.taxon	description	Fig. 13	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFF7332FF23FA20F09C63A1.taxon	materials_examined	Type material. Western Pacific. Japan. Holotype of Leocrates auritus Hessle, 1925 (UUZM 626), Ogasawara, Chichi-jima (27 ° 04 ’ 00 ” N, 142 ° 12 ’ 30 ” E), coral, 25 Jul. 1914, S. Bock, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFF7332FF23FA20F09C63A1.taxon	description	Description. Holotype (UUZM 626) complete, slightly dehydrated along middle region, slightly bent ventrally. Body pale (Fig. 13 A), obconic, slightly wider medially, blunt anteriorly, tapered posteriorly, 39 mm long, 4.5 mm wide (without parapodia), 16 chaetigers; right parapodium of chaetiger 9 previously removed, left parapodium of chaetiger 8 dissected (kept in container). Most tentacular, dorsal and ventral cirri intact. Body pale, eyes dark brown. Prostomium as long as wide, slightly wider anteriorly. Lateral antennae with ceratophores well-defined, antennae about twice longer than prostomium, slightly longer than palps (Fig. 13 B). Palpophores 3 – 4 times longer than palpostyles. Median antenna long, surpassing anterior prostomial margin, inserted centrally on prostomium, between eyes. Eyes dark brown, anterior eyes about twice larger than posterior ones, slightly emarginate, more distant to each other than posterior round eyes (Fig. 13 B). Nuchal organs whitish, L-shaped, lateral lobes extended beyond lateral prostomial margins, lobes medially wider, not parallel-sided; lateral ciliated bands wide, clearly visible dorsally. Tentacular cirri mostly without tips, longer ones reach chaetiger 8. Lateral cushions low, entire, longitudinal striae visible. Pharynx fully exposed, wider distally, without lateral vesicles. Anterior margin smooth, with irregular constrictions, better defined laterally (Fig. 13 C). Dorsal jaw brownish, exposed, inserted below pharynx margin; ventral jaw smaller, exposed. Dorsal cirri longer than body width, including parapodia (Fig. 13 D), complete ones in middle segments half as long as body length. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, about 60 per bundle, delicate, arranged in a transverse fan, notochaetae subdistally denticulate, denticles coarse (Fig. 13 E). Notacicular lobes short, blunt, round (Fig 13 D, inset); neuracicular lobes blunt, projected, tips round, slightly longer than wide; aciculae black, tapered; ventral cirri surpass neurochaetal lobes. Neurochaetae about 30 per bundle, many blades missing, blades decreasing in size ventrally, bidentate, 6 – 9 times longer than wide, guards approaching subdistal tooth (Fig. 13 F). Posterior region tapered, with several dorsal and anal cirri on site. Pre-anal segment with dorsal cirri twice thicker than ventral one. Pygidium with anus terminal; anal cirri reach chaetiger 12 – 13. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFF7332FF23FA20F09C63A1.taxon	discussion	Remarks. Hessle (1925: 18) described Leocrates auritus. His description is complete and the illustrations are very good. However, the type of nuchal organs together with the presence of single fang-shaped, upper and lower jaws of the pharynx indicate it belongs to Lamprophaea and hence the new combination. Pettibone (1970: 213, 219), regarded L. aurita as a junior synonym of Leocrates giardi Gravier, 1900, described from the Red Sea, but this nomenclatural act did not involve the study of types. Further, because the nuchal organs lobes in these two species are so different, they are herein regarded as different species, and belonging to different genera. Consequently, the synonymy must be rejected, and the species name is reinstated. As indicated in the key above, L. aurita resembles L. ockeri n. sp. from the Hawaiian Archipelago because they have notochaetae from chaetiger 4, and anterior eyes round or slightly emarginate. They differ in the shape of prostomial and nuchal organs lobes. In L. aurita the posterior prostomial margins are straight, leaving some space from the posterior eyes, and lateral nuchal organs lobes are medially and distally swollen, whereas in L. ockeri the posterior prostomial margins are curved, leaving almost no space between its margins and posterior eyes, and the lateral nuchal organ branches have parallel sides.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFF7332FF23FA20F09C63A1.taxon	distribution	Distribution. Only known from the Bonin Islands, Japan, in shallow-water, coralline substrates.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFD7333FF23FD02F11063F9.taxon	description	Figs 14, 15 urn: lsid: zoobank. org: act: 1734 A 20 C-A 4 D 3 - 4 E 74 - 90 E 5 - B 0 DCFD 81 D 61 C	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFD7333FF23FD02F11063F9.taxon	materials_examined	Type material. Western Pacific, French Polynesia. Holotype (UF 882), Society Islands, Moorea, E side of Cook’s Bay, opposite to Gump, fringing reef in bay (17 ° 29 ’ 17.88 ” S, 149 ° 49 ’ 09.12 ” W), coral, 1 – 3 m, 1 Nov. 2008, V. Ivanenko & J. Moore, coll. Additional material. Western Pacific, Mariana Islands. One specimen (UF 51), Guam, Urunao Reef (13 ° 30 ’ 00.00 ” N, 144 ° 48 ’ 00.00 ” E), reef flat, under ruble, 1 Jun. 2000, C. P. Meyer & G. Paulay, coll. [24.5 mm long, 3 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFD7333FF23FD02F11063F9.taxon	description	Description. Holotype (UF 882) complete, mature female, bent dorsally. Body obconic, blunt anteriorly, tapered posteriorly, most cirri missing; 30 mm long, 4 mm wide (without parapodia); left parapodium of chaetiger 7 removed for observing parapodial features. Body blackish, from chaetiger 6 brownish because of abundant tubular ovaries in coelom (Fig. 14 A); parapodia pale, midventral depression pale, with blackish margins, along chaetigers 1 – 5. Prostomium wider than long, widest posteriorly, with angular corners, posterior prostomial margins expanded as small horns (Fig. 14 B); anterior margin blackish, pigmentation continued posteriorly from bases of pale lateral antennae. Lateral antennae bent ventrally, longer than prostomium and palps; palpophores three times longer than palpostyles (left one missing). Median antennae blackish, directed posteriorly, short (not reaching prostomial anterior margin), inserted between posterior eyes. Eyes dark brown, anterior ones emarginate anterolaterally, twice larger and slightly more separated than posterior, round eyes; in lateral view, eyes distinct. Nuchal organs lobes L-shaped, lateral projections barely expanded subdistally, with black ridge, anterior and posterior surfaces paler, projected beyond lateral prostomial margins; lateral ciliated bands wide, visible dorsally. Tentacular cirri mostly missing, one ventral remaining pale, barely surpassing chaetiger 1. Lateral cushions project- ed laterally along middle and posterior chaetigers, smooth, entire, longitudinal striae visible, unless too expanded by gonads. Pharynx not exposed, mouth wide open (Fig. 14 C). Lateral vesicles not seen; anterior margin with regular constrictions but not counted. Middorsal jaw single, hyaline, larger than ventral one, inserted below pharynx margin. Dorsal cirri homogeneously blackish, or banded, most missing, a few remaining without tips, shorter than body width. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 60 per bundle, delicate, arranged in transverse fans along most chaetigers, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes long, tapered, three times longer than wide (Fig. 14 D); aciculae black, tapered; ventral cirri tapered, surpassing neuracicular blunt lobes. Neurochaetae about 30 per bundle, blades or similar size (Fig. 14 E), shortest ones missing, bidentate, 5 – 7 times longer than wide, guards approaching subdistal tooth. Posterior end tapered. Prepygidial segment with cirri barely pigmented, dorsal cirri about 4 times longer than ventral ones. Anus terminal, anal cirri reaching chaetiger 13. Oocytes about 100 µm. Spermatids not seen. Variation. A living juvenile specimen from Guam was dark purple (Fig. 15 A), darker medially, paler posteriorly. Upon preservation most cirri were missing, and after almost 20 years in ethanol, most of its pigmentation has disappeared (Fig. 15 B). Remarkably, the prostomial margins as well as the lateral branches of nuchal organs lobes are darker than surrounding areas (Fig. 15 C), and the emarginate, anterior eyes are twice larger than posterior ones. Regarding parapodial features, the notacicular lobes are tapered, and the neuracicular ones are round but about twice wider than long (Fig. 15 D). Neurochaetal blades are 3 – 8 times longer than wide.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFD7333FF23FD02F11063F9.taxon	etymology	Etymology. The specific epithet is made after the Latin adjective cornutus (- a, - um) meaning horned, or provided with horns. The name indicates the posterolateral prostomial squarish projections resembling small horns, and it is in feminine for matching the genus gender (ICZN 1999, Art. 31.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFD7333FF23FD02F11063F9.taxon	discussion	Remarks. Lamprophaea cornuta n. sp. belongs in the group of species having notochaetae from chaetiger 5, and resembles L. pleijeli n. sp. from the Western Indian Ocean by having palpophores three or more times longer than palpostyles, prostomial margins angular, and dorsum blackish. These two species especially differ in the relative size of eyes, and in the length of posterior projections of nuchal organs. In L. cornuta anterior eyes are twice larger than posterior ones, and posterior projections of nuchal organs are 3 – 4 times longer than wide, whereas in L. pleijeli eyes are of similar size, and posterior projections of nuchal organs are about as long as wide.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFFD7333FF23FD02F11063F9.taxon	distribution	Distribution. Western Pacific, from Guam to Moorea, in intertidal to shallow subtidal coralline substrates.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC3730FFF23F930F1646405.taxon	description	Figs 16, 17	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC3730FFF23F930F1646405.taxon	materials_examined	Type material. Western Pacific. Samoa. Lectotype (ZMH V- 1275), and paralectotype (ZMH V- 1274), herein designated, no further field data, H. Godeffroy, coll. [paralectotype complete, dehydrated, pharynx partially exposed, dorsum colorless, midventer brownish; body 12 mm long, 2 mm wide, 16 chaetigers]. Additional material. Samoa. Five specimens (USNM 19194), Pago Pago, May 1920, A. L. Treadwell, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC3730FFF23F930F1646405.taxon	description	Description. Lectotype (ZMH V- 1275) complete, depressed, colorless, partially damaged (Fig. 16 A). Body obconic, wider medially, blunt anteriorly, tapered posteriorly, 20 mm long, 4 mm wide, 16 chaetigers; left parapodium of chaetiger 9 previously removed, left parapodium of chaetiger 7 removed for observing parapodial features. Body pale, eyes barely darker than integument. Some tentacular, dorsal and ventral cirri on site; many missing. Prostomium wider than long, slightly wider anteriorly, with angular corners. Lateral antennae with ceratophores distinct, antennae as long as prostomium, slightly longer than palps (Fig. 16 B). Palpophores 2.0 – 2.5 times longer than palpostyles. Median antenna short, incurved, not reaching prostomial anterior margin, inserted between posterior eyes. Eyes barely darker than surrounding integument, round, anterior eyes slightly larger than posterior ones, slightly more distant to each other than posterior eyes, in lateral view eyes clearly separated. Nuchal organs lobes L-shaped, lateral projections blunt, extended up to lateral prostomial margins, projections expanded anteriorly, ridge whitish, partially concealed by tentacular belt; lateral ciliated bands narrow, visible dorsally. Tentacular cirri without tips, longer ones reaching chaetiger 6. Lateral cushions low, barely projected laterally, entire, longitudinal striae distinct. Peristomium with middorsal tubercle slightly longer than wide, blunt, wider basally to medially, tapered anteriorly (Fig. 16 C). Pharynx with anterior margin smooth, irregularly constricted; upper and lower jaws single, transparent, tapered, upper jaw larger and more anteriorly inserted than lower one. Lateral vesicles not seen. Dorsal cirri shorter than body width. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, delicate, arranged in bundles, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes tapered; neuracicular lobes blunt, truncate, 1.5 times wider than long (Fig. 16 D). Neurochaetae about 20 per bundle, blades decreasing in size ventrally, bidentate, 3 – 8 times longer than wide, guards approaching subdistal tooth (Fig. 16 E). Posterior region tapered. Prepygidial segment with dorsal cirri 3 – 4 times wider than ventral ones; anus terminal, anal cirri missing. Oocytes not seen. Variation. Topotypes (USNM 19194) match the species description. Body 21 – 31 mm long, 2 – 4 mm wide, 16 chaetigers; nuchal organs lobes L-shaped, completely visible in three smaller specimens (up to 26 mm long), almost completely covered by tentacular belt in larger specimens (28 – 31 mm long), but anterior part of lateral lobes exposed. Largest, best preserved topotype (USNM 19194) slightly dehydrated along chaetigers 3 – 14 (Fig. 17 A), slightly bent laterally. Body obconic, slightly wider medially, blunt anteriorly, tapered posteriorly, 31 mm long, 4 mm wide (without parapodia); an anteroventral dissection previously made, irregular, running along chaetigers 1 – 5; left parapodium of chaetiger 8 previously removed, right parapodium of chaetiger 8 dissected (kept in container). Body pale, eyes dark brown. Most tentacular, dorsal and ventral cirri on site. Prostomium slightly longer than wide, slightly wider anteriorly (Fig. 17 B). Lateral antennae with ceratophores distinct, antennae slightly longer than prostomium, barely longer than palps. Palpophores 3 times longer than palpostyles. Median antenna short, not reaching anterior prostomial margin, inserted between posterior eyes. Eyes brownish, round, anterior eyes slightly larger than posterior ones, slightly more distant to each other than posterior eyes, in lateral view eyes clearly separated. Nuchal organs lobes L-shaped, lateral projections extended beyond lateral prostomial margins, projections expanded anteriorly, partially concelead by tentacular belt; lateral ciliated bands narrow, visible dorsally. Tentacular cirri almost complete, tips eroded, longer ones surpass chaetiger 6, not reaching chaetiger 7. Lateral cushions low, entire, longitudinal striae distinct. Pharynx not exposed, inner features observed through dissection. Anterior margin with 20 crenulations or round papillae; lateral vesicles not seen. Middorsal jaw exposed, brownish, falcate, inserted below pharynx margin, midventral jaw not seen, probably destroyed by dissection. Dorsal cirri longer than body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, delicate, arranged in bundles, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes tapered; neuracicular lobes blunt, truncate, wider than long (fig. 17 C). Neurochaetae about 20 per bundle, many blades missing, blades decreasing in size ventrally, bidentate, 2 – 8 times longer than wide, guards approaching subdistal tooth (Fig. 17 D). Posterior region tapered. Preanal segment with dorsal cirri 4 – 5 times longer than ventral ones; anus terminal, anal cirri reach chaetiger 14. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC3730FFF23F930F1646405.taxon	discussion	Remarks. Lamprophaea cuprea Grube, 1867, reinstated, resembles L. aurita (Hessle, 1925), n. comb., reinstated, described from Japan by having L-shaped nuchal organs lobes, and anterior eyes larger than posterior ones. However, L. cuprea has notochaete from chaetiger 5, whereas in L. aurita they are present from chaetiger 4.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC3730FFF23F930F1646405.taxon	description	The largest and best preserved syntype has been selected as a lectotype (ICZN 1999, Art. 74.1). Consequently, in the sections above the term has been employed for this specimen, it has been redescribed and newly illustrated, and it is herein designated because of the poor condition of the other syntype (ICZN 1999, Art. 74.7).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC07309FF23FA94F6876014.taxon	description	Figs 3, 6, 7, 8, 18, Table 2	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC07309FF23FA94F6876014.taxon	materials_examined	Type material. Western Atlantic. Virgin Islands. Holotype of Castalia longicirrata (USNM 15915), off Saint- Thomas, Sail Rock, U. S. Fish Commission Porto Rico Expedition, U. S. S: Fish Hawk, Sta. (151) 6079, coral, 36 – 41 m, 6 Feb. 1899. Additional material. Western Atlantic. Florida, USA. Four specimens (ECOSUR 3075), 800 m SSW off Alligator Reef Light (24 ° 50 ’ 39 ” N, 80 ° 37 ’ 30 ” W), 5 – 8 m, 30 Apr. 1961, pronox fish / dipnet, W. Starck, H. Feddem & T. Starck, coll. [13 – 32 mm long, 2 – 4 mm wide]. One specimen (UF 1890), N off St. Petersburg, hard ground with sponges (28 ° 27 ’ 33.12 ” N, 84 ° 16 ’ 18.48 ” W), 32 m, 13 Mar. 2011, G. Paulay, N. Evans, F. Michonneau, coll. [32 mm long, 4 mm wide]. Two specimens (UF 2531), NNW of St. Petersburg, S of Big Bend area, sponge reef (28 ° 26 ’ 17.88 ” N, 84 ° 16 ’ 21.00 ” W), 35 m, hard bottom, 23 May 2012, J. Slapcinsky, coll. [24 – 32 mm long, 3 – 4 mm wide]. Five specimens (UF 2531 / 2582), NNW of St. Petersburg, S of Big Bend area, sponge reef (28 ° 26 ’ 17.88 ” N, 84 ° 16 ’ 21.00 ” W), 35 m, hard bottom, 23 May 2012, J. Slapcinsky, coll. [19 – 31 mm long, 2 – 4 mm wide]. One specimen (UF 2534), NNW of St. Petersburg, S of Big Bend area, sponge reef (28 ° 26 ’ 17.88 ” N, 84 ° 16 ’ 21.00 ” W), 35 m, hard bottom, 23 May 2012, J. Slapcinsky, coll. [23.5 mm long, 3.0 mm wide]. Panama. One specimen (UF 4933), Bocas del Toro, Punta Puebla (09 ° 22 ’ 01.20 ” N, 82 ° 17 ’ 27.60 ” W), 27 May 2016, M. Leray, F. Michonneau & R. Lasley, coll. [broken into two pieces, 8 + 7 mm long, 2 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC07309FF23FA94F6876014.taxon	description	Description. Holotype (USNM 15915) complete, damaged, bent laterally; right parapodia of chaetigers 3, 10 and 12, and left parapodia of chaetigers 3, 7, 8 previously removed (one in small container, without specification about its position, probably what Pettibone illustrated). Body obconic, blunt anteriorly, tapered posteriorly, colorless (Fig. 18 A), 19 mm long, 3 mm wide, 16 chaetigers; most cirri missing. Prostomium wider than long, lateral margins progressively narrower, round (Fig. 18 B). Lateral antennae (left one complete) longer than prostomium, ceratophores distinct, antennae slightly longer than palps. Palps pale, palpophores three times longer than palpostyles. Median antenna missing, scar inserted between eyes. Eyes brownish, anterior ones reniform, twice larger than posterior, round ones; in lateral view anterior and posterior eyes clearly separated. Nuchal organs lobes L-shaped, lateral branches different in length, right one longer, exceeding lateral prostomial margin, left one smaller, as long as reaching lateral prostomial margin level, ridge whitish; lateral ciliated areas narrow. Tentacular cirri broken, left ones without middle to distal regions, detaching, reaching back up to chaetiger 4. Lateral cushions low, entire, longitudinal striae not visible. Pharynx exposed, anterior margin smooth, with subdistal circle of irregular round constrictions (Fig. 18 C). Upper and lower jaws single, inserted slightly subdistally, brownish. Chaetigers 1 – 4 without notochaetae, notochaetae present along chaetigers 5 – 16, most broken, about 50 per bundle, delicately denticulate subdistally. Notacicular lobes shorter than neurochaetal ones, tapered, blunt, tips swollen (Fig. 18 D, inset). Neuracicular lobes round or blunt, as long as wide (Fig. 18 D). Neurochaetae sparse, about 20 per bundle, blades most broken, decreasing in size ventrally, 3 – 9 times longer than wide (Fig. 18 D, insets), blades bidentate, guard approaching subdistal tooth, sometimes broken. Posterior region damaged, slightly invaginated, preanal segment barely visible, cirri missing. Pygidium damaged, anal cirri missing. Oocytes not seen. Variation. There are slight differences in nuchal organs lobes during ontogeny (Fig. 3). Smaller specimens have lobes slightly divergent, and they proceed by growing laterally, often surpassing the level of lateral prostomial margins; they can be obscured by the tentacular belt anterior margin, but at least the tips of the lateral branches are evident. Parapodial features show some size-related variations (Fig. 6). These variations are subtle regarding notacicular or neuracicular lobes, and they are round in anterior chaetigers, becoming longer in middle and posterior parapodia. The number of chaetae as well as the relative length of neurochaetal blades is size dependent (Fig. 7); neurochaetal blades are longer along anterior chaetigers and in the upper neurochaetal bundle, and become shorter in following chaetigers and in the lower neurochaetal bundle (Table 2). Anatomical features (Fig. 8) were illustrated for a single specimen but no other ones were dissected.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC07309FF23FA94F6876014.taxon	discussion	Remarks. Treadwell (1902) with one specimen collected in Puerto Rico coral bottoms (36 – 41 m depth), described Castalia longicirrata. He was the first to refer to nuchal organs, but called them “ diverging processes ” in the posterior prostomial margin, and indicated size proportions between lateral antennae and palps. His description indicated that anterior eyes were twice larger than posterior ones, and gave some details for chaetae. Treadwell’s original description was enhanced by Augener (1906: 155), by describing Castalia hesionoides (now Dalhousiella hesionoides), with specimens collected in sediments at 300 – 720 m off the Lesser Antilles. Augener compared these two species and clarified the shape of nuchal organs and its lateral lobes, and indicated size proportions between palpophore and palpostyles. Lamprophaea longicirrata (Treadwell, 1902) is newly combined because of the shape of the nuchal organs lobes, and its upper and lower jaws are single, fang shaped. Augener (1906: 156 – 157) provided details about L. longicirrata in his description of Dalhousiella hesionides. Pettibone (1970: 215) regarded L. longicirrata as a junior synonym of L. chinensis Kinberg, 1866, but their nuchal organs are so different that they are herein regarded as belonging to different genera, and consequently, L. longicirrata is reinstated. As indicated in the key above, L. longicirrata belongs in the group provided with notochaetae from chaetiger 5 and with palpophores three times longer than palpostyles. It differs from the other species in the group, L. cuprea Grube, 1867, L. pleijeli n. sp. and L. cornuta n. sp. by being pale and by having convergent, round posterior prostomial margins, instead of being angular, neither round nor convergent.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC07309FF23FA94F6876014.taxon	distribution	Distribution. The holotype was collected in the Virgin Islands, in coral rocks at 36 – 41 m. It has been found from Florida to Panama, in the Western tropical Atlantic, from shallow water (5 m), hard bottoms. The record by Rullier & Amoureux (1979: 159) of Leocrates claparedii for the Abrolhos Archipelago, Brazil, might be conspecific but the specimen was not available for confirmation.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC6730BFF23FA5DF0FB6335.taxon	description	Fig. 19 urn: lsid: zoobank. org: act: 911 F 3 E 10 - 8 E 2 E- 4 FEF-AF 6 B- 3 CD 7532 F 609 B	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC6730BFF23FA5DF0FB6335.taxon	materials_examined	Type material. Central Pacific. Northeastern Hawaiian Islands. Holotype (LACM 10143), Papahãnamokuãkea Marine National Monument, Pearl and Hermes Reef, maze at inside eastern section of atoll, R / V Oscar Elton Sette, Sta. 30103 (27 ° 50 ’ N, 175 ° 55 ’ W), depth unspecified, ghost net and marine debris removal operation, 23 Oct. 2007, L. Harris, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC6730BFF23FA5DF0FB6335.taxon	description	Description. Holotype (LACM 10143) complete, slightly dehydrated along middle and posterior regions, twisted posteriorly. Body obconic, brownish anteriorly (Fig. 19 A), slightly wider medially, blunt anteriorly, tapered posteriorly, 49 mm long, 5 mm wide (without parapodia), 16 chaetigers; left parapodium of chaetiger 8 dissected (kept in container), an oblique anteroventral dissection made for observing pharynx jaws. Most tentacular, dorsal and ventral cirri broken or missing. Body brownish anteriorly, from chaetiger 5 onward pale, eyes dark brown. Prostomium twice wider than long, tapered posteriorly (Fig. 19 B). Lateral antennae with ceratophores distinct, antennae longer than prostomium, as long as palps; palpophores three times longer than palpostyles. Median antenna long, surpassing anterior prostomial margin, inserted centrally on prostomium, between eyes. Eyes dark brown, anterior eyes emarginate anteriorly, slightly larger than posterior ones, emarginate, more distant to each other than posterior round eyes. Nuchal organs lobes L-shaped, lateral projections extended beyond lateral prostomial margins, projections with parallel sides, not expanded medially, ridge slightly brackish; lateral ciliated bands wide, clearly visible dorsally. Tentacular cirri mostly broken, longer ones reaching chaetiger 5. Lateral cushions low, not projected along anterior chaetigers, divided into three sections; swollen in middle and posterior segments, entire; longitudinal striae visible. Pharynx partially exposed, jaws observed by dissection. Lateral vesicles not seen. Anterior margin with about 22 regular furrows, larger ventrolaterally. Middorsal and midventral jaws single, pale, exposed, inserted below pharynx margin; ventral jaw smaller than dorsal one. Dorsal cirri (without tip in chaetiger 4) longer than body width, including parapodia. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, about 60 per bundle, broken in some chaetigers, delicate, arranged in a transverse fan, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes long, tapered (Fig. 19 C), neuracicular lobes projected, blunt, tips round, slightly longer than wide; aciculae black, tapered; ventral cirri tapered, surpass neurochaetal lobes. Neurochaetae about 40 per bundle, some blades missing, blades decreasing in size ventrally, bidentate, 4 – 9 times longer than wide (Fig. 19 D), guards approaching subdistal tooth. Posterior end tapered, distorted by compression. Preanal segment with dorsal cirri twice longer than ventral ones. Pygidum with anus dorsoterminal, ventral surface dark brown, anal cirri breaking appart, anal cirri reaching chaetiger 16. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC6730BFF23FA5DF0FB6335.taxon	etymology	Etymology. The name of this species is a modest homage to Mr. David Ocker in recognition of his three decades of unrestricted support to our research activities, through his unlimited cordiality and generosity by housing my younger colleagues, or my wife and myself in his home. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC6730BFF23FA5DF0FB6335.taxon	discussion	Remarks. Lamprophaea ockeri n. sp. belongs in the group of species provided with notochaetae from chaetiger 4, and resembles L. aurita (Hessle, 1925) n. comb., reinstated, from Japan. There are several differences between these two species, the most important ones are in the prostomial and nuchal organs lobes shape. In L. ockeri the posterior prostomial margins are curved, and the lateral nuchal organ branches have parallel sides, whereas in L. aurita posterior prostomial margins are straight, leaving some space from the posterior eyes, and nuchal organs lobes lateral projections are medially and distally swollen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC6730BFF23FA5DF0FB6335.taxon	distribution	Distribution. Only known from the type locality in the Northeastern Hawaiian Islands.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	description	Fig. 20 urn: lsid: zoobank. org: act: 662 A 4 B 28 - C 433 - 45 F 5 - 96 FF-BA 1489 C 494 E 5	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	materials_examined	Type material. Red Sea, Saudi Arabia. Holotype (UF 3887), Al Wajh, Shaybarah, fore reef (25 ° 21 ’ 41.76 ” N, 36 ° 54 ’ 45.36 ” E), 3 – 30 m, 4 Oct. 2013, G. Paulay & S. McKeon, coll. One paratype (UF 4223), Abu Sahim, backreef slope (22 ° 39 ’ 31.32 N, 38 ° 53 ’ 03.84 ” E), 5 – 9 m, under rocks, 7 Mar. 2014, G. Paulay, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	description	Description. Holotype (UF 3887) complete, bent laterally, posterior region directed upwards (Fig. 20). Body obconic, blunt anteriorly, medially wider by lateral torsion, tapered posteriorly, 24 mm long, 4 mm wide (without parapodia); left parapodium of chaetiger 8 and right parapodium of chaetiger 9 removed for observing parapodial features. Body homogeneously blackish dorsally, parapodia and venter pale; eyes brownish. Prostomium slightly distorted, twice wider than long, tapered posteriorly (Fig. 20 B). Lateral antennae with ceratophores distinct, ceratostyles broken, antennae shorter than palps (slightly shorter than palps in paratype) and twice wider than lateral antennae; palpophores twice longer than palpostyles (four times longer in paratype). Median antenna long, surpassing anterior prostomial margin, inserted centrally on prostomium, between eyes (shorter in paratype, not reaching anterior prostomial margin). Eyes dark brown, anterior ones reniform, markedly emarginate, twice larger than posterior ones, round, more distant to each other than posterior round eyes (Fig. 20 C). Nuchal organs lobes L-shaped, lateral projections blackish along ridges, anterior and posterior surfaces pale, lobes tapered, projected beyond lateral prostomial margins (paler, better defined in paratype); lateral ciliated bands wide, right one more exposed by prostomial distortion. Tentacular cirri mostly broken, remaining ceratostyles broken, damaged, longer ones reaching chaetigers 2 – 3 (reaching chaetiger 8 in paratype). Lateral cushions low, exposure variable, barely projected along chaetigers 1 – 3, following ones more projected, smooth, entire, ridge blackish along internal sides; longitudinal striae visible. Pharynx partially exposed, jaws observed by lateral dissection. Lateral vesicles not seen. Anterior margin with about 22 regular ridges, slightly larger ventrolaterally. Middorsal jaw broken, remaing portion hyaline, round, short (complete in paratype, twice wider than lower jaw); midventral jaw tapered, hyaline; jaws exposed, inserted below pharynx margin. Dorsal cirri missing, relative size regarding body width unknown (paratype with cirri damaged, one complete cirrus longer than body width including parapodia). Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, about 60 per bundle, delicate, arranged in fascicles along chaetigers 1 – 5, and 16, and as longitudinal or oblique fans in chaetigers 6 – 15, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes long, tapered, twice longer than wide (Fig. 20 D); aciculae black, tapered; ventral cirri tapered, missing in several chaetigers, surpass neurochaetal lobes. Neurochaetae about 30 per bundle (about 20 in paratype), some blades missing, blades decreasing in size ventrally, bidentate, 4 – 10 times longer than wide, guards approaching subdistal tooth (Fig. 20 E); denticles delicate; some blades with connecting blades between denticles. Posterior end tapered. Prepygidial segment cirri missing. Anal cirri projected, anal cirri missing. Oocytes not seen. Testis penetrate into parapodial coelom.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	etymology	Etymology. This species name is after Dr. Gustav Paulay, a well-known echinoderm expert, curator or marine invertebrates in the University of Florida Natural History Museum, Gainesville, in recognition of his indefatigable efforts for sampling marine benthic invertebrates worldwide, and by his kind support of my research interests. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	description	Variation. Paratype (UF 4223) grayish, paler medially and posteriorly (Fig. 21 A), without posterior end, 17 mm long, 2 mm wide, 11 chaetigers. Prostomium completely exposed; lateral antennae and palps of similar length; median antenna without tip, not reaching anterior prostomial margin. Anterior eyes reniform, twice larger than posterior round ones. Nuchal organs lobes expanded laterally, slightly beyond prostomial lateral margins, ridges darker than surrounding areas (Fig. 21 B). Lateral ciliated areas barely visible, especially along left side. Pharynx not exposed, laterally dissected; middorsal jaw complete, twice wider than lower jaw. Notacicular lobe digitate, neuracicular lobe conical, blunt (Fig. 21 C). Neurochaetae about 15 per bundle; neuracicular blades 3 – 7 times longer than wide; blades approaching subdistal tooth (Fig. 21 C, insets).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	discussion	Remarks. Lamprophaea paulayi n. sp. belongs in the group with notochaetae from chaetiger 4, and resembles L. poupini n. sp. from the Society Islands because they both have reniform anterior eyes, twice larger than posterior round eyes. These two species differ because in L. paulayi lateral antennae are as long as palps, and neuracicular lobes are blunt, conical, whereas in L. poupini lateral antennae are longer than palps, and neuracicular lobes are rectangular.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC47305FF23FD76F1096231.taxon	distribution	Distribution. Coastal localities in the Red Sea Saudi Arabian region, in rocky substrates in 3 – 30 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCA7306FF23FC78F1386311.taxon	description	Fig. 22 urn: lsid: zoobank. org: act: 89978 CB 2 - F 100 - 4705 - A 7 DF-D 16 AECD 663 CA	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCA7306FF23FC78F1386311.taxon	materials_examined	Type material. Western Pacific. Marshall Islands. Holotype (USNM 23961), Ralik Chain, Bikini Atoll, Bikini Island, outer reef, 178, VI, 14 Aug. 1947, F. M. Bayer & F. C. Zimmerman, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCA7306FF23FC78F1386311.taxon	description	Description. Holotype (USNM 23961), complete, damaged, bent spirally (Fig. 22 A); right parapodia of chaetigers 10 (kept in container), and 13 (not found) previously removed. Body obconic, blunt anteriorly, tapered posteriorly, colorless, 42 mm long, 8 mm wide, 16 chaetigers. Anterior and posterior cirri on site, middle body ones missing. Prostomium wider than long, wider anteriorly. Lateral antennae twisted, 1.5 times longer than prostomium, ceratophores distinct, antennae 1.5 times longer than palps. Palpophores pale, twice longer than palpostyles (Fig. 22 B). Median antenna without tip, twisted, inserted between posterior eyes. Eyes brownish, anterior and posterior eyes round, anterior ones twice larger, more distant to each other than posterior ones; in lateral view, eyes separated from each other (Fig. 22 C). Nuchal organs lobes L-shaped, pale, almost completely concealed by tentacular belt, leaving anterior portions of lateral branches exposed. Tentacular cirri damaged, twisted, longest ones reaching chaetiger 7 (probably longer but too delicate to be pulled backwards to its limit). Lateral cushions low, most distorted, entire, longitudinal striae barely visible. Pharynx fully exposed, anterior margin crenulated. Upper and lower jaws single, submarginal, brownish, upper jaw larger than ventral one (Fig. 22 D). Dorsal cirri of middle chaetigers missing, relative length to body width unknown. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, in bundles, subdistally denticulate, denticles coarse. Notacicular lobes tapered (Fig. 22 E, inset), shorter than neuracicular ones; neuracicular lobes oblique conical to rectangular, as long as wide (Fig. 22 E). Neurochaetae about 30 per bundle, blades decreasing in size ventrally, 3 – 7 times longer than wide, blades bidentate, guard approaching subdistal tooth (Fig. 22 F). Posterior region damaged; prepygidial segment with dorsal cirri three times longer than ventral ones. Pygidium distorted by compression, anal cirri without tips, reach chaetiger 13. Gonads present throughout body. Oocytes not seen, only testis.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCA7306FF23FC78F1386311.taxon	etymology	Etymology. This species name is after the late Dr. Marian H. Pettibone in recognition of her many publications on polychaetes, and especially because of her revision of Leocrates Kinberg, 1866, which has been very useful for my efforts in understanding this group. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCA7306FF23FC78F1386311.taxon	discussion	Remarks. Lamprophaea pettiboneae n. sp. belongs in the group of species provided with notochaetae from chaetiger 5. The dorsal pigmentation and the relative size of palpophores to palpostyles separate this species, such that L. pettiboneae resembles L. longicirrata (Treadwell, 1902) from the Antilles because they are both pale. However, in L. pettiboneae palpophores are twice longer than palpostyles, whereas they are three times longer in L. longicirrata.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCA7306FF23FC78F1386311.taxon	distribution	Distribution. Only known from the type locality in the Marshall Islands, in shallow coralline substrates.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC97300FF23F8D0F5C56289.taxon	description	Fig. 23 urn: lsid: zoobank. org: act: 20 AA 6 D 5 F-EF 54 - 4 FC 9 - A 590 - E 52645074137	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC97300FF23F8D0F5C56289.taxon	materials_examined	Type material. Western Indian Ocean, La Réunion Island. Holotype (UF 659), Saint-Leu, Sec Jaune (21 ° 09 ’ 11.88 ” S, 55 ° 16 ’ 51.96 ” E), rocky slope, basalt blocks, 6 – 15 m, H. Bruggemann & N. Hubert, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC97300FF23F8D0F5C56289.taxon	description	Description. Holotype (UF 659) complete, mature, bent ventrally, and laterally, distorted by dissection of some posterior parapodia. Body obconic, blunt anteriorly, tapered posteriorly, 22 mm long, 3 mm wide (without parapodia); right parapodia of chaetigers 9 – 12 removed for molecular analysis; left parapodium of chaetiger 8 removed for observing parapodial features. Body blackish along anterior four chaetigers (Fig. 23 A) and posterior three ones, middle region brownish; parapodia pale, midventral depression blackish, with abundant transverse oval, whitish points. Prostomium slightly wider than long, tapered posteriorly, anterior margin blackish, posterior margin brownish (Fig. 23 B). Lateral antennae with ceratophores distinct, longer than prostomium and palps; palpophores 2 – 3 times longer than palpostyles. Median antenna long, slightly surpassing anterior prostomial margin, inserted centrally on prostomium, between eyes. Eyes dark brown, anterior ones emarginate anterolaterally, about as large as and slightly more distant to each other than posterior round eyes, in lateral view eyes distinct. Nuchal organs lobes L-shaped, lateral projections slightly progressively expanded, with black ridge, anterior and posterior surfaces paler, projected beyond lateral prostomial margins; lateral ciliated bands narrow, barely visible dorsally. Tentacular cirri banded, blackish, pigmentation fading out, longest ones reaching chaetiger 8. Lateral cushions low, variably projected, smooth, entire, longitudinal striae visible. Pharynx exposed, jaws single, inserted subdistally (Fig. 23 C). Lateral vesicles not seen. Anterior margin with about 20 regular ridges, slightly darker than surrounding pharynx wall, dorsal ones tapered. Middorsal jaw yellowish, larger than ventral one, inserted below pharynx margin. Dorsal cirri with tips eroded, complete ones smaller than body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, delicate, in fascicles along chaetigers 1 – 9, thereafter in longitudinal fans, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes long, tapered, three times longer than wide; neuracicular lobes slightly longer than wide, blunt (Fig. 23 D); aciculae black, tapered; ventral cirri tapered, missing in some chaetigers, surpass neurochaetal lobes. Neurochaetae about 30 per bundle, blades decreasing in size ventrally, bidentate, 4 – 9 times longer than wide, guards approaching subdistal tooth. Posterior region tapered, venter blackish (Fig. 2 F). Prepygidial segment with cirri broken, dorsal one twice wider than ventral one. Anal cirri broken. Oocytes in ovaries, about 100 µm in diameter; spermatids not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC97300FF23F8D0F5C56289.taxon	etymology	Etymology. The specific name is a modest homage to Dr. Fredrik Pleijel, in recognition of his series of publications dealing with taxonomy and phylogeny of hesionids, which have made things easier for anyone interested in the group (including myself), and to provide thanks for his kind support of my research activities during the last 20 years. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC97300FF23F8D0F5C56289.taxon	discussion	Remarks. Lamprophaea pleijeli n. sp. belongs in the group of species with notochaetae from chaetiger 5, and resembles L. cornuta n. sp., from Moorea, by having a blackish pigmentation, palpophores three or more times longer than wide, and peristomial middorsal tubercle truncate. These two species differ, however, especially in the relative size of eyes, and in ventral pigmentation pattern. In L. pleijeli eyes are of similar size, and its midventral depression is blackish, whereas in L. cornuta anterior eyes are twice larger, and midventral depression is paler.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFC97300FF23F8D0F5C56289.taxon	distribution	Distribution. Only known from the Western Indian Ocean, in basalt rocky bottoms in La Réunión Island, in water depths of 6 – 15 m.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCF7301FF23FC20F48C6721.taxon	description	Figs 2 D, 24 urn: lsid: zoobank. org: act: DE 8 BF 0 E 2 - 7217 - 433 E-B 03 F- 1 ADE 1 FBAE 48 C	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCF7301FF23FC20F48C6721.taxon	materials_examined	Type material. Western Pacific, French Polynesia. Holotype (UF 858), Society Islands, Moorea, Temae, northern end, near lighthouse, outer reef slope (17 ° 28 ’ 44.40 ” S, 149 ° 45 ’ 51.48 ” W), rubble cracking, 29 m, 17 Oct. 2008, J. Poupin, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCF7301FF23FC20F48C6721.taxon	description	Description. Holotype (UF 858) complete, bent ventrally, with adsorbed salt particles along integument (Fig. 24 A). Body blunt anteriorly, wider medially, tapered posteriorly, 32 mm long, 4 mm wide, 16 chaetigers; most cirri missing; right parapodium of chaetiger 9 removed for observing parapodial features. Body blackish, darker along chaetigers 1 – 6, progressively paler therafter; parapodia and venter pale; some remaining dorsal cirri blackish to finely banded. Prostomium wider than long, widest medially, tapered posteriorly (Fig. 24 B). Lateral antennae with ceratophores distinct, longer than prostomium and palps; palpophores three times longer than palpostyles. Median antenna long, surpassing anterior prostomial margin, inserted centrally on prostomium, among eyes. Eyes dark brown, anterior ones reniform or markedly emarginate, twice larger and more distant to each other than posterior, round eyes (Fig. 24 B). Nuchal organs lobes L-shaped, lateral projections completely blackish, slightly progressively expanded, projected beyond lateral prostomial margins; lateral ciliated bands wide, visible dorsally. Tentacular cirri almost all missing, one ventral cirrus without tip surpasses chaetiger 2. Lateral cushions swollen in middle and posterior chaetigers; longitudinal striae visible. Pharynx almost fully everted (Figs 2 D, 24 C). Lateral vesicles not seen. Anterior margin with 18 regular wide lobes. Jaws single, dorsal one large, hyaline to yellowish, larger and closer to anterior margin than ventral jaw. Dorsal cirri without tips, as long as body width. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, about 60 per bundle, arranged as transverse fans in most chaetigers; notochaetae subdistally denticulate, denticles coarse. Notacicular lobes tapered, 2 – 3 times longer than wide (Fig. 24 D); aciculae black, tapered; ventral cirri tapered, surpassing neurochaetal lobe. Neuracicular lobe rectangular, slightly longer than wide, neurochaetae about 30 per bundle, some blades missing, blades decreasing in size ventrally, bidentate, 2 – 11 times longer than wide, guards smooth, approaching subdistal tooth. Posterior end tapered. Prepygidial segment with dorsal cirri without tips, three times longer than ventral ones. Pygidium with anus terminal, anal cirri missing. Oocytes not seen. Gonad fragments in parapodial spaces include only spermatids.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCF7301FF23FC20F48C6721.taxon	etymology	Etymology. This species is named to honor Dr. Joseph Poupin, Institut de Recherche, École Navale et Groupe des Écoles du Poulmic, a well-known expert on decapod crustaceans from the tropical Indian and Pacific Oceans, because he collected the specimen used to describe the species. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCF7301FF23FC20F48C6721.taxon	discussion	Remarks. Lamprophaea poupini n. sp. belongs in the group with notochaetae from chaetiger 4, and resembles L. paulayi n. sp. from the Red Sea by having reniform anterior eyes, twice larger than posterior round eyes. The two main differences between these two species are that in L. poupini lateral antennae are longer than palps, and neuracicular lobes are rectangular, whereas in L. paulayi lateral antennae are as long as palps, and neuracicular lobes are blunt, conical.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCF7301FF23FC20F48C6721.taxon	distribution	Distribution. The holotype was collected in rubble, in coralline substrates in 29 m depth in Moorea, Society Islands.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCE7303FF23F98BF79C60D9.taxon	type_taxon	Type species: Leocrates chinensis Kinberg, 1866, by monotypy.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCE7303FF23F98BF79C60D9.taxon	diagnosis	Diagnosis (modified after Wang et al. 2018). Hesioninae with two lateral antennae, and one median antenna on dorsal prostomial surface. Palps biarticulate, palpophores large, massive, palpostyles smaller, blunt. Eyes dark, black or brown, usually anterior ones larger than posterior ones, sometimes approaching each other in lateral view. Nuchal organs lobes horizontal C-shaped, posterior ciliated bands close to each other. Pharynx with single, fang-shaped upper and lower jaws. Parapodia sesquiramous along a few anterior chaetigers, biramous thereafter. Notochaetae from chaetiger 5, subdistally denticulate, delicate, sometimes abundant, size variable, never reaching neurochaetal tips. Neurochaetae compound falcigers, blades bidentate, guards approaching subdistal tooth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCE7303FF23F98BF79C60D9.taxon	etymology	Etymology. Taxonomists of XIX and early XX centuries, as most educated citizens of those times, were well acquainted with Latin as well as with Greek history and mythology. This could explain why the etymology for new taxa was rarely explained in taxonomic publications. Inserting a section for explaining the meaning of the new names started in the late XIX and early XX centuries. Among polychaete specialists, Johnson (1897) and Chamberlin (1919) were the first to introduce explanations for names derived from Greek or Latin, but not for demonyms, or names derived after geographical locations. A few precedent authors such as Grube, Levinsen, Schmarda, and Willey used a Latin diagnosis for their new taxa, and sometimes a complete Latin description, such that these statements might be regarded as indirect etymologies. Consequently, because of this widespread lack of etymological sections, we should often guess about the meaning of the newly proposed taxa names. In Leocrates Kinberg, 1866, we have two alternatives to assess its etymology. One deals with a soldier that had a relevant role during the fight between Athens and Aegina in 458 BC, and probably also in the First Peloponnesian War (431 BC). The other deals with a blacksmith that left Athens in 338 BC, soon after the Macedonians took the city. The blacksmith was charged on treason by Lycurgus, in a speech that was regarded as “ the perfect school text ” or as “ one of the most idiosyncratic and non-representative texts in the classical Athenian oratorical corpus ” (Allen 2000: 6). There were several editions available during Kinberg’s lifetime (some available in Google include Maetzner 1836). One can further speculate that if the holotype was alive, the name might proceed from the latter alternative if Kinberg regarded its fast movements as an escaping behavior. Ehlers (1908) introduced Leocratides to indicate its resemblance to Leocrates by adding the Greek suffix – ide, from – eidos: resembling (Jaeger 1932: 73; Brown 1956: 432). The species of Leocratides have sesquiramous parapodia, instead of biramous ones, as is the case in Leocrates. As indicated above, there are two alternatives for the origin of the name, both based upon male Ancient Greek citizens. Because there are many Greek names used as such for genus-group names, there is no need to modify this name. Gender. Masculine.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCE7303FF23F98BF79C60D9.taxon	discussion	Remarks. Leocrates Kinberg, 1866 with Leocrates chinensis Kinberg, 1866 as its type species, and Tyrrhena Claparède, 1868, with T. claparedii Costa in Claparède, 1868, as its type species, both match the diagnosis for the restricted Leocrates as herein defined. This synonymy was proposed by Gravier (1900: 174), McIntosh (1908: 130), Chamberlin (1919: 190), and Horst (191: 80). Further, Lamproderma Grube, 1877, with L. longicirre Grube, 1877, as its type species was described based on a small specimen from New Britain Island, Papua New Guinea. The genus was diagnosed (Grube 1877: 525) as resembling Hesione - like genera but differed by having biarticulate palps, two antennae, 3 pairs of tentacular cirri, pharynx with upper and lower jaws, sesquiramous and biramous parapodia, with simple notochaetae and compound neurochaetae. Pleijel (1998: 108) studied the type specimen and correctly concluded it was a junior synonym of Leocrates; he also regarded its type species as a junior synonym of Leocrates chinensis Kinberg, 1866. The type specimen of L. longicirre (ZMB 905) is in poor condition (three parapodia in one slide are of little help, one was dried-out before mounting; the two others are mounted vertically). The holotype consists of two fragments, an anterior one 5 mm long, 1.5 mm wide, 6 / 7 chaetigers, and a posterior one 5 mm long, 1.5 mm wide, 6 chaetigers (middle segments are missing). The prostomium has lateral antennae as long as palps, palpophores are slightly longer than palpostyles, which is common in juvenile specimens, and prostomial surface does not show any trace of median antennae (even after Shirlastain-A staining), nor eyes, and the original description indicated anterior eyes were larger than posterior ones. Nuchal organs lobes are horizontal C-shaped lobes, matching Leocrates as herein restricted. There are four pairs of tentacular cirri, indicated by their bases, and notochaetae start in chaetiger 5 (not 4 as originally indicated). The notacicular lobe is tapered with notochaetae long, slightly shorter than neurochaetae, with fine denticulation. Neurochaetal blades are bidentate, with guards approach- ing subdistal tooth, and blades are very long to short, 2 – 15 times longer than wide. These features are also present in other tropical Western Pacific species which are newly described below, but the lack of details for eye size and shape, together with the relative size and position of the median antenna, missing since the original description, indicates this species should be regarded as indeterminable. In addition, Lamproderma Grube, 1877 (Annelida) is a junior homonym of Lamproderma Rostafinski, 1873 (Protozoa), and a junior synonym of Leocrates after Pleijel (1998: 108). The synonyms of Tyrrhena and Lamproderma with Leocrates are herein confirmed especially regarding the type of nuchal organs lobes, pharynx armature, parapodial features, and type of neurochaetal blades. As indicated above, the synonymy of Lamprophaea Grube, 1867, and Dalhousia McIntosh, 1885 must be rejected based on the differences of nuchal organs lobes, and pharynx armature.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCC731DFF23F995F48E6551.taxon	description	Fig. 25 urn: lsid: zoobank. org: act: 9 D 348238 - CAB 6 - 42 E 1 - 8 BED-A 0856444 EDEE	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCC731DFF23F995F48E6551.taxon	materials_examined	Type material. Indian Ocean. India. Holotype (USNM 48942), and paratype (USNM 1548294), Mirkae Wada, Ratnagiri (16 ° 59 ’ 40 ” N, 73 ° 18 ’ 00 ” E), Maharashtra, Mumbai (Bombay) Province, intertidal, mixed shore, oyster and bivalve shells, rocks and white sand, 28 Aug. 1969, U. D. Gaidwad, coll. [paratype releasing oocytes through dissections made for parapodia; right parapodia of chaetigers 7 and 8 previously removed, left parapodium of chaetiger 10 dissected (all kept with specimen). Body slightly dehydrated, 23 mm long, 2.5 mm wide, 16 chaetigers; some body appendages damaged. Antennae without tips, palps twisted. Eyes brownish, reniform, anterior ones slightly larger than posterior ones, directed anterolaterally, posterior eyes directed posterolaterally; a small anterolateral dissection previously made for observation of jaws. First dorsal cirri complete, longer than body width including parapodia (reaching chaetiger 5), others broken; notacicular lobes tapered, neuracicular lobes round, truncate, slightly directed upward. Neurochaetal blades 1 – 8 times longer than wide; oocytes abundant by chaetigers 7 and 8, rare by chaetiger 10, each about 100 µm].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCC731DFF23F995F48E6551.taxon	description	Description. Holotype (USNM 48942) complete, slightly dehydrated medially and posteriorly, bent laterally (Fig. 25 A). Body with parallel sides, blunt anteriorly, tapered posteriorly, 18 mm long, 2 mm wide, 16 chaetigers; left parapodium of chaetiger 7 dissected (kept in container). Tentacular and anterior dorsal cirri almost complete, others broken partially or missing. Body pale, eyes brown. Prostomium wider than long, lateral margins curved, wider medially (Fig. 25 B). Lateral antennae with ceratophores distinct, antennae 1.5 times longer than prostomium, barely longer than palps. Palpophores three times longer than palpostyles. Median antenna missing, inserted between posterior eyes. Eyes dark brown, reniform, anterior eyes each 1 / 12 prostomial width, twice larger than posterior ones, slightly emarginate anterolaterally, more distant to each other than posterior eyes, posterior eyes emarginate posterolaterally. Nuchal organs lobes horizontal C-shaped, completely concealed by anterior margin of tentacular belt; lateral ciliated bands narrow, barely visible. Tentacular cirri without tips, longer ones reaching chaetigers 6 / 7. Lateral cushions low, entire, collapsed by partial dehydration. Longitudinal striae visible along middle and posterior chaetigers. Pharynx fully exposed, slightly expanded distally. Lateral vesicles present on both sides, collapsed, tapered. Anterior margin with 22 regular constrictions, ventrally smaller, a larger one middorsally, three smaller ones midventrally (Fig. 25 C). Dorsal and ventral jaws brownish, exposed, tapered, ventral jaw smaller than dorsal one. Dorsal cirri at least as long as body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, delicate, arranged as transverse or oblique fans, notochaetae subdistally denticulate, denticles fine. Notacicular lobes tapered, neuracicular lobes projected, blunt, slightly longer than wide (Fig. 23); aciculae black, tapered; ventral cirri surpassing neurochaetal lobes. Neurochaetae about 30 per bundle, blades decreasing in size ventrally, bidentate, 2 – 10 times longer than wide, guards approaching subdistal tooth (Fig. 23 E, F). Posterior region tapered, almost without cirri. Preanal segment without cirri. Pygidium with anus terminal, anal cirri missing. Oocytes in ovaries, each about 100 – 150 µm; sperm masses not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCC731DFF23F995F48E6551.taxon	etymology	Etymology. This species name is after Kathryn Ahlfeld, collection manager in the National Museum of Natural History, Smithsonian Institution, Washington, because she has been very helpful during my research visits there, and extremely collaborative in dealing with type materials. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCC731DFF23F995F48E6551.taxon	discussion	Remarks. Leocrates ahlfeldae n. sp. belongs in the group of species with small eyes, although anterior eyes twice larger than posterior ones, and with notacicular lobes tapered, together with L chinensis Kinberg, 1866 from Hong Kong, and L. giardi Gravier, 1900 from the Red Sea. However, L. ahlfeldae differs from the two other species by having lateral prostomial margins curved, expanded, and because the lateral pharynx vesicles are tapered, whereas in the two other species prostomial lateral margins are straight, and pharynx vesicles are blunt.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFCC731DFF23F995F48E6551.taxon	distribution	Distribution. The type specimens were collected in Maharashtra, Mumbai (Bombay) Province, in intertidal environments. Some records of L. claparedii (Costa in Claparède, 1866) for the northern Indian Ocean might correspond to this species but specimens were not available for validation. There are records for the Persian Gulf (Fauvel 1919: 371), the Arabian Sea (Aziz 1938: 24, Mohammad 1972: 555), south of Bombay (Day 1973: 344), the Gulf of Mannar (Fauvel 1930: 12 – 13), the Bay of Bengal and Sri Lanka (Fauvel 1932: 61; Fauvel 1953 b: 106 - 107, Fig. 50 c – g).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD27318FF23FB58F48464AA.taxon	description	Figs 26, 27	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD27318FF23FB58F48464AA.taxon	materials_examined	Type material. Western Pacific. Japan. Holotype of Leocrates anonymous Hessle, 1925 (UUZM 672), Sagami, Misaki, intertidal, 5 Jun. 1914, S. Bock, coll. Additional material. Western Pacific. Hong Kong. Seven specimens (UF 5705), Che Lai Pai, ARMS (autonomous reef monitoring structure), submerged for 2 years (22 ° 27 ’ 46.8 ” N, 114 ° 17 ’ 27.6 ” E), 4 m, sandy bottom, 31 Oct. 2017, D. Baker, coll. [15 – 25 mm long, 2.0 – 3.5 mm wide]. One specimen (UF 5737), juvenile, Tung Pin Chau, ARMS submerged for 2 years (22 ° 32 ’ 34.80 ” N, 114 ° 26 ’ 20.40 ” E), 4 m, patchy corals, 24 Oct. 2017, D. Baker, coll. [15.5 mm long, 1.8 mm wide]. One specimen (UF 5747) Centre Island, ARMS submerged for 2 years (22 ° 28 ’ 01.20 ” N, 114 ° 13 ’ 15.60 ” E), 4 m, sandy bottom, 26 Oct. 2017, D. Baker, coll. [26.5 mm long, 2.5 mm wide]. Two specimens (UF 5748), Centre Island, ARMS submerged for 2 years (22 ° 26 ’ 13.20 ” N, 111 ° 13 ’ 15.60 ” E), 4 m, sandy bottom, 26 Oct. 2017, D. Baker, coll. [data used for variation]. Three specimens (UF 5786), largest specimen without posterior region, Che Lai Pai, ARMS submerged for 2 years (22 ° 27 ’ 46.8 ” N, 114 ° 17 ’ 27.6 ” E), 4 m, sandy bottom, 28 Oct. 2017, D. Baker, coll. [19.5 – 20.5 mm long, 2 mm wide]. Japan. One specimen (ECOSUR 3076), Hachijo Island, Yaene (33 ° 05 ’ 55.3 ” N, 139 ° 46 ’ 16.8 ” E), 5 – 12 m, 18 May 2018, N. Jimi, coll. [19 mm long, 2.5 mm wide]. One specimen (ECOSUR 3077), Kagoshima, Bonotsu (31 ° 15 ’ 15.04 ” N, 130 ° 12 ’ 53.90 ” E), 5 – 15 m, 26 Jul. 2018, N. Jimi, coll. [19 mm long, 3 mm wide]. One specimen (SIO A 4147), Izu Peninsula, East Honshu, Suruga Bay, Cape Koganezaki (34 ° 50 ’ 34.80 ” N, 138 ° 45 ’ 46.44 ” E), 4 – 10 m, 24 Oct. 2008, F. Pleijel, coll. [16 mm long, 2 mm wide]. Indonesia. Two specimens (ZMA V. Pol. 534.6), Irian Jaya, R / V Siboga Exped., Sta. 273, Aru Islands (06 ° 10 ’ S, 134 ° 30 ’ E), Pearl Banks (anchorage off Pulu Jedan), 13 m, trawl, dredge and divers, sand and shells, 23 - 26 Dec. 1899 [17 – 18 mm long, 2 – 3 mm wide]. One specimen (ZMA V. Pol. 1902), Sulawesi, R / V Siboga Exped., Sta. 213, Saleyer (06 ° 00 ’ S, 120 ° 30 ’ E) anchorage, 36 m, coral reef exploration, muddy bottom + sand, 26 Sep. 1899 [17 mm long, 2.5 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD27318FF23FB58F48464AA.taxon	description	Description. Holotype of L. anonymous Hessle, 1925 (UUZM 672) complete, slightly bent laterally (Fig. 26 A), integument variably damaged. Body with parallel sides, blunt anteriorly, barely tapered posteriorly, 29 mm long, 3 mm wide (without parapodia), 16 chaetigers; left parapodia of chaetigers 4 and 8 previously removed, left parapodia of chaetigers 7 and 15 dissected (kept in container). Most tentacular and dorsal cirri on site, most without tips, some ventral cirri missing. Body pale, eyes pale brown (barely pigmented). Prostomium wider than long, slightly wider anteriorly (Fig. 26 B). Lateral antennae with ceratophores welldefined, antennae shorter than prostomial length, slightly longer than palps. Palpophores 3 – 4 times longer than palpostyles. Median antenna broken, probably not reaching anterior prostomial margin, inserted between posterior eyes. Eyes pale brown round, anterior eyes slightly larger than posterior ones. Nuchal organs lobes concelead by tentacular belt, horizontal C-shaped; lateral ciliated bands barely visible. Tentacular cirri mostly on site, without tips, longer ones reach chaetiger 7. Lateral cushions low, entire, longitudinal striae visible. Pharynx fully exposed, slightly expanded distally. Lateral vesicles present on both sides, swollen, round (Fig. 26 B – D). Anterior margin smooth, subdistal circle of irregular constrictions, better defined laterally. Dorsal jaw brownish, exposed, inserted in pharynx margin, in lateral view scimitar-like; ventral jaw exposed, submarginal, smaller than upper one. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, up to 30 per bundle, most broken, delicate, arranged in transverse fans, notochaetae subdistally denticulate, denticles fine. Dorsal cirri as long as body width (including parapodia). Notacicular lobes tapered, neuracicular lobes projected, blunt, tips round, 1.5 times longer than wide (Fig. 26 E); aciculae black, tapered; ventral cirri surpassing neurochaetal lobes. Neurochaetae about 20 per bundle (only one with hood), blades decreasing in size ventrally, bidentate, 4 – 10 times longer than wide, guards approaching subdistal tooth. Posterior region tapered, with several lateral and anal cirri on site, but tips broken. Prepygidial segment with dorsal cirri three times longer than ventral ones. Pygidium with anus terminal, a small lateral fracture, anal cirri long, without tips, reaching chaetiger 15. Oocytes not seen. Variation. Two topotypes (UF 5748) complete, 26 mm long (other topotypes 15.0 – 26.5 mm long), 2 – 3 mm wide, 16 chaetigers. Left parapodium of chaetiger 8 removed. Both specimens with lateral antennae longer than prostomium, slightly longer than palps; palpophores three times longer than palpostyles (Fig. 27 A, D). Eyes brownish, anterior ones each 1 / 10 prostomial width, twice larger, anterolaterally slightly emarginate, and farther apart to each other than posterior ones, round. Median antenna shorter than prostomium. Pharynx partially exposed in smaller specimen; lateral vesicles present only on left side, round; anterior margin with 24 regular constrictions, ventral ones smaller (Fig. 27 B). Dorsal and ventral jaws hyaline, yellowish, exposed, tapered, ventral jaw smaller than dorsal one. Dorsal cirri as long as body width including parapodia (Fig. 27 C, E). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, delicate, denticles coarse (visible in 10 X). Neurochaetae about 30 per bundle, bidentatae, 4 – 11 times longer than wide, teeth of about the same size. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD27318FF23FB58F48464AA.taxon	discussion	Remarks. Leocrates chinensis Kinberg, 1866 resembles L. giardi Gravier, 1900 from the Red Sea because both species have small eyes, but anterior ones larger than posterior ones, and straight prostomial lateral margins. However, there are some differences between these two species, especially regarding length of neurochaetal blades, and notochaetal denticulation. In L. chinensis neurochaetal blades are 3 – 18 times longer than wide, and notochaetae have coarse denticles, whereas in L. giardi neurochaetal blades are 3 – 10 times longer than wide, and notochaetae have fine denticles. Further, L. chinensis resembles L. ahlfeldae n. sp. from India because both have small eyes, notacicular lobes tapered, and neurochaetal blade with teeth of similar size. There is a difference regarding the relative position of eyes when seen from above, that deserves explanation. The anterior eyes are always set farther apart than posterior eyes; however, posterior eyes might be sligthly, or markedly displaced medially. In the first condition, the posterior eyes can have a higher relative overlap along the level of the anterior eyes, and be more distant to each other, whereas if they are more displaced dorsally, they will have a smaller relative overlap with anterior eyes, and be closer to each other. Consequently, the differences between these two species are that in L. chinensis the lateral vesicles are round, and their posterior eyes have a slight overlap with anterior eyes, appearing slightly closer to each other than anterior eyes; whereas in L. ahlfeldae lateral vesicles are tapered, and posterior eyes position falls within anterior eyes level, with a slight overlap, appearing closer to each other than anterior eyes. Leocrates anonymus Hessle, 1925 has been regarded as a possible junior synonym of L. chinensis Kinberg, 1866 by Hessle (1925: 17), Monro (1926: 313), Pettibone (1970: 215), and Pleijel (1998: 160). Their conclusions are herein confirmed. The only difference relies on the shape of the pharynx upper jaw because in the holotype of L. anonymous it is scimitar-shaped, whereas it is subcylindrical and tapered in L. chinensis. This difference can be explained by the size of the specimen, such that larger specimens have a midventral keel projection, and this was also confirmed in some larger topotypes of L. chinensis. Leocrates chinensis Kinberg, 1866 was redescribed recently based upon topotype specimens (Wang et al. 2018); however, these additional topotypes are slightly larger (previously studied ones up to 23 mm long), and some features are indicated and illustrated, such as the pharynx lateral vesicle. The records for the Mediterranean Sea were probably following Pettibone’s (1970) synonymy, which was probably derived from Hartman (1965: 24), such as Sordino (1990: 37 - 38), should be referred to L. claparedii (Costa in Claparède, 1868). On the other hand, there are several records from the Western Pacific, but could not be confirmed because no specimens were available, such as Augener (1922 a: 22, 1927: 131 – 132), Monro (1931: 12), Pleijel (1998: 111, Figs 6 – 8 (partim, perhaps only Fig. 6), and Pleijel & Gustavsson (2010: 95, Fig. 1 d).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD27318FF23FB58F48464AA.taxon	distribution	Distribution. Japan to Indonesia, including Hong Kong, in sandy or mixed bottoms in shallow areas (4 – 15 m depth).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD7731AFF23FA04F7116402.taxon	description	Figs 2 A – C, 28	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD7731AFF23FA04F7116402.taxon	materials_examined	Non-type material. Mediterranean Sea. Naples. One specimen (MCZ ANNa 588), purchased from the Naples Zoological Station, no further data [25 mm long, 3 mm wide]. Two specimens (USNM 5131), purchased from the Stazione Zoologica di Napoli, 1893 [used for description]. Croatia. One specimen (MMS HESI 6), Rovigno d’Istria, no further data [32 mm long, 4.5 mm wide]. Western Africa. Two specimens (NHMD 237479), Dakar, Senegal, rocky coast, 7 Jan. 1928, H. Madsen, coll. [25 – 28 mm long, 2 – 3 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD7731AFF23FA04F7116402.taxon	description	Description. Non-type specimens (USNM 5131) complete, larger one slightly damaged, many cirri missing, with posterior end collapsed by compression, probably after being depressed by labels; some parapodia previously removed (kept in container); both specimens previously pricked middorsally in an anterior chaetiger. Body pale, 25 – 29 mm long, 2.5 – 3.0 mm wide; both specimens with pharynx partially (Fig. 28 A) or fully exposed (Fig. 28 D). Prostomium slightly wider than long (Fig. 28 B), or as long as wide (Fig. 28 E), slightly wider anteriorly, lateral margins straight. Lateral antennae with ceratophores distinct, lateral antennae longer than prostomium, slightly longer than palps. Palpophores 3 – 4 times longer than palpostyles; median antenna short, inserted between posterior eyes, reaching prostomial centre. Eyes small, brownish, anterior ones each 1 / 12 prostomial width, slightly larger, slightly emarginate to reniform than posterior round smaller ones. Nuchal organs lobes almost completely covered by anterior margin of tentacular belt, barely projected posteriorly. Tentacular cirri damaged or broken, largest specimen with tentacular cirri reaching chaetiger 5. Lateral cushions low, entire or divided into two sections, longitudinal striae visible. Basal ring with middorsal tubercle widened basally, tapered into a blunt tip, or damaged; dorsolateral tubercles not seen. Ventrolateral tubercles barely visible. Pharynx partially everted, divided into two rings; one specimen with one left lateral vesicle, the other without vesicles; jaws exposed, upper jaw scimitar-like, lower jaw smaller. Anterior margin smooth or with about 20 irregular constrictions. Jaws single, brownish, upper jaw slightly larger and inserted ahead of ventral jaw. Dorsal cirri broken, shorter than body width (excluding parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 30 per bundle, sometimes expanded as a fan, most broken. Notacicular lobes tapered; neuracicular lobes round, truncate, as long as wide. Ventral cirri reaching or surpassing neurochaetal lobe (Fig. 28 C, F). Neurochaetae about 20 per bundle, bidentate, guards approaching subdistal tooth, blades 2 – 8 times longer than wide, guards approaching subdistal tooth (Fig. 28 C, inset, F, insets). Posterior end tapered into a blunt cone; prepygidial segment (in other specimens) with dorsal cirri 4 – 5 longer than ventral ones; anal cirri without tips, reaching chaetiger 13. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD7731AFF23FA04F7116402.taxon	discussion	Remarks. Leocrates claparedii (Costa in Claparède, 1868) resembles L. giardi Gravier, 1900 from the Red Sea because both have prostomia wider anteriorly, notacicular lobes tapered, and middle chaetigers with about 20 – 30 neurochaetae per bundle. These two species differ especially regarding the shape of notacicular and neuracicular lobes, and by the relative length of neurochaetal blades. In L. claparedii notacicular lobes are blunt, neuracicular ones are wider than long, and neurochaetal blades are 2 – 8 times longer than wide, whereas in L. giardi notacicular lobes are tapered, neuracicular ones are as long as wide, and blades are 3 – 10 times longer than wide. The description of L. claparedii did not indicate the type locality, but Claparède (1868: 537, footnote) explained the name for the genus, Tyrrhena, was “ d’après la mer Tyrrhénienne qu’habite l’espèce-type ”. Despite the lack of precision for the type locality, it should be the Gulf of Naples. Previous publications by Oronzio Gabriele Costa (1787 – 1867), and his son, Achille Costa (1823 – 1898), the latter supposed to fully describe the species (Claparède 1868: 538), and even the corresponding monograph by Claparède himself (1868) were made after his research visits in Naples. Pettibone (1970: 215) listed L. claparedii as a junior synonym of L. chinensis Kinberg, 1866, from Hong Kong, but they differ as indicated elsewhere (Wang et al. 2018). It must be emphasized that L. claparedii was described from the Mediterranean, and Fauvel (1923) documented it. Further, against the synonymy by Pettibone, Kirkegaard (1983: 214) reinstated it. Parapar et al. (2004) redescribed the species and provided camara lucida illustrations for the diagnostic features. During the development of this revision, a short visit was made to the Stazione Zoologica Anton Dohrn in Naples to have a better idea of their collection, but it was under reconstruction, and no specimens were available. However, because the species is currently well-understood, and has been properly and recently illustrated, there is no need for proposing a neotype.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD7731AFF23FA04F7116402.taxon	distribution	Distribution. Mediterranean Sea to Northwestern Africa, in shallow water rocky or mixed bottoms. There are many records for this Mediterranean species for other localities, such as the Indian Ocean by Fauvel (1927: 417; 1930: 12 – 13; 1953 b: 106 – 107, Fig. 50 c – g; 1957 b: 4), Monro (1939: 392), Day (1951: 21; 1967: 230, Fig. 11.2 g – k), Fishelson & Rullier (1969: 59), Mohammad (1972: 555), and Amoureux (1974 c: 435). Likewise, for the Western Pacific by Fauvel (1935: 296 - 297; 1937: 60; 1939: 285; 1947 a: 31 – 32, Fig. 28 a – d), Monro (1939: 392), and Rullier (1972: 58), or even for the Western Atlantic by Rullier & Amoureux (1979: 159). Because their specimens were not available, their identity cannot be confirmed.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD57314FF23FAB8F7F665C2.taxon	description	Figs 29, 30	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD57314FF23FAB8F7F665C2.taxon	materials_examined	Type material. Red Sea, Gulf of Aden. Six syntypes (MNHN 289), some collected in Aden (12 ° 46 ’ 25 ” N, 46 ° 01 ’ 35 ” E) by Dr. Jousseaume in 1893, others in Djibouti (11 ° 36 ’ N, 43 ° 09 ’ E), Djibouti, by Drs Jousseaume and Coutière in 1897, kept in the same container; no further data. Additional material. Red Sea. Gulf of Suez. One specimen (MNHN A 419), no further data [20 mm long, 3 mm wide]. Saudi Arabia. Two specimens (MNHN A 71), Mission Bonin-Perez, Côtes d’Arabie, Sta. 48, 1901, no further data [11 – 13 mm long, 2 mm wide]. Two specimens (MNHN A 71 b), Mission Bonin-Perez, Côtes d’Arabie, Sta. 53, 1901, no further data [15 – 16 mm long, 2 – 3 mm wide]. One specimen (MNHN A 372), Farasan Archipelago, Abulat Island (19 ° 58 ’ N, 40 ° 07 ’ E), Sta. 5, NW off islet North, in Porites, 0 – 5 m, 1951 – 1952, Drach, Cherbonnier & Mercier, coll. [12 mm long, 1.8 mm wide]. One specimen (UF 3586), Farasan Islands, Zahrat Durakah (16 ° 50 ’ 09.24 ” N, 42 ° 18 ’ 22.68 ” E), 2 – 6 m, fringing reef, slope around sand, 11 Mar. 2013, A. Anker, P. Norby & G. Paulay, coll. [21 mm long, 3 mm wide]. One specimen (UF 3499), Farasan Islands, Mahama Island, fringing slope (16 ° 29 ’ 21.12 ” N, 41 ° 56 ’ 39.48 ” E), 4 – 17 m, sand, reef rubble, 9 Mar. 2013, A. Anker, P. Norby & G. Paulay, coll. [data used for variation].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD57314FF23FAB8F7F665C2.taxon	description	Description. Syntypes (MNHN 289) distorted, partially dehydrated, most cirri missing, many chaetae broken (Fig. 29 A – C). Body obconic, blunt anteriorly, tapered posteriorly, dorsum darker along anterior chaetigers (1 – 6 / 8), or dark throughout dorsal surface; some middle parapodia previously removed. Body 18 – 23 mm long, 1.5 – 3.0 mm wide, 16 chaetigers. Prostomium wider than long in smaller syntypes (Fig. 29 A), as long as wide or slightly longer than wide in larger ones (Fig. 29 B, C); slightly wider anteriorly (depending on pharynx eversion), lateral margins straight. Lateral antennae with ceratophores distinct, lateral antennae as long as prostomium (longer in smaller syntypes), as long as palps or slightly longer than them. Palpophores 2 – 3 times longer than palpostyles. Median antenna tapered, short, not reaching prostomial anterior margin, inserted between posterior eyes. Eyes brownish, anterior ones slightly larger, about 1 / 10 as wide as prostomial width, missing in a smaller syntype, circular in another, emarginate in two specimens, and semilunar in two others, larger and more distant to each other than posterior round ones (semilunar in two syntypes). Nuchal organs lobes horizontal C-shaped, visible dorsally in most syntypes; lateral ciliated bands dorsally visible in two syntypes. Tentacular cirri missing (reaching chaetiger 4 in original description). Lateral cushions low, entire, longitudinal striae barely visible. Pharynx fully exposed in one small syntype (Fig. 29 D), partially exposed in another medium-sized one. Anterior margin with about 20 regular constrictions per side. Lateral vesicles not seen. Jaws single, yellowish, upper jaw twice larger than lower one, tapered, inserted slightly ahead of lower jaw. Dorsal cirri remaining in a few segments, without tips, shorter than body width. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 20 per bundle, sometimes arranged in longitudinal fans, or as a bundle, subdistally denticulate, denticles fine. Notacicular lobes tapered; neuracicular lobes blunt, conical slightly longer than wide, sometimes as wide as long (Fig. 29 E); ventral cirri surpassing neurochaetal lobe. Neurochaetae about 20 per bundle, blades bidentate, decreasing in size ventrally, 3 – 10 times longer than wide, guards approaching subdistal tooth (Fig. 29 F, G). Posterior region tapered. Preanal segment with dorsal cirri missing. Pygidium with anus terminal, anal cirri missing. Oocytes about 100 µm in diameter, visible in smaller and larger syntypes. Variation. One specimen (UF 3499) complete, straight. Body pale up to chaetiger 5, thereafter brownish; 16 mm long, 2 mm wide, 15 chaetigers (posterior end removed for molecular analysis; right parapodium of chaetiger 8 removed for parapodial features). Lateral antennae longer than prostomium, slightly longer than palps (Fig. 30 A); palpophores twice longer than palpostyles; median antenna broken, inserted between posterior eyes. Eyes brownish, anterior ones twice larger than posterior ones. Longest tentacular cirri reaching chaetiger 7. Pharynx fully exposed (Fig. 30 A – C), lateral vesicles tapered, upper and lower jaws exposed, hyaline, upper one twice larger than lower one. Notochaetae along chaetigers 5 – 15, many broken, denticulation fine; neurochaetae about 20 per bundle (Fig. 30 D), blades 2 – 9 times longer than wide, guards approaching subdistal tooth (Fig. 30 D, insets). Oocytes not seen, coelom full of damaged testis fragments and spermatids. Additional specimens 11 – 21 mm long, collected in subtidal environments (2 – 17 m depth). Two specimens (UF 3499, 3586) smaller than syntypes (16 – 21 mm long) had pharynxes fully exposed, showing lateral vesicles, each with well-defined tips (tapered).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD57314FF23FAB8F7F665C2.taxon	discussion	Remarks. Leocrates giardi Gravier, 1900 resembles L. claparedii (Costa in Claparède, 1868) from the Mediterranean Sea by having prostomia wider anteriorly, notacicular lobes tapered, and middle chaetigers with about 20 – 30 neurochaetae. They differ, however, by the shape of notacicular and neuracicular lobes, and by the relative length of neurochaetal blades. In L. giardi notacicular lobes are tapered, neuracicular ones are as long as wide, and neurochaetal blades are 3 – 10 times longer than wide, whereas in L. claparedii notacicular lobes are blunt, neuracicular ones are wider than long, and blades are 2 – 8 times longer than wide. The redefinition of L. giardi by Pettibone (1970: 219 – 221, Figs 17 – 19) must be discontinued because she did not study type material or topotype specimens, and the specimens she illustrated do not belong to Leocrates, as herein restricted, but to Dalhousia, as redefined above.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD57314FF23FAB8F7F665C2.taxon	distribution	Distribution. Red Sea, subtidal mixed bottoms (2 – 17 m depth).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDA7316FF23FF6FF1666369.taxon	description	Fig. 31 urn: lsid: zoobank. org: act: D 9 D 73 B 24 - 6 B 3 D- 4 F 44 - 9 CAE- 8 B 9 D 007 F 48 B 5	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDA7316FF23FF6FF1666369.taxon	materials_examined	Type material. Eastern Pacific. Revillagigedo Islands. Holotype (LACM 10146), off Sulphur Bay, Clarion Island, R / V Velero III, Sta. 305 (18 ° 20 ’ 30 ” N, 114 ° 44 ’ 30 ” W), 27 m, sand, coralline algae fragments, 11 Jun. 1934. Additional material. Eastern Pacific. Gulf of California. One specimen (LACM 10147), San Gabriel Bay, Isla Espirítu Santo, R / V Velero III, Sta. 633 (24 ° 24 ’ 10 ” N, 110 ° 21 ’ 55 ” W), 32 m, corallines, 6 Mar. 1937 [15 mm long, 2 mm wide]. Juan Fernandez. Two specimens (ZMB 3722), dried-out, and one slide with 5 parapodia, no further data, L. Plate, coll. [specimens carefully brushed in a 1: 1 white vinegar / ethanol solution to remove excess of salt particles; 11 – 14 mm long, 1.5 – 2.0 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDA7316FF23FF6FF1666369.taxon	description	Description. Holotype (LACM 10146) complete, mature, bent laterally, slightly damaged (Fig. 31 A), branching gonads (ovaries) filling coelom. Body obconic, blunt anteriorly, tapered posteriorly, 22 mm long, 4 mm wide, 16 chaetigers; right parapodium of chaetiger 9 previously removed (missing), left parapodium of chaetiger 7 dissected (kept in container), body wall middorsally broken in chaetiger 14, a midventral dissection from mouth to chaetiger 9 previously made (Fig. 31 C). Body pale, eyes brownish. Prostomium slightly wider than long, slightly wider anteriorly (Fig. 31 B). Lateral antennae directed ventrally, with ceratophores distinct, lateral antennae slightly shorter than prostomium, 1.5 times longer than palps; palpophores twice longer than palpostyles (left palpostyle without tip). Median antenna short, surpassing prostomial centre, not reaching prostomial anterior margin, inserted between posterior eyes. Eyes brownish, anterior eyes each 1 / 10 prostomial width, smaller than posterior ones, slightly emarginate anterolaterally, more distant to each other than posterior, round eyes. Nuchal organs lobes horizontal C-shaped, completely concealed by anterior margin of tentacular belt; lateral ciliated bands thin, visible dorsally. Tentacular cirri broken, longest one reaching chaetiger 2. Lateral cushions low, slightly and irregularly projected throughout body. Longitudinal striae visible along anterior and middle segments, posterior ones too distended. Pharynx not exposed, observed through previous dissection (Fig. 31 C). Lateral vesicles not seen. Anterior margin with about 30 regular constrictions. Dorsal and ventral jaws single, brownish, exposed, tapered, ventral jaw smaller than dorsal one. Most dorsal cirri missing, some remaining without tips, size relation to body width unknown. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, broken off in several parapodia, up to 40 per bundle, most arranged in fascicles, notochaetae subdistally denticulate, denticles fine (not visible in 10 x). Notacicular lobes tapered, blunt, neuracicular lobes projected, slightly longer than wide (Fig. 31 D). Neurochaetae about 20 per bundle, blades decreasing in size ventrally, bidentate, 4 – 11 times longer than wide, guards approaching subdistal tooth, sometimes broken (Fig. 31 E). Posterior region tapered. Preanal segment with cirri broken, dorsal ones 2 – 3 times wider than ventral ones. Pygidium with anus terminal, anal cirri broken, reaching preanal segment. Oocytes in ovaries and loose in coelom throughout body, including parapodial coelomic space, each oocyte about 100 µm. No sperm surrounding oocytes.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDA7316FF23FF6FF1666369.taxon	etymology	Etymology. This species is named as an homage to Leslie Harris, Collection Manager of the Allan Hancock Foundation Polychaete Collection in the Los Angeles County Museum of Natural History, in recognition of her generous help, support, experience and advice during the last 30 yr for our studies on polychaetes. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDA7316FF23FF6FF1666369.taxon	discussion	Remarks. Leocrates harrisae n. sp. is remarkably unique by having anterior eyes of similar size, or smaller than posterior ones. Hartman (1940) recorded this species for Western Mexico as L. claparedii (Costa in Claparède, 1868), originally described from the Mediterranean Sea. The main difference between these two species is the size of palpophores versus palpostyles. In L. harrisae palpophores are smaller, being 1.5 – 2.5 times longer than palpostyles, whereas in L. claparedii palpophores are longer (3 – 4 times longer than palpostyles). In her publication, Hartman (1940) gave no species name after the genus and in the preceding species, she listed first the name she was using, and L. claparedii was inserted above because labels for the specimens and the corresponding jar containing them have L. claparedii written on them. Hartman (1940: 213) made illustrations for prostomial and neurochaetal features, including one with a hood. She also indicated she followed Okuda (1937: 270) who recorded L. claparedii for Japan, but her specimens differ because “ the posterior eyes are as large as the anterior ones, and the middle prostomial antenna is inserted posterior to the eyes. ” The records of L. chinensis for Juan Fernandez by Ehlers (1901) and Augener (1922 b) were based upon two sets of different specimens, each with only two specimens (ZMB 3722), but the Swedish specimens that Augener recorded were not requested for this research. The Berlin specimens are dried-out now. With hesitation, they are regarded as conspecific with L. harrisae on the basis of the illustrations by Ehlers (1901, Pl. 11, Figs 10 – 15), especially regarding the relative size of eyes. These figures show the posterior eyes are as large as the anterior reniform ones, or even slightly larger than them, and because the smaller specimen has some remaining pigmentation and the eye size is similar between anterior and posterior eyes. However, better specimens are needed to clarify their affinities, and concluding about their similarities with L. harrisae.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDA7316FF23FF6FF1666369.taxon	distribution	Distribution. Revillagigedo Islands, off Western Mexico, among corallines in 32 m water depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD97312FF23F892F7C96368.taxon	description	Figs 32, 33	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD97312FF23F892F7C96368.taxon	materials_examined	Type material. Philippines and Samoa. Four syntypes (MNHW 296), in two lots, each with two specimens, number 367 includes a small, dried-out specimen; the others damaged [body 15 mm long, 3 mm wide, 16 chaetigers. Prostomium as long as wide; lateral antennae without tips, as long as palps; palpophores 3 times longer than palpostyles; median antenna without tip, not reaching prostomial anterior margin. Eyes colorless, anterior eyes larger than posterior ones. Nuchal organs lobes horizontal C-shaped. Notochaetae from chaetiger 5; neurochaetae about 30 per bundle, blades long to short], and the other lot contains specimens 714 and 780 [in better condition, best one used for redescription below, the other slightly damaged. Body 14 mm long, 3 mm wide, 16 chaetigers. Eyes colorless, reniform, at least right anterior eye]. Additional material. Philippines. One specimen (ZMA V. Pol. 534.2), Sulu Islands, anchorage off North Ubian, R / V Siboga Exped., Sta. 99 (06 ° 07.5 ’ N, 120 ° 26 ’ E), 16 – 23 m, dredge and tow-net, lithothamnion bottom, 28 - 30 Jun. 1899 [12 mm long, 2 mm wide]. One specimen (USNM 18966), USFS Albatross, Philippine Expedition 1907 – 1909, Sta. 5149, Sulu Archipelago, off Siasi, 4.5 km E off Sirun Island (05 ° 33 ’ 00 ” N, 120 ° 42 ’ 10 ” E), 18 m, corals and shells, 18 Feb. 1908 [16 mm long, 3 mm wide]. Samoa. One specimen (USNM 1548297), Pago Pago (14 ° 16 ’ 46 ” S, 170 ° 42 ’ 02 ” W), May 1920, A. L. Treadwell, coll. [used for redescription]. Indonesia. One specimen (ZMA V. Pol. 534.1), Lesser Sunda Islands, R / V Siboga Exped., Sta. 53, Waingapoe (Waingapu, 09 ° 39 ’ S, 120 ° 16 ’ E), Sumba, Bay of Nangamessi, 36 m, trawl and shore exploration, coral sand, near the shore, mud, 21 - 22 Apr. 1899 [13 mm long, 2.5 mm wide]. One specimen (ZMA V. Pol. 534.4), Sulawesi, R / V Siboga Exped., Sta. 125, Siau Island (02 ° 45 ’ N, 125 ° 24 ’ E), anchorage off Sawan, 27 m, dredge, townet, Monaco trap, reef-exploration, stones, 18 - 19 Jul. 1899 [8.5 mmm long, 1.5 mm wide]. One specimen (ZMA V. Pol. 534.5), Maluku, R / V Siboga Exped., Sta. 231 Ambon (03 ° 42 ’ S, 128 ° 10 ’ E), at anchorage, 40 m, reef expl., coral, sand, 14 - 18 Nov. 1899 [19 mm long, 2.8 mm wide]. Three specimens (ZMA V. Pol. 534.7), Lesser Sunda Islands, R / V Siboga Exped., Sta. 315, Paternoster Island, anchorage E of Sailus Besar, 36 m, dredge, coral, 17 - 18 Feb. 1900 [7 – 11 mm long, 1 – 2 mm wide]. Vietnam. One specimen (MNHN A 71), southern region, Nam K ỳ or Cochinchine, 1906, Capt. St. Jacques & Capt. Modest (no further data) [15 mm long, 2 mm wide]. Marshall Islands. One specimen (LACM 10885), Ralik Chain, Eniwetok Atoll, Lidilbut Island, corals, Sta. 1503 (no further data), May 1952, H. S. Ladd, coll. [15 mm long, 3 mm wideMiddle]. Fiji. Two specimens (USNM 19193), Suva (18 ° 08 ’ 29.76 ” S, 178 ° 26 ’ 30.84 ” E), Apr. 1920, A. L. Treadwell, coll. (12 – 18 mm long, 1.5 – 2.0 mm wide]. French Polynesia. One specimen (UF 3056), Society Islands, Moorea, Vaipahu (17 ° 28 ’ 12.00 ” S, 149 ° 49 ’ 48.00 ” W), barrier reef, 0 – 2 m, algae and rocks, 24 Jan. 2012, M. Leray & L. Penland, coll. [14 mm long, 1.5 mm wide]. New Caledonia. One specimen (MNHN Musorstom Lagon Est- 726), Sta. 726 (21 ° 20 ’ 06 ” S, 165 ° 55 ’ 00 ” E), 51 m, fine sand, Halimeda fragments, 12 Aug. 1986, B. Richer de Forges, coll. [13 mm long, 2.5 mm wide]. Australia. One specimen (AM W 4961), Warangol River mouth, Northeast coast (05 ° 30 ’ S, 151 ° 00 ’ E), New Britain, 5 Sep. 1970, Turner et al., coll. [16 mm long, 3 mm wide]. One specimen (AM 43841), Queensland, Lizard Island, Vicki’s Reef (14 ° 41 ’ 17 ” S, 145 ° 26 ’ 35 ” E), 9 m, calcareous algae 13 Aug. 2013, M. Capa & J. Zanol, coll. [12 mm long, 2 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD97312FF23F892F7C96368.taxon	description	Description (topotype details in brackets). Best syntype (MNHW 296, 714 - 780) complete, obconic, blunt anteriorly, tapered posteriorly (Fig. 32 A); right parapodium of chaetiger 8 removed for observing parapodial features. Body colorless, 14 mm long, 3 mm wide; most cirri missing. [Best preserved topotype (USNM 1548297) complete (Fig. 33 A), with a transverse dissection on the left side between chaetigers 3 and 4, and an oblique ventral dissection from mouth to chaetiger 10, one parapodium previously removed; left parapodium of chaetiger 9 dissected (kept in small container). Body colorless, 17 mm long, 2 mm wide. Many cirri on site]. Prostomium slightly longer than wide, slightly wider anteriorly (Fig. 32 B). Lateral antennae with ceratophores distinct, without tips, slightly shorter than prostomium, as long as palps; palpophores 3 times longer than palpostyles (probably eroded). Median antenna short, tapered, not reaching anterior prostomial margin, inserted centrally. [Prostomium longer than wide, slightly wider anteriorly (Fig. 33 B). Lateral antennae without tips, as long as palps, both directed ventrally; palpophores three times longer than palpostyles. Median antenna broken, scar inserted central to eyes (Fig. 33 C)]. Eyes colorless; other features unclear (Grube indicated anterior eyes reniform, one time larger, and slightly more separated than posterior ones). [Eyes dark brown; anterior eyes each 1 / 5 prostomial width, reniform or halfmoon shaped, twice larger than posterior ones, slightly more distant to each other than posterior round ones (Fig. 33 B, C); in lateral view anterior and posterior eyes distinct]. Nuchal organs lobes as rectangular projections, posterior furrow narrow; lateral cilated bands not visible. Tentacular cirri missing (another syntype in same lot with cirri without tips, longest ones reaching chaetiger 5). [Nuchal organs lobes horizontal C-shaped lobes; lateral ciliated bands not visible. Tentacular cirri without tips, longest ones reaching chaetiger 5. Lateral cushions low, barely projected, entire, longitudinal striae visible]. Pharynx exposed. Anterior margin smooth, slightly eroded (Fig. 32 C). Upper jaw larger and more anteriorly inserted than lower jaw. [Pharynx with about 20 marginal constrictions; upper jaw visible, lower one damaged after dissection (Fig. 33 D)]. Dorsal cirri missing (another syntype with cirri without tips, as long as body width including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, abundant, subdistally denticulate, denticles fine (notochaetae soft, surfaces eroded by long time in formalin solution). Notacicular lobes tapered, longer than dorsal cirrophores in middle segments; neuracicular lobes blunt (Fig. 32 D); aciculae black. Neurochaetae about 30 per bundle, blades bidentate, guards decreasing in size ventrally, blades eroded, soft, fine details difficult to be observed, 3 – 12 times longer than wide (Fig. 32 E, F). [Dorsal cirri with tips broken, as long as body width including parapodia. Chaetigers 1 – 4 without notochaetae. Notochaetae present along chaetigers 5 – 16, abundant, coarsely denticulate subdistally. Notacicular lobes tapered, blunt; neurochaetal lobes blunt (Fig. 33 E); aciculae black. Neurochaetae about 30 per bundle, blades decreasing in size ventrally, 4 – 11 times longer than wide; blades bidentate, guard approaching subdistal tooth (Fig. 33 F)]. Posterior region tapered. Prepygidial segment with dorsal cirri 4 – 5 times longer than ventral ones. Pygidium with anus terminal, anal cirri missing. [Posterior region tapered. Preanal segment with dorsal and ventral cirri, cirrostyles broken medially, or completely. Pygidium tubular, anus terminal, anal cirri missing]. Oocytes not seen. [Gonad fragments seen by transparency in some parapodia; oocytes about 100 µm in diameter]. Variation. More recently collected specimens (USNM 19193) with palps and antennae of similar size, and palpophores twice larger than palpostyles (as indicated in the original description), slightly larger in larger specimen. Anterior eyes are less pigmented but about twice larger than posterior ones, and with a large anterolateral emargination, such that they are reniform, but their contourns are less well-defined. In both, notacicular lobes are tapered, neuracicular lobes conical, blunt, about as long as wide. It is interesting that these specimens were slightly darker than the one from Samoa (USNM 19194 i). Provided they were collected and processed under similar conditions, this difference might be due to the fact that the Samoan specimen was longitudinally dissected, and the fixative might change the basic body wall pigmentation, and its gonads are unmature. No other difference was found. A single small specimen (UF 3056) from the French Polynesia had abundant dark brown spots along body; it is regarded as belonging to this species because this mottled pattern is unexpected, but there was no other diagnostic difference in comparison to other specimens herein regarded as conspecific.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD97312FF23F892F7C96368.taxon	discussion	Remarks. Leocrates iris Grube, 1878 has been regarded as a junior synonym of L. chinensis Kinberg, 1866 by Horst (1924) and by Pettibone (1970), and as a distinct species by Chamberlin (1919), and Treadwell (1926). Leocrates iris deserves to be reinstated because the main difference between these two species is the size of eyes, being large in L. iris (each 1 / 5 prostomial width), and small in L. chinensis (each 1 / 10 prostomial width). On the other hand, L. iris resembles L. rizzoae n. sp. from the Seychelles, because they have notacicular lobes three times longer than wide, and about 30 neurochaetae per bundle in middle chaetigers. These two species differ in the shape of anterior eyes and in the size proportions between palpophores and palpostyles. In L. iris anterior eyes are reniform and palpophores are 2 – 3 times longer than palpostyles, whereas in L. rizzoae anterior eyes are emarginate, and palpophores are twice longer than palpostyles. Hoagland (1920: 604) made no comments about the morphology of the species. Pettibone (1970) reexamined the specimen but made no comments or illustrations after it.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFD97312FF23F892F7C96368.taxon	distribution	Distribution. Philippines, Samoa, Indonesia, Vietnam, Marshall Islands, Fiji, French Polynesia, New Caledonia, Australia, in coralline substrates from the intertidal to 51 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD7312FF23FD5DF0EC656A.taxon	discussion	Remarks. Orrhage (1996: 140, footnote) indicated that this species was “ unintentionally described by Gustafson (1930). ” In order to be available, names published before 1931 must be (ICZN 1999, Art. 12) “ accompanied by a description or a definition of the taxon it denotes, or by an indication ” (12.1), and regarding indication, there must be “ in association with an illustration ” (12.2.7). Gustafson (1930) provided illustrations for the nerve chords in the anterior end, and two histological sections which might be regarded as indications, but no description was provided. Further, because the illustrations are not enough to sort this species out from other species in the genus, it must be regarded as a nomen nudum or indeterminable. The later illustrations by Orrhage (1996: 138, Fig. 1 a) is rather schematic, as the other ones in the same figure, and does not provide diagnostic features either.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD736CFF23FB50F16E6335.taxon	description	Figs 34, 35 urn: lsid: zoobank. org: act: 430 BA 70 F- 6820 - 4 E 50 - B 512 - F 8 ECEC 81 A 074	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD736CFF23FB50F16E6335.taxon	materials_examined	Type material. Western Pacific. New Caledonia. Holotype (UF 5619), Province Sud, Noumea, Baie de Citron, south part (22 ° 18 ’ 14.40 ” S, 166 ° 26 ’ 02.40 ” E), fringing reef, 2 m, under rocks, 31 Oct. 2017, G. Paulay & M. Hoban, coll. Paratype (UF 5021), Province Sud, Baie de Prony, Embouchure Carénage (22 ° 19 ’ 22.80 ” S, 166 ° 50 ’ 24.00 ” E), 0 – 8 m, mangroves and lagoon reef, 6 Dec. 2016, G. Paulay & L. Moroz, coll. (used for variation, 24 mm long, 2 mm wide).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD736CFF23FB50F16E6335.taxon	description	Description. Holotype (UF 5619) complete (Fig. 34 A), bent laterally because of previous left dissection along chaetigers 8 – 10, exposing tubular gonads; anterolateral dissection made for observing pharynx jaws; left parapodium of chaetiger 11 and right parapodia of chaetigers 8 and 15 removed for observing parapodial features. Body pale, finely maculated with brownish transverse thin spots, larger brownish basal spots along posterior parapodial surfaces, 33 mm long, 3 mm wide, 16 chaetigers. Prostomium rectangular, longer than wide (Fig. 34 B). Lateral antennae with ceratophores distinct, as long as prostomium, slightly longer than palps; palpophores 2 – 3 times longer than palpostyles. Median antennae reaching anterior prostomial margin, inserted between posterior eyes. Eyes dark brown; anterior eyes each 1 / 5 – 1 / 6 prostomial width, reniform, twice larger than posterior ones, slightly more distant to each other than posterior round eyes; in lateral view anterior and posterior eyes distinct. Nuchal organs U-shaped, posterior prostomial projections darker than nuchal organs areas. Tentacular cirri faintly annulated, without tips, longest ones surpassing chaetiger 3. Lateral cushions low, barely projected entire, longitudinal striae visible. Pharynx invaginated; anterior margin with about 22 marginal constrictions; upper jaw single, brownish, blunt; lower jaw not seen, probably damaged by dissection. Dorsal cirri broken, shorter than body width. Chaetigers 1 – 4 without notochaetae. Notochaetae present along chaetigers 5 – 16, about 40 per bundle, subdistally denticulate, denticles fine. Notacicular lobes tapered, digitate, with three (Fig. 34 C) or two distal projections (Fig. 34 D). Neurochaetal lobes conical; aciculae black. Neurochaetae about 30 per bundle, blades decreasing in size ventrally, 3 – 10 times longer than wide; blades bidentate with longer basal spines, guards smooth, approaching subdistal tooth (Fig. 34 E, F). Posterior region tapered (Fig. 34 G). Prepygidial segment with dorsal cirri three times longer than ventral ones. Pygidium tubular, anus terminal, anal cirri without tips, reaching chaetiger 14. Gonad tubes exposed by dissection, invading parapodial coelom. Oocytes about 100 µm; spermatids not seen. Variation. Paratype (UF 5021), complete, slightly dehydrated. Body twisted, blunt anteriorly, tapered posteriorly, dorsum paler along first 6 chaetigers, creamy in following ones (Fig. 35 A); right parapodium of chaetiger 9 removed for observing parapodial features. Tentacular and dorsal cirri complete. Body pale, eyes blackish. Prostomium twice wider than long, slightly wider anteriorly (Fig. 35 B). Lateral antennae with ceratophores distinct, antennae longer than prostomium, slightly longer than palps; palpophores 2 – 3 times longer than palpostyles. Median antenna tapered, barely reaching anterior prostomial margin, inserted between posterior eyes. Eyes blackish, anterior ones about 1 / 10 as wide as prostomial width, twice larger than posterior ones, almost round anteriorly, more distant to each other than posterior round eyes. Nuchal organs lobes horizontal C-shaped, concealed by posterior projection of prostomium; lateral ciliated bands narrow, barely visible dorsally. Tentacular cirri almost complete, longest ones reaching chaetiger 8. Lateral cushions low, entire, longitudinal striae not visible (probably after dehydration). Pharynx almost fully exposed (Fig. 35 C); lateral vesicles not seen; anterior margin with about 30 irregular constrictions. Jaws single, yellowish, upper jaw twice longer than lower one, tapered in lateral view, inserted slightly ahead of lower jaw. Dorsal cirri longer than body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 20 per bundle, arranged as bundles, subdistally denticulate, denticles fine. Notacicular lobes tapered, with a small, digitate projection along its upper part; neuracicular lobes blunt, conical slightly longer than wide; ventral cirri surpassing neurochaetal lobe (Fig. 35 D). Neurochaetae about 15 per bundle, blades bidentate, decreasing in size ventrally, 5 – 30 times longer than wide, guards approaching subdistal tooth (Fig. 35 E). Posterior region tapered. Preanal segment with dorsal cirri 2 – 3 times longer than ventral ones. Pygidium with anus terminal, anal cirri reaching chaetiger 13. Oocytes not seen; gonads visible by transparency in posterior region, but not dissected to avoid further damage.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD736CFF23FB50F16E6335.taxon	etymology	Etymology. The specific name is to honor Dr. Jenna Moore, from the University of Florida, Gainesville, as an appreciation for her many efforts to support my research, and for her help in many important details regarding molecular analyses. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD736CFF23FB50F16E6335.taxon	discussion	Remarks. Leocrates mooreae n. sp. belongs in the group of species provided with large eyes, anterior ones often twice larger than posterior ones. However, L. mooreae is the only species having notacicular lobes provided with terminal digitate projections, making it unique among all other species in the genus, although the number of projections vary, being onely one in smaller specimens (paratype), and up to three in larger specimens (holotype).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFDD736CFF23FB50F16E6335.taxon	distribution	Distribution. New Caledonia, in mixed bottoms with mangroves and coral reefs, in 0 – 8 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	description	Figs 36, 37 urn: lsid: zoobank. org: act: B 9292 D 1 D- 04 DA- 41 A 6 - B 4 BA-D 28 B 5 FE 34270	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	materials_examined	Type material. Western Pacific. Marshall Islands. Holotype (LACM 10883), Ralik Chain, Eniwetok Atoll, Lidilbut Island, corals, Sta. 1504 (no further data), May 1952, H. S. Ladd, coll. Paratype (LACM 10884), Ralik Chain, Eniwetok Atoll, Lidilbut Island, corals, Sta. 1503 (no further data), May 1952, H. S. Ladd, coll. [complete. Brownish, posterior region bent dorsally, most notochaetae and cirri missing; a lateral ventral dissection made to observe pharynx jaws; right parapodium of chaetiger 12 dissected (kept in container). Body 19 mm long, 2 mm wide, 16 chaetigers. Anterior eyes emarginate, twice larger than posterior round ones. Notochaetae with coarse subdistal denticulation; notacicular lobe blunt, neuracicular lobe round, about as long as wide; neurochaetae with blades bidentate, guard approaching subdistal tooth. Oocytes not seen; testis with spermatids in parapodia]. Additional material. Western Pacific. Philippines. One specimen (LACM 10142), Palawan, Honda Bay (09 ° 54 ’ 02.5200 ” N, 118 ° 43 ’ 21.3600 ” E), seagrass bed, under coral rubble, 1.2 m, 18 Apr. 1995, K. Fitzhugh, coll. [middle 17 mm long, 2 mm wide]. New Caledonia. One specimen (UF 4016), Poum Pass (20 ° 16 ’ 22.44 ” S, 163 ° 52 ’ 48.72 ” W), back reef, patch coral, 0 – 2 m, 12 Nov. 2013, N. Evans, coll. [18 mm long, 2 mm wide]. French Polynesia. One specimen (UF 1433), Society Islands, Moorea, Atiha Bay, west side of pass (17 ° 35 ’ 44.52 ” S, 149 ° 50 ’ 44.16 ” W), barrier reef, 0.5 – 1.0 m, 8 Dec. 2009, S. McPherson & G. Paulay, coll. [16 mm long, 2 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	description	Description. Holotype (LACM 10883) complete, slightly bent ventrally. Body with parallel sides (Fig. 36 A), blunt anteriorly, slightly tapered posteriorly, 20 mm long, 3 mm wide, 16 chaetigers; right parapodium of chaetiger 7 dissected (kept in container). Most tentacular, dorsal and ventral cirri on site. Body brownish, cirri paler, eyes brown. Prostomium wider than long, slightly wider anteriorly (Fig. 36 B). Lateral antennae with ceratophores distinct, antennae slightly longer than prostomium (1.2 times as long), 1.2 – 1.5 times longer than palps; palpophores 2 – 3 times longer than palpostyles (palpostyles darker along basal half). Median antenna short, surpassing prostomial centre, not reaching prostomial anterior margin, inserted between posterior eyes. Eyes dark brown, right eyes darker; anterior eyes each 1 / 15 prostomial width, twice larger than posterior ones, slightly emarginate anterolaterally, and more distant to each other than posterior, round eyes. Nuchal organs lobes horizontal C-shaped, completely concealed by anterior margin of tentacular belt; lateral ciliated bands narrow, not visible dorsally. Tentacular cirri almost complete, twisted, without tips, longer ones reaching chaetiger 7. Lateral cushions low, entire, slightly projected along middle chaetigers. Longitudinal striae visible along body. Pharynx fully exposed (Fig. 36 C), slightly expanded distally. Lateral vesicles present only on right side, globular, not tapered. Anterior margin with about 30 irregularly defined lobes. Dorsal and ventral jaws single, brownish, exposed, tapered, ventral jaw smaller than dorsal one. Dorsal cirri as long as body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, sparse (about 30 per bundle), delicate (some parapodia with notochaetae broken), some arranged as transverse fans, notochaetae subdistally denticulate, denticles coarse (visible in 10 x). Notacicular lobes blunt, neuracicular lobes projected, blunt, as long as wide (Fig. 36 D). Neurochaetae about 20 per bundle, blades decreasing in size ventrally, bidentate, 3 – 12 times longer than wide, guards approaching subdistal tooth (Fig. 36 D, insets). Posterior region tapered. Prepygidial segment with dorsal cirri 3 – 4 times longer than ventral ones. Pygidium not projected, anus dorsoterminal, anal cirri missing. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	etymology	Etymology. This species is named after the late Dr. Donald J. Reish in recognition of his many publications on several critical localities (including San Quintín and Los Angeles bays in Western Mexico), and especially after his study on the polychaetes from the Marshall Islands (Reish 1968). He made two field trips to the islands and a compilation of all recorded species, and his objective was very important because he tried to prepare a reference collection for the marine lab there (now the Mid-Pacific Marine Lab, University of Hawaii). The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	description	Variation. The paratype (Fig. 37 A, B) has an unusual development of the nuchal organs, because they are completely exposed and projected posteriorly, which differs from the holotype and other additional specimens (Fig. 37 D, E) where the nuchal organs are barely projected posteriorly. Parapodial features do not differ at all (Fig. 37 C, F), as well as other prostomial sensors like palps, antennae and eyes. This explains why they are regarded as conspecific despite the difference in the development of the nuchal organs lobes. An alternative would be to regard this species as belonging in Paradalhousia n. gen. but its species have very large eyes and notochaetae are usually abundant, and these features are not present in the paratype.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	discussion	Remarks. Leocrates reishi n. sp. belongs in the group having small eyes, with anterior eyes twice larger than posterior ones, together with L. ahlfeldae n. sp. from India, L. chinensis Kinberg, 1866 from Hong Kong, and L. giardi Gravier, 1900 from the Red Sea. However, L. reishi is the only species with notacicular lobes blunt (all others have tapered lobes), and with scarce neurochaetae, 15 – 20 per bundle; L. giardi has scarce neurochaetae but its notacicular lobes are tapered.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA2736FFF23FF6FF5FF6254.taxon	distribution	Distribution. Marshall Islands, the Philippines, New Caledonia and the French Polynesia, in coralline substrates, in 1 – 2 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	description	Figs 38, 39 urn: lsid: zoobank. org: act: 2 D 64 FB 95 - 619 F- 47 F 1 - A 19 C- 54 B 6 D 11 CA 155	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	materials_examined	Type material. Indian Ocean. Seychelles Islands. Holotype (USNM 37642), International Indian Ocean Expedition, Seychell Islands Program, Sta. F- 67, Curieuse Island (= Île Rouge; 04 ° 16 ’ 48 ” S, 55 ° 44 ’ 48 ” E), SE end, South point of Laraie Bay, 8 m, 24 Feb. 1964, J. Bohlke, coll. One paratype (USNM 37641), International Indian Ocean Expedition, Seychelles Islands Program, Sta. F- 103, Amirante Islands, D’Arros Island (05 ° 24 ’ S, 53 ° 53 ’ E), off E side, 25 – 29 m, 8 Mar. 1964, J. Bohlke, D. Dockins, R. Rosenblatt, W. Starck, J. Tyler & K. Hill, coll. Additional material. Arabian Sea. One specimen (MNHN A 71 a), Mission Gravier Djibouti, Recif de la Menagerie, in corals, 7 Jan. 1904 [11 mm long, 2 mm wide]. Two specimens (MNHN A 71 b), Mission Gravier 1904, Île Moucha (11 ° 43 ’ N, 43 ° 11 ’ E), Gulf of Tadjoura, Djibouti, dredged, 20 m, 13 May 1904, C. Gravier, coll. [10 – 18 mm long, 2 – 3 mm wide]. Gulf of Suez. One specimen (MNHN A 360), Mission R. P. Dollfus, Sta. 11, 8 Dec. 1928, R. P. Dollfus, coll. [8.5 mm long, 1.5 mm wide]. One specimen (MNHN A 372), R / V Calypso, Sta. unknown (probably Southern Saudi Arabian coast), Drach, Cherbonnier & Mercier, coll. [6 mm long, 1 mm wide]. India. One specimen (MNHN A 419), Nicobar Islands, R / VInvestigator, Indian Survey, Sta. 593, Paway Island (Pahua, 07 ° 22 ’ N, 93 ° 37 ’ E), Bay of Bengal, shore collecting [12 mm long, 2 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	description	Description. Holotype (USNM 37642), complete, slightly damaged, bent, sigmoid, middle and posterior cirri missing. Body obconic, blunt anteriorly, tapered posteriorly (Fig. 38 A), 14 mm long, 2.5 mm wide, 16 chaetigers; left parapodium of chaetiger 9 previously removed (not in container), right parapodium of chaetiger 10 dissected (kept in container). Tentacular and anterior dorsal cirri almost complete, others partially broken or missing. Body pale. Prostomium longer than wide, slightly wider anteriorly (Fig. 38 B). Lateral antennae with ceratophores distinct, antennae directed ventrally, almost as long as prostomium, slightly longer than palps; palpophores twice longer than palpostyles. Median antenna broken, probably not reaching anterior prostomial margin, inserted between eyes. Eyes brownish, anterior eyes each 1 / 6 prostomial width, twice larger than posterior ones, slightly emarginate anterolaterally, more distant to each other than posterior round eyes. Nuchal organs lobes horizontal C-shaped, completely concealed by anterior margin of tentacular belt; lateral ciliated bands narrow, not visible dorsally. Tentacular cirri without tips, longer ones reaching chaetiger 8. Lateral cushions low, entire; longitudinal striae not visible (probably by integument damage). Pharynx barely exposed (Fig. 38 C), jaws observed through previous dissection. Lateral vesicles not seen. Anterior margin with irregular constrictions. Dorsal and ventral jaws brownish, exposed, tapered, ventral jaw smaller than dorsal one. Dorsal cirri as long as body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 50 per bundle, delicate, some arranged in bundles, notochaetae subdistally denticulate, denticles fine. Notacicular lobes tapered; neuracicular lobes triangular, blunt, slightly longer than wide (Fig. 38 D). Ventral cirri as long as neurochaetal lobe. Neurochaetae about 20 per bundle, blades bidentate, most missing, decreasing in size ventrally, remaining blades 5 – 9 times longer than wide, guards approaching subdistal tooth. Posterior region tapered, almost without cirri. Preanal segment with dorsal cirri twice longer than ventral ones. Pygidium with anus terminal, anal cirri missing. Oocytes not seen. Gonad fragments are testis; no oocytes were found.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	etymology	Etymology. This species name is after Alexandra Elaine Rizzo from the Rio de Janeiro State University, a much-appreciated Brazilian colleague, and in recognition of her research activities, especially as a specialist of several polychaete groups, including Hesionidae. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	description	Variation. The paratype (USNM 37641) is complete, anterior half depressed, probably by labels, most cirri and chaetal blades missing; anteroventral dissection along mouth to chaetiger 4 previously made, left parapodia of chaetigers 4 and 7 previously removed (missing), right parapodium of chaetiger 10 dissected (kept in container). Body with parallel sides, blunt anteriorly, tapered posteriorly (Fig. 39 A), 13 mm long, 2 mm wide, 16 chaetigers. Prostomium longer than wide, barely wider anteriorly (Fig. 39 B); left lateral antenna slightly shorter than prostomium, slightly longer than left palp; palpophores twice longer than palpostyles. Eyes dark brown, anterior ones twice larger than posterior ones, slightly farther apart and anterolaterally emarginate. Notochaetae from chaetiger 5, about 50 per bundle, finely denticulate subdistally; notacicular lobes tapered, neuracicular lobes conical, blunt (Fig. 39 C); neurochaetae about 30 per bundle; blades bidentate, 4 – 16 times longer than wide, guards approaching subdistal tooth (Fig. 39 C, insets).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	discussion	Remarks. Leocrates rizzoae n. sp. resembles L. iris Grube, 1878 from the Philippines and Samoa, reinstated, because they have large eyes, anterior eyes larger than posterior ones, notacicular lobes entire, three times longer than wide, and middle chaetigers with about 30 neurochaetae per bundle. These two species differ in the shape of anterior eyes, and by the size proportions of palpophore to palpostyles; in L. rizzoae anterior eyes are emarginate, and palpophores are twice larger than palpostyles, whereas in L. iris anterior eyes are reniform, and palpophores are 2 – 3 times longer than palpostyles. The inclusion of the Red Sea specimens is with hesitation because of the condition of the specimens (one dried out, the other collapsed), and better specimens will help clarify its northwestern distributional range.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA07369FF23FC44F69C60FD.taxon	distribution	Distribution. Arabian Sea, the Seychelles, and Nicobar Islands, Indian Ocean, (Red Sea?), in intertidal to 30 m depth in mixed bottoms, including coralline substrates.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA6736AFF23FB3EF4BF66C2.taxon	description	Fig. 40 urn: lsid: zoobank. org: act: D 3 D 114 CF-D 671 - 4 DC 8 - 811 E- 93718 B 98 C 231	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA6736AFF23FB3EF4BF66C2.taxon	materials_examined	Type material. Western Pacific. Papua New Guinea. Holotype (SIO A 5973), Bismarck Sea, Kimbe Bay, Rest Orff Island, Sta. 8 (05 ° 17 ’ 35.58 ” S, 150 ° 06 ’ 15.4202 ” E), 5 – 10 m, dead Acropora with brittle stars, Breviturma (olim Ophiocoma) pica (Müller & Troschel, 1842), 23 Apr. 2008, F. Pleijel, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA6736AFF23FB3EF4BF66C2.taxon	description	Description. Holotype (SIO A 5973), complete, distorted (Fig. 40 A), bent laterally after a lateral dissection of right parapodia of chaetigers 5 – 7, and posterior region bent dorsally; left parapodia of chaetigers 7 and 9 removed for observing parapodial features. Body 10 mm long, 1.8 mm wide (by chaetiger 4); prostomium and dorsum brownish, trunk pigmentation as about 12 transverse thin lines per segment; venter with two longitudinal brownish bands; many cirri on site, anterior ones banded. Prostomium slightly longer than wide, slightly wider in anterior eyes region (Fig. 40 B). Dorsal surface with a thin, pale ‘ M’ with basal tynes divergent, directed to posterior eyes, connected to an inverted truncate pentagonal area, two larger tear-drop pale areas, narrowing anteriorly, ahead of anterior eyes, and two smaller pale areas before lateral ceratophores. Lateral antennae shorter than prostomium, slightly longer than palps, bent laterally with welldefined ceratophores; palpophores 2 – 3 times longer than palpostyles. Median antenna directed posteriorly, short (not reaching anterior prostomial margin if bent anteriorly), inserted between posterior eyes. Eyes reddish; anterior eyes each 1 / 4 prostomial width, emarginate anteriorly, twice larger than posterior ones, barely more distant to each other than posterior, round eyes; in lateral view anterior and posterior eyes fused. Nuchal organs lobes horizontal C-shaped, slightly projected posteriorly, semicircular, prostomial posterior projections with similar pigmentation as prostomium, ciliateds bands pale; lateral ciliated bands not visible. Tentacular cirri long, some feebly banded, blackish bands thinner than pale ones, longest one reaching chaetiger 10. Lateral cushions low, many distorted, entire, longitudinal striae visible. Pharynx invaginated, observed by a slight dissection to avoid further damage. Anterior margin smooth; jaws single, upper one visible, hyaline, lower one not seen. Dorsal cirri longer than body width (including parapodia). Chaetigers 1 – 4 without notochaetae. Notacicular lobes tapered; neuracicular lobes blunt, slightly longer than wide (Fig. 40 C, inset). Notochaetae present along chaetigers 5 – 16, scarce (about 30 per bundle), subdistally denticulate, denticles fine. Neurochaetae about 30 per bundle, blades decreasing in size ventrally, 3 – 15 times longer than wide; blades bidentate, guard approaching subdistal tooth (Fig. 40 D). Posterior region tapered, distorted (Fig. 40 E), retaining pigmentation pattern. Prepygidial segment with dorsal cirri 2 – 3 times longer than ventral ones, both damaged. Pygidium with anus retracted, terminal, anal cirri missing. Gonads not seen, probably a juvenile.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA6736AFF23FB3EF4BF66C2.taxon	etymology	Etymology. The specific epithet is to honor Dr. Gregory Rouse, University of California, San Diego, Scripps Institution of Oceanography, in recognition of his many contributions to polychaetes systematics, and by his longstanding support of my research activities. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA6736AFF23FB3EF4BF66C2.taxon	discussion	Remarks. Leocrates rousei n. sp. belongs in the group of species with large eyes, and with notacicular lobes tapered with entire tips. However, L. rousei is unique by having reddish to purple eyes (others have brown or black eyes), and a pigmentation pattern over prostomium and dorsum which is unique among all species in the genus.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA6736AFF23FB3EF4BF66C2.taxon	distribution	Distribution. Only known from a single locality in Papua New Guinea, in dead coral fragments, in 10 m water depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	description	Figure 41 urn: lsid: zoobank. org: act: 1 FF 93 A 83 - 5079 - 436 E- 8 B 8 A- 0636 C 2 B 6 BA 42	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	materials_examined	Type material. French Polynesia, Moorea. Holotype (SIO A 4237), W side of Opunohu Pinnacle (17 ° 29 ’ 38.76 ” S, 149 ° 51 ’ 43.20 ” W), 10 m, 11 Nov. 2010. C. Meyer, coll. Additional material. Papua New Guinea. One juvenile (SIO A 5947), Rosemarie’s Reef, Kilu-Tamare reefs, Kimbe Bay, Bismarck Sea (05 ° 25 ’ 05.4000 ” S, 150 ° 05 ’ 30.6600 ” E), 10 – 20 m, coral rubble, 18 Apr. 2008, F. Pleijel, coll. [4.8 mm long, 0.5 mm wide]. Two juveniles (SIO A 5962), Rest Orff Island, Kimbe Bay, Bismarck Sea (05 ° 17 ’ 35.5800 ” S, 150 ° 06 ’ 15.4202 ” E), 3 – 10 m, dead Acropora with brittle stars, Breviturma (olim Ophiocoma) pica (Müller & Troschel, 1842), 25 Apr. 2008, F. Pleijel, coll. [3.2 – 4.8 mm long, 0.4 – 0.7 mm wide]. Australia. One specimen (AM W 29575), Queensland, Lizard Island, west end of Palfrey Island (14 ° 40 ’ S, 145 ° 30 ’ E), 3 – 9 m, rotenone, Acropora head, 3 Nov. 1975, D. Hoese, coll. [15 mm long, 2 mm wide]. One specimen (AM W 43793), Queensland, Lizard Island, Big Vicki’s reef (14 ° 41 ’ 10 ” S, 145 ° 26 ’ 32 ” E), 9 m, coral rubble, 13 Aug. 2013, M. Capa & M. T. Aguado, coll. [13 mm long, 2 mm wide]. One specimen (AM W 43826), Queensland, Lizard Island, front of reef between Bird and South Islands (14 ° 41 ’ 52 ” S, 145 ° 27 ’ 50 ” E), 12 – 18 m, coral rubble and brown flat algae, 14 Aug. 2013, J. Zanol, coll. [10.5 mm long, 1.5 mm wide]. Two specimens (AM 43841 b), Queensland, Lizard Island, Vicki’s Reef (14 ° 41 ’ 17 ” S, 145 ° 26 ’ 35 ” E), 9 m, calcareous algae, 13 Aug. 2013, M. Capa & J. Zanol, coll. [12 – 17 mm long, 2 – 3 mm wide]. One specimen (AM W 44503), Queensland, Lizard Island, North Point, Mermaid Cove (14 ° 38 ’ 46 ” S, 145 ° 27 ’ 13 ” E), 10 m, dead coral, 20 Aug. 2013, J. Zanol et al., coll. [12.5 mm long, 1 mm wide]. Two specimens (AM W 44504), Queensland, Lizard Island, North Point, Mermaid Cove (14 ° 38 ’ 46 ” S, 145 ° 27 ’ 13 ” E), 8 – 12 m, coral rubble in sand, 20 Aug. 2013, M. Capa et al., coll. [Complete 11 mm long, 1.5 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	description	Description. Holotype (SIO A 4237), complete, straight (Fig. 41 A). Body blunt anteriorly, slightly wider medially, tapered posteriorly, 14 mm long, 1.8 mm wide; right body wall and parapodia of chaetigers 11 – 15 removed for molecular studies; right parapodium of chaetiger 8 removed for observing parapodial features. Tentacular and dorsal cirri mostly broken. Body pale. Prostomium slightly longer than wide, slightly wider in anterior eyes region (Fig. 41 B). Lateral antennae with ceratophores distinct, slightly shorter than prostomium, slightly longer than palps; palpophores slightly longer than palpostyles. Median antenna broken, not reaching anterior prostomial margin, inserted centrally, among eyes. Eyes dark brown, anterior eyes each 1 / 5 prostomial width, twice larger than posterior ones, slightly emarginate anteriorly, more distant to each other than posterior round eyes. Nuchal organs lobes C-shaped, completely exposed, pale; lateral ciliated bands narrow, not visible dorsally. Tentacular cirri without tips, longer ones reaching chaetiger 6. Lateral cushions low, entire along anterior chaetigers, bipartite in following ones; longitudinal striae barely visible. Pharynx not exposed, jaws observed through dissection. Lateral vesicles not seen. Anterior margin smooth. Dorsal and ventral jaws single, hyaline, ventral jaw smaller than dorsal one. Dorsal cirri as long as body width, including parapodia (one in right chaetiger 6 reaches prostomium). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, about 30 per bundle, delicate, arranged as transverse fans or bundles, notochaetae subdistally denticulate, denticles coarse. Notacicular lobes tapered, twice longer than wide; neuracicular lobes blunt, about as long as wide (Fig. 41 C); ventral cirri markedly longer than neurochaetal lobes. Neurochaetae about 20 per bundle, blades bidentate, decreasing in size ventrally, 5 – 11 times longer than wide, guards approaching subdistal tooth (Fig. 41 D). Posterior region tapered, bent laterally. Prepygidial segment with dorsal cirri three times longer than ventral ones. Pygidium with anus terminal, anal cirri missing. Gonads without oocytes; either testis or unmature.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	etymology	Etymology. This species is being named after Dr. Charlotte A. Seid, Collection Manager of the Benthic Invertebrate Collection, Scripps Institution of Oceanography, University of California, San Diego, as a means of appreciation of her careful work and by supporting my research activities. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	description	Variation. The Papua New Guinea specimens (SIO A 5947, A 5962) are regarded as juveniles because palpophores and palposytles are of the same length and width, whereas in adults they are usually of different width. These juvenile specimens match the adult one from the French Polynesia.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	discussion	Remarks. Leocrates seidae n. sp. belongs in the group provided with large brown to black eyes, anterior eyes larger than posterior ones, notacicular lobes without terminal projections. However, L. seidae separates from all other species in the group by having short palpophores (up to twice longer than palpostyles), notacicular lobes twice longer than wide, and neuracicular lobes as long as wide.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA47364FF23FF6FF110678B.taxon	distribution	Distribution. Papua New Guinea, French Polynesia and Australia, in coralline substrates in 3 – 20 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAB7365FF23F926F52263F9.taxon	type_taxon	Type species. Leocratides filamentosus Ehlers, 1908 by monotypy.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAB7365FF23F926F52263F9.taxon	diagnosis	Diagnosis. Hesioninae with two lateral antennae, and one median antenna on dorsal prostomial surface. Palps biarticulate, palpophores large, massive, palpostyles smaller, blunt. Eyes dark, black or brown, anterior ones larger than posterior ones, sometimes approaching each other in lateral view. Nuchal organs lobes round, projected posteriorly, parallel to divergent, posterior ciliated bands close to each other. Peristomium with dorsolateral tubercles, often lobate, and ventrolateral ones projected, with a ventral ridge usually with large round papillae. Pharynx with upper jaw double, T-shaped, lower jaw single, fang-shaped; neurochaetal blades bidentate, without guards. Parapodia all sesquiramous. Neurochaetae compound falcigers, blades bidentate, without guards.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAB7365FF23F926F52263F9.taxon	discussion	Remarks. Leocratides Ehlers, 1908 resembles three other Hesioninae genera by having sesquiramous parapodia: Dalhousiella McIntosh, 1908, Elisesione Salazar-Vallejo, 2016, and Hesione Savigny in Lamarck, 1818. Leocratides differs from these genera because it has biarticulate palps (missing in Hesione, simple in Elisesione, biarticulate in Dalhousiella), median antenna (missing in Elisesione and Hesione, present in Dalhousiella), and especially because its pharynx is provided with jaws (missing in the three other genera), being double upper jaws, and single, fang-shaped lower ones. Further, the nuchal organs lobes, as observed in a syntype of L. filamentosus Ehlers, 1908, the type species, are depressed, divergent, markedly separated middorsally (Fig. 44 A, D). There are two other interesting differences in comparison to all other Hesioninae genera, and both might be related to the peristomium. First, there are two complex, dorsolateral tubercles resembling hands by having 3 – 4 thick digitate lobes as in L. filamentosus Ehlers, 1908, or 1 – 2 lobes as in L. ehlersi (Horst, 1921). Second, midventrally there is a continuous ridge with 8 – 10 large, thick, round papillae.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAA7367FF23F9C2F5EA65E1.taxon	description	Figure 42	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAA7367FF23F9C2F5EA65E1.taxon	materials_examined	Type material. Indonesia. Fifteen syntypes in two museums: 13 syntypes (ZMA V. Pol. 533), and two others (USNM 37646), Lesser Sunda Islands, Sumbawa, North coast, Saleh Bay, R / V Siboga Exped., Sta. 312 (08 ° 19 ’ S, 117 ° 41 ’ E), 274 m, trawl, muddy bottom / sand, 14 Feb. 1900. Additional material. Philippines. Two specimens (ECOSUR 3078, Musorstom 3 - 143), Sta. 143 (11 ° 28 ’ N, 124 ° 12 ’ E), 205 – 214 m (190 – 198 m in label), 1 Jun. 1985 [15.5 – 16.0 mm long, 2.5 mm wide]. New Caledonia. One specimen (MNHN Musorstom SMIB 3 - 17), Campagne SMIB 3, R / V Vauban, Sta. 17 (23 ° 40.60 ’ S, 167 ° 59.40 ’ E), 238 m, 23 May 1987 [This looks like a different species. It has entire, smooth dorsolateral tubercles; median antenna more than twice longer than prostomium, lateral antennae about three times longer than prostomium. Nuchal organs lobes markedly projected beyond lateral prostomial margin. Parapodia are in poor shape and better specimens are needed for its formal description, provided they confirm a different species is involved].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAA7367FF23F9C2F5EA65E1.taxon	description	Description. Largest syntype (ZMA V. Pol. 533) complete, slightly damaged, bent laterally. Body blunt anteriorly, wider anteriorly, tapered posteriorly (Fig. 42 A); 28 mm long, 3.5 mm wide, 16 chaetigers; right parapodia of chaetigers 8 and 15 removed for observing parapodial features. Tentacular and dorsal cirri without tips. Dorsal surface pale, venter with a wide longitudinal brownish band along body, triangular in tentacular belt. Becoming roughtly X-shaped along chaetigers 5 – 6, less defined in middle and posterior chaetigers (Fig. 42 B). Prostomium slightly wider than long, slightly wider anteriorly, lateral margins curved, convergent (Fig. 42 C). Lateral antennae with ceratophores distinct, as long as prostomium, slightly longer than palps; palpophores twice longer than palpostyles; median antenna thinner and slightly shorter than lateral antennae, markedly surpassing anterior prostomial margin, inserted centrally among eyes. Eyes barely pigmented, round, anterior ones twice larger and more separated than posterior ones; in lateral view, anterior and posterior eyes separated from each other. Nuchal organs lobes completely exposed, divergent oval ridges, markedly separated middorsaly; lateral ciliated bands not visible dorsally. Tentacular cirri without tips, longest ones reach chaetiger 7. Lateral cushions projected, swollen, bipartite along body, longitudinal striae not visible. Pharynx partially exposed (Fig. 42 D, E). Upper jaw double, missing in largest syntype, present in other ones; crescent shaped, lower jaw single, tapered. Anterior margin smooth, without papillae. Lateral vesicles round, low, not fully projected. Peristomial dorsolateral tubercles projected, narrower than prostomium, each with 2 low, round, wider than long lobes. Ventral ridge with 7 – 8 regular constrictions, barely projected, markedly wider than long. Dorsal cirri mostly broken or without tips, longest one longer than body width including parapodia. Neuracicular lobes tapered, three times longer than wide, tips blunt, directed distally or slightly upwards (Fig. 42 F). Neurochaetae about 18 per bundle, blades most missing, apparently decreasing in size ventrally, bidentate, without guards, blades 3 – 4 times longer than wide (Fig. 42 F, insets). Posterior region tapered. Prepygidial segment short, with dorsal cirri 3 – 4 times longer than ventral ones. Pygidium with anus terminal, anal cirri broken, reaching chaetiger 16. Oocytes visible in posterior chaetigers, each about 120 µm. Gonad fragments in middle chaetigers contain only spermatids. Variation. Syntypes 18 – 28 mm long, 3.0 – 3.5 mm wide. Dorsolateral peristomial tubercles always narrower than prostomium, with up to three lobes each, but sometimes they barely defined by shallow furrows, whereas the ventral ridge can have up to 9 round lobes, each as long as wide. Eyes are round, most barely pigmented, or colorless. Median antenna, or its scar, always inserted centrally between eyes. The lateral pharynx vesicles rarely fully projected, when it occurs, they are globose, blunt. Midventral pigmented longitudinal bands is constant (a badly damaged specimen has a double, longitudinal thin band, leaving a paler middle area). Most neurochaetal blades missing, probably they are delicate, or easily torn off when specimens are removed from the hexactinellid sponge they live in.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAA7367FF23F9C2F5EA65E1.taxon	discussion	Remarks. Leocratides ehlersi (Horst, 1921) reinstated, has been regarded as a junior synonym of L. filamentosus Ehlers, 1908 by Augener (1926: 452) and Pettibone (1970: 230), but others have regarded it as a distinct species (Fauvel 1932: 62, 1953 b: 107; Pleijel 1998: 112, 160). These two species differ, as indicated in the key above, and consequently it is formaly reinstated. In fact, L. ehlersi resembles L. kimuraorum Jimi, Tanaka & Kajihara, 2017 from Japan, by having peristomial dorsolateral ridges with a few to none lobes, and ventral pigmentation. However, these species differ, in the type of neuracicular lobes and pigmentation patterns. In L. ehlersi neuracicular lobes are markedly longer than wide, triangular, non-mucronate, and there is a ventral continuous brownish band, whereas in L. kimuraorum the neuracicular lobes are as long as wide, mucronate, and ventral pigmentation is spotty, never continuous. There are other records for the Andaman Sea by Fauvel (1932: 62, 1953 b: 107 – 108, Fig. 51 a – c), and for the Red Sea by Fishelson & Rullier (1969: 59 – 60). However, because their specimens were not available, they cannot be confirmed.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAA7367FF23F9C2F5EA65E1.taxon	distribution	Distribution. Originally described as living in an hexactinellid sponge, Aphrocallistes sp., in 274 m deep, off Sumbawa, Indonesia.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA87362FF23FAC8F7296335.taxon	description	Figs 43, 44	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA87362FF23FAC8F7296335.taxon	materials_examined	Type material. Malaysia, Western Sumatra. Six syntypes in two lots, five in one lot (ZMB 4431), and another syntype (ZMH PE- 318 b), Nias South Canal, R / V Valdivia, Sta. 198 (00 ° 16 ’ N, 98 ° 07 ’ E), 677 m, in hexactinellid sponge, Aphrocallistes bocagei Wright, 1870 (actually A. beatrix Gray, 1858) [ZMB syntypes slightly dehydrated in two containers. Larger container with four syntypes, including one with an anterior midventral dissection. Body pale, 20 – 36 mm long, 5 – 8 mm wide, 16 chaetigers. Lateral antennae as long as prostomium, as long as or slightly longer than palps; palpophores 2 – 3 times longer than palpostyles; median antennae missing in all these syntypes (still on site when Pettibone examined the specimens), insertion scar visible between posterior eyes. See variation for other features). Smaller container with a large specimen, 44 mm long, 4.5 mm wide, 16 chaetigers. Segments short along chaetigers 1 – 5, less contracted thereafter. Lateral antennae without tips, as long as palps; palpophores 3 – 4 times longer than palpostyles; median antenna thin, long, surpassing anterior prostomial margin, almost reaching tip of palpophore. Eyes brownish, anterior ones reniform, twice larger than posterior round ones. Nuchal organs depressed, lateral ciliated areas visible dorsally. Dorsolateral tubercles with 4 digitate lobes; ventral ridge with 10 large, round tubercles, midventral ones smaller]. Additional material. Philippines. Six specimens (3: MNHN Musorstom 3 - 135 CP, and 3: ECOSUR 3079), Musorstom Expedition, Sta. 135 (11 ° 58 ’ N, 122 ° 02 ’ E), 486 – 551 m, in hexactinellid sponge fragment (Aphrocallistes sp.), 5 Jun. 1985 [5 complete, 17 – 28 mm long, 3 – 4 mm wide]. One specimen (MNHN Halipro 2 - 60), R / V Tangaroa, Sta. 25 (25 ° 16.64 ’ S, 170 ° 23.69 ’ E to 25 ° 17.45 ’ S, 170 ° 23.93 ’ E), 1100 – 1348 m, 11 Nov. 1996 [30 mm long, 3 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA87362FF23FAC8F7296335.taxon	description	Description. Best syntype (ZMH PE- 318 b) complete, distorted by preserving it with other organisms, compressed medially, bent laterally posteriorly. Body obconic, blunt anteriorly, wider anteriorly, tapered posteriorly (Fig. 43 A); 28 mm long, 5 mm wide, 16 chaetigers; left parapodium of chaetiger 9 previously removed, right parapodium of chaetiger 8 removed for parapodial features. Tentacular and dorsal cirri mostly broken. Body dorsal and ventral surfaces pale; eyes dark brown. Prostomium as long as wide, slightly wider anteriorly, lateral margins straight, convergent (Fig. 43 B). Lateral antennae with ceratophores distinct, as long as prostomium, as long as palps; palpophores 2 – 3 times longer than palpostyles; median antenna slightly longer and thinner than lateral ones, markedly surpassing prostomial anterior margin, inserted between posterior eyes. Eyes dark brown, anterior ones semilunar to reniform, twice larger than posterior round ones, slightly more distant to each other than posterior eyes; in lateral view, anterior and posterior eyes close to each other. Nuchal organs lobes completely exposed, pale, depressed, oval, markedly separated middorsally; lateral ciliated bands not visible dorsally. Tentacular cirri without tips, longest ones reach chaetiger 6. Lateral cushions low, roughly bipartite along body; longitudinal striae not visible. Peristomial dorsolateral tubercles wider than prostomium, each with four thick, digitate lobes, each twice longer than wide (Fig. 43 C). Ventral ridge with nine thick digitate, as long as wide papillae, midventral ones slightly smaller. Pharynx exposed (Fig. 43 C, D). Upper jaw double, low, crescent shaped, lower jaw single, tapered. Anterior margin with 16 round, low papillae. Lateral vesicles round, projected laterally. Dorsal cirri mostly broken, a few left as long as body width (Fig. 43 E). Neuracicular lobes tapered, as long as wide or slightly longer than wide, tips sometimes contracted, directed upwards (Fig. 43 E, inset). Neurochaetae about 30 per bundle, blades decreasing in size ventrally, bidentate, without guards; blades 3 – 4 times longer than wide (Fig. 43 E, insets). Posterior region tapered. Prepygidial segment with dorsal cirri 5 – 6 times longer than ventral ones. Pygidium with anus terminal, a single midventral anal cirrus left, reaching chaetiger 14. Oocytes not seen. Variation. The large anterior, 3 – 4 lobed hand-shaped tubercles, together with the ventral ridge of papillae are present even in the smallest syntype (20 mm long), as well as in the largest one (Fig. 44 A, D). Eyes are dark brown, round, emarginate or reniform, sometimes even the posterior eyes are reniform; only one syntype has colorless eyes (and distorted head by compression). Median antenna scar always between posterior eyes. Nuchal organs lobes low diverging, round in all specimens; ciliated bands sometimes visible along lateral prostomial margins, or in the posterior nuchal organ margin. Pharynx features also similar in small to large syntypes (Fig. 44 B, E): upper jaw double, lower one single, tapered. Parapodial features without major differences from small to large syntypes (Fig. 44 C, F); neuracicular lobes tapered, slightly longer than wide, or up to 50 % longer than wide, often with well defined tip, sometimes directed upward or downward (Fig. 44 C, F, insets). Neurochaetae about 20 – 36 per bundle, number size dependent; many blades missing, bidentate, without guards.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA87362FF23FAC8F7296335.taxon	discussion	Remarks. Leocratides filamentosus Ehlers, 1908 resembles L. jimii n. sp. from Japan because they have peristomial dorsolateral tubercles with thick digitate lobes. They differ, however, in their ventral pigmentation pattern and type of neuracicular lobes. In L. filamentosus venter is colorless, without pigmentation, and neuracicular lobes are longer than wide, continued into small mucros, whereas in L. jimii the venter shows a wide brownish band, especially along middle segments, and neuracicular lobes are as long as wide, and globose. These hesionid worms were found in hexactinellid sponges which were recorded as Aphrocallistes bocagei Wright, 1870. This sponge was originally described with deep water specimens from Ireland to Cape Verde Islands (Wright 1870: 5) and it would be difficult to be found also in Malaysia. One of the regional species, A. beatrix Gray, 1858, might correspond to the sponge species originally found having Leocratides specimens. These two species were regarded as synonyms in a recent publication (Reiswig & Kelly 2011: 130), but they were regarded as distinct by Kent (1870: 248) after a comparison of specimens and spicules from both species.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFA87362FF23FAC8F7296335.taxon	distribution	Distribution. Malaysia to the Philippines, in 468 – 1348 m, associated with hexactinellid sponges. The record by Rullier (1972: 58) was based upon poorly preserved specimens collected in the intertidal; his record is herein regarded as questionable because this is a deep-water species.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAD7363FF23FD64F7346634.taxon	description	Fig. 45 urn: lsid: zoobank. org: act: 1 D 277 B 08 - 28 A 2 - 4 EBA-B 52 E- 84 ED 63277 E 25	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAD7363FF23FD64F7346634.taxon	materials_examined	Type material. Madagascar. Holotype (MNHN A 861), 50 km NW off Nosy Mitsio, R / V Vauban, Sta. 9: 44 (12 ° 39 ’ 05 ” S, 48 ° 15 ’ 06 ” E), 450 m, 1 Aug. 1973, A. Crosnier, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAD7363FF23FD64F7346634.taxon	description	Description. Holotype (MNHN A 861), complete, bent ventrally. Body obconic, blunt, wider anteriorly, tapered posteriorly, 34 mm long, 6 mm wide, 16 chaetigers; right parapodium of chaetiger 8 removed for observing parapodial features (kept in container), right parapodium of chaetiger 5 in regeneration. Tentacular and dorsal cirri without tips, or broken. Integument pale, slightly darker along anterior region (Fig. 45 A); venter with a wide, dark brown longitudinal band, from chaetiger 5 to end of body (Fig. 45 B), darker along chaetigers 6 – 9. Prostomium as long as wide, slightly wider anteriorly, lateral margins slightly constricted before anterior eyes (Fig. 45 C). Lateral antennae with ceratophores distinct, as long as prostomium, as long as palps; palpophores twice longer than palpostyles; median antenna about twice longer and wider than lateral ones, inserted centrally among eyes. Eyes dark brown, round, anterior eyes slightly larger and more distant to each other than posterior ones; in lateral view anterior and posterior eyes close to each other (Fig. 45 D). Nuchal organs partially covered by tentacular belt. Lateral lobes oval, markedly separated middorsally, divergent; lateral ciliated bands wide, visible dorsally. Lateral cushions low, bipartite along body; longitudinal striae better defined along anterior third of body. Peristomial dorsolateral tubercles as wide as prostomium, each with four thick, digitate lobes, each about twice longer than wide. Ventral ridge with 8 thick, digitate, as long as wide papillae, midventral ones slightly smaller. Pharynx exposed (Fig. 45 D, E). Upper jaw double, low, crescent shaped, lower jaw single, tapered. Anterior margin with about 20 low papillae. Lateral vesicles not seen. Dorsal cirri broken, as long as body width (Fig. 45 F). Neuracicular lobes globose, as long as wide, or slightly longer than wide (Fig. 45 F, inset). Neurochaetae about 40 per bundle, blades slightly decreasing in size ventrally (Fig. 45 G), 3 – 4 times longer than wide, bidentate, without guards (Fig. 45 H). Posterior region tapered. Prepygidial segment with dorsal cirri without tips, about 5 times longer than ventral ones. Pygidium distorted, anus terminal, one cirri missing, the other broken. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAD7363FF23FD64F7346634.taxon	etymology	Etymology. This species is being named after Naoto Jimi, from the National Institute of Polar Research, Tokyo, a young productive polychaete taxonomist and kind colleague, in recognition of his publications on polychaete taxonomy, and especially because he participated in the description of the most recent Leocratides species. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAD7363FF23FD64F7346634.taxon	discussion	Remarks. As indicated in the key above, Leocratides jimii n. sp. resembles L. filamentosus Ehlers, 1908 by having peristomial dorsolateral tubercles with thick digitate lobes. They differ by their ventral pigmentation pattern and development of neuracicular lobes. In L. jimii there is a wide brownish band, better defined medially, and neuracicular lobes are as long as wide, globose, whereas in L. filamentosus there is no ventral pigmentation, and neuracicular lobes are longer than wide, tapered into small mucros.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAD7363FF23FD64F7346634.taxon	distribution	Distribution. Off Nosy Mitsio, Madagascar, 450 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAC737CFF23F87AF79F63C5.taxon	diagnosis	Diagnosis. Leocratides with peristomial dorsolateral tubercles with two round marginal lobes. Neuracicular lobes as long as wide, mucronate. Venter with black spots along middle and posterior segments, 1 – 2 spots one behind the other, and two larger spots transversely arranged in each neuropodial area.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAC737CFF23F87AF79F63C5.taxon	discussion	Remarks. Leocratides kimuraorum Jimi, Tanaka & Kajihara, 2017 was described recently partially on the basis of specimens previously recorded as L. filamentosus by Imajima (2003: 136 – 138, Fig. 80); such that the species needs no redescription. As indicated in the key above, L. kimuraorum resembles L. ehlersi (Horst, 1921) because both have peristomial dorsolateral ridges with a few to none lobes, and by having some ventral pigmentation. These species differ, however, by the type of neuracicular lobes and pigmentation patterns. In L. kimuraorum neuracicular lobes are as long as wide and mucronate, and ventral pigmentation is spotty, not continuous, whereas in L. ehlersi neuracicular lobes are twice longer than wide, triangular but non-mucronate, and pigmentation is a continuous brownish band.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFAC737CFF23F87AF79F63C5.taxon	distribution	Distribution. Off Shima Peninsula, Japan, in sponges, 103 – 104 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB3737CFF23FCD4F68E670F.taxon	type_taxon	Type species. Leocrates anomalus Chamberlin, 1919.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB3737CFF23FCD4F68E670F.taxon	diagnosis	Diagnosis. Hesioninae with two lateral antennae, and one median antenna on dorsal prostomial surface. Palps biarticulate, palpophores large, massive, palpostyles smaller, blunt. Eyes dark, black, brown or reddish, anterior ones larger than posterior ones, sometimes approaching each other in lateral view. Nuchal organs lobes U-shaped. Pharynx with single upper and lower jaws, and a marginal ring of denticles. Parapodia sesquiramous along chaetigers 1 – 3, biramous thereafter. Notochaetae from chaetiger 4, subdistally denticulate, often abundant, usually very long, reaching neurochaetal tips. Neurochaetae compound falcigers, blades bidentate, guards approaching subdistal tooth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB3737CFF23FCD4F68E670F.taxon	etymology	Etymology. The genus-group name is derived from Dalhousia McIntosh, 1885, with the Greek prefix Para - (= near) to emphasize their close resemblance. Gender. Feminine, as the stem genus-group name.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB3737CFF23FCD4F68E670F.taxon	discussion	Remarks. Paradalhousia n. gen. and Dalhousia McIntosh, 1901 are very similar by having biarticulate palps, pharynx with jaws, U-shaped nuchal organs lobes, and sesquiramous parapodia anteriorly and biramous posteriorly. Their main differences rely in the pharynx armature and in the pigmentation of neurochaetae. In Paradalhousia there are single, fang-shaped upper and lower jaws, and a marginal circle of denticles, and neurochaetae are pale, whereas in Dalhousia the upper jaw is double, T-shaped, and the ventral jaw is a transverse plate, there are no marginal denticles, and neurochaetae are often brownish.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB27379FF23FF6CF5D760D9.taxon	description	Figs 46 – 48	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB27379FF23FF6CF5D760D9.taxon	materials_examined	Type material. Western Pacific. Marshall Islands. Holotype (USNM 19392), USFCS Albatross Pacific Expedition, 1899 – 1900, Unnumb. Sta. (Marshall Islands 9 ° N, 168 ° E), came up on anchor, 21.6 m, Dec. 1899. Additional material. Western Pacific. Micronesia. Two specimens (USNM 29981), Pacific Science Board, Sta. 172, lagoon edge of lagoon reef, Tiatua, Kapingamarangi Atoll, Sta. 137 (01 ° 04 ’ 00 ” N, 154 ° 47 ’ 00 ” E), 13 Jul. 1954, C. Hand, coll. [15 mm long, 3 mm wide]. Two specimens (USNM 37643), Kapingamarangi Atoll, G. F. V. Sta. 137 (01 ° 21 ’ 00 ” N, 154 ° 45 ’ 14 ” E), 12 Aug. 1954, G. F. V., coll. [partially dehydrated, larger one complete, the other without posterior end, some parapodia previously removed; both already dissected midventrally for pharynx details; best specimen 20 mm long, 2.5 mm wide]. Japan, Ryukyu Islands. Two specimens (MCZ ANNa 84731), Horshoe Cliffs, 1 km WNW from Onna Village (26 ° 30 ’ N, 127 ° 50.9 ’ E), 63 m, mixed sand and coral rubble, 1 Jun. 1995, R. F. Bolland, coll. [18.5 – 21.0 mm long, 2.8 – 3.5 mm wide]. Cook Islands. One specimen (UF 393), Ma’uke Island, off Kimiangatau (20 ° 08 ’ 48 ” S, 157 ° 21 ’ 00 ” W), under rocks, 50 – 52 m, 4 Dec. 1984, G. Paulay, coll. [13 mm long, 2 mm wide]. New Caledonia. 10 specimens (SIO A 5582), Îles Loyauté, Baie du Santal, Lifou, Récif Shelter (20 ° 53 ’ 45.00 ” S, 167 ° 03 ’ 30.0024 ” E), 0 – 17 m, 18 Nov. 2000, F. Pleijel, coll. [10 – 14 mm long, 2 mm wide]. French Polynesia, Society Islands, Moorea. One specimen (SIO A 4233), W side of Cook’s Pass, near MPA Marker (17 ° 28 ’ 53.40 ” S, 149 ° 49 ’ 30.00 ” W), no depth given, 9 Nov. 2010, G. Rouse, coll. [13 mm long, 1 mm wide]. One specimen (SIO A 11840), W side of Opunohu Pinnacle (17 ° 29 ’ 38.76 ” S, 149 ° 51 ’ 43.20 ” W), 10 m, 11 Nov. 2010. C. Meyer, coll. [13.5 mm long, 1 mm wide]. One specimen (UF 855), off E of Opunohu Pass, Shark Feeding buoy (17 ° 28 ’ 54.12 ” S, 149 ° 51 ’ 20.88 ” W), outer reef slope, 17 – 18 m, rubble and sand, 16 Oct. 2008, S. McKeon, J. Moore & G. Paulay, coll. [9.5 mm long, 1.5 mm wide]. One specimen (UF 868), between Temae and Afareaitu, outer reef (17 ° 30 ’ 52.20 ” S, 149 ° 45 ’ 41.76 ” W), 30 m, slope base, rubble field, rubble brushing, 23 Oct. 2008, C. Meyer, S. McKeon & J. Moore, coll. [12 mm long, 1.5 mm wide]. One specimen (UF 2150), off Hilton, forereef (17 ° 28 ’ 31.44 ” S, 149 ° 50 ’ 31.20 ” W), 40 m, rubble, 9 Nov. 2010, K. White, coll. [4 mm long, 1 mm wide]. One specimen (UF 3030), off Motu Ahi, north of Afareitu Pass (17 ° 33 ’ 09.36 ” S, 149 ° 46 ’ 24.60 ” W), 57 m, deep reef slope, rubble, 22 Jan. 2012, R. Whitton, coll. [13 mm long, 2 mm wide]. One specimen (UF 3047), off NW Motus (17 ° 29 ’ 10.32 ” S, 149 ° 55 ’ 27.48 ” W), 66 – 72 m, deep reef slope, 28 Jan. 2012, J. Earle, D. Pence, R. Pyle, coll. [10 mm long, 1.5 mm wide, 11 / 12 chaetigers].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB27379FF23FF6CF5D760D9.taxon	description	Redescription. Holotype (USNM 19392) dried-out, distorted; anterior region swollen, middle and posterior regions collapsed, directed upward (Fig. 46 A). Body obconic, anteriorly blunt, posteriorly tapered, 16 mm long, 4.5 mm wide; left parapodia of chaetigers 10 and 12 previously dissected (in container); pharynx previously dissected midventrally. Tentacular cirri, and dorsal and ventral cirri missing. Body pale, eyes dark brown, posterior prostomial projections brownish. Prostomium longer than wide, slightly wider anteriorly (Fig. 46 B, C). Palps and lateral antennae collapsed, not manipulated to avoid further damage. Median antenna broken, basal region visible, inserted between posterior eyes. Anterior eyes twice larger than posterior ones, slightly more distant to each other than posterior ones; in lateral view, eyes almost fused. Nuchal organs lobes divergent, tear-drop shaped; posterior prostomial areas paler than eyes. Lateral cushions low, barely projected, entire or bifid along body, longitudinal striae visible. Pharynx partially exposed (Fig. 46 D). Anterior margin with 20 oval brownish denticles, slightly larger ventrally; dorsal jaw single, brownish, exposed, inserted below pharynx margin; ventral jaw not seen, probably destroyed by previous dissection. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, very abundant, delicate, probably eroded, surface smooth. Dorsal cirri length in relation to body width unknown. Notacicular and neuracicular lobes of similar length, tapered (Fig. 46 E, insets), acicula black, tapered; ventral cirri dehydrated, tip broken, relative length unknown. Neurochaetal tips missing; blades unknown. Posterior region tapered, dehydrated, without lateral or anal cirri. Pygidium with anus terminal. Immature gonad observed with oocytes, each about 25 µm in diameter (Fig. 46 E, inset). Variation. Better preserved, non-type specimens (USNM 29981: 15 mm long, 3 mm wide, MCZ 84731: 18.5 – 21.0 mm long, 2.8 – 3.5 mm wide) slightly dehydrated, bent laterally (Fig. 47 A), or less dehydrated (Fig. 48 E). Body obconic, anterior region blunt, posterior region tapered; pharynx previously dissected midventrally. Many tentacular cirri and dorsal cirri missing. Body pale, eyes dark brown. Prostomium longer than wide, slightly wider anteriorly (Figs 47 B, 48 A, F). Lateral antennae 1 / 3 longer than palps. Palpophores slightly longer than palpostyles. Median antenna short, delicate, apparently complete, not reaching anterior prostomial margin. Eyes brownish, anterior ones slightly larger than posterior ones (Figs 47 A, B, 48 A, F); in lateral view, eyes almost fused. Nuchal organs lobes tear-drop shaped with posterior prostomial areas pale, take Methyl green stain. Pharynx previously dissected (Figs 47 C, 48 C, D). Anterior margin with 28 oval brownish denticles, slightly larger ventrolaterally; dorsal jaw single, brownish, exposed, inserted below pharynx margin; ventral jaw not seen, probably destroyed by previous dissection. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 - 16, very abundant, delicate, subdistal denticles fine. Dorsal cirri as long as body width (without parapodia). Notacicular and neuracicular lobes tapered (Figs 47 D, insets, 48 B, G), aciculae black, tapered; ventral cirri surpassing neurochatal lobes. Neurochaetae with blades decreasing in size ventrally (Fig. 47 E), bidentate, 5 – 22 times longer than wide, guards approaching subdistal tooth (Fig. 47 F). Posterior region tapered, prepygidial segment with dorsal and ventral cirrophores, cirrostyles missing. Pygidium with anus terminal, ventral cirri missing. Oocytes not seen.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB27379FF23FF6CF5D760D9.taxon	discussion	Remarks. Leocrates anomalus Chamberlin, 1919 must be reinstated. Pettibone (1970: 219) regarded it as a junior synonym of L. giardi Gravier, 1900, but these two species belong to different genera, as herein revised, and hence a new combination is proposed below. Chamberlin (1919) made no illustrations for the original description, and Pettibone (1970) did not make any for it despite the fact she studied the holotype. The marginal pharynx denticles present in P. anomala and in two other species also included in this genus (see below), are not present in any Leocrates species. Further, this species is newly combined into Paradalhousia anomala (Chamberlin, 1919) because it has Ushaped nuchal or tear-drop shaped lobes, single upper and lower jaws, and pale neurochaetae. This combination of characters is herein regarded as diagnostic for Paradalhousia. The marginal teeth present in the pharynx of Paradalhousia species resemble a syllid trepan, and the presence of a middorsal jaw at a certain distance of the pharynx margin, makes this resemblance even stronger. It must be stressed that this type of pharynx armature has not been described before in any genus of Hesionidae. Paradalhousia anomala differs from the two other species also newly recombined and reinstated: P. oculata (Treadwell, 1906), and P. papillosa (Monro, 1926), especially in the shape of nuchal organs lobes. In P. anomala nuchal organs lobes are divergent, whereas in the two other species the nuchal organs lobes are parallel. On the other hand, P. anomala was described with a specimen from the Marshall Islands. Chamberlin (1919: 191) indicated that acicular lobes were tapered (‘ acutely pointed’), and although there was no explanation for the specific name, it was probably selected after the observation of ‘ a half circle of conspicuous, chitinized, round papillae’ what separates this species ‘ from other species of the region. ’ He did not call these structures as either denticles or teeth, but he regarded them as sclerotized papillae one page ahead, and by referring to nereidid pharynx he wrote (Chamberlin 1919: 192): ‘ soft or hard papillae, or teeth, the paragnatha (sic), which are of much significance in classification. ’	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB27379FF23FF6CF5D760D9.taxon	distribution	Distribution. Japan (Ryukyu Islands) to Micronesia, French Polynesia and New Caledonia, in coral environments, in 10 – 72 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB6737BFF23FDE2F72B60A5.taxon	description	Figs 49, 50	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB6737BFF23FDE2F72B60A5.taxon	materials_examined	Type material. Hawaii. Holotype (USNM 5200), Maui Island, South of Puhielele Point (21 ° 36 ’ 35 ” N, 156 ° 05 ’ 10 ” W), Aleunihana Channel, between Hawaii and Maui Islands, USFC Steamer Albatross, Sta. 4066, 88 – 317 m, rocky bottom, 18 Jul. 1902. Additional material. Hawaii. One specimen (AMNH 476), no further data [labelled as syntype; 7.1 mm long, 2.3 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB6737BFF23FDE2F72B60A5.taxon	description	Description. Holotype (USNM 5200) complete, bent ventrally, breaking into two (Fig. 49 A), pharynx fully exposed; left parapodia of chaetigers 6 – 9, and right parapodia of chaetigers 6 – 7 previously removed (all in container). Body colorless, 11 mm long, 2 mm wide, 16 chaetigers; eyes brownish, middorsal peristomial tubercle pale; most cirri broken. Prostomium slightly longer than wide, slightly wider anteriorly (Fig. 49 B, C). Lateral antennae with ceratophores distinct, longer than prostomium, slightly longer than palps; palpophores slightly longer than palpostyles. Median antenna broken, inserted between anterior eyes. Eyes brownish, round; anterior eyes emarginate anterolaterally, slightly larger and slightly more distant to each other than posterior round ones; in lateral view anterior and posterior eyes fused (Fig. 49 B, C). Nuchal organs lobes U-shaped, parallel, prostomial posterior projections pale; middle furrow deep. Lateral ciliated bands visible dorsally. Tentacular cirri mostly broken or without tips, longest one reaching chaetiger 6. Lateral cushions low, entire, longitudinal striae visible. Pharynx fully everted; anterior margin with 24 denticles, middorsal and midventral areas bare (Fig. 49 D). Upper and lower jaws single, visible, exposed, brownish in frontal view. Dorsal cirri without tips, about as long as body width (excluding parapodia). Chaetigers 1 – 3 without notochaetae, notochaetae present along chaetigers 4 – 16, about 50 per bundle, as long as neurochaetae, subdistally denticulate, denticles fine, small. Notacicular and neuracicular lobes tapered, blunt (Fig. 49 E, insets); notacicular lobes twice longer than wide, neuracicular ones as long as wide. Neurochaetae about 50 per bundle, blades decreasing in size ventrally, 6 – 7 times longer than wide; blades bidentate, guard approaching subdistal tooth (Fig. 49 E insets). Posterior region tapered. Prepygidial segment without dorsal cirri, ventral cirri medially broken. Pygidium tubular, anal cirri missing. Oocytes about 100 µm.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB6737BFF23FDE2F72B60A5.taxon	discussion	Remarks. Castalia oculata Treadwell, 1906 must be reinstated and newly combined. Pettibone (1970: 219) regarded it as a junior synonym of Leocrates giardi Gravier, 1900, but they belong to different genera. Further, C. oculata deserves a new combination into Paradalhousia because it has U-shaped nuchal organs, single upper and lower jaws, and pale neurochaetae. On the other hand, P. oculata (Treadwell, 1906) n. comb., resembles P. papillosa (Monro, 1926) reinst., n. comb. but they differ in the number of neurochaetae and in the orientation of their teeth. In P. oculata there are about 50 neurochaetae per bundle, and their distal teeth are slightly falcate, whereas in P. papillosa neurochaetae number about 20 (– 30) per bundle, and their distal teeth are straight. The specimen deposited in New York (AMNH 476) is labeled as syntype. However, it was not included in the original description, which is the requisite in order to be included as a syntype (ICZN 1999, Art. 72.4.1), and consequently it is herein regarded as an additional specimen. However, it completely matches the holotype in body shape (Fig. 50 A), anterior end appendages (Fig. 50 B), and pharynx features (Fig. 50 C), rendering it enigmatic why it was not included in the original description.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB6737BFF23FDE2F72B60A5.taxon	distribution	Distribution. Hawaii, rocky bottoms in 88 – 317 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB47375FF23FDF4F4BF61DC.taxon	description	Figure 51	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB47375FF23FDF4F4BF61DC.taxon	materials_examined	Type material. China Sea. Two syntypes (BMNH 1926.4.30.130 – 131), Macclesfield Bank (16 ° 00 ’ N, 114 ° 30 ’ E), no further data [very damaged after chaetal fracture and removal of many parapodia; description based upon both of them]. Additional material. Sibuyan Sea. One specimen (MNHN Musorstom 3 - 137), Philippines, Sta. 137 (12 ° 03 ’ N, 122 ° 06 ’ E), 56 m, 6 Jun. 1985 [8.5 mm long, 1 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB47375FF23FDF4F4BF61DC.taxon	description	Description. Syntypes (BMNH 1926.4.30.130 – 131) complete, smaller syntype better preserved, slightly bent laterally (Fig. 51 A). Body 12 mm long, 2 mm wide, 16 chaetigers; left parapodia of chaetigers 8 – 10 and, right parapodia of chaetigers 6, 7, 9 and 12 previously removed. Larger syntype slightly bent laterally and ventrally. Body 13 mm long, 2 mm wide, 16 chaetigers; left parapodia of chaetigers 8 – 9, and right parapodia of chaetigers 2, 3, 9 – 11 previously removed; right parapodia of chaetiger 8 removed for observing parapodial features (kept in container with syntypes). Pharynx with a longitudinal, and a transverse previously made dissections. Body pale, eyes and peristomial middorsal tubercle brownish; most cirri missing. Prostomium longer than wide, wider anteriorly (Fig. 51 B). Lateral antennae with ceratophores distinct, without tips in smaller syntype, almost as long as prostomium, longer than palps in larger syntype; palpophores slightly longer than palpostyles. Median antenna tiny, probably eroded, tapered, inserted centrally between eyes. Eyes brownish, round; anterior eyes emarginate anteriorly, slightly larger and slightly more distant to each other than posterior round ones; in lateral view anterior and posterior eyes distinct (Fig. 51 B). Nuchal organs lobes U-shaped, parallel to convergent, prostomial posterior projections pale; middle furrow deep. Lateral ciliated bands visible dorsally. Tentacular cirri broken, one ventral incomplete one reaches chaetiger 1. Lateral cushions swollen, projected, entire along body, longitudinal striae visible. Pharynx fully everted in smaller syntype (Fig. 51 C); anterior margin with 20 small, oval denticles, middorsal and midventral areas bare. Upper and lower jaws single, tiny, barely pigmented (an irregular fleshy projection ahead of dorsal jaw). Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, about 50 per bundle, delicate, many broken; if entire, approaching neurochaetal tips, subdistal denticles mostly eroded, visible in a few chaetae, fine. Notacicular and neuracicular lobes tapered, pointed (Fig 51 D); notacicular lobes twice longer than wide, neuracicular ones as long as wide. Neurochaetae broken in most chaetigers, about 20 per bundle, blades bidentate, 11 – 13 times longer than wide, guards approaching subdistal tooth (Fig. 51 D, inset). Posterior region tapered. Prepygidial segment without cirri, dorsal cirrophores twice wider than ventral ones. Pygidium with anus terminal, anal cirri missing. Oocytes not seen. Variation. A non-type, better-preserved small specimen (MNHN Musorstom 3 - 137) has about 30 neurochaeta per bundle, and their blades are 4 – 16 times longer than wide. The illustration by Pettibone (1970, Fig. 19 e) based upon the study of the syntypes indicates they were 5 – 11 times longer than wide.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB47375FF23FDF4F4BF61DC.taxon	discussion	Remarks. Paradalhousia papillosa (Monro, 1926) n. comb. is reinstated because it was regarded as a junior synonym of Leocrates giardi Gravier, 1900 by Pettibone (1970: 219), but they belong to different genera. Further, P. papillosa is newly combined into Paradalhousia because it has U-shaped nuchal organs, single upper and lower jaws, and pale neurochaetae. On the other hand, P. papillosa differs from P. oculata (Treadwell, 1906) especially in the number of neurochaetae, and in the orientation of their distal teeth. In P. papillosa there are about 20 (– 30) neurochaetae per bundle, and their distal teeth are straight, whereas in P. oculata there are about 50 neurochaetae, and their distal teeth are falcate. Monro (1926) included this species from the Macclesfield Bank, South China Sea, as a part of the third contribution from the HMS Alert Expedition samples (Monro, 1926). However, that voyage did not include the China Sea, as can be confirmed elsewhere (Coppinger 1884, Günther 1884). Consequently, the collector attribution, or field data are undefined, but the locality is regarded as precise, albeit the depth was not indicated in the third contribution. For the first two contributions (Monro 1924 a, b), other species were recorded from the same site from stations made at 54 – 65 m, 54 – 81 m, 54 – 90 m, 90 – 108 m. However, sampling was made by two vessels: HMS Rambler in early 1888, and HM Surveying-Vessel Penguin in early 1892, and all specimens were deposited in The Natural History Museum, London (Anonymous 1895). The corals collected during the first survey were reported by Bassett-Smith (1890), who included the description of five new species, and also included a map with depth data and collecting stations.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB47375FF23FDF4F4BF61DC.taxon	distribution	Distribution. Macclesfield Bank (E Paracel Islands) to the Philippine Islands, in mixed bottoms, 54 – 108 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7375FF23FEDDF5D162E3.taxon	type_taxon	Type species. Leocrates diplognathus Monro, 1926.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7375FF23FEDDF5D162E3.taxon	diagnosis	Diagnosis. Hesioninae with two lateral antennae, and one median antenna on dorsal prostomial surface. Palps biarticulate, palpophores large, massive, palpostyles smaller, blunt. Eyes dark, black or brown, anterior ones larger than posterior ones, sometimes approaching each other in lateral view. Nuchal organs lobes L-shaped. Peristomial dorsolateral tubercles large, ventrolateral ones smaller; ventral ridge sometimes with marginal papillae. Pharynx with upper jaws double, lower one transverse plate. Parapodia sesquiramous along anterior 4 or 5 chaetigers, biramous thereafter. Notochaetae subdistally denticulate, delicate, sometimes abundant, size variable, never reaching neurochaetal tips. Neurochaetae compound falcigers, blades bidentate, guards approaching subdistal tooth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7375FF23FEDDF5D162E3.taxon	etymology	Etymology. The genus-group name is derived from Lamprophaea Grube, 1867 reinstated (see above), with the Greek prefix Para - (= near) to emphasize their close resemblance. Gender. Feminine, as the stem genus-group name.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7375FF23FEDDF5D162E3.taxon	discussion	Remarks. Paralamprophaea n. gen. resembles Lamprophaea Grube, 1867 because they have palps biarticulate, pharynx with jaws, nuchal organs lobes L-shaped, and parapodia sesquiramous anteriorly and biramous posteriorly. These two genera differ especially by the type of jaws. In Paralamprophaea the pharynx has upper jaw double, T-shaped, and lower one as a transverse plate, whereas in Lamprophaea the pharynx has upper and lower jaws single, fang-shaped.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7377FF23F948F0DE62C1.taxon	description	Fig. 52 urn: lsid: zoobank. org: act: 11 F 8 BEC 6 - 801 D- 4188 - B 819 - 6 C 263690 CAE 2	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7377FF23F948F0DE62C1.taxon	materials_examined	Type material. Central Indian Ocean, Maldives. Holotype (UF 4141), Capital Reef, S coast of Nilandoo Island (03 ° 02 ’ 55.68 ” N, 72 ° 53 ’ 21.12 ” E), 20 m, 16 May 2014, M. Oliveiro & M. Taviani, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7377FF23F948F0DE62C1.taxon	description	Description. Holotype (UF 4141) complete, slightly bent dorsally, anterior end depressed after pressure upon fixation, mature. Body oboconic, tapered, wider anteriorly, 20.5 mm long, 2.5 mm wide (without parapodia); left parapodium of chaetiger 8 removed for observing parapodial features. Body brownish, wide transverse segmental bands separated by paler intersegmental thinner bands, with similar pigmentation intensity throughout dorsum (Fig. 52 A), but corresponding bands of tentacular belt and chaetigers 1 and 2 fused, bands of chaetiger 16 and preanal segment with intersegmental paler band incomplete; bands better defined along anterior margin, posterior margin middorsaly indented. Dorsal cirri barely banded brown and pale, tapered. Prostomium as long as wide, wider medially by compression, slightly damaged, with a complex pigmentation pattern extending into palps and nuchal organs lateral branches (Fig. 52 B, C). Darker areas connected into an irregularly banded shape, extended anteriorly as an inverted U, leaving a central prostomial area paler, and a posterior W with the lower areas solid, continued into nuchal organs lobes. Lateral antennae with ceratophores distinct, antennae shorter than prostomium, slightly shorter than palps; palpophores 2 – 3 times longer than palpostyles; median antenna distorted by compression, reaching anterior prostomial margin, inserted between anterior eyes. Eyes dark-brown, anterior eyes reniform, twice larger than posterior round ones, and more distant to each other than posterior eyes (Fig. 52 C). Nuchal organs lobes L-shaped, lateral lobes not surpassing level of lateral prostomial margins; lateral ciliated bands not visible, obliterated by compressed lateral prostomial margins. Tentacular cirri banded with brownish bands, each band darked medially, longest ones reaching chaetiger 5. Lateral cushions barely projected laterally, entire, longitudinal striae visible. Pharynx not exposed, observed by dissection. Anterior margin smooth. Jaws hyaline, golden; upper jaw double, lower one single. Lateral vesicles not seen. Basal pharynx ring with 1 – 3 series of tapered papillae, decreasing in number dorsally and ventrally, middorsal and midventral regions smooth. Dorsal cirri smaller than body width. Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, first notochaetal lobe well-developed, slightly smaller than those present in chaetigers 6 – 8 (Fig. 52 D, inset), notochaetae scarce, about 20 per bundle, delicate, arranged in bundles, notochaetae subdistally denticulate, denticles small. Notacicular lobes short, round, slightly longer than wide; neuracicular lobes blunt, wider than long. Neurochaetae about 20 per bundle; neurochaetal blades bidentate, 2 – 5 times longer than wide, guards approaching subdistal tooth (Fig. 52 E, insets). Posterior region tapered. Prepygidial segment with cirri banded, dorsal ones 4 – 5 times longer than ventral ones. Pygidium with anus terminal; anal cirri banded reaching chaetiger 15. Oocytes not seen; testis visible through chaetiger 8 are testis.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7377FF23F948F0DE62C1.taxon	etymology	Etymology. The specific epithet is derived after Miss Amanda Bemis, Collection Manager of the University of Florida Museum of Natural History, Gainesville, in recognition of her help and support for my research activities. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7377FF23F948F0DE62C1.taxon	discussion	Remarks. The pigmentation pattern of Paralamprophaea bemisae n. sp. differs from the two other similar species, P. diplognatha (Monro, 1926) n. comb. from the South China Sea, and P. leslieae n. sp. from Kiribati, because in P. bemisae the dorsal pigmentation is homogeneous throughout the body, not progressively paler posteriorly as shown in the two other species. The species might have been already recorded from the Eastern Indian Ocean by Fauvel (1932: 62, 1953 b: 107), as Leocrates diplognathus, from the Mergui Archipelago, Birmania. He indicated the dorsal pigmentation as “ dark chestnut-brown traversed by intersegmental bands of white ”. However, in the original description, Monro (1926: 313) indicated that the “ colouring is more intense in the anterior segments, and in two examples it disappears altogether behind the first half-dozen segments. ” Because Fauvel (1932) did not indicate any variation in pigmentation intensity, the affinities of these two Indian Ocean records would be clarified after a comparison with his specimens, but currently they are not available.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBA7377FF23F948F0DE62C1.taxon	distribution	Distribution. Only known from the Maldives, in the Central Indian Ocean, in 20 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB87371FF23FBE8F0F361F1.taxon	description	Fig. 53 urn: lsid: zoobank. org: act: BD 71540 B- 2 B 16 - 409 A-A 57 E- 2690 A 68 E 6 D 91	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB87371FF23FBE8F0F361F1.taxon	materials_examined	Type material. Madagascar. Holotype (MNHN A 861), Sta. P 4 (15 ° 24 ’ 30 ” S, 46 ° 02 ’ 00 ” E), 250 – 265 m, 7 Nov. 1972, A. Crosnier, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB87371FF23FBE8F0F361F1.taxon	description	Description. Holotype (MNHN A 861) slightly damaged, complete. Body obconic, 34 mm long, 4 mm wide; dorsum with transverse brownish bands with anterior and posterior margins well defined (Fig. 53 A), pigmentation paler posteriorly. Venter pale. Right parapodium of chaetiger 8 removed for observing chaetal features. Prostomium slightly wider than long, lateral margins parallel (Fig. 53 B). Lateral antennae with distinct ceratophores, markedly longer than prostomium and palps; palpophores slightly longer than palpostyles. Median antenna directed posteriorly, short, not reaching anterior prostomial margin, inserted between eyes. Eyes brownish, round, anterior eyes twice larger and more distant to each other than posterior eyes (Fig. 53 A). Nuchal organs lobes markedly surpassing lateral prostomial margins. Lateral ciliated bands visible dorsally. Longest tentacular cirri without tips, reaching chaetiger 8. Lateral cushions barely projected, bifid along most body segments; longitudinal striae visible. Peristomium with three dorsal tubercles, dorsolateral ones truncate, not reaching the level of middorsal tubercle; ventrally with shorter lateral tubercles and about 16 ventral, round cushions (Fig. 53 C, D). Pharynx fully exposed (Fig. 53 C). Anterior margin smooth. Jaws brownish, upper jaw double, lower one transverse plate, without lateral transparent blades. Lateral vesicles not seen, but left pharynx side laterally expanded into a blunt cone. Basal ring smooth, without papillae. Dorsal tentacular cirri damaged, some without tips, middle segments with cirri at least twice longer than body width (including parapodia). Chaetigers 1 – 4 without notochaeetae; notochaetae present along chaetigers 5 – 16; notacicular lobes basally swollen, slightly projected into a hemispheric short lobe (Fig. 53 E); neuracicular lobes blunt, wider than long. Notochaetae about 20 per bundle, subdistally denticulate, denticles coarse. Neurochaetae about 30 per bundle, some blades missing, blades decreasing in size ventrally, bidentate, 2 – 7 times longer than wide, guards approaching subdistal tooth (Fig. 53, insets). Posterior end damaged, tapered into a truncate lobe. Prepygidial segment with dorsal cirri 5 – 6 times longer than ventral ones. Pygidium with anus terminal, anal cirri damaged, reaching chaetiger 11. Oocytes not seen, perhaps there were only testis, or gonads damaged.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB87371FF23FBE8F0F361F1.taxon	etymology	Etymology. This species name is after Alain Crosnier, a famous French carcinologist, who kindly invited me in 1999 to process the material in the Paris Museum from the Indian and Western Pacific regions, the so-called Musorstom Expeditions (Salazar-Vallejo 1999). He was personally involved in many ambitious sampling programs in Western Africa, Madagascar, and several archipelagos in the Western Pacific. The 1999 visit allowed me to start studying the extensive and impressive Paris collections, and because of his personal support and with funding from the museum, a series of research visits have helped me do a better taxonomic work. This is a modest means to thank him for his kind support, and especially because he collected the holotype.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB87371FF23FBE8F0F361F1.taxon	discussion	Remarks. Paralamprophaea crosnieri n. sp. resembles P. diplognatha (Monro, 1926) n. comb. from the China Sea, and P. leslieae n. sp. from Kiribati because they have transverse brownish bands along body, each with welldefined anterior and posterior margins. Their main difference is the shape of the peristomial dorsolateral tubercles. In P. crosnieri they are truncate, whereas they are lobate in P. diplognatha, and hemispherical in P. leslieae. The suspicion that these three species would belong to the same one, with these tubercles reflecting a progression in size depending on body size, cannot be sustained. The smaller species, P. diplognatha and P. leslieae, have bodies of similar size (13 – 22 mm vs 12 – 19 mm), but the shape of the dorsolateral tubercles does not change with size; further, the holotype of P. crosnieri is the largest specimen (34 mm long) but has truncate tubercles.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFB87371FF23FBE8F0F361F1.taxon	distribution	Distribution. Only known from the type locality, in Madagascar, in 250 – 265 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBE7373FF23FEB8F01261F1.taxon	description	Figs 54, 55	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBE7373FF23FEB8F01261F1.taxon	materials_examined	Type material. South China Sea. Four syntypes (BMNH 1926.4.30.123 – 125), Macclesfield Bank, no further data [largest syntype used for description; others used for variation]. Additional material. Japan. One specimen (ECOSUR 3080), off Yakushinsone, TR / V Toyoshio, Sta. 3 ’ - 2 (29 ° 49.073 ’ N, 130 ° 22.549 ’ E), 172 – 182 m, 21 Oct. 2016, N. Jimi, coll. [27 mm long, 4 mm wide]. Sulu Sea. Six specimens (MNHN Musorstom 3 - 131), Philippines, Sta. 131 (11 ° 37 ’ N, 121 ° 46 ’ E), 111 – 113 m, 5 Jun. 1985 [7.5 – 11.0 mm long, 1.0 mm wide]. Philippines. Nine specimens (MNHN Musorstom 3 - 117), Timor Sea, R / V Coriolis, Sta. 117 (12 ° 31 ’ S, 120 ° 39 ’ E), 92 – 97 m, 3 Jun. 1985 [7.5 – 16.5 mm long, 1 – 2 mm wide]. One specimen (MNHN Musorstom 3 - 121), Timor Sea, R / V Coriolis, Sta. 121 (12 ° 08 ’ S, 121 ° 17 ’ E), 73 – 84 m, 3 Jun. 1985 [15 mm long, 2 mm wide]. Five specimens (MNHN Musorstom 3 - 131), Timor Sea, R / V Coriolis, Sta. 131 (11 ° 37 ’ S, 121 ° 43 ’ E), 120 – 122 m, 5 Jun. 1985 [10 – 15 mm long, 1 mm wide]. Two specimens (MNHN Musorstom 3 - 134), R / V Coriolis, Sta. 134 (12 ° 01 ’ S, 121 ° 57 ’ E), 92 – 95 m, 5 Jun. 1985 [17 – 18 mm long, 2.0 – 2.5 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBE7373FF23FEB8F01261F1.taxon	description	Description. Largest syntype (BMNH 1926.4.30.123 – 125) complete; body slightly wider medially, first four chaetigers with dark wide transverse band, interrupted by an irregular, barely paler intersegmental thin band (Fig. 54 A); middle and posterior chaetigers without banding (probably faded out). Body 22 mm long, 2.5 mm wide, (without parapodia); pharynx exposed, previously cut transversely and laterally; left parapodium of chaetiger 8 and right parapodia of chaetigers 7 and 12 previously removed (some kept in container, unspecified), an irregular fracture of the body wall by chaetigers 12 – 13. Venter with a longitudinal middle pale area and two longitudinal darker bands along body. Prostomium slightly wider than long, wider anteriorly, laterally constricted before anterior eyes, laterally expanded along eyes region (Fig. 54 A). Lateral antennae with distinct ceratophores, slightly shorter than prostomium, longer than palps; palpophores slightly longer than palpostyles. Median antenna missing, inserted between anterior eyes. Eyes round, barely pigmented, anterior eyes twice larger and more distant to each other than posterior eyes. Nuchal organs lobes L-shaped, lateral branches markedly surpassing lateral prostomial margins; lateral ciliated bands visible dorsally (Fig. 54 A). Longest tentacular cirri without tips, reaching chaetiger 6. Lateral cushions projected, bifid along first body half, entire along posterior one, longitudinal striae visible in some parapodia. Peristomium with three dorsal tubercles, lateral ones projected anteriorly into 5 short lobes, middle one larger, more projected; ventrally with two triangular, blunt cones; papillae minute (visible under dorsolateral tubercles); lateral tubercles blunt, smaller than dorsolateral ones. Pharynx fully exposed (Fig. 54 B). Anterior margin smooth. Jaws hyaline, core brownish; upper jaw double, lower one transverse plate (Fig. 54 C, D), exposed blades transparent, core brownish. Lateral vesicles present on both sides, globose (right one damaged by dissection). Middle ring corrugated, without papillae. Dorsal tentacular cirri as long as body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, first notaciular lobes minute, barely visible, notacicular lobes basally swollen, tapered, slightly capitate (Fig. 54 E, inset); notochaetae up to 20 per bundle, delicate, denticles fine. Neurochaetae about 20 per bundle, most blades missing, blades decreasing in size ventrally (and in posterior chaetigers), bidentate, 4 – 8 times longer than wide (3 – 4 times longer than wide in posterior chaetigers), guards approaching subdistal tooth (Fig. 54 E, insets). Posterior region tapered into a truncate lobe. Prepygidial segment with dorsal cirri 3 – 4 times longer than ventral ones. Pygidium with anus terminal, anal cirri missing. Oocytes grouped in a gonad fragment in a posterior chaetiger, each oocyte about 100 µm in diameter. Variation. Syntypes (BMNH 1926.4.30.123 – 125) 13 – 16 mm long, 2 – 3 mm wide, 16 chaetigers; smallest one almost broken in two, larger one with transverse bands barely visible dorsally. Prostomium with lateral margins curved, wider anteriorly and in ocular region. Lateral antennae damaged; palpophores up to twice longer than palpostyles; median antenna present in one syntype, short, tapered, not reaching anterior prostomial margin, inserted between anterior eyes. Nuchal organs lateral branches reaching prostomial lateral margins or slightly surpassing them (Fig. 55 A, D). Peristomium with three dorsal tubercles, lateral ones projected anteriorly into a blunt lobe; ventrally with two tubercles projected into a larger lobe (Fig. 55 B). Upper jaw double (Fig. 55 C), ventral one transverse plate, transparent margin eroded in some syntypes. Lateral vesicles visible in larger syntypes, blunt, damaged by dissection in two of them. Basal pharynx ring smooth. Notochaetae start in chaetiger 5.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBE7373FF23FEB8F01261F1.taxon	discussion	Remarks. Paralamprophaea diplognatha (Monro, 1926) n. comb. is newly combined because of the development of nuchal organs lobes and the type of pharynx armature. Further, P. diplognatha resembles P. leslieae n. sp. from Kiribati by having a similar pigmentation pattern, with transverse brownish entire (rarely indented) bands. The main differences between these two species is the shape of the dorsolateral peristomial tubercles, and the size of papillae in the basal pharynx ring. In P. diplognatha dorsolateral tubercles are anteriorly projected, and papillae are minute, whereas in P. leslieae the tubercles are semicircular, not projected, and papillae are large. Some remarks about the wrong inclusion of this and other species as part of the HMS Alert Expedition by Monro (1926) were made for Paradalhousia papillosa (see above). Field data deserve improvement, but solving that is beyond my current objectives. Suffice it to say that the specimens from the Macclesfield Bank were collected in 1888 by the HMS Rambler, or in 1892 by the HM Surveying Vessel Penguin, but nothing can be assured about precise dates, localites or depths.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBE7373FF23FEB8F01261F1.taxon	distribution	Distribution. South China Sea to the Philippines, in sediments, in 73 – 182 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBC734EFF23FA6FF02264C9.taxon	description	Figs 56, 57	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBC734EFF23FA6FF02264C9.taxon	materials_examined	Type material. Gulf of Guinea, São Tomé and Príncipe Islands. Holotype (ZMH V- 5700), Ilhéu das Rolas (00 ° 00 ’ 00 ” N, 06 ° 31 ’ 25 ” E), near Ilha de São Tomé, R. Greeff, coll. Additional material. Cabo Verde. One specimen (MNHN A 372), Campagne de la Calypso, Iles du Cap Vert, Fogo Island (14 ° 57 ’ 00 ” N, 24 ° 20 ’ 33 ” W), Sta. 35, 0.6 km SW off Encarnacao, 45 – 55 m, 20 Nov. 1959 [14 mm long, 3 mm wide]. Two specimens (MOM 180817), Albert 1 er de Monaco Expeditions, Sta. 1203 (15 ° 54 ’ 00 ” N, 22 ° 54 ’ 45 ” W), 6.2 km off SW Boa-Vista Island, 91 m, hard bottom, 18 Aug. 1901 [14 mm long, 2.5 mm wide]. Gulf of Guinea. One specimen (MNHN A 732 a), Príncipe Island, R / V Calypso, Sta. 97 (01 ° 43 ’ 10 ” N, 07 ° 28 ’ 20 ” E), 73 m, 1 Jul. 1956 [19 mm long, 3.5 mm wide]. One specimen (MNHN A 372 b), R / V Calypso, Sta. 110 (01 ° 20 ’ 45 ” N, 07 ° 17 ’ 37 ” E), Tinhosa Grande Islet, 25 – 40 m, 7 Jul. 1956 [17 mm long, 3 mm wide]. One specimen (MNHN A 475), Benin (Dahomey), 06 ° 10 ’ 30 ” N, 02 ° 32 ’ 00 ” E), 55 m, corals, 24 Oct. 1963, M. Faubert, coll. [26 mm long, 3 mm wide]. One specimen (BMNH 1930.10.8.1128), RSS Discovery I, Sta. 283, 1.3 – 1.6 km N 12 ° E off Pyramid Rock (01 ° 20 ’ 03 ” S, 05 ° 38 ’ 09 ” E), 18 - 30 m, 4 Aug. 1927 [data used for variation]. Fifty-six specimens (NHMD 237476), R / V Galathea, Sta. 49, Ilhéu das Rolas (00 ° 00 ’ N, 06 ° 32 ’ E), near Ilha de São Thomé, corals, 42 m, 29 Feb. 1950 [6 – 24 mm long, 1 – 3 mm wide]. Five specimens (NHMD 237477), R / V Galathea, Sta. 50, Ilhéu das Rolas (00 ° 00 ’ N, 06 ° 32 ’ E), south Ilha de São Tomé, 5 – 8 m (0 m in Kirkegaard 1983: 214), 29 Feb. 1950 [8 – 22 mm long, 1.0 – 3.5 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBC734EFF23FA6FF02264C9.taxon	description	Description. Holotype (ZMH V- 5700) complete, slightly damaged (Fig. 56 A). Body obconic, blunt anteriorly, tapered posteriorly, 10 mm long, 1.5 mm wide, 16 chaetigers; right parapodium of chaetiger 8, and left parapodia of chaetigers 2, 7, 8 previously removed. Previously midventrally dissected up to chaetiger 3 to study pharynx and jaws; previously pinned through chaetiger 8. Most tentacular, and parapodial cirri on site. Body pale. Prostomium as long as wide, slightly wider anteriorly (Fig. 56 B, C). Lateral antennae with ceratophores welldefined, antennae about as long as prostomium, slightly longer than palps. Palpophores 1.5 times longer than palpostyles. Median antenna with tip broken, not reaching anterior prostomial margin, inserted centrally on prostomium. Eyes colorless (pigmentation and shape after original description and redescription); eyes blackish, anterior eyes semilunar, 3 – 4 times larger than posterior ones; posterior eyes round (original description), to semilunar (redescription). Nuchal organs lobes L-shaped, laterally projected, not reaching lateral prostomial margins, lobes expanded distally, not parallel-sided; lateral ciliated bands narrow, not visible dorsally. Tentacular cirri almost complete, longest one reach chaetiger 10. Lateral cushions swollen, entire, longitudinal striae not seen. Peristomium with two round, smooth dorsolateral tubercles, ventral tubercles slightly longer than wide, smooth. Pharynx details observed by previous dissection, anterior margin smooth. Upper jaw double, colorless, inserted subdistally; lower jaw not seen, probably destroyed by dissection. Dorsal cirri broken, shorter than body width. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16, about 10 per bundle, delicate, arranged in a bundle (Fig. 56 D), notochaetae subdistally denticulate, denticles fine. Notacicular lobes short, blunt, round (Fig. 56 D, inset); neuracicular lobes blunt, wider than long; aciculae dark brown, tapered, surround by browinish pigmented areas. Neurochaetae about 20 per bundle, some blades missing or severely eroded, decreasing in size ventrally (Fig. 56 D, inset), bidentate, 2 – 5 times longer than wide (2 – 4 times longer than wide in Pettibone 1970, Fig. 22 f), guards approaching subdistal tooth. Posterior region tapered, prepygidial segment and pygidium without cirri. Pygidium with anus terminal, anal cirri size unknown. Oocytes not seen. Variation. A large specimen collected close to the type locality (BMNH 1930.10.8.1128) is 25 mm long, 4 mm wide, with feebly pigmented, wide transverse brownish bands along chaetigers 1 – 4 (Fig. 57 A), chaetigers 5 – 10 pale (damaged), posterior chaetigers with indistinct transverse bands, darker than preceding region. Prostomium slightly longer than wide, wider anteriorly and along eyes’ region (Fig. 57 B); lateral antennae longer than prostomium, slightly longer than palps; palpophores twice longer than palpostyles. Median antenna complete, tapered, surpassing anterior prostomial margin, inserted centrally between eyes. Eyes slightly darker than surrounding integument, anterior eyes reniform, twice larger than posterior round ones. Eyes barely pigmented. Nuchal organs lobes slightly projected beyond lateral prostomial margins, with a basal constriction and progressively swollen; lateral ciliated bands barely visible dorsally. Peristomium with dorsolateral tubercles round, slightly wider than long, smooth, ventral tubercles blunt triangular. Pharynx anterior margin smooth, lateral vesicles present, globose, right one with a transverse constriction looking like an upper and a lower vesicle (Fig. 57 C). Upper jaw double, brownish, ventral one single transverse plate. Tentacular cirri broken, longest one reaching chaetiger 4. Notochaetae from chaetiger 4, about 20 per bundle, subdistally denticulate, denticles coarse. Notacicular lobe digitate, twice longer than wide (Fig. 57 D). Neuracicular lobe short (about twice wider than long). Neurochaetae about 20 per bundle, blades bidentate, decreasing in size ventrally, 4 – 6 times longer than wide, guards approaching subdistal tooth (Fig. 57 D, insets). Other non-type specimens (ZMUC 237476) were studied for nuchal organs lobes variations. They are slightly divergent, not surpassing lateral prostomial margins in smallest specimen (Fig. 57 E), progressively growing laterally surpassing lateral prostomial margins and with lobes medially furrowed in larger specimens (Fig. 57 F). Eyes are more emarginate or reniform in smaller specimens, and become round in larger ones. Dorsal pigmentation usually involves a thin transverse pale band corresponding to parapodia, but becomes middorsaly wider, and less defined medially and posteriorly.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBC734EFF23FA6FF02264C9.taxon	discussion	Remarks. Paralamprophaea greeffiana (Augener, 1918) n. comb. is newly combined because the species match the diagnostic generic features of Paralamprophaea, instead of those of Leocrates, where it was originally included, especially after the type of nuchal organs lobes and pharynx armature. The declination for the specific epithet has to be modified to match the gender of the genus, feminine (ICZN 1999, Arts 34.2, 48). On the other hand, L. greeffiana is unique among similar species in the genus because its notochaetae start in chaetiger 4, against being present from chaetiger 5, as in the case in other similar species: L. bemisae n. sp., L. diplognatha (Monro, 1926) n. comb., and L. meyeri n. sp. The record by Monro (1930) for the Gulf of Guinea, as L. diplognathus corresponds to L. greeffiana. Monro (1930: 92) indicated that his specimen (BMNH 1930.10.8.1128) has notochaetae from chaetiger 4, as is the case for L. greeffiana, instead of having them from chaetiger 5, as in L. diplognatha. The difference in the development of the nuchal organs lobes’ lateral branches, being better developed in the Gulf of Guinea’s specimen, can be explained by size differences; Augener’s type specimen was 12 mm long, a juvenile, wheras Monro’s one was twice as large (24 mm long). The development of the nuchal organs lobes of the holotype lies somewhere between the smallest juvenile and larger specimens herein illustrated.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FFBC734EFF23FA6FF02264C9.taxon	distribution	Distribution. Cabo Verde to the Gulf of Guinea, in sediments at 5 – 55 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	description	Figs 58, 59 urn: lsid: zoobank. org: act: F 2483 FC 9 - 6 CEC- 45 B 6 - A 6 EC- 95 D 70 E 0 E 3 D 0 C	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	materials_examined	Type material. Kiribati. Holotype (LACM 10144), Phoenix Islands, Kanton Island, outer reef, Sta. 34 (02 ° 48 ’ 57.54 ” S, 171 ° 42 ’ 59.22 ” W), 39 m, rubble, 21 Jun. 2002, M. J. Adams, coll. Additional material. Kiribati. One specimen (LACM 10145, Phoenix Islands, Oroma Island (04 ° 30 ’ 17.10 ” S, 178 ° 52 ’ 57.72 ” W), 13 m, dead coral, 3 Jul. 2002, M. J. Adams, coll. [15 mm long, 2 mm wide]. One specimen (UF 4030), Line Islands, Millenium Island (09 ° 54 ’ 36.00 ” S, 150 ° 12 ’ 36.00 ” E), dead Pocillopora sp., 12 m, 5 Nov. 2013, F. Michonneau & N. Knowlton, coll. [15 mm long, 2 mm wide]. One specimen (UF 4043), Line Islands, Malden Island, Tent site (04 ° 01 ’ 12.00 ” S, 154 ° 55 ’ 12.00 ” E), dead Pocillopora sp., 10 m, 1 Nov. 2013, F. Michonneau & N. Knowlton, coll. [14 mm long, 2 mm wide]. French Polynesia. One specimen (UF 3051), Society Islands, Moorea, off Motu Ahi, north of Afareity Pass (17 ° 33 ’ 10.80 ” S, 149 ° 46 ’ 22.80 ” W), reef slope, rubble, 63 – 70 m, 21 Jan. 2012, J. Earle, D. Pence, R. Pyle & R. Whitton, coll. [12 mm long, 2 mm wide]. One specimen (UF 3059), Society Islands, Moorea, north of Vaire Pass, off Sofitel (17 ° 30 ’ 15.84 ” S, 149 ° 45 ’ 30.24 ” W), deep reef slope, rubble, 65 – 66 m, 26 Jan. 2012, J. Earle, D. Pence, R. Pyle & R. Whitton, coll. [resembling UF 3051 in pigmentation but slightly darker; 15 mm long, 1.5 mm wide]. Philippines. One specimen (MNHN Musorstom 3 - 104), Timor Sea, Sta. 104 (13 ° 56 ’ S, 120 ° 22 ’ E), 13 m, 1 Jun. 1985 [17 mm long, 3 mm wide]. Christmas Island. One specimen (BMNH 1933.10.11.21), Flying Fish Cove (10 ° 25 ’ 18 ” S, 105 ° 40 ’ 41 ” E), no further data, F. Harius, coll. [19 mm long, 2.5 mm wide].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	description	Description. Holotype (LACM 0000) complete. Body obconic, tapered, wider anteriorly, wide transverse bands along body (Fig. 58 A), interrupted intersegmentally by pale thin bands; bands with anterior and posterior margins well defined, progressively decreasing in intensity in middle and posterior chaetigers; 12 mm long, 1.8 mm wide (without parapodia); pharynx exposed, previously cut longitudinally; left parapodium of chaetiger 9 dissected (kept in container). Prostomium as long as wide, lateral margins straight, wider anteriorly and in ocular area; pigmentation brownish, leaving pale areas, one roughly X-shaped ahead of median antenna, two ahead of anterior eyes anteriorly ta- pered, and two others behind posterior eyes continued laterally and posteriorly (Fig. 58 B). Lateral antennae with distinct ceratophores, slightly longer than prostomium, about twice longer than palps; palpophores as long as palpostyles. Median antenna short, not reaching anterior prostomial margin, inserted between anterior eyes. Eyes dark brown, anterior eyes twice larger, slightly emarginate anterolaterally and more separated from each other than posterior round eyes. Nuchal organs lobes L-shaped, brownish, lateral branches barely surpassing lateral prostomial margin level, slightly swollen subdistally. Lateral ciliated bands visible dorsally. Tentacular cirri almost complete, tips eroded, longer ones reaching chaetiger 11. Lateral cushions swollen, projected, entire, longitudinal striae visible. Peristomium with three dorsal and two ventral tubercles, dorsolateral ones hemispherical; ventral ones hemispherical. Pharynx fully exposed (Fig. 58 C). Anterior margin smooth. Jaws hyaline, core golden; upper jaw double, lower one transverse bar. Lateral vesicles present on both sides, small, globose. Basal pharynx ring with 1 – 4 series of blunt, conical papillae, lateral, under lateral vesicles, decreasing in number dorsally and ventrally, middorsal and midventral regions smooth, bare dorsal surface smaller than ventral one. Dorsal tentacular cirri broken, as long as body width (excluding parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 16, first notochaetal lobes minute, barely visible, notochaetae up to 10 per bundle, delicate, denticles fine. Notacicular lobes digitate, blunt; neuracicular lobes truncate (Fig. 58 D, inset); neurochaetae about 15 per bundle, some blades missing, blades decreasing in size ventrally, bidentate, 3 – 7 times longer than wide, guards approaching subdistal tooth (Fig. 58 D, inset, E). Posterior region tapered into a blunt cone. Prepygidial segment with dorsal cirri 3 – 4 times longer than ventral ones. Pygidium with anus terminal, anal cirri reaching chaetiger 14. Oocytes included in ovaries, oocytes about 80 µm each.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	etymology	Etymology. This species is named after my good friend and teacher, Leslie Harris, Collection Manager of the Allan Hancock Foundation Polychaete Collection (LACM), in recognition of her sustained support during the last 40 years, and as an appreciation of her phenomenal experience with polychaetes from all over the world. She was captivated by the gorgeous specimen that became the holotype, and it is my pleasure to name this species after her. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	description	Variation. The morphological features of the species are rather conservative. However, another specimen from the type locality (UF 4030) shows smaller middorsal papillae in 2 transverse series, in its middle pharynx ring (Fig. 59 A). The midventral surface of the same pharynx ring is smooth (Fig. 59 B, C). The specimen from Christmas Island (BMNH 1933.10.11.21) has a similar pigmentation pattern (Fig. 59 D) and shows three differences: first, its anterior venter surface is mottled (Fig. 59 E); second, neuracicular lobes are round, barely projected (Fig. 59 F), not truncate as in the type specimen; and third, neurochaetal blades are slightly shorter, 3 – 4 times longer than wide (Fig. 59 F, insets) than those present in other specimens. These differences, however, are not regarded as substantial for proposing a separate species.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	discussion	Remarks. Paralamprophaea leslieae n. sp. resembles P. diplognatha (Monro, 1926) n. comb. from the South China Sea by having dorsal transverse bands brownish with margins entire (rarely indented), as indicated in the key above. These two species have two main differences between them regarding the shape of the peristomial dorsolateral tubercles, and the size of papillae in the basal pharynx ring. In P. leslieae the dorsolateral tubercles are semicircular, not projected, and papillae are large, whereas in P. diplognatha tubercles are anteriorly projected, and papillae are minute.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF817349FF23F9E0F1296254.taxon	distribution	Distribution. Kiribati, the Philippines, and Christmas Island, in coralline substrates, in 10 – 70 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF86734AFF23FC45F06A63A1.taxon	description	Fig. 60 urn: lsid: zoobank. org: act: 14 A 7714 F- 6 D 3 D- 4 EC 3 - 83 F 4 - CF 34 CA 9 CC 864	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF86734AFF23FC45F06A63A1.taxon	materials_examined	Type material. Western Pacific, French Polynesia. Holotype (UF 2158), Society Islands, Moorea, West side of Opunohu Pinnacle (17 ° 29 ’ 38.76 ” S, 149 ° 51 ’ 43.20 ” W), rubble and sponges, 10 m, 11 Nov. 2010, C. Meyer, coll.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF86734AFF23FC45F06A63A1.taxon	description	Description. Holotype (UF 2158) complete, slightly dehydrated especially along posterior region, ventrally bent, posterior region twisted. Body tapered, wider anteriorly, 17 mm long, 2 mm wide (without parapodia), 14 chaetigers; chaetigers 15 – 16 and pygidium removed for molecular analysis; an anterolateral dissection for observing pharynx jaws, right parapodium of chaetiger 8 removed for observing parapodial features. Body blackish, pigmentation progressively fading off along body; transverse bands interrupted middorsally along chaetigers 1 – 5, following bands complete, anterior and posterior margins diffuse (Fig. 60 A); dorsal cirri with brownish cirrophores; midventral dark brownish longitudinal band interrupted by transverse oval pale spots; dark brown spots on parapodial bases (Fig. 60 B). Most tentacular, dorsal and ventral cirri on site. Prostomium brownish, slightly longer than wide, slightly wider medially; pale areas include two anterior tapered ones ahead of anterior eyes, three round areas at the level of posterior eyes, and two laterally and posteriorly continued areas behind posterior eyes (Fig. 60 C). Lateral antennae with ceratophores distinct, antennae longer than prostomium and palps; palpophores 2 – 3 times longer than palpostyles. Median antenna short, almost reaching anterior prostomial margin, inserted between anterior eyes. Eyes brownish, anterior eyes semilunar, slightly darker, larger and more distant to each other than posterior round eyes, in lateral view eyes clearly separated (Fig. 60 C). Nuchal organs lobes L-shaped, fully exposed, mostly unpigmented, with blackish areas basally in the outer sides, and as a thin longitudinal line along anterior margin; lateral branches reaching prostomial lateral margins, expanded subdistally, slightly directed anteriorly; lateral ciliated bands barely visible dorsally. Tentacular cirri twisted, almost complete, longer ones reaching chaetiger 8. Lateral cushions low, entire, longitudinal striae distinct. Pharynx not exposed, inner features observed through dissection. Anterior margin smooth, lateral vesicles not seen. Jaws golden, exposed, upper jaw double, lower one single transverse bar. Basal ring with series of large papillae, collapsed by invagination, about 6 papillae per series. Dorsal cirri with tips eroded, as long as body width (including parapodia). Chaetigers 1 – 4 without notochaetae; notochaetae present along chaetigers 5 – 14, up to 20 per bundle, delicate, arranged in bundles, notochaetae subdis- tally denticulate, denticles coarse. Notacicular lobes small, blunt, digitate; neuracicular lobes semicircular, twice wider than long (Fig. 60 D). Neurochaetae about 30 per bundle, blades decreasing in size ventrally, bidentate, 3 – 6 times longer than wide, guards denticulate, approaching subdistal tooth (Fig. 60 E). Posterior region tapered. Preanal and pygidium unknown. Whitish gonad fragments expelled by posterior end dissection. Oocytes in ovary, each about 100 µm.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF86734AFF23FC45F06A63A1.taxon	etymology	Etymology. This species name is to honor Dr. Christopher P. Meyer, Director of the Moorea Biocode Project, National Museum of Natural History, Smithsonian Institution, in recognition of his initiatives in generating COIbarcode libraries for different localities, and because he collected the holotype. The species-group name is a noun in the genitive case (ICZN 1999, Art. 31.1.2).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF86734AFF23FC45F06A63A1.taxon	discussion	Remarks. Paralamprophaea meyeri n. sp. differs from the other species having notochaetae from chaetiger 5, and especially by its pigmentation pattern. Its transverse bands are blackish, not brownish, despite being in ethanol for nearly a decade, and the bands are not continuous, especially along the first few chaetigers, being middorsally interrupted by large oval pale areas connecting to each other middorsally. Further, the venter has a longitudinal middle band progressively thinner and paler, and some ventral round spots in the base of neuropodia.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF86734AFF23FC45F06A63A1.taxon	distribution	Distribution. Moorea, French Polynesia, in mixed bottoms at 10 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF84734BFF23FF6CF0C26274.taxon	type_taxon	Type species. Leocrates wesenberglundae Pettibone, 1970.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF84734BFF23FF6CF0C26274.taxon	diagnosis	Diagnosis. Hesioninae with two lateral antennae, and one median antenna on dorsal prostomial surface. Palps biarticulate, palpophores large, massive, palpostyles smaller, blunt. Eyes dark, black or brown, anterior ones larger than posterior ones, distant to each other in lateral view. Nuchal organs lobes horizontal C-shaped, posterior ciliated bands close to each other. Pharynx with single, fang-shaped upper and lower jaws. Parapodia sesquiramous along a few anterior chaetigers, biramous thereafter. Notochaetae from chaetiger 4, smooth, delicate, sometimes abundant, size variable, never as long as to reach neurochaetal tips. Neurochaetae compound falcigers, blades unidentate, guards, if present, aristate, surpassing distal tooth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF84734BFF23FF6CF0C26274.taxon	etymology	Etymology. The genus-group name is derived from Leocrates, by adding the Greek prefix para- to indicate its close similarity to the stem genus-group name. Gender. Masculine, as in the stem genus-group name.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF84734BFF23FF6CF0C26274.taxon	discussion	Remarks. Paraleocrates n. gen. resembles Leocrates Kinberg, 1866 by having C-shaped nuchal organs lobes, and single, fang-shaped upper and lower jaws in their pharynx. They differ in three features: the start of notochaetae, the surface of notochaetae, and the type of neurochaetal blades. In Paraleocrates notochaetae start in chaetiger 4, their surface is smooth, and neurochaetal blades are unidentate, sometimes with guards surpassing the denticle, whereas in Leocrates notochaetae start from chaetiger 5, they are subdistally spinulose, and neurochaetal blades are bidentate. There are only two species in this genus that can be separated with the key below.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF847345FF23FB20F0DE63A0.taxon	description	Figure 61	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF847345FF23FB20F0DE63A0.taxon	materials_examined	Type material. Philippines. Holotype (ZMA V. Pol. 534.3), Sulu Islands, off NE Gujangan Island, R / V Siboga, Sta. 105 (06 ° 08 ’ N, 121 ° 19 ’ E), 275 m, dredge, coral, 4 Jul. 1899. Indonesia, paratype (ZMA V. Pol. 2104), off Djangkar, Java Island, R / V Siboga, Sta. 5 (07 ° 46 ’ S, 114 ° 30.5 ’ E), 330 m, deep-sea trawl, muddy bottom, 10 Mar. 1899 (used for variation).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF847345FF23FB20F0DE63A0.taxon	description	Description. Holotype (ZMZ V. Pol. 534.3), complete, slightly distorted, dehydrated, most body cirri missing (Fig. 61 A). Body obconic, wider anteriorly, tapered posteriorly, 12 mm long, 2 mm wide, 16 chaetigers; right parapodium of chaetiger 3, and left parapodia of chaetigers 3, 8, 9, and 14 previously removed (three ones removed by Pettibone in a small vial, others missing); a slightly lateral anteroventral dissection for observing pharynx features previously made. Body brownish, venter with two wide longitudinal brown bands (Fig. 61 C), leaving a paler midventral region (purple in recently preserved specimens after Horst). Prostomium slightly wider than long, slightly wider anteriorly (Fig. 61 B). Lateral antennae with ceratophores distinct, left one corrugated, right one missing, slightly longer than left palp (right palp distorted, directed laterally); palpophore 2 – 3 times longer than palpostyles. Median antenna missing, scar inserted centrally, between eyes. Eyes barely pigmented, round, anterior ones twice larger and slightly more distant to each other than posterior ones. Lateral peristomial tubercles round, as long as wide. Nuchal organs lobes horizontal C-shaped, completely concealed by anterior margin of tentacular belt; lateral ciliated bands visible dorsally. Tentacular cirri almost completely missing, one ventral cirrus without tip reaching chaetiger 3. Lateral cushions low, projected, most tripartite; longitudinal striae visible. Pharynx not exposed; jaws observed through previous dissection. Lateral vesicles not seen. Anterior margin with about 20 irregular constrictions. Dorsal and ventral jaws single, fang-shaped brownish, exposed, tapered, ventral jaw smaller than dorsal one. Peristomial dorsolateral tubercles as long as wide, slightly projected anteriorly, smooth. Dorsal cirri broken, some remaining shorter than body width. Chaetigers 1 – 3 without notochaetae (Fig. 61 D); notochaetae present along chaetigers 4 – 16, about 30 per bundle, delicate, arranged in bundles, notochaetal surface smooth. Notacicular lobes tapered, blunt; neuracicular lobes projected, blunt, as long as wide (Fig. 61 E). Ventral cirri surpassing neurochaetal lobes. Neurochaetae about 30 per bundle, decreasing in size ventrally, blades unidentate, 4 – 65 times longer than wide, guards projected beyond tooth as long aristae, tooth slightly falcate (Fig. 61 D, E, insets). Posterior region tapered, almost without cirri. Prepygidial segment with dorsal cirrophores 4 – 5 thicker than ventral ones. Pygidium with anus terminal, anal cirri missing. Oocytes not seen. Variation. Paratype (ZMA V. Pol. 2104) damaged, soft, brownish. Body 35 mm long, 3 mm wide, 16 chae- tigers; cirri and many chaetae missing; left parapodia of chaetigers 4 and 6, and right parapodia of chaetigers 9 and 10 previously removed. An anteroventral dissection for observing pharynx features previously made. Notopodia from chaetiger 4, most notochaetae missing, a few remaining, smooth. Neuropodia with a few chaetae remaining (7 – 15), blades unidentate, most straight, a few bent dorsally, guards projected far beyond tip of tooth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF847345FF23FB20F0DE63A0.taxon	discussion	Remarks. Paraleocrates djankarensis (Augener & Pettibone in Pettibone, 1970) is newly combined into Paraleocrates by having unidentate blades, as opposed to bidentate blades present in Leocrates, as herein restricted. Paraleocrates djankarensis differs from the other species in the genus, P. wesenberglundae (Pettibone, 1970) n. comb. from the Gulf of Oman, in the size of eyes and in the type of neurochaetal blades. In P. djankarensis anterior eyes are twice larger than posterior ones, and neurochaetal blades are unidentate, with teeth round and have long, aristate guards, whereas in P. wesenberglundae eyes are of similar size, and neurochaetal blades have no guards at all, but their teeth are tapered but without aristae. The paratype has neurochaetal blades slightly different because instead of having slightly falcate tooth, as in the holotype, it is rather straight, and even bent dorsally; because of the poor condition of the specimen, no other differences were found and consequently it is regarded as conspecific pending the finding of better specimens.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF847345FF23FB20F0DE63A0.taxon	distribution	Distribution. Philippines and Indonesia, in mixed or muddy bottoms at 275 – 330 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF8A7347FF23FD09F71860FD.taxon	description	Fig. 62	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF8A7347FF23FD09F71860FD.taxon	materials_examined	Type material. Iranian Gulf. Holotype (NHMD 109183), 27 km NW of Kuh-i-Mubarak, Jask Bay, G. Thorson Sta. 78 B (26 ° 03 ’ N, 57 ° 06 ’ E), 70 m, tough grey clay; 21 Apr. 1937. Two paratypes (NHMD 109184, NHMD 109185), 3.2 km NW by North off the Light-buoy at Jask, G. Thorson Sta. 72 B (25 ° 41 ’ N, 57 ° 43 ’ E), 12 m, tough grey clay, 20 Apr. 1937 [data used for variation].	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF8A7347FF23FD09F71860FD.taxon	materials_examined	One paratype (USNM 37647), 1.6 km SE from Hormuz Island, G. Thorson Sta. 80 (27 ° 02 ’ N, 56 ° 31 ’ E), 15 m, soft brown clay, 22 Apr. 1937 (damaged, without cirri, slightly dehydrated, bent dorsally. Body pale, eyes colorless; left parapodia of chaetigers 7 and 8 previously dissected (five parapodia kept with paratype). Prostomium with left lateral antennae, shorter than prostomium, as long as palps; palpophores 4 times longer than palpostyles. Pharynx exposed, anterior margin with crenulations and right lateral vesicle, upper and lower jaws visible, exposed. Notacicular lobes tapered, neuracicular lobes blunt. Chaetigers 1 – 3 without notochaetae, present along chaetigers 4 – 16, broken in many parapodia; about 50 per bundle, delicate, smooth. Neurochaetae with blades entire, without guards, blades decreasing in size ventrally, some with a few tiny denticles along its cutting edge).	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF8A7347FF23FD09F71860FD.taxon	description	Description. Holotype (NHMD 109183), complete, slightly distorted, posterior region bent upward, posterior gut expelled through anus, most body cirri missing (Fig. 62 A). Body obconic, wider anteriorly, tapered posteriorly, 12 mm long, 1.5 mm wide, 16 chaetigers; right parapodium of chaetiger 2, and left parapodia of chaetigers 8 and 13 previously removed (not available in container). Body brownish anteriorly up to chaetigers 4 – 5, paler afterwards, venter with two yellowish longitudinal wide bands along parapodial bases, midventral region pale (Fig. 62 C). Prostomium wider than long, slightly wider anteriorly (Fig. 62 B). Lateral antennae with ceratophores distinct, slightly shorter than prostomium, as long as palps; palpophores 3 – 4 times longer than palpostyles. Median antenna tiny, probably broken, inserted towards the posterior prostomial margin (between posterior eyes after original figure). Eyes without pigmentation, not visible (tiny, slightly darker than surrounding integument, round, anterior ones slightly larger and more separated than posterior ones in paratype NHMD 109184). Nuchal organs lobes horizontal C-shaped, not concealed by anterior margin of tentacular belt; lateral ciliated bands visible dorsally. Tentacular cirri missing, dorsal cirrophores progressively and gradually larger posteriorly. Lateral cushions reduced, entire along anterior and middle regions (two-thirds of body), bipartite in posterior region; longitudinal striae barely visible. Pharynx partially exposed (Fig. 62 C, D). Lateral vesicles not seen (paired, globose in NHMD 109184, 109185). Anterior margin with abundant, irregular contractions, each with smaller folds complicating their counting. Dorsal and ventral jaws single, fang-shaped brownish, tapered, ventral jaw smaller than dorsal one. Peristomial dorsolateral tubercles as long as wide, slightly projected anteriorly, smooth. Dorsal cirri missing. Chaetigers 1 – 3 without notochaetae; notochaetae present along chaetigers 4 – 16. Parapodial features observed in paratypes (NHMD 109184, 109185). Notochaetae delicate, abundant (about 50 per bundle), boken in many parapodia, arranged in bundles, notochaetal surface smooth. Notacicular lobes tapered, with a dorsal middle projection or ridge (NHMD 109184), or tapered (NMHD 109185) (Fig. 62 E). Ventral cirri surpassing neurochaetal lobes (Fig. 62 E, F). Neurochaetae about 30 per bundle, blades unidentate, 12 – 50 times longer than wide, tapered into long aristae, without guards (Fig. 62 E, inset, 62 F, inset). Posterior region tapered into a truncate tip, without cirri. Prepygidial segment reduced, dorsal cirrophores slightly wider than ventral ones. Pygidium with anus terminal, anal cirri missing. Oocytes not seen; present in one paratype (NHMD 109184), some loose, most grouped in gonad fragment; oocytes about 100 µm in diameter. Variation. Paratype NHMD 109184 distorted, completely bent laterally. Body 17 mm long, 1.5 mm wide, 16 chaetigers. Left lateral antenna broken, right one as long as palps; palpophores 4 times longer than palpostyles; lateral peristomial tubercles granulose, probably eroded, left one oval, wider than long, right one as long as wide. Pharynx completely exposed, depressed; jaws single, fang-shaped brownish, upper one larger than lower one; lateral pharynx vesicles globose. Notochaetae from chaetiger 4; most neurochaetal blades on site. Paratype NHMD 109185 distorted, bent laterally; pharynx completely exposed, upper jaw previously removed. Body obconic, colorless, 12 mm long, 1.5 mm wide, 16 chaetigers; all antennae missing; palps present, palpophores 4 times longer than palpostyles; lateral prostomial tubercles round, wider than long; lateral pharynx vesicles globose; notochaetae from chaetiger 4; many neurochaetal blades missing.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF8A7347FF23FD09F71860FD.taxon	discussion	Remarks. Paraleocrates wesenberglundae (Pettibone, 1970) n. comb. belongs in Paraleocrates because it has unidentate blades, instead of bidentate blades present in Leocrates, and hence the new combination. Paraleocrates wesenberglundae differs from P. djankarensis (Augener & Pettibone in Pettibone, 1970) from Indonesia and the Philippines especially in two features: the size of eyes, and the type of neurochaetal blades. In P. wesenberglundae anterior eyes are slightly larger than posterior ones, and neurochaetal blades have no guards at all, with teeth tapered into long aristae, whereas in P. djankarensis anterior eyes are twice larger than posterior ones, and neurochaetal blades are unidentate, with round teeth, and with long, aristate guards.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
2D1987E4FF8A7347FF23FD09F71860FD.taxon	distribution	Distribution. Gulf of Oman, in sediments, in 12 – 70 m depth.	en	Salazar-Vallejo, Sergio I. (2020): Revision of Leocrates Kinberg, 1866 and Leocratides Ehlers, 1908 (Annelida, Errantia, Hesionidae). Zootaxa 4739 (1): 1-114, DOI: 10.11646/zootaxa.4739.1.1
