Hesione picta Müller, 1858

Salazar-Vallejo, Sergio I., 2018, Revision of Hesione Savigny in Lamarck, 1818 (Annelida, Errantia, Hesionidae), Zoosystema 40 (12), pp. 227-325 : 289-296

Hesione picta Müller, 1858

( Figs 39-42 )

Hesione picta Müller, 1858: 213 , 214, pl. 6, fig. 3. — Fauvel 1953d: 7. — Hartman 1951: 35. — Jones 1962: 180. — Nonato & Luna 1970: 67, fig. 9. — Nonato & Amaral 1979: 49, fig. 100 (same fig. as in Nonato & Luna, 1970). — Fauchald 1977b: 16, 17. — Rullier & Amoureux 1979: 159. — Dueñas 1980: 86, pl. 5, figs A-D. — Ibarzábal 1988: 3, 4, fig. 2. — San Martín & Gómez Esteban 1992: 104.

Hesione proctochona Schmarda, 1861: 79 , 80, pl. 28, fig. 226, plus one unnumb. textfig. — Treadwell 1902: 184; 1928: 473; 1939: 217, 218, fig. 45A-B. — Hoagland 1919: 571 — Horst 1922: 200, 201. — Augener 1927b: 49; 1933b: 224.

Fallacia proctochona – Quatrefages 1866: 99. — Webster 1884: 311, pl. 8, fig. 21.

Hesione margaritae Hansen, 1882: 6 , pl. 1, figs 18-22.

Hesione vittigera Ehlers, 1887: 143-147 , pl. 41, figs 1-4. — Hartman 1938: 6.

Hesione splendida – da Costa 2013: 30-41, figs 1, 3-5 (non Savigny in Lamarck, 1818).

TYPE MATERIAL. — Brazil, Florianópolis. Neotype, ZMB 3815 , and one specimen labelled paraneotype, ZMB 3815 p, Grube Collection, Desterro (olim Santha Catharina Island , now Florianópolis), no further data, F. Müller coll. [paraneotype 36 mm long, 5 mm wide, slightly swollen medially, tapered towards both body ends, some parapodia previously removed and preserved as permanent slides; body grayish, without pigmentation in ethanol; prostomium as long as wide, subpentagonal, projected anteriorly into a round lobe; antennae digitate, 3 times as long as wide; eyes brownish, anterior eyes slightly larger than posterior ones; dorsal cirrophore 2-3 times as long as wide; cirrostyle basally smooth, cylindrical, annulated medially, articulated distally; neuraciculae black, tapered; acicular lobe double, upper tine digitate about 4 as times long as wide, lower tine rounded, bifid, about half as long as upper tine; neurochaetae about 20 per bundle, blades bidentate, subdistal tooth variable, usually smaller than distal one, guard approaching distal tooth, usually broken; posterior end tapered, pygidium granulose; anus open with 7 rounded anal papilla, three superior, smaller, and four laterals, larger].

Brazil, Rio de Janeiro. Four syntypes of Hesione margaritae Hansen, 1882 , RMNH 1275 , 1872 (fide de Bont 2008), no further data, E. van Beneden coll. [18-34 mm long, 3-6 mm wide; slightly distorted by pressing them in small container; many cirri lost, many chaetal blades broken, at least one parapodium removed per specimen; acicular lobe double, blunt, upper tine twice longer than lower one, better developed in larger specimens; neurochaetal blades bidentate, guard approaching distal tooth].

Gulf of Mexico, Florida. Four syntypes of Hesione vittigera Ehlers, 1887 , MCZ 837 , RV Blake, Sta. unnumb., Key West , 2-4 m depth, A. Agassiz coll. [15-26 mm long, 3-4 mm wide; macerated, most chaetal blades lost; body pale, largest syntype depressed, probably for observation; pharynx barely exposed; eyes almost colorless, anterior ones slightly larger than posterior ones; acicular lobe double, digitate, upper tine about twice longer than lower one but in smaller syntypes upper one 3-4 times longer than lower one; pygidium broken with 7 marginal, short digitate anal papillae] .

ADDITIONAL MATERIAL. — Brazil. 1 specimen, LACM 8581 , juvenile, Ilha do Mel, Paranagua Bay , Parana, rocks, 30 m depth, 15.VIII.1998, G. Rouse, F. Pleijel & A. Nygren coll. [11 mm long, 2 mm wide; pigmentation lost; eyes barely pigmented; acicular lobe double, only one tine visible over chaetal lobe margin] . — 2 specimens, MNHN-IA-PNT101a (formerly jar 888a), RV Calypso, Sta. 17 (03°48’35”S, 33°24’50”W), 52 m depth, 18.XI.1961 [20-22 mm long, 4 mm wide; macerated, colorless; neuraciculae tapered, largest one black, smallest one brownish; acicular lobe double, blunt, upper tine digitate, four times longer than lower one; neurochaetal blades about 20 per bundle, blades bidentate, distal tooth larger, guard approaching distal tooth]. — 1 specimen, MNHN-IA-PNT101b (formerly jar 888b), RV Calypso, Sta. 19 (03°49.7’S, 32°26.0’W), 31 m depth, sand, 18.XI.1961 [dried out, probably under coverslip]. — 1 specimen, NHMW unnumb., Desterro ( Florianópolis ), F. Müller coll., no further data (39 mm long, 6 mm wide; macerated, pigmentation lost, eyes colorless, after methyl-green staining anterior ones twice as large as posterior ones; neuraciculae blackish tapered; acicular lobe double, upper tine slightly longer; about 30 neurochaetae but handles damaged, distorted by dissolution or other chemical damage, no blades left; body depressed, progressively wider up to chaetiger 14; without chaetal features it cannot be proposed as a neotype) . — 1 specimen, ZMUC 2424, Rio de Janeiro, van Benyon coll., no further data [23 mm long, 3 mm wide; colorless, sligthly bent backwards, most cirri and neurochaetal blades lost; some parapodia previously removed; acicular lobe double, blunt, upper tine 2-3 times longer than lower one] .

Northwestern Atlantic. Virginia. 1 specimen, LACM 10163 , collected near Washington, D.C., 9.I.1938, O. Hartman coll. [35 mm long, 3.5 mm wide; splendid, perfectly relaxed specimen; eyes barely pigmented; chaetal lobes slightly invaginated; acicular lobe double, upper tine longer, tapered, lower tine blunt; neurochaetal blades medium-sized or short, bidentade, guard approaching distal tooth; Hartman indicated its pigmentation resembled the one given for H. vittigera Ehlers, 1887 ].

Gulf of Mexico. Florida, United States. 1 specimen, UF 685 , Palm Beach Co., Peanut Island (26.7, -80.03; 26°42’00.0000”N, 080°01’48.0000”W), fixed dead, but on ice, 0 -2 m depth, 4.III.2008, G.Paulay coll. [28 mm long, 4 mm wide; pigmentation visible as thick pale brown bands transversely separated by whitish lines into 6-8 narrow bands; posterior end regenerated, bent posteriorly; antennae minute; anterior eyes slightly larger and darker than posterior ones; most tentacular and dorsal cirri lost; dorsal cirri with cirrophores 3 times as long as wide; cirrostyle cylindrical basally, smooth (cells arranged into longitudinal lines); chaetal lobe slightly tapered; acicular lobe double, digitate, upper tine about twice longer than lower one; blades of chaetigers 3, 7, and 9 progressively smaller, teeth lateral; subdistal tooth smaller; guards mostly broken, some reaching distal tooth] . — 1 specimen, UF 825 , Tampa Bay , 3 km E of Sunshine Skyway Bridge (27.631, – 82.628; 27°37’51.6000”N, 082°37’40.8000”W), 8-9 m depth, 6.II.2009, G. Paulay coll. [18 mm long, 2 mm wide; acicular lobe double, blunt, upper one twice as long as lower one; pharynx not exposed; blades of chaetigers 3, 7, and 9 progressively smaller, teeth lateral; 4-8: 1 in chaetiger 3, 3-6: 1 in chaetiger 7 and 3-5: 1 in chaetiger 9]. — 1 specimen, UF 826 , Tampa Bay , 3 km E of Sunshine Skyway Bridge (27.631, – 82.628; 27°37’51.6000”N, 082°37’40.8000”W), 8-9 m depth, 6.II.2009, G. Paulay coll. [26 mm long, 4 mm wide; pigmentation fading out, darker along anterior chaetigers; posterior region distorted by dissection and muscular anterior enteron exposed; longest tentacular cirri reaching chaetiger 5; dorsal cirri as long as body width (excluding parapodia); blades of chaetigers 3, 7, and 9 progressively smaller, teeth lateral; 5-8: 1 in chaetiger 3, 3-5: 1 in chaetigers 7 and 9] . — 1 specimen, UF 1590 , Dade County, Biscayne Bay National Park, Elliott Key (25.4517, -80.1972; 25°27’06.1200”N, 080°11’49.9200”W), bay side off jetty, 30.IV.2010, F. Michonneau, G. Paulay, S. McPherson, M. Bemis, H. Lin, J. Moore & N. Evans coll. [34 mm long, 5 mm wide; slightly distorted, body wall dissected for molecular studies; gonads partially exposed; dorsal transverse brownish bands separated into 6-8 thinner bands by whitish lines; acicular lobe double, upper tine about twice as long as lower one; body] . — 1 specimen, UF 1591 , Monroe County, Florida Keys, Tennessee Reef (24.7651, -80.7542; 24°45’54.3600”N, 080°45’15.1200”W), 6 m depth, 3.V.2010, F. Michonneau, G. Paulay, S. McPherson, M. Bemis, H. Lin, J. Moore & N. Evans coll. (18 mm long, 2.5 mm wide; pharynx not exposed; pigmentation pattern still visible]. — 1 specimen, UF 1593 , Dade County , Biscayne Bay National Park , Elliott Key, bay side off jetty (25.4517, -80.1972; 25°27’06.1200”N, 080°11’49.9200”W), 30.IV.2010, F. Michonneau, G. Paulay, S. McPherson, M. Bemis, H. Lin, J. Moore & N. Evans coll. (32 mm long, 4 mm wide; pigmentation pattern visible; transverse white band complete only in chaetigers 2 and 16, chaetigers 3-6 with a middorsal spot, following ones with bands extended along the whole dorsal segmental surface, 6-8 per segment; body distorted by two lateral dissections to remove parapodia for molecular studies; one complete Harmothoe sp. in its stomach; blades of chaetigers 3, 7, and 9 progressively smaller, teeth lateral; 3-8: 1 in chaetiger 3, 3-5: 1 in chaetiger 7 and 3-4: 1 in chaetiger 9]. — 1 specimen, UF 1594 , Monroe County , Florida Keys, Long Key, LONF1 tower dive site, W of Florida Keys Marine Laboratory (24.843, -80.862; 24°50’34.8000”N, 080°51’43.2000”W), 2 m depth, 2.V.2010, F. Michonneau, G. Paulay, S. McPherson, M. Bemis, H. Lin, J. Moore & N. Evans coll. (28 mm long, 5 mm wide; body wall originally dissected for molecular studies; stomach and ovaries partially exposed (photos), now opened ventrally for study of digestive caeca (one copepod inside one of the caeca: given to E. Suárez); dorsal transverse brownish bands separated into 6-8 thinner bands by whitish lines; body; oocytes about 100 µm]. — 2 specimens, ECOSUR 2913, 800 m SSW off Alligator Reef Light, 5-7 m depth, 30.IV.1960, WA Starck, T. Starck & HA Feddern coll. [28-29 mm long, 4.0- 5.5 mm wide; larger specimen fusiform, not subcylindrical; no pigmentation; pharynx papilla as long as wide; dorsal cirri smooth; acicular lobe double, digitate, most with one lobe larger than the other]. — 1 specimen, LACM 10164 , Thornton Island , near Englewood, Florida, in crevices, 15.I.1938, O. Hartman coll. (22 mm long, 3 mm wide; juvenile, body posterior region laterally collapsed in vial; antennae minute].

Veracruz, México. 1 specimen, UANL 3988 , Bajo La Galleguilla , 4.XI.1992, J. A. de León-González coll. [without pigmentation; antennae minute; pharynx invaginated; dorsal cirri smooth basally; acicular lobes double, of about the same length; anal cirri low, blunt; body 29 mm long, 4 mm wide] . — 1 specimen, UANL 4119 , La Galleguita , 9.VIII.1999, J. A. de León-González coll. [29 mm long, 3 mm wide; prostomium posteriorly covered by tentacular segment, no pigmentation; antennae minute; pharynx invaginated nuchal organs connected middorsally, only the posterior depression visible; acicular lobe double, both truncate triangular, upper one 1.2 times longer than lower one] . — 1 specimen, UANL 4120 , Isla Sacrificios , 11.VIII.1999, J. A. de León-González coll. [25 mm long, 4 mm wide; macerated, no pigmentation, probably dead before fixation; prostomium fully exposed, nuchal organs connected middorsally; antennae minute; pharynx invaginated; most cirri and chaetal blades lost; acicular lobe double, both digitate, upper one longer than lower one] .

Campeche, México. 1 specimen, ECOSUR 2914 , Laguna de Términos , Sta. 27 A, July, 1984, E. Escobar coll. [40 mm long, 4.5 mm wide; no pigmentation; antennae minute; pharynx papilla not seen; dorsal cirri smooth basally; acicular lobes double, of about the same length; anal cirri low, blunt] . — 1 specimen, ECOSUR 2915 , Champotón , 1 km 5 of river mouth, 4 m depth, rock, 16.II.1999, J. R. Bastida & SISV coll. [20 mm long, 3 mm wide, slightly distorted; no antennae; pharynx everted; dorsal cirri multiarticulated; acicular lobes double, upper tine digitate, longer, lower tine blunt, shorter; pygidium granulose, slightly smashed, anal cirri blunt, short, barely seen] . — 1 specimen, UANL 127 , Isla Pérez, 7.VII.1977, U. Garza coll. [34 mm long, 4 mm wide; partly dehydrated, without pigmentation; dorsal cirri basally smooth; acicular lobe double, of about the same length; most cirri and chaetal blades lost; prepygidial segment granulose; anus projected with blunt cirri, 6 upper, shorter, two lower wider] .

Yucatán, México. 1 specimen, ECOSUR 2916 , Celestún, 1 m depth, 17.II.1991, SISV coll. [12.5 mm long, 2.5 mm wide; juvenile, pharynx invaginated; dorsal cirri basally smooth, medially and distally multiarticulated; acicular lobes double digitate, upper lobe twice as long as lower one] . — 1 specimen, ECOSUR OH-486, San Felipe , 500 m W off lagoon’s mouth, 2 m depth, 10.VI.2009, L. F. Carrera-Parra & SISV coll. [36 mm long, 4 mm wide; pharynx not exposed, some pigmentation left 7 years after collection, chaetal lobes slightly invaginated] . — 1 specimen, ECOSUR 2917 , Ría Lagartos , 1 km E of channel mouth, 2 m depth, rock, 18.II.1999, J. R. Bastida & SISV coll. [24 mm long, 3 mm wide, chaetigers; no antennae; dorsal cirri multiarticulated; acicular lobes double, one digitate longer, the other blunt, shorter; anal cirri not seen] .

Quintana Roo, México. 1 specimen, ECOSUR 2918 , Holbox, in seagrasses, 1 m depth, 4.V.2000, C. Campos coll. [17 mm long, 2.5 mm wide; pharynx everted, dorsal papilla slightly as long as wide; dorsal cirri multi-annulated; acicular lobes double, one digitate, longer, the other blunt, rounded; anal cirri not projected] . — 1 specimen, ECOSUR OH-468, Cabo Catoche , 800 m N off Lighthouse, 10.VI. 2009, 4 m depth, L. F. Carrera-Parra & SISV coll. [35 mm long, 4 mm wide; pigmentation still visible] .

Caribbean Sea. México. 1 specimen, ECOSUR OH-371, under coral rock, 2 m depth, 30.VIII.2004, L. F. Carrera-Parra coll. (21.5 mm long, 3 mm wide; no pigmentation; body incurved; eyes dark brown, anterior ones slightly larger than posterior ones; acicular lobes double, upper tine digitate, longer, lower one shorter, rounded to digitate].

West Indies. Undefined locality. 1 specimen, ZMUC 2430 , 1860, Marboc coll., no further data [38 mm long, 6 mm wide; body macerated, pharynx partially exposed; chaetal lobes invaginated; acicular lobe double, blunt, upper tine larger, 2-3 times larger than lower one].

Puerto Rico. 5 specimens, MCZ 46504, La Parguera, 25.I.1971, L. S. Roberts coll. (25-36 mm long, 3 mm wide; slightly dehydrated; prostomium projected anteriorly into a blunt, tapered lobe; eyes of similar size, anterior ones slightly more separated; pharynx not exposed; dorsal cirri basally smooth; acicular lobe double, upper tine digitate, lower tine rounded to triangular, ¼ to 2/3 as long as upper one; neurochaetal blades long and short, guard reaching distal tooth; pygidium granulose, with 4-7 short, thick digitate anal cirri]. — 6 specimens, USNM 15972 , RV Fish Hawk, Sta. unnumb., Port Real , 1898 [juveniles, 8-17 mm long, 1.5-2.0 mm wide; one with pharynx exposed; partly dehydrated, most chaetae broken]. — 1 specimen, ZMH-P 1295 , Ponce, Dr Bock coll., no further data [18 mm long, 3 mm wide; dorsum darker but without transverse dark bands; antennae digitate, tiny, twice as long as wide; eyes brownish, anterior ones slightly larger than posterior ones; acicular lobes double but chaetigers 14-16 with lower tine barely visible, blunt, digitate, upper tine often twice longer then lower one].

Haiti. 1 specimen, ZMH-P 1292b, Port au Prince , Kepperschmidt coll. No further data (42 mm long, 5 mm wide; bent laterally, without pigmentation; acicular lobe double, digitate, of similar size]. — 5 specimens, ZMH-P 2887, Port au Prince, A. Gagzo coll., no further data [18-21 mm long, 2.5-4.0 mm wide; colorless, two bent ventrally, two others with pharynx exposed, dorsal papilla rounded, as long as wide; body anterior eyes slightly larger or up to twice larger than posterior eyes; antennae minute, digitate, 3-4 times as long as wide, or about as long as posterior eyes diameter; longest tentacular cirri reaching chaetiger 7; dorsal cirri as long as body width (excluding parapodia); acicular lobe double, digitate, upper tine twice as long as lower one].

Jamaica. 5 specimens, BMNH 1912.2.2.18/22, Duerden coll., no further data [23-58 mm long, 4-8 mm wide; colorless, variably distorted; two large specimens with a ventral dissection through chaetigers 1-14 (or 16); antennae 3-4 times as long as wide; eyes almost colorless, anterior ones slightly larger than posterior ones; acicular lobe double; most chaetae without blades; pharynx exposed in one specimen, dorsal papilla slightly as long as wide] . — 1 specimen, ZMB 6313 , Kingston, Kükenthal & Hartmeyer [23 mm long, 4 mm wide; partially dehydrated, some cirri lost; pharynx slightly exposed; prostomium projected anteriorly; antennae minute, slightly as long as wide; eyes brownish, anterior ones slightly larger than posterior ones; dorsal cirri partially dehydrated; dorsal cirrophore twice as long as wide; neuraciculae blackish, tapered; acicular lobe double, blunt, upper tine slightly longer than lower one; about 20 neurochaetae per bundle; blades bidentate, guard approaching distal tooth] . — 1 specimen, ZMH-P 6752, Kingston, 4.VI.1905, O. Gagzo coll. [32 mm long, 3.5 mm wide; complete, bent dorsally, eyes colorless, anterior lenses twice as large as posterior ones; antennae minute, about twice as long as wide; acicular lobe double, digitate, blunt, lower tine slightly smaller than upper tine; most neurochaetal blades lost] . — 1 specimen, ZMH-P 9043, Kingston, Hartmeyer & Kükenthal coll., identified by Augener, no further data [complete, bent dorsally; dorsum brownish, lateral cushions pale; eyes brownish, of similar size; antennae minute, digitate, about twice as long as wide; acicular lobe double; many chaetae broken, many neurochaetal blades lost] .

Virgin Islands. 1 specimen, LACM LHarrisJul2000, Beef Island , Trellis Bay, coral rubble, 1-2 m depth, 12.VII.2000, G. Hendler, J. Martin, K. Fitzhugh & R. Ware coll. [36 mm long, 3 mm wide, splendid specimen; some neurochaetae without blades; body pale, laterally contracted; antennae not visible; anterior eyes slightly larger than posterior ones; anterior margin slightly projected, lateral margins rounded, posterior margins with a deep furrow, running close to the posterior eyes; dorsal cirri smooth basally, annulated medially, multiarticulated distally; chaetal lobe truncate; acicular lobe double, upper tine slightly longer than inferior one; ventral cirri surpassing chaetal lobe; neurochaetal blades medium-sized to short, bidentate, guards broken; pygidium granulose, anus projected, with six anal papillae, short, rounded]. — 1 specimen, UF 670 , St John Island , T-kai (18.339097, -64.676144; 18°20’20.7492”N, - 064°40’34.1184”W), 0.5-6.0 m depth, 30.XI.2002, V. Bonito coll. [18 mm long, 3 mm wide; a dark brown spot in posterior surfaces of parapodial bases from chaetiger 9, not visible in anterior chaetigers; acicular lobes double, blunt, upper tine twice as long as lower one].

Saint-Thomas. 2 specimens, ZMUC 2428, 1860, Krebs coll., no further data [15-34 mm long, 3.0- 4.5 mm wide; body damaged, colorless, most cirri and chaetal blades lost, larger specimen distorted, acicular lobe double, blunt, upper tine larger than lower one] . — 2 specimens, ZMUC 2429 , 1860, Krebs coll., no further data [26-31 mm long, 3-6 mm wide; larger one in better condition, smaller one macerated; colorless, without most cirri and chaetal blades, larger one distorted, acicular lobe double, blunt, upper tine 1-2 times longer than lower one]. — 5 specimens, ZMUC 2431 , coral, no further data [32-34 mm long, 3.5-4.0 mm wide; macerated, chaetal lobes invaginated, cirri and neurochaetal blades variably lost; acicular lobe double, upper tine 2-3 times longer than lower one]. — 3 specimens, ZMH-P 5457, C. Calwood coll., no further data [26-29 mm long, 3.0- 3.5 mm wide; partially dehydrated, colorless, one bent dorsally, another one laterally; acicular lobe double, upper tine and lower tine of similar size, or upper tine larger, up to twice as large as lower one].

Saint-Martin. 1 specimen, UF 2679 , Creole Rock (18.118, -63.056; 18°07’04.8000”N, 063°03’21.6000”W), 3 -10 m depth, 18.IV.2012, G. Paulay, J. Slapcinsky, M. Bernis, F. Michonneau, A. Anker, J. - P. Marechal coll. [36 mm long, 4 mm wide; pigmentation pattern clearly defined; mature female, oocytes released through body wall cut, about 100 µm in diameter each] .

Panama. 1 specimen, UF 1176 , Bocas del Toro, Isla Bastimentos, Salt Creek , 1-2 m depth, A. Anker coll. [35 mm long, 4 mm wide, 16 chaetigers; mature, bent ventrally, pigmentation pattern fading off, visible only along anterior 9 chaetigers; a dark brown spot in the basis of all parapodia; oocytes about 100 µm in diameter] . — 1 specimen, USNM 61643 , Galeta Reef , Colón, Acanthophora zone, 8.VIII.1972, A. Reimer coll. [24 mm long, 2 mm wide; body without pigmentation; distorted, laterally contracted, anterior prostomial margin truncate; anterior eyes slightly larger than posterior ones; acicular lobe double, blunt, upper tine longer; neurochaetal blades bidentate, guard approaching distal tooth] .

Colombia. 1 specimen, USNM 58277 , Caño de Loro, Cartagena , rocky bottom, subtidal, X.1977, P. R. Dueñas coll. [24 mm long, 4 mm wide; most cirri on site; several neurochaetal blades lost; no pigmentation; prostomium barely projected forward; antennae minute, ovoid; eyes almost without pigmentation, anterior ones larger than posterior ones; neurochaetal blades bidenate, guard approaching distal tooth; pygidium granulose; anal tube collapsed, only ventral papillae visible, rounded, slightly as long as wide].

Venezuela. 1 specimen, ECOSUR 2919 , E Margarita Island, Pilsbury Sta. 712 (11°09’N, 63°18’W), 26 m depth, 19.VII.1968 [32 mm long, 4 mm wide; no pigmentation; antennae tiny; pharynx papilla not visible; dorsal cirri multi-annulated; acicular lobes double, upper tine digitate, longer, lower tine blunt, shorter; anal cirri short, blunt, 5-6] .

Curaçao. 1 specimen, RMNH 1276 , 1920 , van der Horst coll., no further data [35 mm long, 5 mm wide; complete, slightly distorted by pressing it in small container; colorless, eyes barely pigmented, anterior eyes larger than posterior ones; antennae minute, smashed over prostomial surface; acicular lobe double, digitate] . — 3 specimens, ZMH-P 10343 , Spaansch Water, in Porites fucata , 7.IV.1920 (18.V.1920), van der Horst coll. [27-34 mm long, 4-5 mm wide; smaller one bent laterally, smaller and largest with pharynx exposed; acicular lobe double, digitate, upper tine up to twice as long as lower one; most neurochaetal blades lost] .

DISTRIBUTION. — Western Atlantic , from Virginia, United States to southern Brazil (Florianópolis), including the Gulf of Mexico and Caribbean Sea, in mixed bottoms, sea grasses, macroalgae, or rocks from the intertidal to 52 m depth; sometimes with other invertebrates as bryozoans, brittle-stars, and mussels (da Costa 2013: 41).

DIAGNOSIS. — Hesione with prostomium laterally curved; parapodia with dorsal cirri basally cylindrical, dorsal cirrophore twice as long as wide; larger acicula blackish; acicular lobe double, tines long, digitate, upper tine twice longer than lower one; neurochaetal blades bidentate, 4-5 times as long as wide; subdistal tooth smaller; guards approaching distal tooth.

DESCRIPTION

Neotype, ZMB 3815, complete, pharynx fully exposed, without pigmentation pattern ( Fig. 39A ) in ethanol; most tentacular or dorsal cirri on site; left parapodium of chaetiger 10 removed for observation (kept in vial), right parapodia of chaetigers 10 and 13 previously removed, preserved in separate permanent slide. Body bent laterally, becoming wider posteriorly (widest about chaetigers 12-13), 36mm long, 5 mm wide.

Prostomium as wide as long, anterior margin truncate, lateral margins rounded, posterior margin deeply cleft, about as long as ¼ prostomial length, reaching about posterior eyes, without longitudinal depression ( Fig. 39B ; specimen labelled paraneotype with pharynx invaginated, prostomium anteriorly projected, antennae positioned slightly ventrolaterally, appearing smaller than diameter of posterior eyes, Fig. 39E ). Antennae minute, digitate, directed laterally, twice as long as wide, as along as diameter of posterior eyes. Eyes brownish, anterior ones darker, twice larger than posterior ones.

Tentacular cirri long, variably twisted, longest ones reaching chaetiger 4 (right) or 6 (left) (reaching chaetiger 5 in syntypes of H. margaritae ). Lateral cushions slightly projected, surface entire in first three and last three chaetigers, other ones divided into anterior and posterior sections.

Parapodia with chaetal lobes as long as wide, truncate; dorsal cirri with cirrophores about twice as long as wide; cirrostyle smooth, cylindrical basally, annulated medially, articulated distally, as long as body width (without parapodia). Ventral cirri smooth, surpassing chaetal lobes. Ventral cirri smooth, surpassing chaetal lobe tip, as long as chaetal lobe width ( Figs 39D , 42F ).

Neuraciculae blackish, single, tapered. Acicular lobe double, digitate, upper tine twice longer than ventral one.

Neurochaetae about 30 per bundle, blades bidentate, blades at a certain angle from handle, decreasing in size ventrally, 4-5 times as long as wide, each with usually smaller subdistal tooth, guard approaching distal tooth.

Posterior region tapered into a blunt cone; pygidium subspherical, surface granulate, anus projected ( Fig. 39C ), with 7 round, blunt papillae, four lateral, three dorsal (5-7 in syntypes of H. margaritae ).

Pharynx fully exposed, with two muscular rings, anterior ring 1/3 as long as posterior one, with granulose margin; dorsal papilla rounded, twice as wide as long. Some oocytes within parapodia, about 100 µm in diameter.

Pigmentation

Body with transverse brown wide bands ( Figs 40 A-C, 41A). Each band with similar intensity throughout body, sometimes slightly darker along a wide middorsal region, espe- cially along posterior segments ( Figs 40C , 41B ). Dark bands present along body, interrupted dorsally along chaetal lobes by a wide pale band (1/2-1/3 as long as dark bands, sometimes poorly defined as in Fig. 40B ), missing in chaetiger 1, and in pygidium. Each band interrupted by 6-8 thin, pale transverse lines along a wide middorsal region, sometimes incomplete or fused to others, but dark bands continued over lateral cushions without being interrupted by pale transverse lines. This pigmentation pattern modified over anterior region as follow: prostomium with an inverted irregular pentagonal dark area, projected posteriorly beyond the level of posterior eyes, with paler areas ahead of anterior eyes, and over anterior middorsal prostomial surface, and projected laterally along peristomial dorsolateral surfaces ( Fig. 41E ). Tentacular segment and first chaetiger with several thin transverse pale lines but towards its anterior margin, lines becoming directed anteriorly, not middorsally connected. Eyes dark-brown to reddish, usually anterior eyes larger than posterior ones ( Fig. 40C, D ), but subequal also commonly found ( Fig. 41C ). In less than 10% of living specimens, pale transverse bands sometimes reduced to rounded middorsal spots (da Costa 2013: 39, fig. 5), or even disappearing completely. Pygidium with a wide band leaving a paler anal region ( Fig. 40A, E , 41D ).

REMARKS

Hesione picta Müller, 1858 , as indicated in the key below, is very similar to H. reticulata von Marenzeller, 1879 . There are two main differences between them regarding their integument and pigmentation. In H. picta the dorsum is smooth, annulated without longitudinal striae, and living specimens have transverse dark brown bands, whereas in H. reticulata the back is rugose because there are longitudinal striae, and living specimens have a complex, reticulate pigmentation pattern. Müller (1858) described H. picta from Santa Catharina Island (now Florianópolis), Brazil, but no specimens were deposited and no type specimen is listed in the museums where Grube used to deposit his material: Wroclaw, Poland (Viktor 1980), or Berlin, Germany ( Hartwich 1993). Hartman (1951: 35) regarded H. picta as a valid species and included two other species as junior synonyms: H. proctochona Schmarda, 1861 , and H. vittigera Ehlers, 1887 (she misspelled it as vittata ). Later, Hartman (1959: 185) expanded the list of synonyms by including H. praetexta Ehlers, 1887 , which was followed by Pleijel (1998: 159) who regarded it as a “possibly junior synonym of H. picta Müller, 1858 ”. This latter species is regarded below as distinct, but the two other names are confirmed as junior synonyms.

The original description by Müller (1858) included a figure for the prostomium, and the description is very short but contains enough detail including the distinctive pigmentation pattern. In contrast, the description by Schmarda (1861) for H. proctochona is much more detailed and richly illustrated.

It must be taken into account that there are several differences regarding the shape of prostomium, presence of antennae and about the position and size of the eyes in the species currently regarded as synonyms. Müller (1858) described H. picta from Southern Brazil; the single illustration shows a prostomium slightly as wide as long, depressions along the anterior and posterior prostomial margins, no antennae, and four large eyes, all of similar size, placed towards the anterior prostomial surface. As indicated above, H. margaritae Hansen, 1882 , has a transversely banded dorsum, and was described from Southern Brazil too, and Horst (1921: 80) regarded it as a junior synonym of H. picta ; however, the original illustration indicates a prostomium as long as wide, its posterior margin is entire, antennae are shown as tiny projections, and the four large eyes are placed over the central prostomial region. These differences in prostomial shape might be explained by variations in pharynx eversion, whereas the size of eyes is not as constant and specimens from the same locality and date (see characteristics of materials) have eyes of similar or different size. Further, because pigmentation pattern and eye pigments fade off quite soon once specimens are preserved in alcohol, although eye pigments last longer, and because the prostomium can be modified depending on the contraction state of the specimen, or the degree of pharyngeal eversion, prostomial shape and eye size are informative but barely relevant as diagnostic features. The most detailed illustration for H. picta was made by Nonato & Luna (1970: fig. 9), with specimens from Northern Brazil, and their specimen shows several differences: the prostomium is slightly as long as wide, and there are furrows over its anterior and posterior margins, there are two minute antennae, and the anterior eyes are twice as large as posterior ones, being placed towards posterior prostomial surface. Other Brazilian specimens, resembling H. picta , were recorded as H. splendida by Costa et al. (2008), and their illustration shows a as wide as long prostomium with anterior and posterior furrows, antennae minute, and anterior eyes larger than posterior ones, placed over the medial prostomial surface. These differences do not warrant specific status and consequently H. margaritae is herein confirmed as another junior synonym of H. picta .

On the other hand, H. vittigera was separated from other similarly pigmented species such as H. picta Müller, 1858 , H. proctochona Schmarda, 1861 , and H. margaritae Hansen, 1882 because it was regarded as having only 15 chaetigers. However, the largest syntype (and the three other ones), which was probably used for the description since it lacks the right parapodium of chaetiger 1, has 16 chaetigers as any other member of the genus. In the first chaetiger the left parapodium has no chaetae remaining, and on the right side only one chaeta remains (now the body wall is transparent and neuraciculae are visible). All other illustrated features (herein reproduced as Figure 42 ), match the features shown in H. picta and the other species described from the Eastern Atlantic, such as overall pigmentation pattern ( Fig. 42A ) prostomial features ( Fig. 42B ), parapodia including the acicular lobe ( Fig. 42C ), and the size and dentition of blades. Another difference that Ehlers (1887: 147) regarded as diagnostic was the presence and number of anal cirri, and he stated none was visible in his material; however, at least in the largest syntype, they are visible despite the poor state of the pygidial integument.

Augener (1934: 123) studied the type collection of Hansen. Augener could not find the type specimen for one species each from the families Amphinomidae , Aphroditidae , Chaetopteridae , Terebellidae and Hesionidae , including the type of H. margaritae . At least for the last species, it seems that it was overlooked because it was reidentified by Horst as H. proctochona ( Horst 1922: 200) , who also corrected the relative size, originally stated as up to 75 mm long, by indicating that it “likely is a mistake, for the largest specimen found by myself in the collection of van Beneden only has a length of 36 mm.” Consequently, the Leiden specimens must be the syntypes that Augener could not find, and have been regarded as such above.

The original description of H. vittigera completely matches the description of H. proctochona , including the intense lead-oxyde (reddish orange) color of dorsal cirrophores that both Schmarda and Ehlers observed in freshly collected or living specimens. On the contrary, the specimens Hansen (1882: 6) saw had been about 10 years in ethanol before he studied them, such that he did not see the dark brown transverse bands and instead found that: “la surface dorsale des anneaux transversalement striée avec un éclat brunâtre” (Transl.: the dorsal surface of the segments transversely streaked by a brownish burst).

Because of the prevalining confusion, the neotype and specimen labelled paraneotype have been designated for H. picta . In full agreement with the Code ( ICZN 1999: art. 75), this designation will clarify the taxonomic status of the species (art. 75.3.1), and the above description and remarks provide its diagnostic features together with a comparison with its junior synonyms (art. 75.3.2-3). A formal request for searching its type material was submitted to the curatorial staff in the museums of Berlin and Wroclaw, where most of Grube specimens were deposited, and none was found (art. 75.3.4). The neotype and pareneotype specimens were collected by Fritz Müller from the type locality (art. 75.3.5-6), and for selecting the neotype, the specimen with a round prostomial shape was preferred because it resembles what Müller illustrated (1858: pl. 6, fig. 3); these specimens might be the type material, but they were not designated as such.

Figures

FIG. 39. — Hesione picta Müller, 1858, neotype, ZMB 3815: A, anterior end, dorsal view; B, head, close up (arrow points to left antenna); C, posterior end, dorsal view; D, chaetiger 10, left parapodium, anterior view (insets: acicular lobe, and long and short neurochaetal blades); E, specimen ZMB 3815b, head, close-up (arrow points to left antenna). Scale bars: A, 1.5 mm; B, 0.3 mm; C, 1.7 mm; D, 0.7 mm; E, 0.2 mm.

FIG. 40. — Hesione picta Müller,1858, living specimens: A, Virgin Islands,LACM 10165,dorsal view;Brazil;B, Juvenile, LACM 8581,dorsal view, posterior region omitted; C, anterior end, dorsal view; D, head, dorsal view; E, posterior end, dorsal view. Scale bars: A, E, 1.3 mm; B, 0.8 mm; C, 2.4 mm; D, 1.7 mm (Photos: A, B, Leslie Harris; C-E, Vinicius da Rocha Miranda).

FIG. 41. — Hesione picta Müller, 1858, non-type specimen, ECOSUR OH 468: A, anterior region, dorsal view; B, posterior region, dorsal view; C, head, close-up, dorsal view; D, pygidium,close-up,dorsal view; E, anterior region,ventral view; F, chaetiger 8,left parapodium, anterior view (insets upper and lower neurochaetal blades). Scale bars: A, 1.5 mm; B, 2 mm; C, D, 0.6 mm; E, 1.1 mm; F, 0.7 mm.

FIG. 42. — Hesione picta Müller, 1858: A, dorsal view; B, anterior end, dorsal view; C, parapodium from a middle-body segment,anterior view; D, same, close-up of neurochaetal tip; guard tip broken (modif. after Ehlers 1887, scale bars missing in original).