Thalassas 25

Thalassas, 26 (2): 47-73 An International Journal of Marine Sciences

REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

CRISTIAN ALDEA(1,2) & JESÚS S. TRONCOSO(1)

Key words: Trophon, distribution, bathymetry, morphology, Antarctica, South-America, Sub-Antarctic islands.

ABSTRACT of this genus, no summarizing data from these areas are known and some records are confused by using of Among the several groups of molluscs in the combining genera/subgenera (i.e. Coronium Simone, southern hemisphere, the genus Trophon Montfort, 1996, Pagodula Monterosato, 1884, Nodulotrophon 1810 has a particular importance, because it is a highly Habe & Ito, 1965 and Fuegotrophon Powell, 1951). In diversified taxon in the Southern Ocean and Sub- this work we gathered data of the distribution, shell Antarctic waters. In the middle of the XX century, 27 morphology and taxonomic remarks of 46 species species were known, which increased to 33 species at of Trophon (s.l.) starting from a performed database the beginning of the XXI century, but more than 100 with all records toward the pole from about 20ºS species were described under this genus along the time, in South-American waters, and from about 45ºS in most of them being synonyms or belonging to other the Eastern Atlantic, Indian and Western Pacific genera at the moment. Despite the great diversification Oceans. Seventeen species were found inhabiting exclusively in South-America, three in Antarctica, five in Western Sub-Antarctic waters, and five in Eastern Sub-Antarctic waters; 16 species presented

(1) Departamento de Ecología y Biología Animal, a wide range of distribution. Bathymetrically, 22 Facultad de Ciencias del Mar. species were exclusive of the continental shelf, 23 Campus Lagoas Marcosende, 36310. reached the slope and one the deep-sea (>3,000 m). Universidad de Vigo, ESPAÑA. E-mail: [email protected] (C. Aldea), [email protected] From the shell morphology, five main morphotypes were recognized. Affinities between species and with (2) Fundación Centro de Estudios del Cuaternario de Fuego- Patagonia y Antártica (CEQUA), combining genera were discussed from the obtained Avenida Bulnes 01890, Punta Arenas, CHILE. data.

47 CRISTIAN ALDEA & JESÚS S. TRONCOSO

Figure 1: Distribution of species of Trophon (s.l.) in South-America: T. acanthodes (ac), T. amettei (am), T. bahamondei (ba), T. columbarioides (cb), T. ceciliae (ce), T. clenchi (cl), T. condei (co), T. declinans (de), T. fasciolarioides (fa), T. geversianus (ge), T. iarae (ia), “T.” malvinarum (ma), T. minutus (mi), T. ohlini (oh), T. pallidus (pa), T. pelseneeri (pe), T. plicatus (pl), T. parodizi (pr), T. patagonicus (pt), T. triacanthus (tr), T. vangoethemi (va) and T. veronicae (ve).

48 Fig.�1/10 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Figure 2: Distribution of species of Trophon (s.l.) around Antarctica: T. coulmanensis coulmanensis (cc), T. coulmanensis multilamellatus (cm), T. cuspidarioides (cu), T. drygalskii (dr), T. echinolamellatus (ec), T. enderbyensis (en), T. leptocharteres (le), T. longstaffi (lo), T. minutus (mi), T. nucelliformis (nu), T. poirieria (po), T. shackletoni paucilamellatus (sp), T. shackletoni shackletoni (ss) and T. scotianus (st).

INTRODUCTION the morphologic pattern with all fusiform and/or lamellate –usually with spiral ornamentation– The genus Trophon Montfort, 1810 represents muricids (Pastorino and Scarabino, 2008) are particular importance among the several genera highlighted points of shell morphology in the of molluscs in the southern hemisphere since it identity of this genus. has a high diversification, mainly in the Southern Ocean and adjacent waters. The austral origin, From the proposal of the genus for Buccinum being the older species traced as far back as the geversianus Pallas, 1774, numerous records with Oligocene (Griffin and Pastorino, 2005), and taxonomic remarks and new species from research

Fig.�2/10 49 CRISTIAN ALDEA & JESÚS S. TRONCOSO

Figure 3: Distribution of species of Trophon (s.l.) in western Sub-Antarctica: T. acanthodes (ac), T. albolabratus (al), T. arnaudi (ar), T. brevispira (br), T. coulmanensis coulmanensis (cc), T. cribellum (cr), T. cuspidarioides (cu), T. distantelamellatus (di), T. drygalskii (dr), T. echinolamellatus (ec), T. emilyae (em), T. leptocharteres (le), T. longstaffi (lo), T. minutus (mi), T. nucelliformis (nu), T. shackletoni paucilamellatus (sp), T. shackletoni shackletoni (ss) and T. scotianus (st). expeditions were published (e.g. Rochebrune and species have been noticed (e.g. McLean and Mabille, 1889; Strebel, 1908; Powell, 1951; Powell, Andrade, 1982; Dell, 1990; Numanami, 1996; 1958), giving a first account of 27 species by Houart, 1997), considering morphological aspects, Powell (1960). Later on, new records and new but regardless of their real affinities.

50 Fig.�3/10 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Figure 4: Distribution of species of Trophon (s.l.) in eastern Sub-Antarctica: T. albolabratus (al), T. coulmanensis coulmanensis (cc), T. declinans (de), T. eversoni (ev), T. geversianus (ge), T. macquariensis (mc), T. minutus (mi), T. mawsoni (mw), T. pallidus (pa), T. scolopax (sc), T. septus (se) and T. shackletoni shackletoni (ss)..

Then Pastorino (2002a) revised the species from more than 100 proposed species names morphological patterns of the systematics under the genus for South America and Antarctica, and phylogeny of the genus in Patagonia and many of which are now junior synonyms or belong Antarctica, pointing out the existence of 33 to other genera (i.e. Xymenopsis Powell, 1951). The

Fig.�4/10 51 CRISTIAN ALDEA & JESÚS S. TRONCOSO

Figure 5: Bathymetry of species of Trophon (s.l.). Three bathymetric boundaries were marked at 400 m, 800 m and 3,000 m; in brackets the depth range of the species; species with large distribution with black bars. T. acanthodes (ac), T. albolabratus (al), T. amettei (am), T. arnaudi (ar), T. bahamondei (ba), T. brevispira (br), T. columbarioides (cb), T. coulmanensis coulmanensis (cc), T. ceciliae (ce), T. clenchi (cl), T. coulmanensis multilamellatus (cm), T. condei (co), T. cribellum (cr), T. cuspidarioides (cu), T. declinans (de), T. distantelamellatus (di), T. drygalskii (dr), T. echinolamellatus (ec), T. emilyae (em), T. enderbyensis (en), T. eversoni (ev), T. fasciolarioides (fa), T. geversianus (ge), T. iarae (ia), T. leptocharteres (le), T. longstaffi (lo), “T.” malvinarum (ma), T. macquariensis (mc), T. minutus (mi), T. mawsoni (mw), T. nucelliformis (nu), T. ohlini (oh), T. pallidus (pa), T. pelseneeri (pe), T. plicatus (pl), T. poirieria (po), T. parodizi (pr), T. patagonicus (pt), T. scolopax (sc), T. septus (se), T. shackletoni paucilamellatus (sp), T. shackletoni shackletoni (ss), T. scotianus (st), T. triacanthus (tr), T. vangoethemi (va) and T. veronicae (ve).

same author (Pastorino, 2005) developed a review The last works known on the genus are the review of all living species from both coasts of southern of the last described species of Trophoninae from South America, involving those taxa living in Chilean waters (Houart and Sellanes, 2006) and environments associated with the continental the description of two species from Argentinean shelf, but not considering deeper water species. deep-sea by Pastorino and Scarabino (2008).

Fig.�5/10 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Despite the extensive information existing at RESULTS the present time, no summarizing data are available mainly in the Antarctic and East Sub-Antarctic areas Distribution and bathymetry and some records are confused by using combining genera/subgenera (i.e. Coronium Simone 1996, Forty-six species of the genus Trophon (s.l.) were Pagodula Monterosato, 1884 and Fuegotrophon found (Table 1) inhabiting in the study area. Twenty- Powell, 1951). Therefore, the aim of this paper is two species were distributed in South America (Fig. to gather the scattered literature of all Trophon 1), of which 17 species presented an exclusive South- (s.l.) species in the Southern Ocean and adjacent American distribution (T. amettei, T. bahamondei, T. areas, standing out the morphology of the shell and ceciliae, T. clenchi, T. columbarioides, T. condei, T. geographical, bathymetrical and some systematics fasciolarioides, T. iarae, “T.” malvinarum, T. ohlini, aspects to help to the identification of the species. T. parodizi, T. patagonicus, T. pelseneeri, T. plicatus, T. triacanthus, T. vangoethemi and T. veronicae). MATERIALS AND METHODS Fourteen species were found in Antarctic waters (Fig. 2), from which three are exclusive from there (T. A complete database with about 700 records coulmanensis multilamellatus, T. enderbyensis and T. of Trophon (s.l.) and related genus (Coronium and poirieria). Eighteen species are from western Sub- Pagodula) of species distributed in the Southern Antarctic islands and waters (Fig. 3), five of which Ocean and adjacent areas was performed. Concretely, being endemic from there (T. arnaudi, T. brevispira, those records toward the pole from about 20ºS in the T. cribellum, T. distantelamellatus and T. emilyae). western Atlantic Ocean and eastern Pacific Ocean Twelve species are from eastern Sub-Antarctic waters –South America–, and from about 45ºS in the eastern (Fig. 4), five from which being endemic from there (T. Atlantic, Indian and western Pacific Oceans were eversoni, T. macquariensis, T. mawsoni, T. scolopax considered. The database included the geographic and T. septus). location, bathymetry, and some taxonomic remarks of all records of the species of the genus found in Sixteen species have a distribution that exceeded the the literature and in the National Collection of the limits established here: T. acanthodes, T. geversianus Smithsonian National Museum of Natural History, and T. pallidus have a large South-American USNM, available in Internet (http://invertebrates. distribution (Fig. 1), but the first reaches western si.edu/, accessed in March, 15th, 2009). Sub-Antarctic waters (Fig. 3) while T. geversianus and T. pallidus were found in eastern Sub-Antarctic Data were processed geographically and islands (Fig. 4); T. coulmanensis coulmanensis, T. bathymetrically to characterize the species by their drygalskii, T. longstaffi, T. minutus, T. scotianus and distributions. The bathymetrical boundaries were T. shackletoni shackletoni have a circum-Antarctic established following Aldea et al. (2008). For each distribution (Fig. 2) that reaches the western Sub- species, taxonomic remarks and morphological Antarctic waters (Fig. 3), while T. minutus also reaches information of the shell were processed, and images South-America (Fig. 1) and eastern Sub-Antarctic or draws of representative specimens were redrawn to waters (Fig. 4) and T. coulmanensis coulmanensis point out the main conchological features and to split and T. shackletoni shackletoni also reach eastern the species in morphological groups. A dichotomic Sub-Antarctic waters (Fig. 4); T. cuspidarioides, T. key for all species was performed by means of the echinolamellatus, T. leptocharteres, T. nucelliformis division of the groups and using the terminology and T. shackletoni paucilamellatus are species shared given in the original descriptions of the species or between the West Antarctica (Fig. 2) and western subsequent revisions for the sculpture. Sub-Antarctic waters (Fig. 3); T. albolabratus and T.

53 CRISTIAN ALDEA & JESÚS S. TRONCOSO

rr rr wc rl l rr l lc c

B D A C

l F bc gs l c lc lc l

G I E

Figure 6: Redraws of Trophon (s.l.) species: A, T. parodizi, paratype from Pastorino (2005), 21.1 x 10.6 mm; B, T. minutus, holotype from Melvill and Standen (1907), 7.3 x 4.0 mm; C, T. pallidus, specimen from Strebel (1908), 27.1 x 12.9 mm; D, T. condei, holotype from Houart (2003), 61.4 x 27.7 mm; E, T. macquariensis, USNM 886110, 19.6 x 9.3 mm; F, T. declinans, from Watson (1886), 20.3 x 8.1 mm; G, T. emilyae, holotype from Pastorino (2002b), 12.1 x 3.2 mm; H, T. cuspidarioides, holotype from Powell (1951), 13.0 x 5.7 mm; I, T. fasciolarioides, holotype from Pastorino and Scarabino (2008), 11.2 x 4.8 mm; bc, blunt cord; c, cord; gs, growth stria; l, lamella; lc, low cord; rr, rounded ridge; rl, rolled lamella; wc, weak cord.

declinans are species from eastern Sub-Antarctic species the lower slope (above 800 and 3,000 m waters (Fig. 4), being the first found in the western depth, respectively), and one species (T. shackletoni Sub-Antarctic islands (Fig. 3) as well and the second shackletoni) reached the deep-sea (below 3,000 m in the waters of South-America (Fig. 1). depth). Fig.�6/10 In terms of vertical distribution (Fig. 5), 22 species External morphology and dichotomic key had a distribution above 400 m depth, being been able to consider them as exclusive of the continental Five main morphotypes were recognized: (a) shelf, 10 species reached the upper slope zone and 13 fusiform shells of rounded shape with rounded –or

54 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

l os c os os l l os l

D B A

wc os l l l c t

E F G

Figure 7: Redraws of Trophon (s.l.) species: A, T. acanthodes, from Watson (1886), 38.1 x 16.3 mm; B, T. columbarioides, holotype from Pastorino and Scarabino (2008), 14.7 x 8.4 mm; C, T. septus, from Watson (1886), 23.1 x 10.9 mm; D, T. vangoethemi, holotype from Houart (2003), 16.9 x 7.9 mm; E, T. veronicae, holotype from Pastorino (1999), 52.2 x 19.6 mm; F, T. arnaudi, holotype from Pastorino (2002b), 11.0 x 5.8 mm; G, T. scolopax, from Watson (1886), 24.1 x 10.7 mm; c, cord; l, lamella; os, open spine; t, thread; wc, weak cord. weakly angulose– shoulders and external lip, and (1) Fusiform shells of rounded shape with (2) siphonal canal variable in length (Fig. 6); (b) fusiform rounded –or weakly angulose– shoulders shells of angulated shape with angulose shoulders and external lip (Fig. 6) and external lip, and a long (about 25–43% total shell (1’) Shells with other shape: fusiform shells of (10) length) almost straight or curved strong siphonal angulated shape or oval shells canal (Fig. 7), or (c) short to medium (about 13–23% (2) Axial sculpture with lamellae and/or (3) total shell length) siphonal canal (Fig. 8); (d) oval growth striae shells with a short to medium siphonal canal (Fig. 9) (2’) Axial sculpture only of irregular and low or (e) with a long, thin and brittle prolonged siphonal rounded ridges. canal and longFig.�7/10 and straight open abaxial spines along T. parodizi Pastorino, 2005 (Fig. 6A) the shoulders (Fig. 10).

55 CRISTIAN ALDEA & JESÚS S. TRONCOSO

(3) Siphonal canal medium to long and (4) (11’) Short to medium (about 13–23% total (18) no strong axial lamellae adaperturally shell length) siphonal canal (Fig. 8) rolled (12) Axial lamellae producing open spines (13) (3’) Siphonal canal short and axial lamellae only along the shoulders adaperturally rolled. Some specimens (12’) Axial lamellae producing open spines (16) can be more shouldered. T. minutus that tend to be short –or spine-like Strebel, MS. Melvill and Standen, 1907 expansions– along the spiral sculpture (Fig. 6B) or weak projections along the periphery (4) Axial rounded ribs with lamellae (5) when crossing the spirals (4’) Without axial rounded ribs, only lamellae (7) (13) The spines tend to be long and abaxial (14) or growth striae (13’) The spines tend to be short (or triangular) (15) (5) Few axial lamellae located on top of the (6) and adapical ribs crossing low, almost obsolete, spiral (14) Irregular spiral cords, mainly visible at cords the base. (5’) Many axial lamellae arranged tightly, T. acanthodes Watson, 1883 (Fig. 7A) crossing narrow and strong spiral cords (14’) Without spiral sculpture. and forming reticulate sculpture. T. columbarioides Pastorino and T. pallidus (Broderip, 1833) (Fig. 6C) Scarabino, 2008 (Fig. 7B) (6) The lamellae are weak and the aperture (15) Spire low (about 28% total length) and is expanded. “sub-quadrate” expanded aperture. Some T. condei Houart, 2003 (Fig. 6D) specimens have weak expansions on the (6’) The lamellae tend to be strong and the base. aperture is not expanded. T. septus Watson, 1882 (Fig. 7C) T. macquariensis Powell, 1957 (Fig. 6E) (15’) Spire high (about 34% total length) and (7) Lamellae clearly visible and they are (8) less expanded aperture. arranged in order T. vangoethemi Houart, 2003 (Fig. 7D) (7’) Very weak lamellae that are not arranged (9) (16) Lamellae that tend to produce spines; (17) in order, or growth striae spire tending to be shorter (25–35% total (8) Protruding lamellae and without spiral length) with less that seven whorls sculpture. (16’) Weak lamellae that tend to produce weak T. declinans Watson 1882 (Fig. 6F) projections along the periphery; spire (8’) Not protruding lamellae and low rounded tending to be higher (about 40% total spiral cords. shell length) with seven or more whorls. T. emilyae Pastorino, 2002 (Fig. 6G) T. veronicae Pastorino, 1999 (Fig. 7E) (9) Weak lamellae crossing a few blunt spiral (17) Four to five real spiral cords on last whorl. cords. T. arnaudi Pastorino, 2002 (Fig. 7F) T. cuspidarioides Powell, 1951 (Fig. 6H) (17’) Two to three feeble rounded threads on Growth striae crossing many rounded and last whorl. (9’) well defined spiral cords. T. scolopax Watson, 1882 (Fig. 7G) T. fasciolarioides Pastorino and (18) Shell sculptured only by axial low varices, (19) Scarabino, 2008 (Fig. 6I) or spines not produced by striae neither (10) Fusiform shells of angulated shape with (11) lamellae; with spiral ribs or cords angulose shoulders and external lip (Figs. (18’) Shell sculptured by axial lamellae or (20) 7 and 8) growth striae, which can produce spines; (10’) Oval shells (31) with or without spiral sculpture (11) Long (about 25–43% total shell length) (12) (19) Low varices crossing about three spiral almost straight or curved strong siphonal cords in the last whorl. Six whorls. canal (Fig. 7) “T.” malvinarum Strebel, 1908 (Fig. 8A)

56 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

(19’) Spines on periphery arranged in three (26) Axial lamellae crossing many spiral (27) spiral ribs in the last whorl. Striae at the cords, weak ribs or threads base. Four whorls. (26’) Many thin and tight axial lamellae crossing “T.” triacanthus Castellanos, Rolán and three thick spiral cords in the last whorl Bartolotta, 1987 (Fig. 8B) beginning at the periphery, which are (20) Spiral sculpture almost invisible, rarely (21) more visible along the external lip. present and very weak or lacking T. drygalskii Thiele, 1912 (Fig. 8I) (20’) Evident spiral sculpture of cords, ribs or (26) (27) Axial lamellae crossing many strong (28) threads cords producing knobs with short spines along the shoulders or a cancellated (21) Well defined long and open axial spines (22) sculpture only along the shoulders, which tend to be abaxial. Spiral sculpture lacking (27’) Axial lamellae crossing many weak ribs (29) or threads, only producing open spines or (21’) Axial lamellae producing open spines (23) expansions along the shoulders that tend to be adapical. Spiral sculpture lacks or very weak and rarely present (28) Ten axial lamellae producing knobs and sometimes short open spines along the (22) The spines are connected to growth striae. periphery. T. poirieria Powell, 1951 (Fig. 8C) T. ceciliae Houart, 2003 (Fig. 8J) (22’) The spines are connected to evident axial (28’) Up to 13 axial arranged lamellae producing lamellae. a cancellate sculpture when crossing the T. iarae Houart, 1998 (Fig. 8D) spirals. Sometimes the lamellae producing (23) Relatively thick shells with lamellae that (24) a short expansion on the shoulder of the tend to be thick, attached to the shell or last whorl. sometimes almost lack T. distantelamellatus Strebel, 1908 (Fig. 8K) (23’) Delicate shells with lamellae that tend to (25) (29) The axial lamellae tend to be strong and (30) be thin and sharp adaperturally curved, producing adapical spines or expansions along the shoulders. (24) Well defined thick lamellae that can Spiral sculpture of tight weak threads or produce open adapical curved spines cords along the shoulders; the lamellae being sometimes adaperturally cur- (29’) The axial lamellae tend to be weak and ved or sometimes almost lack. Spiral shallow producing feeble expansions ornamentation almost invisible, some- along the shoulders. Spiral sculpture of times six weak cords at the base. distant weak ribs. Some specimens can be T. plicatus (Lightfoot, 1786) (Fig. 8E) a more rounded shape. T. ohlini Strebel, 1904 (Fig. 8L) (24’) Thick lamellae that tend to be not protruding, but producing open adapical (30) The expansions produced by the lamellae short spines along the shoulders. Lack of are rounded and concave. Spiral sculpture spiral ornamentation. of 6–10 cords on lower part of the last T. bahamondei McLean and Andrade, whorl, others obsolete. 1982 (Fig. 8F) T. amettei Carcelles, 1946 (Fig. 8M) (25) The lamellae produce open adapical (30’) The expansions produced by the lamellae spines only along the shoulders. Spiral tend to be angulose to true open spines, ornamentation of very fine striae. although some specimens can be a more T. coulmanensis coulmanensis Smith, rounded shape. Spiral sculpture of weak 1907 (Fig. 8G) threads. T. pelseneeri Smith, 1915 (Fig. 8N) (25’) The lamellae produce open adapical spines along the shoulders and the base. (31) Oval shells with siphonal canal short to (32) Spiral ornamentation unknown. medium and without long and straight T. coulmanensis multilamellatus open abaxial spines along the shoulders Numanami, 1996 (Fig. 8H) (Fig. 9)

57 CRISTIAN ALDEA & JESÚS S. TRONCOSO

(31’) Oval shell with a long, thin and brittle (39) About 8–10 well defined and regular prolonged siphonal canal and axial axial lamellae crossing strong spiral cords lamellae (7–9) producing long and beginning at the periphery, and producing straight open abaxial spines along the adapical short open spines along the shoulders (Fig. 10). shoulders; whilst some specimens can T. clenchi (Carcelles, 1953) vary in thickness and from almost smooth (32) Shells with well defined spiral sculpture (33) with more prominent spiral sculpture to of cords or ridges very lamellated with weak spirals T. geversianus (Pallas, 1774) (Fig. 9F) (32’) Shells without spiral sculpture, missing, (42) or very weak either spaced lines, low (39’) About 17–22 irregular axial lamellae ridges or very fine striae crossing weak spiral ridges, and sometimes producing rounded expansions (33) Axial sculpture of regular lamellose cords (34) along the shoulders. crossing strong spirals cords, giving a T. leptocharteres Oliver and Picken, cancellated sculpture 1984 (Fig. 9G) (33’) Axial sculpture of true lamellae, which (35) (40) More than six thin axial lamellae regularly (41) can produce reticulation when crossing spaced the spirals in some species (40’) With about one shallow lamellate varix (34) Spire very low (about 24% total length) in one-third whorl and a closely spaced of four whorls. group of three associated with the labial T. brevispira Martens, 1885 (Fig. 9A) varix. (34’) Spire higher (about 36% total length) of T. enderbyensis Powell, 1958 (Fig. 9H) five whorls. (41) About 12–13 adaperturally curved T. cribellum Strebel, 1908 (Fig. 9B) lamellae producing weak adapical (35) Numerous axial lamellae producing (36) rounded expansions along the shoulders reticulated sculpture when crossing in some specimens. conspicuous spiral cords or ridges T. eversoni Houart, 1997 (Fig. 9I) (35’) Axial lamellae that, being more distant, (40) (41’) About 6–11 more straight lamellae do not produce reticulation when crossing producing adapical coronated processes low rounded spiral cords along the shoulders. T. scotianus Powell, 1951 (Fig. 9J) (36) Without expansions nor open spines along (37) the shoulders (42) Axial ornamentation of lamellae (in some (43) cases almost obsolete), spiral lacking, (36’) Expansions or adapical short open spines (39) missing or very fine striae, and expanded along the shoulders produced by axial aperture lamellae (42’) Axial ornamentation of irregular growth (37) Axial lamellae not producing sharp (38) striae, spiral of very weak spaced lines or recurved hollow spines when crossing low ridges in some shells, and oval less rounded spiral cords expanded aperture. (37’) Axial lamellae producing sharp recurved T. nucelliformis Oliver and Picken, 1984 hollow spines when crossing flat spiral (Fig. 9K) cords. (43) Less than about 14 solid lamellae (44) T. echinolamellatus Powell, 1951 (Fig. 9C) producing adapical spines or conspicuous (38) About 46–50 axial lamellae crossing expansions along the shoulders, lacking about 13–18 spirals cords. or missing spiral sculpture T. albolabratus Smith, 1875 (Fig. 9D) (43’) About 10–27 thin lamellae sometimes (38’) About 20–28 axial lamellae crossing producing weak sharp and rounded about 17–18 spirals cords. Some half- expansions along the shoulders. Very grown specimens can present some weak fine spiral striae that are rubbed off in expansion in lamellae. some specimens. T. mawsoni Powell, 1957 (Fig. 9E) T. longstaffi Smith, 1907 (Fig. 9L)

58 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

os v s os c r l gs

D A B C

l os os c os l l fs os l l

F H G E I

cs c l l lc k wr os l l c l M K

J N L

Figure 8: Redraws of Trophon (s.l.) species: A, “T.” malvinarum, from Strebel (1908), 7.2 x 3.6 mm; B, T. triacanthus, holotype from Castellanos et al. (1987), 3.1 x 2.1 mm; C, T. poirieria, holotype from Powell (1951), 15.5 x 8.8 mm; D, T. iarae, holotype from Houart (1991), 74.6 x 61.0 mm; E, T. plicatus, from Pastorino (2005), 51.0 x 25.9 mm; F, T. bahamondei, holotype from McLean and Andrade (1982), 49.4 x 26.1 mm; G, T. coulmanensis coulmanensis, holotype from Smith (1907), 13.0 x 7.0 mm; H, T. coulmanensis multilamellatus, from Numanami (1996), 9.0 x 4.6 mm; I, T. drygalskii, from Thiele (1912), 7.0 x 3.7 mm; J, T. ceciliae, holotype from Houart (2003), 41.5 x 21.6 mm; K, T. distantelamellatus, from Strebel (1908), 15.1 x 7.9 mm; L, T. ohlini, USNM 901764, 8.7 x 4.3 mm; M, T. amettei, holotype from Pastorino (2005), 26.8 x 13.7 mm; N, T. pelseneeri, from Strebel (1908), 20.5 x 9.5 mm; c, cord; cs, concave spine; fs, fine stria; gs, growth stria; k, knob; lc, low cord; l, lamella; os, open spine; r, Fig.�8/10 ridge; s, spine; v, varix; wr, weak rib.

59 CRISTIAN ALDEA & JESÚS S. TRONCOSO

llc c llc l c fc l hs c

A B C D

os r l l c l c lv lc

E F G H

c l gs l l lc fs

I J K L

os l l os

N O M

Fig.�9/10 Figure 9: Redraws of Trophon (s.l.) species: A, T. brevispira, from Martens and Pfeffer (1886), 29.0 x 21.0 mm; B, T. cribellum, from Strebel (1908), 17.7 x 8.8 mm; C, T. echinolamellatus, paratype from Powell (1951), 64.0 x 39.0 mm; D, T. albolabratus, from Smith (1879), 40.0 x 18.0 mm; E, T. mawsoni, USNM 896644, 27.0 x 17.0 mm; F, T. geversianus, from Pastorino (2005), 61.8 x 40.5 mm; G, T. leptocharteres, USNM 901770, 18.5 x 10.8 mm; H, T. enderbyensis, holotype from Powell (1958), 70.0 x 38.5 mm; I, T. eversoni, holotype from Houart (1997), 75.8 x 43.8 mm; J, T. scotianus, holotype from Powell (1951), 34.0 x 24.0 mm; K, T. nucelliformis, holotype from Oliver and Picken (1984), 21.6 x 12.6 mm; L, T. longstaffi, from Smith (1907), 41.0 x 25.5 mm; M, T. patagonicus, syntype from Pastorino (2005), 54.2 x 38.4 mm; N, T. shackletoni shackletoni, from Hedley (1911), 26.0 x 18.0 mm; O, T. shackletoni paucilamellatus, holotype from Powell (1951), 31.0 x 21.0 mm; c, cord; fc, flat cord; fs, fine stria; gs, growth stria; hs, hollow spines; l, lamella; lc, low cord; llc, lamellose cord; lv, lamellate varix; os, open spine; r, ridge.

60 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

(44) Axial lamellae thin but solid, producing (45) adapical open spines that tend to be long. Lacking spiral sculpture c (44’) Axial lamellae strong producing adapical l conspicuous expansions that tend to be short or in some specimens the lamellae ss become obsolete or lacking. Spiral sculpture missing and only present in first whorls of some specimens. T. patagonicus (d’Orbigny, 1839) (Fig. 9M) (45) About 12–14 axial lamellae. T. shackletoni shackletoni Hedley, 1911 (Fig. 9N) (45’) About 7–10 more prominent axial lamellae. T. shackletoni paucilamellatus Powell, 1951 (Fig. 9O)

DISCUSSION AND CONCLUSIONS Figure 10: Redraw of Trophon clenchi, holotype from Pastorino (2005), 52.3 x Of the numerous well-known works for the genus 32.8 mm; c, cord; l, lamella; ss, straight spine. Trophon in the Southern Ocean and Sub-Antarctic adjacent waters, the work of Pastorino (2005) can that many specimens lose the protoconch and firsts be considered as a true revision, which constitutes whorls. However, there are some taxonomical remarks a review of all living species from both coasts and distributional observations important to detail. of southern South America living in environments associated with the continental shelf. Other good Taxonomical remarks accounts where some species were synonymized were developed by Powell (1951), focused mainly on Well documented species and some of them with Magellan, Sub-Antarctic and West Antarctic species, a long list of accepted synonyms are those revised and by Dell (1990), focused mainly on Antarctic in South America by Pastorino (2005), but he did Ross Sea species, but none of both studies was a not include in his work T. mucrone Houart, 1991 true revision. The work of Pastorino (2002a) revised (not considered here, see below) and T. veronicae the morphological patterns of the systematics and since they seem to belong in a different group when phylogeny of the genus in Patagonia and Antarctica. more data are available, T. ohlini for resembling the All fragmented information generated contrasting boreal Boreotrophon truncatus, and T. triacanthus with the high number of species found in this work for belonging probably to the muricid genus Apixystus that shared distribution between Antarctic and Sub- (see Pastorino, 2005, p. 55). However, Pastorino in Antarctic areas, being at the moment more viable a his work included Trophon wilhelmensis Ramírez- work of compilation to gather the scattered information Bohme, 1981, from off Chiloé Island, Chile, as a and data of this genus, without taken account species valid species, which later on was located under the distributed from median/low latitudes toward the muricid genus Coronium by Houart and Sellanes equator (i.e., South African species and from 20ºS (2006) based on radular and protoconch characters; in South America), and starting from the aim to therefore, we did not consider the currently Coronium achieve a quick identification of the species using wilhelmensis in this paper. Pastorino also included macroscopic conchological characters, considering “Trophon” malvinarum under dubius genus awaiting

Fig.�10/10 CRISTIAN ALDEA & JESÚS S. TRONCOSO

more data to confirm the true identity, and included areas as southern South America and Sub-Antarctic T. varians (d’Orbigny, 1839) as a junior synonym of areas (e.g. T. acanthodes), or high Antarctic and Sub- T. geversianus, but Houart (1998) figured a syntype Antarctic islands (e.g T. drygalskii), there are some from MNHN (Paris) that Pastorino did not list species with doubtful records (see Table 1) such as T. as synonym. Furthermore, there are species whose geversianus at Western Antarctic Peninsula and South generic position is pending of definitive verification, Orkney Islands that was questioned by Pastorino like T. acanthodes, which possess a shell similar to (2005), or T. poirieria at South Georgia Islands, south Atlantic Coronium coronatum (Penna-Neme & questioned by Zelaya (2005). In addition, the strongly Leme, 1978) according Pastorino (2005), being able discontinuous pattern of distribution of some species to belong to that genus. caught our attention, such as the South American T. geversianus and T. pallidus that were found at eastern T. pallidus was referred as Fuegotrophon pallidus Sub-Antarctic islands, and the eastern Sub-Antarctic by Pastorino (2002a), but we do not have data T. albolabratus and T. declinans that were found at to consider it under that genus, which originally western Sub-Antarctic islands and in waters of South- proposed as the subgenus Fuegotrophon by Powell America, respectively (see Table 1). All those records (1951) based on the protoconch and radula. need a comparison by means of future revision works and more data –and specimens– available from Houart and Sellanes (2006) described Pagodula intermediate areas. concepcionensis from off Concepción, Chile, and one specimen assigned as P. cf. concepcionensis, that Extralimital species of this study were the South according the same authors may be referable to T. American T. aculeatus Watson, 1882, from Brazil concepcionensis (Houart and Sellanes, 2006, p. 61), (9–23ºS), T. verrillii Bush, 1893, from Brazil (19ºS), but more data and specimens are necessary. and T. mucrone Houart, 1991, from Brazil (18–38ºS), presenting more affinity with species from the northern Finally, T. pileus Lamy is a doubtful species from hemisphere (see Houart, 1991), excepting T. mucrone Southwest Atlantic Ocean (55ºS, 64.8ºW) housed in that presented affinity with T. ceciliae as well, that USNM and not figured. differs in having a larger and broader shell with more numerous primary spiral cords and secondary cords, Distributional observations which are not present in T. mucrone (Houart, 1998).

In this work the geographic and bathymetric ACKNOWLEDGMENTS range of the species is gathered and actualized. In this sense, Pastorino (2002a) showed three groups This paper was generated from the data gathered of species arranged according to their geographical and information developed starting from the Spanish distributions: Magellanic, Circumantarctic and cruises BENTART 2003/2006 that were carried out insular ones, and that there is no interchange between under the auspices of the Spanish Government through Patagonia and Antarctica species. However we found the Antarctic Programmes REN2001-1074/ANT and some species recorded in both areas –Magellan and GLC2004-01856/ANT of the Ministry of Education Antarctica– (e.g. T. minutus) and that only three are and Science (MEC). We thank Linda Ward of the truly Antarctic species, most species reaching insular Department of Invertebrate Zoology, Smithsonian and Sub-Antarctic areas at deeper waters. Institution, for kindly sending the Trophon records of USNM. We also thank two anonymous referees for Although a definite pattern of distribution was improving the manuscript and Cristina Vertan for her observed in most species, inhabiting contiguous kindly revision of the English language.

62 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Table 1. Species of Trophon (s.l.) arranged alphabetically showing their distribution and Table 1: Species of Trophonbathymetry. (s.l.) arranged Coordinates alphabetically of geographic showing their toponym distributions areand bathymetry.indicated Coordinatesin the Appendix; of geographic USNM, toponyms are indicated in the SmithsonianAppendix; National USNM, Museum Smithsonian of Natura Nationall History; Museum of n.d., Natural no History; data were n.d., noobtained. data were obtained.

Species - Distribution Bathymetry (m) and source T. acanthodes Watson, 1883 Río Grande do Sul n.d. (Rios, 1994) Off Uruguay n.d. (Pastorino, 2005) Off Punta Médanos 104–111 (Pastorino, 2005) Off Mar del Plata 192 (Pastorino, 2005) Off Bahía Blanca 82–95 (Pastorino, 2005) East of Falkland/Malvinas Islands 855–866 (Pastorino, 2005) (56.7ºW) Between South Georgia and South n.d. (Houart, 1998) Orkney Islands (59ºS, 38ºW) Tierra del Fuego Island n.d. (Pastorino, 2005) Off Desolación Island (75.3ºW) 119–319 (Pastorino, 2005) Between Manuel Rodriguez and 188–544 (Pastorino, 2005), 448 (Watson, 1886) Desolación Islands Wellington Island 320 (Watson, 1886) Boca del Guafo to Ninualac Channel 160–200 (Osorio et al., 2006) Gulf of Corcovado 130–169 (Cárdenas et al., 2008) T. albolabratus Smith, 1875 South Georgia Islands n.d. (Martens and Pfeffer, 1886) (1) Kerguelen Islands 0 (Smith, 1879), 0–30 (Powell, 1957), 25–50 (Troncoso et al., 2001), 0–104 (Cantera and Arnaud, 1985), 46–110 (Watson, 1886) T. amettei Carcelles, 1946 Bustamante Bay 11 (Pastorino, 2005) T. arnaudi Pastorino, 2002 South Sandwich Islands 355–468 (Pastorino, 2002b) T. bahamondei McLean and Andrade, 1982 Off Coquimbo to off Pichilemu 200–400 (McLean and Andrade, 1982) SW Coquimbo 370 (Houart, 2003) Off Coquimbo 400 (Pastorino, 2005) Gulf of Ancud to Gulf of Corcovado 130–252 (Cárdenas et al., 2008) T. brevispira Martens, 1885 South Georgia Islands 0–16 (Martens, 1885), 2–15 (Strebel, 1908), 18–37 (Carcelles, 1953), 18–270 (Powell, 1951), n.d. (Lamy, 1911, Zelaya, 2005) T. ceciliae Houart, 2003 Antofagasta 1000–1300 (Houart, 2003) Off Caldera 434 (Houart and Sellanes, 2006) Off Coquimbo 600 (Houart and Sellanes, 2006) Off Concepción Bay (73.7°W) 900 (Houart and Sellanes, 2006) T. clenchi (Carcelles, 1953) Off Mar del Plata (55.6ºW) 90 (Pastorino, 2005) Off Gulf of San Matías (57.6ºW) 1062 (Pastorino, 2005) Off Gulf San Jorge (57.3ºW) n.d. (Pastorino, 2005)

25 63 CRISTIAN ALDEA & JESÚS S. TRONCOSO

Species - Distribution Bathymetry (m) and source East of Falkland/Malvinas Islands 646–845 (Pastorino, 2005) (56.6ºW) Staten Island n.d (Pastorino, 2005) Tierra del Fuego Island 342–353 (Pastorino, 2005) T. columbarioides Pastorino and Scarabino, 2008 Off Mar del Plata (54–55ºW) 209–382 (Pastorino and Scarabino, 2008) T. condei Houart, 2003 Ancud 1350 (Houart, 2003) Off Concepción Bay (73.7°W) 728–930 (Houart and Sellanes, 2006) T. coulmanensis coulmanensis Smith, 1907 South Sandwich Islands 93–121 (USNM) (2) Weddell Sea n.d. (USNM) (2) South Shetland Islands 220–1120 (Dell, 1990) (2) Bransfield Strait 809–1116 (USNM) (2) Western Antarctic Peninsula 294 (Aldea and Troncoso, 2008) Ross Sea 183 (Smith, 1907) (2), 351–1674 (Dell, 1990) (2) Cape Adare 329–366 (Smith, 1915) (2) George V Land 527 (Hedley, 1916) (2) Davis Sea n.d. (Thiele, 1912) (2) Kerguelen Islands 125–1218 (Cantera and Arnaud, 1985) (2) T. coulmanensis multilamellatus Numanami, 1996 East Antarctica (24ºE) 270–289 (Numanami, 1996) T. cribellum Strebel, 1908 South Georgia Islands 20–75 (Strebel, 1908), 22 (Zelaya, 2005), 44 (Carcelles, 1953) (3) T. cuspidarioides Powell, 1951 South Georgia Islands 120–204 (Powell, 1951) Peter I Island 410 (Aldea and Troncoso, 2008) T. declinans Watson 1882 Northwest of Heard Island (52.1ºS, 274 (Watson, 1886) 71.4ºE) Kerguelen Islands 30–190 (Cantera and Arnaud, 1985) Crozet Islands 115–235 (Cantera and Arnaud, 1985) Marion Island 99–113 (Powell, 1951), 126 (Watson, 1886), 0–527 (Branch et al., 1991) East of Falkland/Malvinas Islands 229–236 (Powell, 1951) (57ºW) Drake Passage (59.8ºS, 68.8ºW) 1043–1208 (USNM) T. distantelamellatus Strebel, 1908 South Georgia Islands 64–74 (Strebel, 1908), 18–110 (Powell, 1951), 46–101, (USNM) T. drygalskii Thiele, 1912 South Sandwich Islands 161–210 (USNM) Weddell Sea 588 (Hain, 1990), 119–2315 (Gutt et al., 2000) South Shetland Islands and Bransfield 210–426 (Dell, 1990) Strait Bransfield Strait n.d. (USNM)

64 26 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Species - Distribution Bathymetry (m) and source Western Antarctic Peninsula 126 (USNM) Bellingshausen Sea 1426–1814 (Aldea and Troncoso, 2008) Ross Sea 311–348 (USNM), , 265–392 (Dell, 1990), , 292–457 (Smith, 1915) Davis Sea n.d. (Thiele, 1912) Enderby Land 193–300 (Powell, 1958) East Antarctica (24ºE) n.d. (Numanami, 1996) T. echinolamellatus Powell, 1951 South Sandwich Islands 93–228 (USNM) South Georgia Islands 220–320 (USNM) Weddell Sea 504 (Gutt et al., 2000) South Orkney Islands 284 (USNM) South Shetland Islands 115 (Aldea and Troncoso, 2008), 342 (Powell, 1951), 120–462 (USNM) Bransfield Strait 210–1373 (USNM) Western Antarctic Peninsula 37–800 (USNM) T. emilyae Pastorino, 2002 NW Amundsen Sea (54.8ºS, 129.8ºW) 549–1153 (Pastorino, 2002b) T. enderbyensis Powell, 1958 Enderby Land 193 (Powell, 1958) T. eversoni Houart, 1997 Kerguelen Islands 185 (Houart, 1997) T. fasciolarioides Pastorino and Scarabino, 2008 Off Bahía Blanca 1000 (Pastorino and Scarabino, 2008) Burdwood Bank 286–292 (Pastorino and Scarabino, 2008) T. geversianus (Pallas, 1774) Buenos Aires Province n.d. (Pastorino, 2005) South Atlantic Ocean (37.8ºS, 36.2ºW) 100 (Strebel, 1908) San Antonio Oeste 0 (Pastorino, 2005) Sierra Grande 0 (Pastorino, 2005) Puerto Lobos n.d. (Pastorino, 2005) Puerto Pirámides n.d. (Pastorino, 2005) Puerto Madryn n.d. (Pastorino, 2005) Gulf San Jorge n.d. (Pastorino, 2005) Puerto Deseado 5 (Pastorino, 2005) Punta Peñas 0–2 (Pastorino, 2005) Río Santa Cruz n.d. (Pastorino, 2005) Off Rio Santa Cruz (67.7ºW) 56–84 (Powell, 1951) Rio Gallegos n.d. (Pastorino, 2005) Falkland/Malvinas Islands 0–11 (Melvill and Standen, 1907) (4), 1–16 (Powell, 1951), 22 (Watson, 1886), 10–40 (Strebel, 1908), n.d. (Melvill and Standen, 1898), (Pastorino, 2005) Burdwood Bank 102 (Melvill and Standen, 1912) (4), n.d. (Pastorino, 2005) Staten Island 0 (USNM), 0–641 (Pastorino, 2005) Usuahia n.d. (Strebel, 1908, Pastorino, 2005) Usuahia and Beagle Channel n.d. (Rochebrune and Mabille, 1889) Beagle Channel 0 (Pelseneer, 1903, Dell, 1971), 15–53 (Osorio, 1999)

27 65 CRISTIAN ALDEA & JESÚS S. TRONCOSO

Species - Distribution Bathymetry (m) and source Cape Horn n.d. (Rochebrune and Mabille, 1889) Magellan Strait 0 Mutschke et al., 1998), 2–3 (USNM), 14–22 (Osorio, 1999), 27 (Powell, 1951), 0–296 (Pastorino, 2005), n.d. (Rochebrune and Mabille, 1889, Lamy, 1906, Houart, 1998, Linse, 2002) Tierra del Fuego Island 0–1 (USNM), 0–31 (Pastorino, 2005), 27–38 (Powell, 1951) (4), n.d. (Smith, 1905, USNM) Heard Island 137 (Watson, 1886) Western Antarctic Peninsula 30 (Lamy, 1906) (5) South Orkney Islands 16–18 (Melvill and Standen, 1907) (5) T. iarae Houart, 1998 Off Albardão 55 (Houart, 1998) Off Uruguay n.d. (Houart, 1998) T. leptocharteres Oliver and Picken, 1984 South Orkney Islands 15–30 (Oliver and Picken, 1984) South Shetland Islands 64–82 (USNM) Western Antarctic Peninsula 18–130 (USNM) T. longstaffi Smith, 1907 South Sandwich Islands 148–201 (USNM) South Georgia Islands 1299–1400 (USNM) Weddell Sea 123–446 (Gutt et al., 2000) Peter I Island 90–126 (Aldea and Troncoso, 2008) Ross Sea 47–75 (Smith, 1907), 8–124 (USNM), 37–146 (Hedley, 1911), 457 (Smith, 1915), 8–1080 (Dell, 1990) George V Land 22–732 (Hedley, 1916) Adelie Land 5–140 (Arnaud, 1972) Wilkes Land 128–146 (USNM) East Antarctica (40ºE) 5 (Numanami, 1996) T. macquariensis Powell, 1957 Macquarie Island 69 (Powell, 1957), 110–124 (USNM) T. mawsoni Powell, 1957 Macquarie Island 69 (Powell, 1957), 29–549 (USNM), n.d. (Hedley, 1916) (6) T. minutus Strebel, MS. Melvill and Standen, 1907 South Sandwich Islands 148–201 (Dell, 1990), 122–210 (USNM) South Georgia Islands 24–52 (Strebel, 1908) Drake Passage (56ºS, 61.5ºW) 47–54 (USNM) Magellan Strait 27–73 (USNM) South Orkney Islands 16–18 (Melvill and Standen, 1912), 20 (Oliver and Picken, 1984), 16–27 (Melvill and Standen, 1907) Weddell Sea 119–170 (Gutt et al., 2000) South Shetland Islands 38–82 (USNM) Bransfield Strait 210–240 (USNM), 210–265 (Dell, 1990) Western Antarctic Peninsula 6–37 (Dell, 1990), 53 (Aldea and Troncoso, 2008), 93–130 (Powell, 1951), 21–305 (USNM) Ross Sea n.d. (Dell, 1990) George V Land 46 (Hedley, 1916) (7) Adelie Land 31 (Arnaud, 1972) (7)

66 28 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Species - Distribution Bathymetry (m) and source Kerguelen Islands 24–158 (Cantera and Arnaud, 1985) (7) T. nucelliformis Oliver and Picken, 1984 South Orkney Islands 3–6 (Melvill and Standen, 1907) (8), 2–20 (Oliver and Picken, 1984) South Shetland Islands 44 (USNM) Western Antarctic Peninsula 4–6 (USNM) T. ohlini Strebel, 1904 South Atlantic Ocean (47.6ºS, 60.8ºW) 133–219 (Powell, 1951) West of Falkland/Malvinas Islands 141–162 (Powell, 1951) (63ºW) Burdwood Bank 124 (USNM) Magellan Strait 27 (Strebel, 1904), 38–92 (Linse, 2002) (9), 91–110 (USNM) Beagle Channel 13–135 (Linse, 2002) (9) Cape Horn 40 (Linse, 2002) (9) T. pallidus (Broderip, 1833) Falkland/Malvinas Islands 11 (Melvill and Standen, 1907) (10), 10–146 (Powell, 1951) Burdwood Bank 102 (Melvill and Standen, 1907) (10), 124–128 (USNM), 137–150 (Strebel, 1908) (12) Staten Island 118 (Powell, 1951) Off Tierra del Fuego Island (65–67.2ºW) 79–124 (USNM) Beagle Channel 15–53 (Osorio, 1999), 15–135 (Linse, 2002) Cape Horn 40–66 (Linse, 2002), 115 (USNM), 24–357 (Rochebrune and Mabille, 1889) (10) (11) Drake Passage (56.3ºS, 67.2ºW) 121 (Powell, 1951) Magellan Strait 0 Mutschke et al., 1998), 14–35 (Osorio, 1999), 45–92 (Linse, 2002), 95 (Powell, 1951) Off Desolación Island (74.9–75ºW) 64–101 (USNM) Gulf of Corcovado 130 (Cárdenas et al., 2008) Gulf of Ancud 252 (Cárdenas et al., 2008) Crozet Islands 505 (Cantera and Arnaud, 1985) T. parodizi Pastorino, 2005 Le Maire Strait 229–265 (Pastorino, 2005) T. patagonicus (D'orbigny, 1839) Rocha n.d. (Houart, 1998) (13) Off Rocha 100 (Pastorino, 2005) Off Punta Rasa (54.6–54.7ºW) 26–54 (Pastorino, 2005) Mar Azul 55 (Pastorino, 2005) Mar del Plata n.d. (Pastorino, 2005) Miramar n.d. (Pastorino, 2005) Necochea 30–50 (Pastorino, 2005) T. pelseneeri Smith, 1915 Macaé 30–55 (Pastorino, 2005) Río de Janeiro 50–100 (Houart, 1991), 55–225 (Ríos, 1994) Off Rocha (34.8ºS, 54.4ºW) 25 (Pastorino, 2005) Buenos Aires n.d. (Pastorino, 2005) Necochea n.d. (Carcelles, 1944) Falkland/Malvinas Islands n.d. (Pastorino, 2005)

29 67 CRISTIAN ALDEA & JESÚS S. TRONCOSO

Species - Distribution Bathymetry (m) and source West of Falkland/Malvinas Islands 229 (Smith, 1915) T. plicatus (Lightfoot, 1786) Río Grande do Sul n.d. (Ríos, 1994) Necochea 0 (Carcelles, 1944) (14) Off Bustamante Bay (65ºW) 82–88 (Pastorino, 2005) Comodoro Rivadavia n.d. (Pastorino, 2005) Cabo Blanco 82–88 (Powell, 1951) (14) Punta Peñas 2–3 (Pastorino, 2005) Falkland/Malvinas Islands 25 (Strebel, 1908), 79 (Powell, 1951) (14), 79 (Houart, 1998) (16), n.d. (Melvill and Standen, 1898) (14) West of Shag Rocks (43.4ºW) 160 (Strebel, 1908) Burdwood Bank 137–150 (Strebel, 1908) Off Tierra del Fuego Island (65–67.2ºW) 82–128 (Pastorino, 2005) Tierra del Fuego Island n.d. (Pastorino, 2005) Staten Island 84–91 (Pastorino, 2005) Cape Horn 0 (Lamy, 1906) (14), 12–28 (Rochebrune and Mabille, 1889) (14), n.d. (Pastorino, 2005) Usuahia 8–10 (Pastorino, 2005), 12–28 (Rochebrune and Mabille, 1889) (14) Beagle Channel 24–65 (Osorio, 1999), 55–82 (Pastorino, 2005) Magellan Strait 0 Mutschke et al., 1998), 24 (Osorio, 1999), 13–46 (Pastorino, 2005) Off Desolación Island (75ºW) 92–101 (Pastorino, 2005) Puerto Edén 0–18 (Dell, 1971) (15) Gulf Elefantes 0–15 Reid and Osorio, 2000) Off Gulf of Penas (83.9ºW) 298 (Pastorino, 2005) Estero Elefantes 0–15 Reid and Osorio, 2000) Pulluche Channel 62 (Osorio et al., 2006) Gulf of Ancud 252 (Cárdenas et al., 2008) T. poirieria Powell, 1951 South Shetland Islands 342 (Powell, 1951) Western Antarctic Peninsula 93–130 (Powell, 1951) South Georgia Islands n.d. (Carcelles, 1953) (17) T. scolopax Watson, 1882 Northwest of Heard Island (52.1ºS, 274 (Watson, 1886) 71.4ºE) Kerguelen Islands 60–620 (Cantera and Arnaud, 1985) Crozet Islands 330–600 (Cantera and Arnaud, 1985) T. scotianus Powell,1951 South Georgia Islands 0 (Carcelles, 1953), 97–101 (Dell, 1990), 107 (Powell, 1951), 18– 144 (Zelaya, 2005), 84–218 (USNM) Weddell Sea 437–519 (USNM), 202–617 (Hain, 1990) (18), 462–620 (Gutt et al., 2000) Western Antarctic Peninsula 20–106 (USNM) Ross Sea 256–474 (Dell, 1990) East Antarctica (39.5ºE) 147 (Numanami, 1996) T. septus Watson, 1882 Kerguelen Islands 52 (Watson, 1886), 48–65 (Troncoso et al., 2001), 0–150 (Powell, 1957), 30–620 (Cantera and Arnaud, 1985)

68 30 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

Species - Distribution Bathymetry (m) and source Crozet Islands 62–355 (Cantera and Arnaud, 1985) Marion and Prince Edward Islands 140–200 (Branch et al., 1991) T. shackletoni paucilamellatus Powell, 1951 South Georgia Islands 18 (Carcelles, 1953), 75 (Strebel, 1908) (19), 97–101 (Dell, 1990), 82–138 (Zelaya, 2005), 100–178 (Powell, 1951), 66–366 (USNM) South Sandwich Islands 121–228 (USNM), 278–329 (Powell, 1951) Southeast of South Orkney Islands 593–598 (Dell, 1990) (61.4ºS, 41.9ºW) South Orkney Islands 97–305 (USNM), 298–403 (Dell, 1990) South Shetland Islands 421–462 (USNM) Western Antarctic Peninsula 128–165 (Dell, 1990), 50–375 (USNM) Ross Sea 347–358 (USNM) T. shackletoni shackletoni Hedley, 1911 South Georgia Islands 102–137 (USNM) (21) Weddell Sea 123 (Gutt et al., 2000), n.d. (Hain, 1990) (20) South Orkney Islands 35 (Oliver and Picken, 1984) Ross Sea 13–37 (Hedley, 1911) (21), 293–549 (Smith, 1915) (21), 188–1890 (Dell, 1990) West of Macquarie Island (151ºE) 2746–3248 (Dell, 1990) George V Land 100–110 (Hedley, 1916) (21) Shackleton Ice Shelf 43 (Egorova, 1982) Davis Sea 110 (Hedley, 1916) (21) Amery Ice Shelf 456 (Powell, 1958) (21) Enderby Land 193–220 (Powell, 1958) (21) East Antarctica (24ºE) 275–310 (Numanami, 1996) T. triacanthus Castellanos, Rolán and Bartolotta, 1987 Off Gulf San Jorge (60°W) 600 (Castellanos et al., 1987) Falkland/Malvinas Islands 646–866 (USNM) T. vangoethemi Houart, 2003 Gulf of Corcovado 169 (Cárdenas et al., 2008) Off Concepción Bay (73.1–73.6°W) 350 (Houart, 2003), 530–613 (Houart and Sellanes, 2006) T. veronicae Pastorino, 1999 Northwest of South Georgia Islands 298–1281 (Pastorino, 1999) (50.5ºS, 43.5ºW) Desolación Island 675 (Pastorino, 1999) Off Southern Chile (46–46.1°S, 83.9– 298–742 (Pastorino, 1999) 84°W) Boca del Guafo 200 (Osorio et al., 2006) "T." malvinarum Strebel, 1908 Falkland/Malvinas Islands 197 (Strebel, 1908) Beagle Channel 83 (Pastorino, 2005) (22)

(1) Cited as T. cinguliferus Pfeffer, 1886. Synonymized after Strebel (1908) and Cernohorsky (1977). (2) Cited as T. coulmanensis. (3) Misidentified as T. cinguliferus Pfeffer, 1886 according Zelaya (2005).

31 69 CRISTIAN ALDEA & JESÚS S. TRONCOSO

(4) Cited as T. philippianus Dunker, 1878. Synonymized after Pastorino (2005). (5) According Pastorino (2005) are probably T. nucelliformis Oliver and Picken, 1984 or T. maquariensis Powell, 1957, or T. echinolamellatus sensu Powell (1951). (6) Cited as T. albolabratus, according Powell (1957). (7) Cited as T. condensatus Hedley, 1916. Synonymized after Dell (1990). (8) Misidentified as T. cinguliferus Pfeffer, 1886 according Oliver and Picken (1984). (9) Conferred specimens. (10) Cited as T. crispus (Couthouy M.S. Gould, 1849). (11) Cited as T. fasciculatus (Hombron and Jacquinot, 1848). Synonymized after Powell (1951). (12) Cited as T. crispus var. burwoodianus (Couthouy M.S. Gould, 1849). Synonymized after Powell (1951). (13) Misidentified as T. plicatus according Pastorino (2005). (14) Cited as T. laciniatus (Martyn, 1784) (Pastorino, 2005). (15) Cited as T. lamellosa (Gmelin, 1791) (Pastorino, 2005). (16) Misidentified as T. iarae according Pastorino (2005). (17) Zelaya (2005) doubts this record since no reference in collections was neither found nor well documented. (18) Cited as “T. sp.1” according Numanami (1996). (19) Cited as T. laciniatus, but according Powell (1951) are probably T. shackletoni paucilamellatus. (20) Cited as T. shackletoni, conferred. (21) Cited as T. shackletoni. (22) Established under dubious genus.

APPENDIX Caldera, CL (27.1ºS, 70.8ºW) Cape Adare, AN (69.7ºS, 163.4ºE) List of toponyms used in the Table 1 and their Cape Horn (56ºS, 67.2ºW) locations. Comodoro Rivadavia, AR (45.9ºS, 67.5ºW) AR, Argentina; AN, Antarctica; BR, Brazil; CL, Concepción Bay, CL (36.6–36.7ºS, 73–73.1ºW) Chile; UR, Uruguay. Coquimbo, CL (30ºS, 71.4ºW) Crozet Islands (46–46.5ºS, 50.2–52.3ºE) Adelie Land, AN (136.2–142ºE) Davis Sea, AN (80–105ºE) Albardão, BR (33.2ºS, 52.7ºW) Desolación Island, CL (52.8–53.4ºS, 73.1–74.7ºW) Amery Ice Shelf, AN (70–75ºE) Drake Passage (56–62ºS, 58–69ºW) Amundsen Sea, AN (99–126ºW) Enderby Land, AN (44.6–59.6ºE) Ancud, CL (41.9ºS, 73.8ºW) Estero Elefantes, CL (45.8º–46.3ºS, 73.7ºW) Antofagasta, CL (23.6ºS, 70.4ºW) Falkland/Malvinas Islands (51–52.5ºS, 57.7–61.3ºW) Bahía Blanca, AR (39ºS, 62ºW) George V Land, AN (142–153.8ºE) Beagle Channel (54.9–55.1ºS, 66.5–70.2ºW) Gulf Elefantes, CL (46.3–46.6ºS, 73.8ºW) Bellingshausen Sea, AN (70–99ºW) Gulf of Ancud, CL (41.8–42.3ºS, 72.6–73.4ºW) Boca del Guafo, CL (43.7ºS, 75ºW) Gulf of Corcovado, CL (42.3–43.5ºS, 72.8–73.5ºW) Bransfield Strait, AN (63ºS, 59ºW) Gulf of Penas, CL (46.8–47.7ºS, 74.1–75.7ºW) Buenos Aires (Province), AR (34.1–41ºS, 58.4–62.8ºW) Gulf of San Matías, AR (40.8–42.3ºS, 63.5–65.2ºW) Burdwood Bank (53.8–55.3ºS, 55.7–61.5ºW) Gulf San Jorge, AR (44.9–47.1ºS, 65.5–67.6ºW) Bustamante Bay, AR (45.1ºS, 66.5ºW) Heard Island (53–53.2ºS, 73.3–73.8ºE) Cabo Blanco, AR (47.2ºS, 65.7ºW) Kerguelen Islands (48.5–49.8ºS, 68.7–70.6ºE)

70 32 REMARKS ON THE GENUS TROPHON (S.L.) MONTFORT, 1810 (MOLLUSCA: GASTROPODA: MURICIDAE) IN THE SOUTHERN OCEAN AND ADJACENT AREAS

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ADDENDA

During the edition of this volume of Thalassas, was publish the following paper: Harasewych MG, Pastorino G (2010). Trophonella (Gastropoda: Muricidae), a new genus from Antarctic waters with description of a new species. The Veliger 51(1): 85- 103.

The authors describe a new species under the genus Trophonella from Laurie Island (South Orkney Islands ) called Trophonella rugosolamellata. Besides four species (Trophon scotianus, T. echinolamellatus, T. enderbyensis and T. eversoni) previously cited by us in this paper like Trophon was translated to the new genus Trophonella.