THE NAUTILUS 121(4);163-181. 2007 Pase 163 Late Cenozoic Tegulinae (Gastropoda: Trochidae) from southern Peru Thomas DeVries' J. Burke Museum ol Naluial llistorx aud Culture Unhersih- ofWashiH^ton Seattle, \\"A 98195 USA ABSTRACT ince (Keen, 1971; Alamo and Valdivieso, 1997), although with mostly different species, and is more speciose tlian Four new fossil tegulines (Gastropoda: Trocliidae) are de- the teguline fauna of the Magellanic Faunal Fro\4nce s(lcartiebeMdiofcreomnesotuothlaetrenPPleiorcuen[eC)h,loIrnotsitsotioimuiiiqpuiiijiujiuaanneewwsgpeencuise,s (four species), which is a subset of the Peiinaan fauna newspecies(earlvPliocene),Cantallocostomapanistostiim new (Forcelli, 2000). genus, new species (late Miocene to earK- Pliocene), Teguhi The geological record ot tegulines from the Peruvian (s.l.) masiasi new species (earls' to middle Miocene)], as are Faunal Province is meager. Tegula luctuosa, T. atra, and Pliocene and Pleistocene occurrences of the extant CJiloro- T. tridentata are listed in Herm's (1969) study of stoma atnim (Lesson, 1830), C. hicttiosum (d'Orbigny, 1841), Pliocene and Pleistocene mollusks from Chile. Tegula Cantallocostoma qticidricostatum (Wood, 1828), Aiialhisioiua luctuosa and 7'. melaleucos were reported from upper patagoniciim (d'Orbigny, 1835), T. (s.l.) melaleticos (Jonas, Pliocene and Pleistocene beds of northern Peru 1844), and T. (s.l.) tridentata (Potiez and Michaud, 1838). (DeVries, 1986). Four new species ot tegulines, consid- TanhceiseentdattoabsehoawPtlHiaotcdeineePseirsutveriagnrcohulportoostCoamriinbebegarno-uAptliasnttioco ered to have early to middle Miocene age (DeVries and agadiistomines; indicate additional eastern Pacific groups of Frassinetti, 2003; Finger et al, 2007), have been de- teguUnes e.^ist with roots reaching into die Miocene; and fiu- scribed from the Navidad F'ormation of central Chile by dier demonstrate the success of A. patagoniciim as a wide- Nielsen et al. (2004). spread and long-h\ed teguline in australwaters. A small radia- This paperdocuments fournew fossil species and sev- tion ofPeru\ian clilorostomines during the late Pliocene coin- eral previously known Recent species of tegulines in cided with a nioUuscan mass extinction event in the Penu-ian Neogene deposits ofsouthern Pena, including the oldest Fauna] Pro\ince. knovvai teguline from Peru, the early Miocene Tegida Additional Keywords: Mollusk, Tegula, Chlorostoina, Agathi- (s.l.) masiasi new species; creates two new genera of stnina. Miocene. Pliocene. Cenozoic, Pisco Basin, Taxnnoin\- teguhnes, Cantallocostoma, new genus, and Intistoma, new genus, each with a newly described Neogene fossil species in the Pemvian Faunal Province, Cantallocos- IXTRODUCTIOX toma panistosum ne'w species and Intistoma pirqua new species, and each having a modem representative in the Se\"en e.xtant teguline species (Gastropoda: Trochidae) eastern Pacific Ocean, the Penivian C. quadrieosiatum inhabit the cool coastal waters of the Peru\iaii Faunal and Californian /. mireotinctum (Forbes, 1852); adopts a Pro\ince.The\-are, accordingto theirtraditional nomen- full generic status for Tegula (Chlorostoma); and pro- clature, Tegula atra (Lesson, 1S30), T. luctiiosa vides evidence for a late Miocene origin ot a westem (d'Orbigny, l841), T. etin/omphala (Jonas, 1844), T. ig- South American group ot Chlorostoma species. nofa Ramirez-Bohme, 1976, T. tridentata (Potiez and Michaud, 1838). T. quadricostata (Wood, 1828), and T. patagonica (d'Orbigny, 183.5) (Alamo and GEOLOGY \'aldi\ieso, 1997; \'eliz and \'asquez, 2000). An eighth species, Tegula melaleucos (Jonas, 1844), a species from warmer waters of northern Peru, does appear rarelv at The late Cenozoic marine stratigraphy ot southern Pe- soutliem Peruvian latitudes. This teguhne fauna is as mvianforearcbasinshasbeendescribedbyDunbaretal. diverse as that ofthe warm-water Fanamic Faunal Pn)\- (1990) and DeVries (1998). Teguline-bearing deposits crop outwest ot Nazca and throughout the Sacaco Basin (Fig. 1). These bioclastic conglomerates and sandstones, Mailing address: P.O. Box 13061. Burton, WA 9S013 USA which were assigned to the La Planchada and Pisco for- Page 164 THE NAUTILUS, Vol. 121, No. 4 Subfamily Tegulinae Kuroda, Habe and 0\ama, 1971 Genus Tcg^iila Lesson, 1835 Type Species: Trochus clcgans Lesson, 1835 (by monot\iw) (= Trochiis pelUsseij>entis Wood, 1828). Re- cent, Pacific coast of Central America. Remarks: Resolving the difficult subfamilial place- ment of Tegulinae \\ithin Trochoitlea (Hickman and McLean, 1990; Bouchet and Rocroi, 2005) is beyondthe scope ol this paper. Teguki itself has been assigned to three ditferent subfamilies o\er the past centuiy, but most taxonomists now refer it to the undiagnosed Teg- idinae Kuroda, Habe, and Oyama, 1971 (Hickman and McLean, 1990). Onlyone extantteguline species, Tegula pclUsscrpcntis (Wood, 1828), pertains to Tegula (s.s.) (Keen, 1971). Its combination of features (densely packed and heavily beaded spiral cords, tightly t\\isted Camana Basin columella, and nimierous teeth extending from the col- _L _L I , umella across the apertural floor) is not seen in other Figure 1. Location ofonshore portions ofCenozoic iorearc species assigned to Tegula, so T. pelUsserpentis is herein basiu.s in southern Peni. Newfossil species oi Tegula are from considered to be the monotypic representative ofTegula Cenozoic deposits west and south of Nazca. (s.s.), an opinion shared bv ]. H. VIcLean (pers. comm., 2006). Taxa often considered as subgenera of Tegula, mations by Beaudet et al., 1976, and Muizoii and e.g., Chtnivstoiua and Agathistoma, are elevated to ge- DeVries, 1985, respectively, I'epresent high-energs' fore- neric status in this paper in accord with the practice of shore and intertidal environments lying close to moun- some audiors (e.g., Arnold, 1907; Higo et al., 1999) and tainous paleo-shorelines and cjuieter foreshore and inner the opinitjn of ]. H. McLean (pers. comm., 2006). shelf environments lying hundreds to thousands of meters from paleo-shorelines defined h\' the beginning Genus Chlorostonia Swainson, 1840 ofthe Andean foothills. TApe Species: Trochus argi/rostonuis Gmelin, 1791 (In subsL'(|uent designation, Herrmarmsen, 1846). Re- MATERIALS AND METHODS cent, [apan. Most Penivian specimens described in this stucK were Remarks: Swainson (1840) erected Chloro.stoma as a found by the author. Comparative material was studied subgenus of Trochus Linnaeus, 1758, to include species at the Natural History Museum of Los Angeles County, with a "remarkably oblique" aperture, a very deep um- Los Angeles, California, USA (LACM). Localitv and bilicus, and one or two teeth on the outer (= lower part sample descriptions are listed in the appendix. Lengths of inner?) lip. He assigned t^vo species to Clilorostoma: (L) and widths (W) are measured in millimeters, with Trochus (Chlorostoma) argijrostonia (= Tegula argi/ros- dimensions ofbroken specimens enclosed b\' parenthe- toma ol authors) and Trochus (Chlorostoma) unibilicaris ses. A non-standard orientation for plicjtograplis ofsome [= Gibbula umhilicaris (Linnaeus, 1758)]. Herrmannsen specimens has been necessan- to reveal important char- (1846) iniplicitK-limitedthe definition oiChlorostoma by acters. Some figured specimens were coatedwith ammo- choosing T. argi/rostoma as the type species. Examina- nium chloride. Types and numbered specimens, includ- tion of LACM material from tlie western North Pacific ing those figured, are deposited at the Orton Geological Ocean [C. argi/rostoma, C. lischkei (Tapparone-Canefri, Museum, Ohio State University, Columbus, Ohio USA 1874), C. nisticum (Gmelin, 1791), C. turbinatum (A. (OSU); the Departamento de PaleontologiadeVertebra- Adams, 1853), C. xaiuhostigiita (A. Adams, 1853)] shows dos, Museo de Historia Natural, Universidad Nacional that adult chlorostoniine umbilici can be either open or Mayor de San Marcos, Lima, Peru (MUSM INV): and closeil. In the view of Grant and Gale (1931), Chloros- the Burke Museum ofNatural HistoiAand Cultine, Uni- toma shouldbe further restricted to toothed species hav- versity of Washington, Seattle, Washington (UWBM). ing \'entricose whorls, a nacreous interior, and a dark The prefixes "DV" refers to DeVries localities, "JM" to puiplish exterior. This diagnosis makespossible the iden- localities of Machare (Instituto Geologico Minero y tificationof Recentchlorostoniinetaxafrom bothsidesof Vletalurgico,J.Lima, Peiii), and "WJZ" to localities ofW. the North Pacific Ocean and in the eastern South Pacific (. Zinsmeister (Purdue Universif-y, Indiana, IKSA). Ocean. Chlorostoma atrum (Lesson, 1830) SYSTEMATICS (Figures 2-4, 7-9, 13) Superfamily Troclioidea Rafinesque, 1815 Trochusliter Lesson, 1830, vof 2, pt. 1, p. 344, Mollusques, pi. Familv Trocliidae Ra(iiics(|iii', 1815 16, fig. 2; d'OHii^nv, ISI(» 101) Pliilippi ISIt Ahhildun- T. De\"ries, 2007 Pasje 165 Figures 2-4, 7-9, 13. Ciduro.stoiiui atriiiii (Lesson, 1830). 2. UWBM 97863, DV 1372-1, Recent, oLilique spire view, width = 9.8 mm. .3. UWBM 97855. D\-398-1, Recent, basal view, widtli = 30.7 mm. 4. UWBM 97863, oblique basalview. 7. MUSM INV 126, DX 1252-1. earl\- Pleistocene. obUque spire view, width = 27.1 mm. 8. UWBM 97855, apertural \iew, 9. I'WBM 97860, Paracas Hotel, Recent,lateralview,length =22.2mm. 13. UWBM97860,obliquebasalview,widdi = 23.4 nmi. Figures5,6. Chlomsfoma funebralis (A.Adams, 1855). L^VBM 97862.southofLaJoUa,CaHfomia, Recent,width = 13.4mm. 5.ObUquespireview. 6.ObUque basal\ie\\-. Figures 10-12, 14-19. Clilorostoma luctuosum (d'Orbigny, 1841). 10. LTWBM 97864, Paracas Hotel. Recent, lateral \iew, lengdi = 20.2 mm. 11. UWBM 97865, Paracas Hotel, Recent, apertural view, length = 20.5 mm. 12. UWBM 97866, Hueco LaZorra. Peru. Recent, apertural view, length = 21.6 mm. 14. UWBM 97864, oblique basal view, v\idth = 23.7 mm. 15. UWBM 97S65. basal view, width = 24.4 nun. Figure 16. l^WBM 97866. obUque spire view, width = 28.5 mm. 17. UWBM 97867, Lomas, Peru, Recent, oblique spire view, width = 26.4 nun. 18. UW'BM 97867, oblique basal view. 19. UWBM 97868, Chile, Pleistocene, obUque basal view", vviddi = 37.3 mm. Figures 20, 21. Cldonistoma cun/ompluihiin (Jonas. 1844). UWBM 97871, DV 1599-1, Recent, lengdi = 26.0 mm. 20. .Apertural view. 21. Lateralview. Figures 22, 2.3. Clilorostoma ignotmn (liainirez-BoUme. 1976). Ur\MBM 97872, Pellehue, Cliile, Recent, width = 17.5 mm. 22. Oblique spue view. 23. Basal view. Figures 24-26. Chlorosloirui qiiipua new species. 24. UA\'BM 97873, DV 1254-Bal 6, late early Pliocene, basal view, width = 17.3 mm. Note faint protractive stripes on base. 23. MUSM IX\' 136. D\' 571-1, svntvpe, late Miocene, oblique spire view, width = 17.0 mm. 26. UWBM 97873, oblique spire view. Page 166 THE NAUTILUS, Vol. 121, No. 4 gen und beschreibungen neuer oder wenig gekaiinter beneath the umbilical callus and terminates on the edge Conchylien, v. 1, p. 188, pi. 5, fig. 6; Philippl 1846, Die of the columella as a thickened tooth. A weakly devel- Kreiselschnecken oderTrochoideen, p. 198, pi 30, fig. 1: oped white outer cord traces the boundaiy of the um- Hupe, 1854, p. 142,—Malacologia, pi 4, fig. 2. bilical area and becomes flattened on the columella, not Monodonta atra Lesson. Potiez and Michaud, 18.38: 319, pi quite protiiiding far enough to produce a tooth. Rarely, 29. figs. 14, 15. one or two weak spiral spurs develop between the two Tegula atra Lesson.—Dall, 1909: 239, pi 24, fig. 4; Cairelles and Williamson, 1951: 262;—AldeaandValdovinos, 2005: fig. SB. umbilical cords in a nacreous area that lies adaperturally CJilorostoma ater Lesson. Morch, 1850: 20. — of a thin, glossy, umbilical veneer. They, too, do not Tegula (Chlorostoma) atra (Lesson, 1830). Marincovich, extend far enough to produce columellar teeth. 1973: 24, fig. 42; Alamo and Valdivieso, 1997: 13, fig. 25; Chlorostoma luctuosum (d'Orbignv, 1841) Guzman etal., 1998: 35, fig. 22; Forcelli, 2000: 61, fig. 87; (Figures 10-12, 14-19) Veliz and Vasquez, 2000: 759, fig. IB. Trochiis moestusJonas, 1844: 113; PhiHppi, 1846, Abbildungen Trochus- hictuosu.i d'Orbignv, 1841, v. 5, p. 409, pi 76, figs. und besehreibungen neuer oder wenig gekannter Con- 16-19; Philippi, 1846, Die Kreiselschnecken oder Tro- chylien, V. 2, pi. 6, fig. 5; Philippi, 1846, Die Kre- choideen, p. 153, pi. 25, figs. 4, 5; Hupe, 1854: 143. iselschnecken oder Trochoideen, p. 199, pi. 30, fig. 2; Tegula luctuosa Orbigny.—Dall, 1909: 239; DeVries, 1986: Hupe, 1854; 147, Malacologia, pi. 4, figs. 3, 3a, .3b. 512,pi. 27,figs. 3,4; Guzmanetal., 1998; 36,fig. 23;Veliz Tegula moesta (Hupe, 1854).—Dall, 1909: 239; Alamo and and Vasquez, 2000: 762, fig. ID; Alamo and Valdivieso, Valdivieso, 1997: 14. 2000: 14: Aldea and Valdovinos. 2005: fig. 8E. Chlorostoma minor Morch, 1850; 20. Diagnosis: Shellwidth to 35 mm. Lastwhorlwith one Diagnosis: Shell width about 40 mm; last whorl to three spiral cords or keels. Adult umbilicus open; ju- broadly rounded, mcluding shoulder; keels lacking. Um- venile umbilicus usually open; imibilicus with two well bilicus oi juvenile and adult shells white, closed; umbih- exposed umbilical spiral cords, the adaxial cordterminat- cus with two well exposed spiral cords, the ackixial cord ing in a columellar tooth. WtocMUeefanWrtt2me,;BirnLUiMaatW=li9nB72mEg85Mx.5Vi0a8n,,Bm9aiW7MD8nc5oVe7=ld,u:13m40eD.1l78Vl;U-a1DrWU1,2VtBW5loa2oMBt-de1iMs,.t97PL98l75i85=o5,c6e1,n1D.eDV8,.VL3W938=9-8=1-(,1212,8.R.2le6)o-;,t PrHMHaloaucitteaoeescclre,oHinoeaRLt,leeaclLeE,ZnxotR=ar,ermca1iLe,9nn.5te=R,,ed2c:WL0e.n3=t,=,O2W0S2L.3U5.,==9;3W2723.U1.5.=796W;6,2,B4UW.MD4W;V=BU97M22W8846.0BW549-;,72M83U6,P95aW7,rl8aaBt6cPe6aaMs,s-t WoUM(=1f9UW22.==;32SB.)3U,34MM06;.WW.26U;9IB;7=NWM8MV26B7U39.,M1S712;86MD6,M90V79,UI8D76NP18SV13a5V,M7r92a.11-cI2I21ap58N,su,2Vn-HR,1Do4e,1t6Vc2Ce3e7hel-n,4ia,1tl6r,D,e3lR,-ylVe1aLct,RPe1elen=4lecPt1oiel,t88sen.-tiot0L1os,f,,ct=e2oWL.lnco2eet2=,n.=e2o3,L;1f9..lW482o=,;;t W9m(M=270UV48)36=,SB.71W,M3M;7.ML3I=9o;UN7m28VSU8a6.sM84W1,;3B0cIUW,lMNJuWJVmZpMB9.173M28849R62,5e299,0c,D7,e8CVnDh7Pti0Vl.,l,e3e8i,W3L2s-8JtP12ol,Z=-ce1eLi,n3s(e42t=,P50o.l,c26lee0)oiLn.t,se6t,,o=ofcWL(e23n.5=e==.,0)232,61L6...W418=;;, Occurrence: Late Pliocene to upper Pleistocene: Occurrence: Late Pliocene to middle Pleistocene: northern Pem to Tongoy, central ChUe. Recent: Galapa- southern Peru to southern Chile. Late Pleistocene: gos Islands to Concepcion, central Chile (southern limit southern Peru, Chile, southern Argentina. Recent: from L,\CM collections). northern Pem to southern Chile, southernArgentina (G. Pastorino, pers. comm., 2002). Remarks: Specimens of Chlorostoma luctuosum are large and puip)le-black, either entirely or dorsally, only. Remarks: Specimens of CJilorostoma atruin can ex- mm Adult specimens are generally distinguished from speci- ceed 40 in width and are generally smoodi-shelled mens of C. atrum hvha\ing an open umbilicus and from and purple-black, either entirely or dorsally, only. The both C. atnim and C. eunjomphalum by having one to last whorl is always broadly rounded; it lacks the keeled three primary spiral cords or keels: one near the base of spiral cords present on specimens of C. luctuosum The . the whorl, forming the periphery (Figure 12); another base on some specimens of C. atrnm has weak spiral about one quarter ofthe distance anteriorly from suture threads; the spire ofsome also has one or more narro\v to suture (Figure 12); and athird occasionally developed spiralgroovesthatproduce as manyas 15inten-eninglow just anterior to the periphery (Figure 19). Some speci- broad spiral cords (Figure 7), not unlike the spiral sculp- mens ofC luctuosum are alsocoveredwithtertiaryspiral ture of the Cahfomian C. funebralis (A. Adams, 1855) threads (Figures 17, 18). The thin umbilical veneer and (Figures 5, 6). columellar teeth are identical to those on specimens of The adult shell of Chlorostoma atrum is usually dis- C. atnini, as are the umbilical cords, except that theyare tinguishedfrom thatofotherPeruvianchlorostominesby exposed coiling deep into the umbilicus. its closed umbilicus. [G. Collado (pers. comm., 2005) notes that juveniles of C. luctuosum and adults of the Chlorostoma curijiunpluilum (Jonas, 1844) small Tegula (s.l.) triclentata occasionally have closed (Figures 20, 21)' umbilici.] The white umbilical area on specimens of C. Trochus cuii/oiiijilinliis Jonas. 1844: .113; Philippi, 1844. Abbil- alrum has two spiral cords. A white innercord rises from dimgen luid beschieibungen neuer oder weniggekannter ' T. De\'ries. 2007 Pa£i;e 16" spiral cortis or keels. On some specimens oi C. luctiio- stiin, however, including Recent Peruvian and Chilean LACM examples from collections, Pleistocene Chilean specimens from WJZcollections, andupperPliocene Pe- ru\ian specimens from northern PeiTi, the spiral cords are soweaktluit assigningthe materialtoC.luctuosiim or C. ciini()mj)h(iltn)i is problematic. Chlomstoma ignotuni (Rarnfrez-Bohme, 1976) (Figures 22, 2.3) Tegula ignola Runu'rez-Bolniu-, 1976: ,3, figs. 1-6; Forcelli, 2000: 61, fig. 88; Veliz and Vasquez, 2000: 762, fig. IF; Aldea and Valdovinos, 2005; fig. 8D; CoUado and Brown, 2005: 131. Diagnosis: Shell width to 30 mm. Outer layer slate colored. Sculpture consists ofseveralwelldeveloped, un- beaded, primaiy spiral cords. Umbilicus open. UWBM Material Examined: 97872, La Rinconada, W UWBM Pelluhue, Chile, Recent, L = 11.1, = 17.5; W 97906, Ipun, Chile, Recent, L = 29.4, = 32.6. Occurrence: Recent; centra! to southern Chile. Remarks: Chlomstoma ignotum is faiown only as a Recent species from Chile (e.g., Collado and Browai, 2005). Specimens of C. ignotum differ from those of other extant chlorostomines of the Peru\ian Faunal Province in twosignificant respects: theylackthepui-ple- blackouter shell layerofC. atnim, C. luctuosum, and C. SC\LE LOMAS eun/omphalum, beingratherslate colored, andtheyhave Conlourinlenalistndm numerous, prominent, well-differentiated primary and secondan' spiral cords between the periphen' and suture Figure 41. T\pe localih (DN"571-1) ol Chhinysfoiua qtiijuui and to a lesser extent on the base. The well-e.xposed ne\\' species. umbilical cords, thin umbilical veneer, and columellar teeth ai"e identical, however, witla tliose ofother species Conchvlien. \-. 2, p. 27, pi. 6, fig. 4; Pliilippi, 1846, Die of South American Clilorostoma, and the strong spiral TegulKareeiusne/losmcpllmieaclkae(nJoondaesr.T1r84o4c)h.o—idCeaernc.elpl.es15a.5n,dpiW.il2H5a,mfsiog.ii7,. cords (Figrn-e 22) are like those seen on rare specimens ofearlv Pliocene C. qiiipua new species (Figures 25, 26; 1951: 262. — Tegula eunjompliiilus [sic] (Jonas). Dull, 1909: 2.39; Alamo see below). Tegulaandcu\n'jalodmip\iheaslo,a1(9J9o7ne:s,141.844) [sic].—Guzmanetal, 1998: Clilorosloiiui (juipuu new species 36, fig. 25; VeUz and \'asquez, 2000: 762, fig. IE; Aldea (Figures 24-40) and \'aldo\inos, 2005: fig. 8C. Diagnosis: Adult whorls purple-black dorsally; spire Trochus kieneh Hupe. 18.54, p. 144, ,\lalacologia, pi. 4, figs. 1, andbase usuallylightbrown, commonlywith darkbrown la. lb. protractive stripes or mottling. Umbilicus narro\\', open. Diagnosis: Shell width to 35 mm. La.st whorl l^ioadK Description: Shell up to 30 mm wide. Spire angle rounded. Umbilicuswhite, open; umbilicuswithtwowell about 80 degrees. Peripheiy near base, shai"ply rounded exposed spiral cords, the adaxial cord terminating; in a to slightly angular. Sutures appressed. Protoconch un- columellar tooth. known; teleoconch with five flat-sided to slightly convex Material Examined: UW'BM 97871, D\' 1599-1, Re- whorls. Axial sculpture absent or rarelywith mgose pro- W cent, L = 26.0. = 29.9. tracti\-e ribs. Thin growth lines stronglyprosocline. Spiral sculpture of20 muted spiral threads posterior to peiiph- Occurrence: Late Pleistocene; Northern to central eiy; rarely with three to five spiral grooves separating Chile. Recent: Southern Peru to central Chile. four to six broad low spiral cords orwithout spiral scuIjd- Remarks: Specimens of ChJorostoma etirijonipluituin ture. Twenty to 30 evenly spaced spiral threads on base are large, purple-black, and characterized b\- a fuoad of juvenile specimens, muted or obsolete on adult open umbilicus and broadl\- roundedwhorls. The\- differ whorls. Outei' shell layer pui^ple-black on adult \\'horls, C from specimens of atnim. which have a closed umbi- tan Ol' light brov\ai on spire whorls and base. Protractive hcus. and C. luctuositm, which have one or more angular and rarely retrotractive wrinkled brown stripes usually Page 16S THE NAUTILUS, \'ol. 121, No. 4 Figures 27^0. Clilomsfoma cjuipua new species. 27. MUSM INV 131, DV 571-1, svnhpe, spire view, width = 28.4 mm. 2>S. MUSM I.W ]3:3. D\' 16.35-2, earivPliocene, basalview, uidth = 16.0 mm. 29. UWBM 978S0, DV 16.35-2, basal\ie\\, width = 16.6 mm. 30. UWBM 97S79, DV 16.35-'2,basalview,width = 18.5mm. 31. UWBM 97890, DV 1598-1,earlyPliocene,oblique.spire,width = 11.2 mm. 32. UWBM 97883, DV 1284-], Pliocene, oblique basal \iew, width = 8.9 mm. 33. UWBM 97876, DV 571-1, s)ait)pe, apertural view, len2;t]] = 22.7 mm. 34. UWBM 978S0, oblique spire view. 35. MUSM INV 133, apertural view. 36. UWBM 978S1, DV 1029-1, earlv- Pliocene, spire view, vvidtli = 17.4 mm. 37. UWBM 97879, oblique spire view. 38. MUSM IN\' 135, DV 1029-1, obliquespireview,width = 11.3 nun. 39. UWBM 97878, DV809-1, Pliocene, spire\iew,width = 22.1 mm. 40. MUSM INV 132,D\' 809-1, basal view, width = 19.0 mm. UWBM present on lia,se and less often on spire. Inside edge of TApe Material: W(All D\' 571U-1W,BalMl svntvpes) noueteerrtlhiipn.smCoooltuhm.eUlmlbailwiicthustohpicekn,innnaerrrotwo;otuhmbialticeanldveo-f W9787=6,(25L.1)=;2M2.U7,SM =IN2V7.31;31, L = (9187.897)7,,WL == 2(81.74);, \vell-e,xposed white spiral umbilical cord. Weak outer MUSM INV 136, L = (1(».2), \\' = 17.0. tooth adjacent to floor of aperture at end of thin spiral Other Material Examined: UWBM 97873, D\T254- TcsriooyeurtptdaelhffLoolfloalcnioagnwletiientbrgysa:eroceutltiyeDorXno'vmwei.atr5rl7hgi1a-ina1n.gbioaAnfnlgtdlouounnmGbberiildalinicpdacaleuvs,ae.rdaebaro.ouaStmdoaitlkol'Spkaamn- 92B1407a..l8447;;64,,UUlaWWDteBVBeMMa1r2l959v7478-P87lB78ia5,ol,ceDD1n0VeV,,8lL10a29te-5=14,P(-l1BP1ia.lol0ic)oe,1cn0eeW,n,eL,L=L=1=7=(.1(344;1..581.U))4,,,WWWWBM=== fuan de Marcona, on south-facing hillside west ol Pan- 22.1; UWBM 97879, DV 1635-2, early Pliocene, L = american Highway; one of several shell banks ol the 12.2, W = 18.5; UWBM 97880, DV 1635-2, L = (11.3), W UWBM DV Pisco Formation (Figure 41). Locality inaccurately re- = 16.6; 97881, 1029-1, earK' Pliocene, ferred to as El Jaliuay in Muizon and DeVries (19S5). L = (12.3), W = (17.4); UWBM 97882, DV 1029-1, L = 15°26'57"S, 74°52'06"W (Acari 1:10(),()()0 quadrangle). (12), W = 16.7; UWBM 97883, DV 1284-1, Pliocene, W UWBM Middle U[)per Vliocene. L = (5.2), = 8.9; 97890, Y)\ 1.598-1, early T. De\Vies, 2007 Page 169 PLli=oc1e6n,e8,, LW==S.(41,9.W0):=M1U1.S2:MMUINS\'M1I3N3,N'D1V32,16D35V-2S,()9L-l=, \Deenstcrriicpotseiotno:quaSdhrealtle:uppetroip3h5enm'mweainkldyiabmiectareirn.atWeh.oSrplis- W MUSM M11U.2S, M IN=\'161.305;, D\' 1029-I1N,\'L1=346,.9.16W.35-=2,11.l3o.t of 2; kraelnsciunltpotubie"aedso:fitnhtreeresptaocefsivweitphritmwaoiytospfiirvael ccoonrtdisnuboruos- Occurrence: Middle late Miocene to earlv late or weakly beaded tertiaiy threads. Base with fom- to five beaded or non-beaded primaiy spiral cords and in- Pliocene: .southern Peiii. ten'euing secondaiy ctn'ds and tertian' threads. Um- Et>inoIogA': '\)nipua. Latinized \ersion ot "quipu," bilical areawhite, tabulate, sliaiply defined, with margin Inca conntiiii:; cle\ice oi braided and knotted .string.s. of umbilical area flaring towards aperture as steep- e\"oked h\ the wrinkled brown .stripes on the base and K int'lined wall. Llmbilicus open, broad, with two spiral spire ofthis species. umbilical cords situated ada.xially, the innermost Remarks: Specimens ot ClilorosloiiKi (jiiipiiii dillei- thicker: each cord terminating in columellar tooth. from those of C. cunjomplialinn and C. Iiictuosinn l)\ Third tooth sometimes present atbase ofcolumella. Um- bilical veneer variablv developed. Columella thin, ha\ing a narrow'er umbilical area, smallerparietal tlangc, and protracti\e browai stripes. Specimens of C. quipiKi upright, without parietal flange (nerhanging umbili- lack the keeled spiral cords of C. hictnosiim and closed cus. Floor ofaperture with ledge but without teeth. In- umbilicus of C. atnnn. Some specimens of C. quiptia, ner lip sometimes with lour to six closelv spaced low- both Mioceneand Pliocene,havebroadspiral cords (Fig- teeth. ure 25) like those seen on the ju\'enile whorls oi some Occurrence: Late Miocene to Pleistocene: southern specimens of C. atnim (Figure 7). A single specimen l^eru. Recent: northern Peru to Chile. from upper Pliocenebetls abo\e Pla\a Huacllaco (Figure 26) has spiral cords as pronoimced as the raised spiral Etymology: "Cantalloc," site near Nazca, Peru, where cords on specimens of the modern Chilean C. i^notiini subterranean aqueducts are I'eached from gromid level (Figm'e 22). Some lower Pliocene specimens nearYanca h\ pre-Incaic stonewall-lined spiral paths that resemble (Figure 31) have coarse protractive axial I'ibs like those the spiial imibilical coixls of tliis genus. on some .Asian chlorostomines. Remarks: Specimens of CantaUocostoma differ from Specimens of Cljlorostoma ipiipiui superficially re- thoseofAsian, Californian, andPermdanChlorostoma by semble those ofC. gallina (Forbes, 1852), a Pliocene-to- having beaded spiral cords and two adaxiallv situated Recent species from California and Baja California umbilical spiral cords. They differ from specimens of (Grant and Gale, 1931: McLean, 1978), and C. rugixsinH Iiiti.stiiina newgenus, byhavingcloselyspacedbeads and [A. Adams, 1853). a Recent species from the Gulf of lacking a subsutural band ofwell-developed protractive Cahfomia (Keen. 1971). Specimens of all three species nodes. Specimens of CantaUocostoma differ from teg- ha\e some degree ofpuiple-black color and protractive ulines traditionally assigned to Agatlnstoma b)- lacldng stripes on the spire and/or base. Specimens ofC. ^(iJUiui hallmarks of that genus: "narrow open vunbilicus, a and C. nigosum, however, are more ventricose later:ill\ smooth or fineK beaded spiral sculpture, and a \arie- andbasalK' and have\veakto prominent protractive axial gated surface coloration" (Olsson and Hai-bison, 1953: ribs and stripes posterior to the base. Specimens of C. 351). gallina usuaU\- ha\e a closed umbilicus. Shells ofChlorostoino qnipua are found in upper Mi- CantaUocostoma qiiaclricostatnm (Wood, 1828) ocene beach deposits near .Alto Grande (D\' 571-1: see (Figures 42-45, 47-49) Muizon and De\'ries. 1985) udth specimens of Chorus frassinetti De\'ries, 1997, and Acanthina obesa DeVries, TTmrcolcnhiussqiujmilKdicdorsictoisittaatuWsooWdo.odL.S2—<S:PhIifli,ipppii.,5,18fi4g6.,16D.ie Krei- 2of00C3. (qDnei\p'uiiaeso,cc1u9r97t,og2e0t0h3e).r wLdotwhesrpePchiomceennseosfpethceimmenn-s TcgiilsaeLqsiicahdnreicckoesntaioade(rWoTordo,ch1oSi2d8e)e.n—pV.el1i5z4,anpdi.V2a.s5,qufiegz.,6.2000: ricid gastropods, ConchoJepas kiencri Hupe, 1854: Xan- 759, fig. lA; Aldea and \'aldo\inos, 2005: SF. thochonis oclmroma DeVries. 2005: and Hcrminespina Tegida quadricostata Grav.—Dull, 1909: 240; Alamo and saskiae De\'ries and\^ermeij, 1997: and tlie turbinidgas- " Valdivieso, 1997: 14. tropod, Piisoga.ster mcleani DeVries. 2006 (DeVries. Moiiodnntii catcnifcni Potiez and Mieli:iud, IS3H: 318, pi. 29, 2005. 2006: De\'ries and \'ermeij. 1997). I'igs. 12-13. Tnichtis tondosiis I'liilippi. 1S43, ,\bliil(lnngcii niul Be.scluei- Genus Cantallocastoiim new genus liungen neuer oder vvenig gekannter Conch\lien, \. 1, pi. Tjpe species: Trochus ipiddricostatus NN'ood, 1828. 2, fig. 12. Recent. Peni and Chile. Diagnosis: Low spire, white to cream-colored with puq^le along edges of sculptural elements. Sculpture of Diagnosis: \\Tiite to brown outer shell laver. Three to three to live prominent primaiy beaded spiral cords. fi\"e beadedprimar\'.spiral cords. Lhnbilicus open, jjroad. Two adaxialK' situated spiral umbilical cords terminating Material Examined: UWBM 97884, D\' 1372-1, Re- ientalcoflluamneglelaorxetreheatnhg.inPcarriuemtbailliwcaulsl.\ertical, witliout pari- cLen=t,(1L3.3=),(1W4.4=),(2W0.8=):2U3.W9:BUMW9B78M869,78D8V5,17D1V3-11,37e2a-r1l,v Pase 170 THE NAUTILUS, Vol. 121, No. 4 Figures42^5, 47^9. Caiilalloaisfoiiuiquadricostatiim Wood, 182S. 42. UWBM 978S4, D\' 1.372-1, Recent. ol)li(jue.spireview, widtli = 2,3.9 rnni. 43. UWBM 978S6, DV 1713-1, early Pleistocene, apcrtnal view, width = 21.5 mm. 44. MUS.M IN\' 137, DV 1.3.55-1, late Pliocene, basal view, width = 16.0 mm. 45. UM'BM 97884, oblique basal \iew. 47. UWBM 97885. D\' 1372-1, oiiliijue lateralview,width = 20.8 mm. 48. UWBM 97887, DV 1355-1,obliquespireview,width = 18.1 mm. 49. UWBM 97886,obliqueiiasal view. Figures 46, 50-.54. CaiilalhiciKlinna panistostnm new species. 46. MUSM INV 141, DV 1635-2, early Pliocene, oblique .spireview, width = 14.3 irnn. 50. UWBM 97891, DV571-1, late Miocene, obliquespireview, width = 14.9 mm. 51. UWBM 97S89, DV 1598-1, syntype, oblique lateral view, width = 33.7 mm. 52. UWBM 97889, oblique basal view. 53. UWBM 97889, apertural view. 54. MUSM INV 139, DV 1598-1, synty^je, basal view, width = 24.9 mm. Figures 55-57. Intistoina pirqiui new species. UWBM97892,DV470-1,syntype,earlyPliocene,length = 38.9mm. 55. Lateralview. 56. Aperturalview(lightingfromupperright). UWBM .57. Obliquebasalview. Figures .58,.59. Intistonmmireatincium {Forbes, 1852). South ofLa Jolla, California, Recent. 58. 97896, lateral view, length = 20.0 umi. 59. UWBM 97897, oblicjue basal view, vsidth = 17.8 mm. Figures 60-63. A'^atliistoimi UWBM patugonicum (d'Orbigny, 1835). 60. UWB.Vl 97893, Argentina. Recent, oblique .spire view, width = 15.4 nun. 61. 97893, oblique basal view. 62.'UWBM 97895, DV 1032-2, late Phocene, lateral view, width = 12.7 mm. 63. UWBM 97895, oblique .spire view. T. De\'i-ies, 2007 Page 171 Pleistocene, L = (14.5), W = 21.5; UWBM 97887, DV 1355-1, late Pliocene, L = 12.6, \V = 18.1; UWBM 917378,88D,VDX135153-515,-1L, =L(W=8.9()1,0.W0),=W16.=0;1M6.U1;SMMUISNMV I13W8, DV 1355-1, L = 14.1, = (19). OccuiTence: Late Pliocene: southern Peiu. Hecent; northern Peni to Chile. .^ Remarks: The nrnnbei'ofpriman'beaded spiral cords on thelastwhorloi CantaUocostoma quadricostatum var- iesbetweendireeandfi\e. Mostmodern specimenshave t^^o wideK" spaceil priman' spiral cords on the anterior h;ili of the whorl and t^\•o closely spaced primary spiral cords adjacent to the posterior suture (Figure 42). Some specimens ha\e an additional primaiy spiral cord be- tween the t\\'o anterior spiral cords (Figure 47); other Figure 64. Tvpe localit)- (DV 1598-1) of CantiillocDsloina specimens ha\e one of the two posteriormost priman' jxinistostniii new species and other teguline localities between spiral cords missing (Figure 43). The six knowii late Sacaeo and Yauca. Pliocene specimens from southern Peiii (Figure 48) and psriinmgalie"\e"arsKp'irPalleciosrtdosc.ene specimen (Figure 43) have tln-ee D15V98-115,98L-1,= L(13=.78).,4,WW==241.19.;2;MMUUSSMMIINNVV14103,9,PaDnV- Wamericau roadcut at Yauca, early Pliocene, L = (17.9), Cantallocostoma panistostiiin new species = 33.3. (Figures 46, 50-541 Other Material Examined: UWBM 97891, DV 571- Dsciualgptnious'eiso:fdirEe.extbereiaodredunsipfiroarlmcloyrdsp.alBeasbalroswpinr;alscpoirrdasl D1,Vlat1e63M5i-2o,ceenaer,lyLPl=ioc10e.n4e,,WL == 81.48.,9;WM=US14M.3.INV 141, bunched towards mnbilical area. Occurrence: Late Miocene to early Pliocene: soudi- ern Peru. Description: Shell nearh- .35 umi in diameter. Spire Etjinolog)': "Panis." Latin noun meaning"bread," and angle about 70 degrees. Periphen" at base, sharply "tostum," Latin neuter past participle-adjective meaning rounded. Sutures impressed. Protoconch unknowai. Te- "toasted," referring to the bread-cnist color ofthis spe- leoconch with five quadrate whorls. Axial sculpture ab- cies. sent e.xcept for intermittently iTjgose prosochne growth lines. Spiral sculpture ol closely beaded spiral cord at Remarks: Tlu- light brown color of the outer shell base and hvoposteriorspiral rows ofmorewidelyspaced layer on specimens ot Cantallocostoma paiiistostuiii re- beads, about 25 per whorl, bracketing shoulder. Poste- sembles that of specimens ofTegiila heinphilU Oldroyd, rior beads sometimes slightly protractiyely elongate. In- 1921, a late Pliocene-to-Pleistocene species from Cali- terspaces rarelyvvidi beaded secondaryspiralcords; usu- fornia (Grantand Gale, 1931). Specimens ofT. hemphilli aUv with wa\"ering tertiarv spiral threads. Base weakly and mmierous other Miocene and Phocene Califomian con\ex. with six to ten closely spaced spiral cords adja- teguline species with similar coloration lack the two um- cent to umbilical area. Outer shell layer pale brown, bilical spiral cords close to the axis and are coyered by without color pattern. Inner edge of outer lip smooth. numerous closely spaced priman' spiral cords, none of Umbilicus open, wide, with two ada.\ially situated spiral wiiich are beaded. cords. Umbihcal \eneer thin. Innermost umbilical cord Specimens of Cantallocostoma panistostiim are found prominent, second cord thin; each terminates in col- in upper Miocene nearshore sandstones with Cliloros- umeUar tooth. Third tooth at base of cohunella nearly toma quipiia; Clionis frassinctti DeVries, 1997; Acan- obsolete. ColumeOa thin; parietal flange absent. thina obcsa; and Xanthochorus stephanicus DeVries, 2005; and in lower Pliocene cobbly bioclastic gravels as- T}pe Localitv: DV 1598-1. roadcut along Panameri- sociated wth the mouth of the paleo-Rio Yauca with can Highwa\-, shell beds exposed along descent from disarticirlated valves ofan undescribed Anadaro species, nordi into Rio Yauca valley (Figure 64). 15°39'49"S, venerid bivalves, Chlorostoma cpiipua, Xanthocluniis 74°31'50"W (Yauca 1:100,000"quadrangle). Lower ocluiroma, and Concholepas nodosa Miiricke, 1896. Pliocene. Genus Intistoma new genus T97)8^8e9.MDa\t'e1r5i9a8l-:1.(LD\='(2155.928)-,1W. a=ll33s^y7n;t\U-pWesB)MU9W78B90M, Tplyepiestoscpeenceiteos:ReceTnrto,chCuaslifaourrncioat.inctiis Forbes. 1852. Page 172 THE NAUTILUS, Vol. 121, No. 4 Diagiiosi.s: Spiral .sculpture oi suli.sutural baiul of thick bitistoma pinjiid new species protracti\e nodes and peripheral and suh-peripheral pri- (Figures 55-57) maiT spii'al cords. Base with three thick, primaiy spiral cords. Umbilicus open. Diagnosis: Shell large, weakly bicarinate, with poste- rior row oi protractive nodes. Base of shell \^'ith three broad spiral cords. Umbilicus open. Description: Sliell up to 45 nun wide, spire angle about 75 degrees. Whorls tour to fix'e in number, quad- Description: Shell large, width more than 40 mm; rate to caiinate; sutures weakly impressed. Pi'otoconcli spire angle about 60 degrees. Whorls ventricose, weakly miknowii. Sculpture oi thick romided protractive axial bicarinate; sutures deeph' impressed. Protoconch un- ribs intersecting wdth an equally thick spiral cords, pi^o- known; early spire whorls missing; teleoconch ofat least ducing a broad subsutural spiral band of elongate pro- three whorls. Lastwhorl \v\\\\ sculpture ofabout IS elon- tracti\'e nodes, a near-basal peripheral bantl oi stubbv gate protractive ribs on posterior half of whorl; wdth pj-otractive nodes more numerous than nodes in the sub- about 21 circular to slightly protractively elongate nodes sutural band, and a sub-peripheral primaiy spiral cord atperipherv, two-thirds the distance from suture tobase; with little axial modification. Tertiary threads sometimes and wdth a continuous low liroad spiral cord at edge of present; often cornigated h\' slightK' raised strongK' ob- base. Base \v\(\\ three low broad spiral cords, innermf)st lique lamellar growth lines. Umbilicus open. Columella cord bordering umbilical area and twice as wade as otiier thin: parietal (lange erect to slightly overhanging umbi- two cords. Faint traces of secondaiy spiral cords in in- licus; parietal callus small. Umbilical veneer thick, cov- terspaces laterally and basally. Aperture oblique, outer ering all but wedge-shaped adapertural portion oi um- lip and colabral growth lines moderately prosocline (40 bilicalwall. Umbilical spiralcord submerged in umbilical degrees). Umbilicus open. Columella with at least one wall, emergentterminallyas prominentcolumellartooth. tooth, chrected basally; anteriorportion partly excavated, Smaller second columellar tooth sometimes present partly missing. Floor ofaperture partlymissing; no teeth abaxialK' adjacent to first tooth. evident. T\pe Locality: D\' 470-1, above rock-\' road from Ha- Etymology: "Inti," the Inca sun god, «ath a nod to cienda Tunca to Quebrada Huaricangana; hillside of Caliiornia's sunshine and the sunset-orange color inside browii sandstone (Figure 65). 14°56' S, 75°09' W (Palpa the umbilicus of the tx'pe species, Inti.stoma aureotinc- 1:100,000 qu:ulrangle). llpper Miocene or lower tiiiii. Pliocene. Occurrence: Late Miocene or early Pliocene: south- T)pe Malcrial: {D\ 470-1, syntypes, late Miocene or ern Peru. Earlv Pleistocene to Recent: California. P T Remarks: The new genus, Intistoma. is proposed ior 9O TDoVwnlsocoarlivtiyl-lsaagmepsles p PALPA^ _\*^. ^-'^"k30'S^ t\vo \eiy similar species: the Pleistocene-to-Recent C^ali- fornian Intistoma aiircotinctum and the earh Pliocene Pemvian /. pirijua new species. Specimens oi both spe- cies differ irom those oi nearly all otherteguline taxa by possessing three thick prinian' spiral cords on the base, rather tlian cords that are more numerous and thinner. Specimens oi Intistoma additionally differ iiom those properly assigned toAfiatliistoma bylacking teeth on the iloor oi the aperture and the inner edge ofthe outer lip and by lacking closely spaced beaded pri?nan' spiral cords. Intistoma anrcotiinium lias been considered the ex- tant representative of a lineage of Calilornian Neogene tegulines (Addicott, 1970) that includes the early Mi- ocene Tegida clalli arnoldi Addicott, 1970, the late Mi- ocene Tegula nasliae Clark, 1915, and PBocene Tciuila hemphiUi Oldroyd, 1921. Specimens offossil Caliloiiii:ui \ PUERT species do have asubsutural band ofelongate protractive 6cABALLAS^i> nodes, as do specimens of Inlistoma, and some have thick peripheral spiral cords, but none have the distinc- tive intistomine combination of tliick basal spiral cords i5°n()'S and tertiaiyspiral threads across the entire surface ofthe Figures 05. ']"\pe loculiU (l)\' 470-1 i ol liilishniiii inhjiui wiir)rls. new .species.