Hesione panamena Chamberlin, 1919: 188-190 Treadwell 1937: 149 Hesione intertexta Hartman 1940: 212 Monro 1928: 79 1933: 26 Kudenov 1975: 78 Hesione genetta Fauvel 1941: 9 Hesione pantherina Fauvel 1941: 9 Revision of Hesione Savigny in Lamarck, 1818 (Annelida, Errantia, Hesionidae) Salazar-Vallejo, Sergio I. Zoosystema 2018 2018-06-19 40 12 227 325 3L3YL Chamberlin, 1919 Chamberlin 1919 [191,604,568,594] Polychaeta Hesionidae Hesione Animalia Phyllodocida 54 279 Annelida species panamena reinstated      Hesione panamena Chamberlin, 1919: 188-190, pl. 22, figs 9, 10. —  Treadwell 1937: 149.    Hesione intertexta–  Hartman 1940: 212, pl. 33, figs 30, 31 ( partim, nonfig. 30). —  Monro 1928: 79;  1933: 26. —  Kudenov 1975: 78( non Grube, 1878).    Hesione genetta–  Fauvel 1941: 9( non Grube, 1867).    Hesione pantherina–  Fauvel 1941: 9( non Risso, 1826).   TYPE MATERIAL. —  Eastern Tropical Pacific, Panama. Holotype, USNM 19379, RV Albatross, unnumb. Sta., Perico Island( 08°55’00”N, 79°31’00”W), shore,  12.III.1891. ADDITIONAL MATERIAL. — Eastern Tropical Pacific, Panamá.  2 specimens, BMNH1928.9.13.54/55, Taboga Island( 08°47’00”N, 79°33’00”W),  8-10 mdepth,  8.II.1916, T. Mortensencoll. [both complete, one lateraly bent; left parapodia of chaetigers 8 and 10 (one of each specimen) previously removed; body 28-29 mmlong, 3-4 mmwide; anterior eyes twice larger than posterior ones, darker in larger specimen; acicular lobe single, tapered, variably contracted; if fully extended, about 1/3 as large as neurochaetae]. —  1 specimen, BMNH1932.12.24.130, Balboa( 08°57’N, 79°34’W), St. George Pacific Expedition,  1923-1924, C. Crosslandcoll. [complete, splendid, body 46 mmlong, 5.5 mmwide; anterior eyes slightly larger than posterior ones; acicular lobe single, tapered, if fully extended, about 1/3 as large as neurochaetae; pharynx fully exposed, dorsal papilla round]. —  2 specimens, ZMUC2423, Taboga Island( 08°47’00”N, 79°33’00”W),  8-10 mdepth,  8.II.1916, T. Mortensencoll. [one complete 21 mmlong, 3 mmwide; the other reduced to fragments; colorless, acicular lobe single, tapered].  Western   Baja California Sur. 1 specimen, LACM85732, RV Velero IV, Sta. 2024 ( 27°48’33”N, 114°42’30”Wà 27°49’00”N, 114°42’09”W),  30 kmE off Punta Eugenia,  16 mdepth, rocks and sand,  18.IV.1951[ 35 mmlong, 4 mmm wide; upper acicular lobes tapered or digitate, 4-6 times longer than rounded lower ones]. —  1 specimen, LACM8556, Punta Abreojos( 26°42’N, 113°35’W), Knelpenshoal,  18 mdepth, in kelp holdfast,  2.III.1959, D. Millercoll. [ 21 mmlong, 5 mmwide; dehydrated, compressed, ventrally bent; pharynx exposed, prostomium collapsed; acicular lobe double, upper one tapered, very long, lower one rounded, short; two neuraciculae; neurochaetal blades bidentate, subdistal tooth small, guard approaching distal tooth].  Gulf of California.   Baja California. 1 specimen, LACM8565, RV VeleroIII, Sta.719 ( 30°53’00”N, 114°28’35”W), Consag Rock,  18-45 mdepth, basket stars,  24.III.1937[ 48 mmlong, 6 mmwide; laterally bent; antennae minute, as long as interocular distance; longest tentacular cirri reach chaetiger 5; upper acicular lobe long, tapered, lower one rounded, barely visible].   Sonora. 1 specimen, LACM8569, Punta Cholla, Puerto Peñasco,  9.V.1941, S. A. Glassellcoll. [ 32 mmlong, 4 mmwide; prostomium not collapsed, pharynx partially exposed; several parapodia previously removed; most dorsal cirri on site; upper acicular lobe contracted, digitate, 3-4 times longer than lower one, rounded; in chaetiger 10, left parapodium, upper lobe bifid into two similar sized ones, but placed at the same plane; other parapodia with lobes as indicated before]. —  1 specimen, LACM8567, Puerto Lobos,  16.IV.1965, P. Pickenscoll. [ 38 mmlong, 4 mmwide; partially dehydrated; upper acicular lobe tapered, 4-6 times longer than lower one rounded]. —  1 specimen, LACM8559, Playa Miramar, Guaymas,  10.II.1948, G. W. & N. MacGinitiecoll. [ 22 mmlong, 2.5 mmwide; prostomium with a longitudinal depression throughout its length; most dorsal cirri lost; acicular lobe with upper tine tapered, 4-5 times longer than lower one, rounded]. —  4 specimens, LACM8562, RV VeleroIII, Sta.1042 ( 28°43’30”N, 112°19’05”W), Turner’s Island, S of Tiburon Island, rocky shore,  24.VI.1940[macerated, probably very long; neuropodia with a long tapered acicular lobe].    Baja California Sur. Isla Tortuga. 1 specimen, LACM 8571, RV VeleroIII, Sta. 576 ( 27°25’30”N, 111°53’25”W),  38 mdepth, volcanic sand,  13.III.1936[ 27 mmlong, 2.5 mmwide; acicular lobe with upper tine tapered, 4-5 times longer than lower rounded ones].    Bahía Concepción. 2 specimens, ECOSUR 2186, Laguna Terminal,  1 mdepth, rocky bottom,  8.V.1981, R. Ríoscoll. [ 34-40 mmlong, 4 mmwide; smaller one with pharynx exposed, without dorsal papilla].—  3 specimens, LACM8574, RV VeleroIII, Sta.585 ( 26°43’25”N, 111°54’05”W), Coyote Bay,  3.6-7.2 mdepth, sand and kelp,  14.III.1936[ 37-38 mmlong, 4.5-5.0 mm wide; macerated; pharynx exposed in 1 specimen, margin integument ciliated, partially eroded; chaetal lobe invaginated in many parapodia; acicular lobe with upper tine long tapered, lower one inconspicuous]. —  1 specimen, LACM 8570, RV VeleroIII, Sta.688 ( 26°41’40”N, 111°51’05”W),  22 mdepth, mud and sand,  16.III.1937[body macerated; pharynx barely exposed; chaetal lobe invaginated in many parapodia; acicular lobe with upper tine tapered, 4-5 times longer than lower one]. —  1 specimen, LACM 8560, RV Velero IV, Sta.1771 ( 26°42’03”N, 111°53’34”Wto 26°42’17”N, 111°53’33”W),  13 mdepth, rocky bottom,  27.III. —  1 specimen, UANL 641,  18.VII.1985, J. A. de León-Gonzálezcoll. [ 5.5 mmlong, 1 mmwide; partially dehydrated; pigmentation lost; body cylindrical, pharynx everted, pharynx papilla covered by prostomial anterior margin; prostomium with posterior depression barely visible, eyes not seen; dorsal cirri multi-annulated basally; acicular lobe single, tapered].—  1 specimen, UANL642,  20.VII.1985, J. A. de León-Gonzálezcoll. [ 21 mmlong, 3 mmwide; poorly preserved; most integument detached from body wall; cirri macerated; body depressed, parallel sided pharynx invaginated; prostomium with posterior depression visible, eyes visible, brownish, very close to each other, anterior ones slightly more separated and larger than posterior ones; most chaetae lost; posterior end cirri lost].    Bahía de La Paz. 1 specimen, UANL128,  14.XI.1984, U. Garza coll. [ 21 mmlong, 3.5 mmwide; pharynx everted, dorsal papillae digitate, twice as long as wide; prostomium with posterior depression visible, eyes visible, brownish, very close to each other; dorsal cirri basally multiarticulated; acicular lobe double, upper tine digitate, lower lobe conical, blunt, 1/5 as long as upper one; posterior end cirri lost, anus in an 8-shaped area].—  5 specimens, LACM8566, RV  VeleroIII, Sta.503, off Lighthouse, coralline algae,  9 mdepth,  21.II.1936[ 19-25 mmlong, 2.0- 4.5 mmwide; largest specimen mature ( 25 mmlong, 4.5 mmwide), oocytes about 100 µm; acicular lobe double, upper tine tapered, sometimes basally contracted, 5-8 times longer than lower, rounded one; neurochaetal blades longer in smaller specimens (series of slides and photos in 3 specimensof different length and width)]. —  4 specimens, LACM8575, RV VeleroIII, Sta.1111 ( 24°21’55”N, 110°15’15”W), San Lorenzo Channel,  11 mdepth, sand, shell, coralline,  14.II.1940[ 29-41 mmlong, 3.5-5.0 mm wide; macerated; chaetal lobes invaginated; acicular lobe with upper tine tapered, 5-8 times longer than lower, rounded one].   Espíritu Santo Island. 2 specimens, USNM 35566, 24 mdepth,  30.III.1934, E. & C. Berkeleycoll. [damaged, most cirri and many chaetal blades lost, partially dehydrated, stiff, many parapodia with longer upper acicular lobe and smaller, rounded lower lobe; not measured to avoid further damage].  AD E B C  FIG. 33. —  Hesione panamena Chamberlin, 1919, holotype, USNM 19379: A, anterior end, dorsal view; B, close-up of head (arrow points to right lateral antenna); C, chaetiger 8, right parapodium (inset: close-up of acicular lobe, upper tine eroded); D, same, blade of neurochaeta (inset: close-up of another neurochaetal blade tip); E, posterior region, dorsal view. Scale bars: A, 1.3 mm; B, 0.3 mm; C, 70 µm; D, 30 µm; E, 1.7 mm.   San José Island. 2 specimens, MNHN-IA-PNT 91f(formerly jar 70), in different containers; one from the southern lagoon, among corals; the other without precise locality, 1904, L. Diguetcoll. [ specimenin better condition 24 mmlong, 3 mmwide; the other distorted, dried-out; acicular lobe single, can be seen in several parapodia despite their condition; acicular lobes single throughout body].   Ensenada de Muertos. 1 specimen, LACM 8564, RV VeleroIII, Sta.627 ( 23°58’55”N, 109°49’25”W),  9 mdepth, sand,  5.III.1937[ 27 mmlong, 2.5 mmwide; many cirri lost; body with circular adsorbed salts throughout the body; acicular lobe with upper tine tapered, long, sometimes digitate, lower one minute].  Gulf of California,   Baja California Sur(?). 1 specimen, MNHN-IA- PNT 91g(formerly jar 70), no locality, 1894, L. Diguetcoll. [ 13 mmlong, 2 mmwide; distorted, partially dehydrated, acicular lobes single, can be seen in several parapodia]. —  1 specimen, MNHN- IA-PNT91h (formerly jar 70b), no locality,  II.1895, L. Diguetcoll. [ 32 mmlong, 4 mmwide; distorted by compression; body almost without pigmentation, one middorsal pale spot between chaetigers 2 and 3].—  2 specimens, MNHN-IA-PNT  91i(formerly jar 70c), no locality,  II.1895, L. Diguetcoll. [ 31-36 mmlong, 5-7 mmwide; macerated; both with pharynx fully everted; acicular lobes single].  Western   Mexico. Sinaloa. Sixteenspecimens, LACM7101, Topolobampo( 25°36’N, 109°04’W),  26.I.1971, R. C. Bruscacoll. [ 17- 28 mmlong, 2-3 mmwide; upper acicular lobe tapered, 4-5 times longer than lower rounded one; 1 specimenmature, 25 mmlong, 3 mmwide, dissected for anatomical features]. — 1 specimen, UANLEMU 442, Estero de Urías, Mazatlán, intertidal, 1980, A. Rutgerscoll. [ 11.5 mmlong, 2 mmwide; bent ventrally; colorless; acicular lobe single, tapered].    Nayarit. 1 specimen, LACM8572, RV VeleroIII, Sta.745 ( 21°54’10”N, 105°53’05”W), Isabel Island,  18-32 mdepth, corallines,  2.IV.1937[ 37 mmlong, 4 mmwide; slightly macerated; acicular lobe rounded, upper tine contracted, 3-4 times longer than lower ones].  Guerrero.  1 specimen, ECOSUR OH-374, Hornos Beach, Acapulco, on oysters (  Spondylus calcifer),  4 mdepth,  19.IV.2008, L. F. Carrera-Parra& SISV coll. [ 23 mmlong, 3 mmwide; irregular, thin longitudinal lines still visible on tentacular segment only, rest of body colorless; pharynx not exposed; prostomial anterior margin depressed; anterior eyes twice as large as posterior ones; parapodia with acicular lobes single, tapered; posterior end smashed, integument smooth]. —  5 specimens, LACM 8555, RV Velero, Sta. 2596, Santa Lucíabay ( 16°50’33”N, 99°55’28”W), Acapulco,  2-8 mdepth, sponges, algae, rock, muddy sand bottoms,  1.II.1954[four in better condition, 14-35 mmlong, 2.5-5.0 mm wide; dorsal cirri longer than body width, including parapodia; one right parapodium (chaetigers 7-9) removed (kept in container); oocytes 100 µmin diameter].  FIG. 34. —  Hesione panamena Chamberlin, 1919, non-type specimens: A, dorsal view of living specimen, depressed by glass slide, ECOSUR OH 374, reddish hue due to blood vessels fracture after pressure and osmotic shock; B, same, anterior end, dorsal view; C, same, after some time in ethanol, pharynx barely ex- posed; D, chaetiger 6, right parapodium, anterior view (insets: blade tips); E, another specimen, LACM 8556, chaetiger 10, left parapodium, anterior view. Scale bars: A, 3 mm; B, 1 mm; C, 0.7 mm; D, 0.4 mm; E, 0.5 mm (photos: A, B, H. Bahenas).    Revillagigedo Islands, México. 1 specimen, LACM 134 34, RV VeleroIII, Sta.134 ( 18°20’35”N, 114°44’20”W),  25 mdepth, rock, nullipores,  5.I.1934[extremely distorted, body twisted over itself, posterior region compressed; 17 mmlong, 2 mmwide; acicular lobe with upper tine tapered, 4-5 times longer than rounded lower one]. —  1 specimen, LACM 8563, RV VeleroIII, Sta.289 ( 18°41’50”N, 110°57’20”W),  8-27 mdepth, sand and nullipores,  8.VI.1934[ 16 mmlong, 2.5 mmwide; mature, gonads visible through body wall from chaetiger 10, as homogeneous masses; acicular lobe digitate, upper tine 5-7 times longer than rounded lower one]. —  1 specimen, LACM 8561, RV VeleroIII, Sta.304 ( 18°20’25”N, 114°44’30”W),  36 mdepth, nullipores, algae,  11.VI.1934[ 26 mmlong, 3 mmwide; mature; body distorted, gonads exposed after body wall rupture; acicular lobe digitate, upper tine 4-5 times longer than rounded lower one; sperm spherical, about 5 µmin diameter].   Ecuador. 1 specimen, LACM 8583, RV VeleroIII, Sta.213 ( 01°15’25”S, 81°05’15”W), off La Plata Island,  13-18 mdepth, rock, nullipores,  10.II.1934[ 14 mmlong, 2 mmwide; macerated, body soft, semitransparent, including cirri; acicular lobe with upper tine very long, about 10 times longer than lower, rounded one]. —  1 specimen, LACM10150, RV VeleroIII, Sta.59 ( 01°14’31”S, 90°26’30”W), off Cormoranat Bay, Charles Island, Galápagos Islands,  23 mdepth,  6.II.1933[ 27 mmlong, 4 mmwide; macerated; chaetal lobes invaginated, acicular lobe with upper tine 4-5 times longer than lower, rounded one].—  1 specimen, LACM148-34, RV VeleroIII, Sta.148 ( 00°16’41”S, 91°22’39”W), Tagus Cove, Albermarle Island, Galápagos Islands,  22-45 mdepth,  13.I.1934[ 20 mmlong, 2.8 mmwide; macerated, with abundant adsorbed particles on body and chaetae; chaetal lobes invaginated, acicular lobe with upper tine 4-5 times longer than lower, rounded one].—  5 specimens, LACM10151, RV VeleroIII, Sta.167 ( 01°14’37”S, 90°28’08”W), Post Office Bay, Charles Island, Galápagos Islands,  27 mdepth, rocks,  19.I.1934[ 20-32 mmlong, 2-4 mmwide; macerated, most cirri and neurochaetal blades on site; chaetal lobes variably invaginated; right parapodium of chaetiger 9 in 3 specimensremoved (kept in vial); acicular lobe with upper tine 3-4 times longer than lower one; largest specimen mature; oocytes about 100 µm]. —  1 specimen, LACM10151, RV VeleroIII, Sta.336 ( 00°16’30”S, 90°35’20”W), Sullivan Bay, James Island, Galápagos Islands,  36 mdepth,  12.XII.1934[ 25 mmlong, 3 mmwide; macerated; pharynx fully everted; chaetal lobes slightly invaginated, acicular lobe with upper tine 4-5 times longer than lower, rounded one].—  1 specimen, LACM10161, RV VeleroIII, Sta.811 ( 00°51’35”S, 90°02’00”W), Barrington Island, Galápagos Islands, shore,  Pocillopora,  26.I.1938[ 30 mmlong, 4 mmwide; macerated; chaetal lobes invaginated; acicular lobe with upper tine tapered or digitate, 3-5 times longer than lower, rounded one].  DISTRIBUTION. — Easterntropical Pacific, From Punta Eugenia, Baja California Sur, Méxicoto the Galápagos Islands, from intertidal to  45 mdepth, in sandy or mixed bottoms.  DIAGNOSIS. —  Hesionewith prostomium slightly curved laterally; parapodia with dorsal cirri basally cylindrical, dorsal cirrophore twice as long as wide; larger acicula blackish; acicular lobe single, long, tapered, lower tine rounded, small; neurochaetal blades bidentate, 3-8 times as long as wide; subdistal tooth smaller than distal one, with guards approaching distal tooth.  DESCRIPTION  Holotype, USNM 19379, complete, tapered, colorless in ethanol, slightly macerated with several dorsal cirri and ventral cirri damaged or lost; chaetae from chaetiger 1 lost; body integument finely annulated dorsally, annulations not continued into lateral cushions, especially behind chaetal lobes ( Fig. 33A). Body 39 mmlong, 5 mmwide. Prostomium as long as wide, anteriorly narrower, expanded towards posterior margin; anterior margin projected, posterior margin with a small furrow ( Fig. 33B). Antennae minute, blunt, slightly as long as wide. Eyes barely pigmented, anterior ones placed by the middle of the prostomium, slightly larger and farther apart than posterior ones, displaced posteriorly. Tentacular cirri damaged, longest ones reaching chaetiger 4. Lateral cushions projected, corrugated in anterior region, divided into two sections in posterior region. Parapodia with dorsal cirri damaged; cirrophore annulated, about twice as long as wide; cirrostyle macerated, looking smooth. Ventral cirri with cirrophores small, indistinct; cirrostyle longer than chaetal lobe. Chaetal lobe thick, with parallel sides, slightly tapered, annulated basally ( Figs 33C, 34D, E). Neuraciculae blackish, two, one markedly thicker than the other. Acicular lobe single, tapered, rarely directed upwards, mostly projected laterally, sometimes with a round basal projection, upper tine (eroded) 3 times longer than lower one (basal tine not visible in other specimens, Fig. 34E). Neurochaetae about 16 neurochaetae per bundle, blades bidentate, slightly shorter ventrally, 8 times as long as wide, with a small subdistal tooth, guard usually passing it ( Fig. 33D) (other specimens with blades 3-7 as long as wide, Fig. 34D). Pygidium with integument smooth; anus projected as a short, distally expanded cylinder ( Fig. 33E); about 8 short, digitate anal papillae. Pharynx not exposed. Oocytes not seen.  Pigmentation Body with longitudinal, dorsal, irregular, discontinuous thin dark green or pale brown bands ( Fig. 34A), restricted to first 2-3 anterior chaetigers, alternating with small, round spots, and completely replaced by them in medial and posterior chaetigers, expanded into lateral cushions, but not into pygidium.Tentacular and dorsal cirri colorless; cirrophores pale in anterior chaetigers, yellowish in posterior ones; chaetal lobes pale. Prostomium with thin irregular marginal lines. Longitudinal bands progressively thinner ( Fig. 34B); all chaetigers with an irregular middorsal spot, irregularly ovoid, as long as wide, along chaetigers 1-6, becoming as long as wide in following chaetigers, connected by a thin, darker middorsal line continued to the last achaetous segment. This thin middorsal line bordered by paler areas, short in anterior and medial segments, becoming longer, running throughout all dorsal segmental surface in posterior segments. Pigmentation remaining visible after 8 years in ethanol ( Fig. 34C).  REMARKS   Hesione panamena Chamberlin, 1919, reinstated, resembles  H. pantherina Risso, 1826in the key below. Their main difference is the typeof acicular lobe. In  H. panamenaacicular lobes are long, tapered, whereas in  H. pantherinathey can be short or long but their tips are blunt to distally swollen. Living specimens also differ because in  H. panamenathere are middorsal blackish, round spots whereas in  H. pantherinathere are middorsal pale areas.   Hesione panamenawas originally spelled as such for the description ( Chamberlin 1919: 188) and  H. panamicafor the legends to figures, and although both epithets are correct, the former has been followed by subsequent authors. Chamberlin (1919: 189)indicated that antennae were broken off, but they are present in the type, although because their bases are not exposed, they are difficult to observe. The details of the neurochaetal blades, as indicated in the original illustrations (his plate 22, figures 9-10) are accurate by showing long blades, with a small subdistal tooth, and its guard usually passing subapical tooth.   Hesione panamenaresembles  H. intertexta Grube, 1878by the small size of the eyes and antennae, and they also have similar neurochaetal blades. However, in  H. panamenathe blade length/width proportions are smaller and the antennae are ovoid, whereas in  H. intertextathere is a wider variation in blade size, and antennae are tapered. Further, although pigmentation fades quite soon in ethanol, their patterns show some differences; in  H. panamena, segmental longitudinal lines are discontinuous whereas they are continuous in  H. intertexta,and in  H. panamena, there is a blackish, as long as wide middorsal spot just before a paler area, and this is not so evident in  H. intertexta. Further, in  H. intertextaacicular lobe is single, without any lower one, whereas in  H. panamenathere can be a smaller, rounded lower tine.  Fauvel (1941: 9)recorded two  Hesionespecies for the Gulfof California:  H. pantherina Risso, 1826(this record is herein regarded as belonging to  H. panamena), and  H. genetta Grube, 1867because of its pigmentation pattern: “à la face dorsale des traces de mouchetures brunes, ovales, alternées, qui rappellent la robe de la Genette.” (Transl.: on the dorsal surface there are remains of brown oval, alternating spots, resembling the genette skin). This specimen was found in the Paris museum, it differs from  H. hartmanae n. sp.described above, and initially regarded as having potential affinities because acicular lobes are double in  H. genettaand  H. hartmanae n. sp., but the Paris specimen has acicular lobes single. Fauvel’s specimen had been in ethanol for about 50 years before he saw it, and he referred to a pigmentation pattern, but after other half a century spent before I studied the specimen, this pigmentation is now completely faded out. This specimen is also regarded as conspecific with  H. panamena.  Monro (1926: 312)made a comparison of the size of guards and separated  Hesionespecies into three groups; among those having guards approaching the apical tooth he listed  H intertexta,  H. reticulata von Marenzeller, 1879,  H. praetextaEhlers, 1885and  H. panamena Chamberlin, 1919. Further, in the previous page, Monro rejected, following Fauvel (1923a), the use of pigmentation pattern and the development of acicular lobes. As a consequence, in two of his publications on Panamian polychaetes ( Monro 1928: 79; 1933: 26) he regarded  H. panamenaas a junior synonym of  H. intertexta, and recorded it for the Pacific coast of Panama. Hartman (1940: 212)followed this synonymy and listed specimens from Western Mexico to the Galápagos Islands, but her material includes two species: one with a very long upper acicular lobe, herein retained as  H. panamena, and another one with two acicular lobes, which were illustrated, and were based on Galápagos specimens, which are regarded as belonging to an undescribed species and described above as  H. hartmanae n. sp.The record of  Hesione pantherina Risso, 1826for the Gulf of California ( Fauvel 1941: 9) is regarded as belonging to  H. panamena.Consequently, there are two species in the Galapagos ( Blake 1991: 78) but their names should be changed to  H. panamenaand  H. hartmanae n. sp.(see above). 2595817461 1891-03-12 RV Albatross & Sta. Panama 8.916667 Eastern Tropical Pacific 21 -79.51667 Perico Island 54 279 1 holotype 2595817436 1916-02-08 BMNH T. Mortensen 9 8.783334 Taboga Island 21 -79.55 54 279 2 2595817484 1923 1924 1923 BMNH C. Crossland 8.95 Balboa 1300 -79.566666 St. George Pacific Expedition 54 279 1 2595817476 1916-02-08 ZMUC T. Mortensen 9 8.783334 Taboga Island 21 -79.55 54 279 2 2595817441 1951-04-18 LACM RV Velero IV & Punta Eugenia 16 27.816668 Baja California Sur 20 -114.7025 54 279 1 Baja California Sur 2595817434 1959-03-02 LACM Knelpen & D. Miller 18 26.7 Punta Abreojos 1240 -113.583336 54 279 1 Baja California Sur 2595817409 1937-03-24 LACM RV Velero & Sta. & Consag Rock 32 30.883333 Baja California 20 -114.47639 54 279 1 Baja California 2595817426 1941-05-09 LACM S. A. Glassell Punta Cholla Puerto Penasco 54 279 1 2595817466 1965-04-16 LACM P. Pickens Puerto Lobos 54 279 1 2595817381 1948-02-10 LACM G. W. & N. MacGinitie Playa Miramar Guaymas 54 279 1 2595817419 1940-06-24 LACM RV Velero & Sta. Isla Tortuga 28.725 Turner's Island 20 -112.318054 54 279 4 2595817389 1936-03-13 Baja California Sur. Isla Tortuga 38 28.725 Isla Tortuga 20 -112.318054 54 279 1 2595817473 1981-05-08 ECOSUR R. Rios 1 Laguna Terminal Bahia Concepcion 54 279 ECOSUR 2186 2 2595817365 1936-03-14 LACM RV Velero & Sta. 5 26.723612 Coyote Bay 20 -111.90139 54 279 4 2595817425 1937-03-16 LACM RV Velero 26.694443 Sta. 20 -111.85139 Sta. 54 279 LACM 8570 23 22 2595817377 27. III LACM RV Velero IV & Sta. 13 26.704723 20 -111.892494 54 279 LACM 8560 1 2595817459 1985-07-18 UANL J. A. de Leon-Gonzalez 54 279 UANL 641 1 2595817369 1985-07-20 UANL J. A. de Leon-Gonzalez Bahia de La Paz 54 279 1 2595817361 1984-11-14 UANL J. A. de Leon-Gonzalez Bahia de La Paz 54 279 1 2595817452 1936-02-21 LACM Velero & Sta. 9 Lighthouse 54 279 8 2595817373 1940-02-14 LACM RV Velero & Sta. 11 24.365278 San Lorenzo Channel 21 -110.254166 54 279 4 2595817411 1934-03-30 USNM E. & C. Berkeley 55 280 Island Island 54 279 USNM 35566 26 24 2595817416 L. Diguet Island Island 55 280 94 91 2595817440 1937-03-05 LACM RV Velero & Sta. 9 23.981945 21 -109.82361 55 280 LACM 8564 1 2595817397 L. Diguet Baja California Sur 55 280 1 Baja California Sur 2595817340 1895-02 L. Diguet 55 280 1 2595817437 1895-02 1991-01 1895-02 L. Diguet 55 280 2 2595817400 1971-01-26 LACM, UANL R. C. Brusca & Mazatlan & A. Rutgers Mexico Sixteen 25.6 Estero de Urias 1245 -109.066666 Topolobampo 55 280 2 Sinaloa 2595817401 1937-04-02 LACM RV Velero & Sta. 25 21.902777 Isabel Island 21 -105.88472 55 280 1 Nayarit 2595817354 2008-04-19 L. F. Carrera-Parra 4 Hornos Beach Acapulco 55 280 1 2595817339 1954-02-01 RV Velero Saint Lucia 5 56 281 16.8425 Acapulco 21 -99.92444 55 280 5 2595817485 1934-01-05 RV Velero & Sta. Mexico 25 18.343056 Revillagigedo Islands 21 -114.738884 56 281 1 Revillagigedo Islands 2595817380 1934-06-08 LACM RV Velero & Sta. 18 18.697222 21 -110.95555 56 281 LACM 8563 1 2595817391 1934-06-11 LACM RV Velero 36 18.340279 Sta. 21 -114.74166 Sta. 56 281 LACM 8561 1 2595817382 1934-02-10 RV Velero & Sta. Ecuador 16 -1.2569444 La Plata Island 21 -81.0875 56 281 1 2595817344 1933-02-06 LACM RV Velero & Sta. Ecuador 23 -1.2419444 Charles Island 21 -90.441666 Cormoranat Bay 56 281 1 Galapagos Islands 2595817417 1934-01-13 LACM RV Velero & Sta. Ecuador 34 -0.27805558 Albermarle Island 21 -91.3775 Tagus Cove 56 281 1 Galapagos Islands 2595817468 1934-01-19 LACM RV Velero & Sta. Ecuador 27 -1.2436111 Charles Island 21 -90.46889 Post Office Bay 56 281 8 Galapagos Islands 2595817446 1934-12-12 LACM RV Velero & Sta. Ecuador 36 -0.275 James Island 21 -90.58889 Sullivan Bay 57 282 1 Galapagos Islands 2595817394 1938-01-26 LACM RV Velero & Sta. Ecuador -0.85972226 Barrington Island 21 -90.03333 57 282 1 Galapagos Islands