— THE VELIGER © CMS, Inc.. 2003 The Veliger46(4):332-350 (October6, 2003) Acanthina Fischer von Waldheim, 1807 (Gastropoda: Muricidae), an Ocenebrine Genus Endemic to South America THOMAS DeVRIES J. Burke Museum of Natural History and Culture, University of Washington, Seattle, Washington 98195, USA' Abstract. The endemic South American muricid genus, Acanthina Fischer von Waldheim, 1807. is re-evaluated using characters of fossil and Recent shells. Three new taxa from Peru and Chile are described; A. triangularis, sp. nov. (late Pliocene), A. obesa. sp. nov. (late Miocene to early Pliocene), and A. rugosa, sp. nov. (late Miocene). These species and the late Oligocene to middle Miocene A. katzi Fleming, 1972, are found in deposits that indicate nearshore envi- ronments like those favored by the extant A. unicornis (Bruguiere, 1789) and A. monodon (Pallas, 1774). The pace of acanthinine evolution matches that of contemporaneous muricid genera in western South America. INTRODUCTION GEOLOGY Acanthina Fischer von Waldheim, 1807, comprises spe- The Cenozoic stratigraphy and paleontology of forearc basins with Acanthina-hearmg deposits was described by cies of ovate ocenebrine muricids (Kool, 1993) with a Muizon & DeVries (1985). DeVries (1988, 1998), and labral tooth that range from southern Peru to southeastern Argentina (Dall, 1909; Carcelles, 1954; Figure 1). Most CDhuinlbe.arTehtealg.eo(l1o9g9i0c)alforsetPteirnug aonfdLboy ATbaavrecraa,(1a9n79u)ppfeorr authorities recognize two modern subspecies or species Miocene Chilean locality with Acanthina, was discussed (Carcelles, 1954; Dell, 1971; Wu, 1985). Absent biolog- by Covacevich & Frassinetti (1990). ical data, the two taxa, largely disjunct geographically, are herein treated as separate species.Acanthina monodon JVIATERIALS and METHODS (Pallas, 1774) [synonym: A. calcar (Martyn, 1784)], the type species, encompasses specimens from Argentina and Specimens described in this study were found by the au- southern Chile with imbricate spiral cords and arelatively thor, V. Alleman, and J. Machare. Comparative material thin outer lip. Acanthina unicornis (Bruguiere, 1789) was provided by the CaliforniaAcademy ofSciences, Los [synonym: A. crassilabrum (Lamarck, 1816)] includes Angeles County Museum of Natural History, and Museo specimens from southern Peru to southern Chile that gen- Nacional de La Plata (Argentina). erally lack imbrication and possess a thick, often dentate Locality descriptions are in the appendix. Lengths (L) outer lip. are measured from the apex ofthe spire to the end ofthe Fossils ofAcanthina have long been known from Pleis- siphonal canal, parallel to the axis ofcoiling. Widths (W) tocene and upper Pliocene strata ofChile (Philippi, 1887; are measured at right angles to the length between the Herm, 1969). Most were assigned to one ofmany Recent farthest points left and right of the coiling axis. Both are reported in millimeters (mm). Dimensions diminished by forms (Sowerby, 1835) that are best synonymized with A. monodon or A. unicornis. An older species, A. katzi bsrpeeackiamgeensaraereendcelpoossietdedbyatpatrheenBthuerskees.MTuyspeeusmanofdNfaitguurraeld Fleming, 1972. occurs in lower Miocene deposits ofChi- loe, Chile (Watters & Fleming, 1972) and lowerto middle HWiassthoirnygtaonnd);CuDletpuarreta(mUneinvteorsidtey VoefrtWeabsrhaidnogst,onM,usSeeaottldee, Miocene deposits of southern Peru (DeVries & Vermeij, Historia Natural (Universidad de San Marcos, Lima, 1997). Peru); the Museo de Historia Natural, Universidad Ricar- This paper describes three new species of Acanthina do Palma (Lima, Peru); and the Museo Nacional de His- from upper Miocene and Pliocene strata in Peru and toria Natural (Santiago, Chile). Chile, documents a range extension of the genus into Abbreviations—used for museums and collections are as northernmost Peru during the late Pliocene, and summa- follows: LACM Los Angeles County Museum of Nat- rizes the evolutionary history of the genus. ural—History, Los Angeles, California, USA; MUSM INV Departamento de Vertebrados, Museo de Historia Natural, Universidad de San Marcos, Lima, Peru; OSU ' Mailing address: Box 13061, Burton, Washington 98013, Orton Museum,—Ohio State University, Columbus, Ohio, USA. USA; SGO.PI. Museo Nacional de Historia Natural, T. J. DeVries, 2003 Page 333 — UCMP Santiago. Chile; University of California at labral teeth at the edge of the outer lip, and/or broad pri- Berkeley, Mu—seum ofPaleontology, Berkeley, California, mary spiral cords not seen in species from South Amer- USA; UNLP Museo Nacional d—e La Plata, Universidad ica. Hence, they have been assigned by others to different de La Plata, Argentina; USNM Department of Paleo- genera or subgenera: A. brevidentata (Wood, 1828) to bDi.oCl.o,gyU,SAU;niUteWdBSMta—tesBuNrakteioMnaulseMuumseoufmN,atuWraaslhiHnigsttoonr,y ATchaanitshRaoidsinVge,rme1i7j98,&bKyooClo,ok1e994(,19b1a9s)edanodnWauna(t1o9m8y5)anodr and Culture, Un—iversity ofWashington, Seattle, Washing- radular characters; A. tubercidata (Sowerby, 1835) to ton, USA; VE Private collection, V. Alleman, Facultad Mancinella Link, 1807, by Wu (1985) based on similar de Biologfa, Universidad Ricardo Palma, Lima, Peru. criteria (Vermeij & Kool, 1994); and Acanthina angelica Oldroyd, 1918, A. punctulata (Sowerby, 1835), A. higub- SYSTEMATIC PALEONTOLOGY ris (Sowerby, 1821), and A. pauciliiata (Stearns, 1871) to Acanthina (Accmthinncella) or simply Acanthinucella Family Muricidae Rafinesque, 1815 Cooke, 1918, based on shell morphology and radularfea- Subfamily Ocenebrinae Cossman, 1903 tures (Cooke, 1918; Wu, 1985; Vermeij, 1993). Acanthina Genus Acanthina Fischer von Waldheim, 1807 spirata (Blainville, 1832) has both been grouped with and Acanthina Fischer von Waldheim, 1807:174. separated from A. punctulata (e.g.. Grant & Gale, 1931; McLean, 1969). Acanthina emersoni Hertlein & Allison, Type species: Buccinum monodon Pallas, 1774:33, pi. 3, 1959, a Pliocene species from California, was thought by figs. 3, 4. its authors to be related to A. spirata. A. angelica andA. Original description: "Coquille ovale. Ouverture se ter- alcugaunhtrhiisnahaMvaerrkeocen&tlyVebremeenijp,la1c9e9d9.in the new genus Mex- minant inferieurement en un canal tres court, oblique, peu echancre a I'extremite. Bord gauche calleux, formant un Toothed specimens from Chile and Peru with a deeply bourrelet distinct de la columelle. Levre droite epaisse, constricted base assigned to Nucella (Acanthinucella) by munie pres de la base d'une epine tres longue." H(1e9r8m6)(1ar9e69n)oawndplAaccaendthiinnHaen(AnciannetshpiinnucaelDleaV)ribeysD&eVrVieers- Diagnosis: Shell with short, unconstricted orslightly con- meij, 1997. stricted base. Two primary spiral cords on early whorls, one at shoulder, the other anterior to first; these and ad- Occurrence: Uppermost Oligocene to upper Miocene: ditional spiral cords evident on adult body whorl, if not southern Peru to southern Chile. Pliocene: northern Peru subdued or obsolete. Columella excavated. Thin spiral to central Chile. Pleistocene to Recent: southern Peru to groove near base ending in labral tooth inside outer lip. southeastern Argentina (Figure 1). Additional description: Shell to 60 mm long; trigonal, Acanthina monodon (Pallas, 1774) biconic, or fusiform; anterior slightly constricted or un- (Figures 2-7) constricted. Spire 10 to 30 percent ofshell length. Sutures impressed to appressed, usually adherent just below Buccinum monodon Pallas, 1774:33, pi. 3. figs. 3, 4. shoulder. Protoconch heterostrophic, submerged, absent Buccinum calcar Bruguiere, 1789:253, volume 1. figure 10; on fossils. Teleoconch with three to five whorls. Shoulder volume 2, figure 50 [aUributed to Martyn (1784)]. angulate or rounded. Periphery near or posterior to mid- Nucella (Acanthina) calcar (Martyn). Carcelles, 1954:pl. 1, point ofbody whorl. Growth lines prosocline, sometimes figs. 5-11; pi. 2, figs. 12-22. (Seeextensive synonymy.) Not Nucella (Acanthina) crassilabnun calcar (Martyn). lamellose. Axial sculpture lacking or, rarely, with ineg- Herm, 1969:pl. 17, figs. 7-9. (correct identification is ular swellings. Spiral sculpture of two rounded primary Acanthina unicornis). cords on juveniles, one at shoulder, the other anterior to Acanthina monodon monodon (Solander, 1786). Wu. 1985: first; both sometimes obsolete or joined by other spiral 56, figs. 13-14, 24, 34, 52-58, 71. cords of equal or lesser strength on adults. Thin spiral Acanthina monodon (Pallas, 1774). Kool, 1993:229, figs. 27A-D. groove near base, ending in labral tooth emerging from within outer lip. Aperture ovate. Outer lip sometimes Diagnosis: Sculpture typically imbricate; primary and thickened, beveled, dentate within, and/or crenulate at secondary spiral cords numerous. Outer lip usually not outer edge. Parietal callus weak. Columella concave to thickened. straight, smooth, excavated along entire length. Fasciole mm Description: Shell to 60 long, thin to moderately short, arched, usually bordered posteriorly by sharp ridge thick-shelled; adults ovate, spire 20-30 percent oflength. of growth-line chevrons. Siphonal canal short, narrow, Sutures slightly to moderately impressed. Shoulder even- open. ly, broadly rounded, rarely with weak sutural platform. Discussion: Species from Central and North America Spiral sculpture imbricate; about 13 primary cords be- once placed inAcanthina are notable for combinations of tween shoulder and spiral groove, variably alternating prominent and regular axial sculpture, glossy apertures. with secondary cords of varying strength. Five to seven Page 334 The Veliger, Vol. 46, No. 4 secondary spiral cords on base, three to four posterior to shoulder, with or without a single primary spiral cord. mm Outer lip in adults generally less than 4 thick, thin or slightly thickened and weakly beveled; outer edge crenulate; inner edge weakly dentate (seven to 12 teeth) or smooth. Labral tooth well developed. Columella straight to concave: excavated but not broadly. Fasciolar ridge poorly developed. Whorls of small juveniles with five to seven primary spiral cords posterior to spiral groove. Two spiral cords, one at shoulder and another anterior to shoulder, some- times more prominent (LACM 72-157-2e, Figure 5). Discussion: Acanthina monodon at present occupies in- tertidal and shallow subtidal habitats from the Chonos Archipelago of southern Chile to Santa Cruz Province, Argentina (Carcelles, 1954). The species has been found on upper Pleistocene terraces in Argentina (UNLP 1407b and UNLP 1407c, Figures 6, 7), but not in older deposits (Pastorino, 1994). Some Recent specimens from southern Chile have a slightly thickened and dentate outer lip (LACM 54679a, Figure 2). characters shared with many specimens ofA. unicornis. Material: LACM 72-157.2a. L 52.7, W 39.0; LACM 72- 157.2b, L 39.4, W 28.6; LACM 72-157.2c, L 30.6, W 22.4; LACM 72-157.2d, L 28.0, W 21.1; LACM 72- 157.2e, L 11.1, W 7.5; LACM 72-157.2f, L 9.5, W 5.7; LACM 54679a, L 50.0, W 35.3; LACM 54679b, L 46.0, W W 31.4; UNLP 1407a, L 42.9, 28.9; UNLP 1407b, L W W 42.9. 28.8; UNLP 1407c, L 27.5. 18.4; UNLP 1398, lot of six fragments. Occurrence: Upper Pleistocene to Recent. Southern Chile to southeastern Argentina. Acanthina unicornis (Bruguiere, 1789) (Figures 8-22) Buccinum imicorne Bruguiere. 1789:254. Monoceros crassilabrnm Lamarck. 1816:pl. 396, figs. 2a,b. Monoceros acuminata Sowerby, 1835:50. Figure 1. Distribution of Recent Acanthina unicornis and A. Monoceros citrinum Sowerby, 1835:51. monodon. Pacific NeogeneforearcbasinswithAca;zf/7;>zi3-bearing Monoceros costatum Sowerby, 1835:50. marine sediments are also shown. Monoceros globulus Sowerby, 1835:50. MonocerosambiguusSowerby, 1846:261, pi. 4, figs. 66, 67. Monoceros unicorne Sowerby. Hupe, 1854:194. Monoceros ambiguus Sow. Philippi. 1887:56, pi. 7, fig. 1. Figures 2-7. Acanthina monodon. Figure 2. LACM 54679a. apertural view, Xl.O. Figure 3. LACM 54679a, abapertural view, Xl.O. Figure 4. LACM 72-157.2c, apertural view, XI.12. Figure 5. LACM 72-157.2e, apertural view, X2.69. Figure 6. UNLP 1407b, apertural view, XO.96. Figure 7. UNLP 1407c, apertural view, X1.25. Figures 8-22. Acanthina unicornis. Figure 8. UWBM 97373, apertural view, Xl.O. Figure 9. UCMP 3102-d, apertural view, x2.18. Figure 10. UCMP 3102-a. apertural view, XI.87. Figure 11. LACM 75-32.4-b. apertural view, Xl.02. Figure 12. LACM 75-32.4-c, apertural view. Xl.30. Figure 13. MUSM INV 017, apertural view, Xl.O. Figure 14. UWBM 97373, abapertural view, Xl.O. Figure 15. LACM 75-32.4-d, apertural view, Xl.O.Figure 16. LACM 75-32.4-d. abapertural view, X1.07. Figure 17. UCMP 3102-b, abapertural view. X1.81. Figure 18. T. J. DeVries, 2003 Page 335 UNLP 26668. apertural view. XO.92. Figure 19. UCMP D-3735a. apertural view. XO.96. Figure 20. UCMPD-3735a. abapertural view, X0.96. Figure 21. UCMP D-3735f, apertural view, X1.25. Figure 22. UCMP D-3735f, abapertural view, Xl.25. 1 Page 336 The Veliger, Vol. 46, No. 4 Monoceros crassilabris Brug. Philippi, 1887:56, pi. 6, fig. 6. est acanthinine specimens that are unequivocally A. uni- Not Monoceros costatus ? Sow. Philippi, 1887:56, pi. 5, fig. cornis. 9 (correct identification is Chorus sp.). Herm (1969: pi. 17, figs. 7-9) illustrated two Pliocene Nucella (Acanthina) crassilabnim (Lam.). Carcelles, 1954: 257, pi. 1, figs. 1-4 (with lengthy synonymy). ^'acutninata-type'' specimens of.4. unicornis from Chile Nucella (Acanthina) crassilabrum crassilabnim (Lamarck). but identified them as Nucella (Acanthina) crassilabrum Herm, 1969:138, pi. 17, figs. 4a,b 6a,b. calcar, reserving the name 'W. (A.) crassilabrum acu- Nucella (Acanthina) crassilabrum acuminata (Sowerby). minata" for an early Pleistocene specimen that should be Herm, 1969:140, pi. 17, figs. 5a,b. assigned to his N. (A.) crassilabrum costata. Nucella (Acanthina) crassilabrum calcar (Martyn). Herm, Pleistocene specimens of non-imbricate, high-spired 1969:pl. 17, figs. 7-9. Acanthina crassilabrum (Lamarck, 1816). Dell, 1971:210. Acanthina from marine terraces in Chubut Province, Ar- Acanthina monodon crassilabrum (Lamarck, 1789). Wu, gentina (UNLP 26668, Figure 18) resemble some Chilean 1985:58, figs. 15-17, 23, 35-37, 50-51, 59, 71. specimens erroneously referred to Nucella (Acanthina) acuminata by Carcelles (1954). They have two primary Diagnosis: Thick-shelled; shoulder evenly rounded. Spi- spiral cords on earlier whorls, one at the periphery and ral cords numerous, unequally and irregularly developed, the other anterior to the first. These are joined on the usually subdued over entire body whorl. Inner edge of anterior of later whorls by two or three subdued primary outer lip sometimes strongly dentate. spiral cords. The outer lip is slightly thickened and some- Description: Shell to 60 mm long, broadly fusiform to times finely dentate. There is no evidence on 17 Chubut ovate, base of body whorl usually slightly constricted. specimens ofthe imbricate sculpture that characterizesA. Adults thick-shelled. Spire 25-30 percent of length. Su- monodon from Recent beaches or Pleistocene marine ter- tures appressed to impressed. Shoulder evenly rounded, races in Santa Cruz Province, Argentina. The Chubut specimens were found with specimens of Tegula atra rarely with sutural platform; periphery at or slightly an- terior to midpoint of body whorl. Spiral sculpture on (Lesson, 1830) and fissurellids now encountered only in adults of 20-25 spiral cords, either equally prominent or Chilean waters (G. Pastorino, personal communication. alternatingly primary and secondary, a few accentuated 2002). with chocolate brown color in modern specimens; usually Modern specimens ofAcanthina imicornis from Chile subdued or obsolete but sometimes strongly imbricate or occur in intertidal and sublittoral environments between rarely with two well developed primary spiral cords be- 29°S and 42°S (Carcelles, 1954; Dell. 1971). Fresh beach tween shoulder and periphery. Three to five subdued spi- specimens of A. unicornis (MUSM INV 018) from San ral cords anterior to spiral groove. Outer lip thick, strong- Juan de Marcona (15°27'S) confirm reports (Dall, 1909) ly beveled, sometimes strongly dentate, with eight to 1 ofthe species in southern Peru, as do observations oflive elongate teeth evenly spaced along aperture. Labral tooth intertidal individuals at Matarani (17°S; M. Rabi, written well developed. communication, 1997). Late Pliocene and Pleistocene Early whorls in juveniles with five to seven primary specimens are known from coarse-grained bioclastic de- spiral cords between suture and spiral groove; cord at posits throughout northern Chile (Herm, 1969) and south- periphery and another anterior to peripheral cord some- ern Peru (Muizon & DeVries, 1985; DeVries, 1986; Ort- times more prominent (UCMP 3102-d and UCMP 3102- lieb et al., 1990; Ortlieb & Machare, 1990). a. Figures 9, 10). Material: LACM 75-32.4-b. L 37.0. W 28.4; LACM 75- Discussion: The morphological diversity illustrated by 32.4c, L 30.5, W 21.8; LACM 75-32.4-d, L44.3, W 29.8; LACM MUSM Carcelles (1954) and lengthy synonymy generated by 75-32.4, lot of 14 additional specimens; stSs"hhoAoeal.wltvleeoardfocbf/uty4h.mAii(smcn1oapa8nnrt3too5ahdb)"iolnntea(e.msStouSibnwfoyyiewcrretobeorrycnb,otiyhgsen1(i8ad1z3in8i5fd3nf)5gi,)cdauiwlansitntytiidhnnotCgfeaubrirodscmteheehsildcnlirngeaiustbmeiit(en1fgrf9oor5rtuo4mhsm),e U3DI1,1NVC0LV2M-45P0b641,8.7-23,L,21,0WD12L8-V.d48,43,36.8.L8W27;-,11U13,W3.C6.L,1M;3PW73U.98.3C;991,.M0M22;WP-UaUS,33C10ML0.M28-;P1Ic5N,.MD1V,-UL3W0S73113M9.59,2a.,4I,D;WNVLUVC248020M11..2889P-,;, imbIrnidciavtiedusaplisralofcoArcdasntahnidnaa tuhniicckordneinstawtiethouttehre l"ipa.cumi- W37352c9,.2;L U(3C4.M3)P, WD-32763.56b;,ULCM45P.1,D-W373531d.,0;LU(C18M.4P), DW- nata-type" combination of imbricate spiral cords and a 16.6; UCMP D-3735e, L 37.9, W 27.5; UCMP D-3735f, ctehlilceks,de1n9t5a4t)e aountderspleipcilmievenstoodcacyurininsoPultihoecrenneChainldeP(lCeairs-- ULC2M8.P4W, DW-58221.68a;,ULC4M4P.5,D-W373351,.0;lotUoCfMWtPwoD-sp5e8c2i6mbe,nsL; tocene deposits of southern Peru (MUSM INV 017, Fig- 27.3, 22.3; UNLP 26668, L 43.0, 25.6; UNLP ure 13; also UWBM 97374) and Chile (LACM D-3735- 26669, lot of eight specimens; UNLP 26670, lot of nine a; Figures 19, 20). In Peru, imbricate adult specimens of specimens; UWBM 97372, DV 468-2, L 36.0, W 27.1; Acanthina occur in upper Pliocene strata (UWBM 97373, UWBM 97373, DV 812-1, L 48.0, W 34.1; UWBM UWBM W Figures 8, 14; also 97374) together with the old- 97374, JM84 319B, L 35.1, 25.7. T. J. DeVries, 2003 Page 337 ^-^^^^^ Figures 23-35. Acanthina triangularis DeVries, sp. nov. Figure 23. UWBM 97375, apertural view, XO.98. Figure 24. OSU 37358, apertural view, X1.04. Figure 25. OSU 37358. abapertural view, Xl.04. Figure 26. UWBM 97375. abapertural view, xO.98. Figure 27. OSU 37359, apertural view (suture distorted by growth ofbalanids), X1.05. Figure 28. UWBM 97376, apertural view, X1.32. Figure UWBM UWBM UWBM 29. 97376, abapertural view, X1.32. Figure 30. 97377, apertural view. XO.94. Figure 31. 97378, abapertural view, Xl.70. Figure 32. UCMP D-3735-d, apertural view, X1.83. Figure 33. OSU 37360, apertural view, X2.23. Figure 34. UCMP D- UWBM 3735, abapertural view, Xl.83. Figure 35. 97379, abapertural view, X1.62. Occurrence: Upper Pliocene to Recent. Southern Peru to Nucella {Acanthina) crassilabrum costata (Sowerby).Herm, southern Chile. 1969:139, pi. 17, figs. 2a, 2b, 3. Not Monoceros cos- tatum Sowerby, 1835. Acanthina triangularis DeVries, sp. nov. Diagnosis: Body whorl angulate or biangulate in both (Figures 23-35) adults and juveniles. Sutural platform planar or slightly concave. Two primary spiral cords or keels; other spiral Acanthina triangularis DeVries, 1986:592, pi. 33, figs. 9, sculpture usually subdued or absent. 12; pi. 34, figs. 2, 6, 7, 10, 12; pi. 35, figs. 1, 2, 3 mm (unpublished dissertation). Description: Shell to 70 long; biconic. Adults thin ' Page 338 The Veliger, Vol. 46, No. 4 to thick shelled. Spire 25-30 percent of length. Sutures 81°10'W 8I°00'W appressed near shoulder. Sutural platform planaror slight- ly concave, broad, inclined 30-45 degrees. Shoulder an- gulate; periphery located one-third to one-half length of body whorl from suture, usually coincident with shoulder. Spiral sculpture of primary spiral cord at shoulder, sub- dued to keeled; another primary cord anterior to first, of- ten reduced to spiral swelling or obsolete. Additional weakly developed secondary spiral cords and tertiary threads, usually obsolete or nearly so; rarely, imbricate. Outer lip angulate or biangulate; sometimes moderately thickened, beveled, neither crenulate nor dentate. Labral tooth deep within outer lip, often projecting from internal spiral ridge. Columella concave, deeply but not broadly excavated. Pseudumbilical callus thick, with arching ridge parallel to fasciole. - 4°50'S Juveniles with three subequal primary cords on first whorls of teleoconch; cords differentiating on later whorls, one becoming secondary cord on expanded su- tural platform, another a primary cord on shoulder, and ILocality - - third a primary cord anterior to shoulder cord. Additional secondary cords on largerjuveniles and subadults insert- SCALE ed on sutural platform (one cord), between two primary cords (one cord), between anterior primary cord and spi- Figure 36. Type locality (DV 272/DV 273) oi Acanthina tri- ral groove (four to five cords), and on base (two to three angularis DeVries. sp. nov. Outcrops of basal bioclastic sand- cords) (OSU 37360, Figure 33). stones with A. triangularis occur in quebradas between La Brea and Amotape. Discussion: Acanthina triangularis is not the same cos- tate taxon as Sowerby's (1835) Acanthina costata from Chile, which has been synonymized herein with A. uni- late Pliocene age for specimens of A. triUanWguBlMaris from cornis. Costate specimens ofA. unicornis, modern (Car- a marine terrace in sUouWthBerMn Peru (e.g., 97375, celles, 1954:figs. 36-43; UCMP 3102-a, unfigured) and Figures 23, 26, and 97376, Figures 28, 29) is fossil (Herm, 1969:pl. 17. figs. 5a, b, termed Nucella also indicated by the presence of the mollusks Trachy- (Acanthina) acuminata: UCMP 3735-f, Figures 21, 22) cardium procerum domeykoanum (Philippi, 1887), have shoulders that are more rounded, outer lips that are Amiantis domeykoana (Philippi, 1887), Concholepas no- more often dentate, anterior profiles that are less con- dosa Moricke, 1896. and Thais chocolata (Duclos, 1832) stricted, and spiral cords that are more numerous than is (Herm. 1969; DeVries, 1995). typical for specimens ofA. triangularis. Type locality: Strata between Quebrada Songora, 14 km Adults ofAcanthinatriangularis are distinguished from southeast of La Brea, and Quebrada Cardo Grande, 10 broadly rounded specimens ofthe more recent A. unicor- km north of Amotape, northwestern Peru (localities DV nis and smooth-shelled, tightly rounded specimens ofthe 272/273; Figure 36). older A. obesa by their planar sutural platform aUndWBtwMo Material: OSU 37358, holotype, DV 272/273, L 38.0, W prominent primary spiral cords (or rarely, one: 28.7; OSU 37359, paratype, DV 272/273, L38.1,W28.4; 97378, Figure 31). Juveniles of A. triangularis and A. OSU 37360, paratype, DV 273-1, L 15.3, W 11.8; omboersea eavreenliydenrtoiucanld,edwhaenrdeashavtehosseevoefralA.pruniimcaorrynisspiarrael MINUVS0M14,INDVV041233,-3D,VL4(2337.-23),,LW(2360..55);,MWUS24M.6;INMVU0S15M, ctoherTdhsR,ei'ortaytCphheeirrlato,chaalnniottrywtohf.eorrnAcPcemrtuhi(n4a°4t0r'iSa)n,guilsarais,signnoirftihcaontf D(3,1V7L.64)2263.W-63,W1L7.262(;.503U;.5UW)CBWMM4P1.9D47-;337M753,U5-SDdM,VCI4oN2q3Vu-i30m,1b6oL,,D6C0hV.i9l,4e2,W3L- n(oOrStUhw3a7r3d58r,anFgiegureexste2n4s,io25n; fOoSrUAc3a7n3t5h9i,naF.iguSrpeec27i)meanrse 4U4.W5;BMUW9B73M77,97D3V76,42D3-V3, 4L23-431,.6,LW(313.26).,0;WUW2B3.M1; found in the lowest pebbly sandstone beds of the Taime 97378, DV 423-3, L (19.8), W 16.3; UWBM 97379. DV formation (DeVries, 1986, 1988) with other mollusks, W 423-3, L 21.1, 17.6. e.g., Chorus blainvillei (d'Orbigny, 1842) and Hermines- pina mirabilis (Moricke, 1896), that suggest a late Plio- Occurrence: Late early to late Pliocene, northern Peru to cene age (DeVries, 1997a; DeVries & Vermeij, 1997). A northern Chile. T. J. DeVries, 2003 Page 339 Acanthina obesa DeVries, sp. nov. Type locality: Aguada de Lomas, east of intersection of Panamerican Highway with road to Lomas. East slope, (Figures 37-52) DV south of abandoned HierroPeru road (locality 369; Acanthina obesa DeVries, 1986:591, pi. 34, figs. 1, 2 (un- Figure 53). published dissertation). W Material: OSU 37361, DV 361-5, L 25.3, 19.2; OSU Diagnosis: Shell broadly trigonal; shoulder tightly round- 37362, DV 361-5, L 27.4, W 21.7; OSU 37363, DV 361- ed; usually smooth. Spire usually low. Juveniles with two 5, L 24.9, W 20.4; OSU 37365, holotype, DV 369-3, L prominent spiral cords at and anterior to shoulder 40.9, W 36.3; OSU 37366, paratype, DV 370-2, L 35.5, W W Description: Shell to 50 mm long; broad, width and 31.3; OSU 37368, paratype, DV 370-2, L 16.2, height of body whorl usually about equal. Base barely 13.1; MUSM INV 010, DV 571-1, L (39.3), W 31.9; constricted. Shell thin to moderately thick. Spire usually MUSM INV Oil, DV 1230-1, L 38.4, W 31.3; MUSM 10 to 20 percent of shell length. Sutures appressed. Su- INV 012. DV 57W1-1, L 19.6 W 14.8; SGO.PI.5750, Lo tural platform broad, slightly convex, typically inclined Abarca, L W18.7, 15.0; VE09261a, Aguada de Jahuay, less than 30°. Shoulder usually tightly rounded, periphery L (46.2)W, 40.9; VE09261b, Aguada de Jahuay, L atmidpoint ofbody whorl. Spiral sculpture usually absent W(40.4), 32.1; VE09261c, Aguada de Jahuay,WL 29.7, or consisting of weakly developed spiral cords separated 25.8; VE09313, Aguada de WJahuay, LU2W9.B4,M 23.5; by thin grooves. Rarely, sculpture with mix of 25—35 VE09262, Cerro Alto, L 34.3, 27.9; 97380, ousrcmaetllhyliacske,bcrooofnatddelanyr,ybedcveoeerlpdelsdyaienfxdclaattveterartt,ieadir.nysitdhereeaddsg.eOsumtoeorthl.ipCtohli-n D3(41V9.)8,;57UW1-W13,B5.LM4;290U7.32W83WB,MD15V.987;336U802-W,2,BDMLV29917.23838,01-W,1,D2V4L.653.761.0-,1,WL Juveniles with three primary or secondary spiral cords Occurrence: Upper Miocene to lower Pliocene, southern on sutural platform; primary cord on shoulder; primary Peru to central Chile. cord anteriorto shouldercord with intercalated secondary spiral cord; five or six secondary cords on body whorl; Acanthina riigosa DeVries, sp. nov. three secondary spiral cords on base (OSU 37368, Fig- ures 38 and 40; UWBM 97380, Figure 41; MUSM INV (Figures 54-57) 012, Figure 43). Diagnosis: Shell globose. Axial sculpture ofeight to nine Discussion: A low spire, broad sutural platform, and weak collabral folds; nodes at intersections of axial folds tightly rounded shoulder give specimens of Acanthina and two primary spiral cords. obesa a profile with a distinctive posterior inflation. mm Description: Shell less than 40 long, thick, globose. Adults are usually entirely smooth, in contrast with other Base slightly constricted. Spire moderately elevated, 15— species of Acanthina. Large specimens with high spires (UWBM 97381, Figure 49) can be distinguished from 20 percent of shell length. Sutures appressed. Sutural platform broad, slightly convex, inclined 40 to 50 de- specimens ofA. unicornis by their smooth shell and thin- grees. Shoulder angulate to sharply rounded, coincident ner, non-dentate outer lips. Specimens with more fully (UWBM with periphery, about two-fifths distance from suture to developed spiral sculpture 97382, Figures 42, base of shell. Axial sculpture of eight to nine collabral 44) can be separated from those of older A. katzi and folds, forming nodes at intersections with primary spiral youngerA. unicornis by their posterior inflation. Strongly cords. Spiral sculpture oftwo subdued primary cords, one sculptured early Pliocene specimens (OSU 37361, 37362, at periphery, second anterior to first; two weak secondary 37363, Figures 50-52), however, do resemble those of cords, one on sutural platform, another at midpoint of the younger A. triangularis. body whorl; and numerous tertiary threads between su- The oldest Peruvian specimens of Acanthina obesa come from beach coquinas on the eastern flank of Que- ture and base. Outer lip thick, beveled, usually smooth, rarely dentate. Labral tooth short. Columella broadly ex- bradaRiachuelo (Figures 42, 44) that overliemiddle Mio- cavated. cene strata containing specimens of A. katzi. Farther Juveniles biconic; axial and spiral sculpture as above, south, specimens ofA. obesa (Figures 37, 39) overlie ash but more sharply defined and generally imbricate beds at Alto Grande and Aguada de Lomas with ''^K- (UWBM '"'Ar dates of 9.5 Ma and 8 to 8.8 Ma, respectively (Mui- 97385, Figures 56, 57). zon & DeVries, 1985; Muizon & Bellon, 1986). Numer- Discussion: Specimens of Acanthina rugosa are distin- ous specimens of A. obesa (UWBM 97383, Figures 45, guished from all other species ofAcanthina except those 46) are found in lowermost Pliocene strata at Sud-Sacaco of the oldest, A. katzi, by possessing numerous collabral (Muizon & DeVries, 1985). The single specimen of A. folds. Like specimens of A. katzi, they have well devel- obesa known from Chile (SGO PI 5750, Figures 47, 48) oped, well differentiated, and imbricate primary and less- was found by the author in upperMiocene shell and grav- er spiral cords, but unlike those ofA. katzi, they have a el banks at Lo Abarca (Covacevich & Frassinetti, 1990). broadly excavated columella and a broader sutural plat- Page 340 The Veliger, Vol. 46, No. 4 Figures 37-52. Acanthiua obesa DeVries, sp. nov. Figure 37. OSU 37365, aperturai view, Xl.21. Figure 38. OSU 37368, apertural view, XI.91. Figure 39. OSU 37365, abapertural view, Xl.21. Figure 40. OSU 37368, abapertural view, X1.91. Figure 41. UWBM UWBM MUSM 97380, apertural view, XI.71. Figure 42. 97382, apertural view, XI.37. Figure 43. 012, apertural view, XI.77. Figure UWBM UWBM UWBM 44. 015, abapertural view, X1.37. Figure 45. 97383, apertural view, Xl.32. Figure 46. 97383, abapertural UWBM view, Xl.32. Figure 47. SCO.PI.5750, apertural view, X2.10. Figure 48. SGO.P1.5750, abapertural view, X2.10. Figure 49. 97381, abapertural view, XO.98. Figure 50. OSU 37361, apertural view, XI.57. Figure 51. OSU 37362, apertural view, X1.41. Figure 52. OSU 37363, apertural view, xl.56. T. J. DeVries, 2003 Page 341 Figure 53. Type locality (DV 369) ofAcanthina obesa DeVries, sp. nov. form, characters shared with specimens ofthe youngerA. (Moricke, 1896)] suggest a late Miocene age (DeVries. & obesa. 1997a; DeVries Vermeij, 1997). The only collection of Acanthina rugosa comes from Type locality: South side of Quebrada Huaricangana, bioclastic conglomerates very near the base ofa Neogene above basement platform. Sample DV 387-3 (Figure 58). Hsubaeruapceasetreini5moi8cenn)ans.notugnlTaatthnrhhaeaetb,uscrfwoeofhuaenitrrgshclbhos5emi0tsde0ewerreaovmetfeenedtQetubarhetesebdtdruheaeprpdeotsatstioisHmtteuhcaaelornofidspcedetaeahnptkegoosaooinpiftageiCnno(eenFroPiruagao-s-s WM210a27..t654e.;;r1iM;MalUUU:SSWUMMBWMIIBNNM9VV7300980075893,,,84ppD,aarrVhaaottl3yyo8ppt7eey-,,p3DDe,,VVloDt33V88o77f3--8335.7,,-LL3,((L2260..(502))9,,.6WW), cific Ocean. Other mollusks from the same horizon [Cho- Occurrence: Upper Miocene to lower Pliocene, southern rus frassinetti DeVries, 1997; Herminespina philippi Peru.