ARCTIC VOL. 41, NO.3 (SEPTEMBER 1968) P. 167-172 14ewn Species of Limpets from the Neogene of Alaska (Patellogastropoda: Mollusca) DAVID R. LINDBERG’ and LOUIE MARINCOVICH, JR.’

(Received 1I March 1987; accepted in revisedform 4 February 1988)

ABSTRACT. Twonew species of patellogastropod limpets from the Miocene-Plioceneof Alaska are described-Putelloidu grudutus new species from the Unga Conglomerate Memberof the Bear Lake Formationat Cape Aliaksin, Alaska Peninsula, southwestern Alaska, and Niveotecturu myrukeenue new species from the Yakataga Formation in the northeastern ofGulf Alaska. A third species from the Narrow Cape Formationof Kodiak Islandmay be referrableto Putelloidu sookensis Clarkand Arnold, 1923, a species previously known only from VancouverIsland, Canada. These and other speciesof Patelloidinae dominated the northeastern Pacific patellogastropod fauna for over60 m.y. The presence in Alaska of these three warm-water limpet species may be related to the middle Miocene warm-water event that is well documented elsewhere in the NorthPacific. However, regional cooling during the late Neogene drove this predominantly tropicalgroup from higher latitudes, leaving them poorly represented in the Holocene borealfauna. Key words: paleontology, historical biogeography, paleoecology, fossil gastropods, Miocene, Patellogastropoda,Patelloidu, limpets

RÉSUMÉ. On ddcrit deux nouvelles esfices de patelles gastdropodes trouvdes dans lemioctne-plioctne de l’Alaska: la nouvelle esp&cePutelloidu grudurus venant duconglodrat de I’Unga, membre lade formation Bear Lakeau cap Aliaksin dans la péninsule del’Alaska, au sud-ouestde l’Alaska, et la nouvelle esfice Niveotecturu myrukeenue venant de la formation Yakataga dans la partie nord-est du golfel’Alaska. de Une troisitme esptce de la formation Narrow Cape de l’îleKodiak peut se rattacherà la Putelloida sookensis Clark etAmold, 1923, une esfice qui n’avaitkt6 trouvée auparavant que dans l’île de Vancouver au Canada. Ces esfices et d’autres appartenant aux Patelloidinae ont domind la faune des patelles gastéropodes du Pacifique Nord-Est pendant60 millions d’annees.On peut codler la presence en Alaska de ces troisesfices de patelles d’eaux tempérkesau fait, bien documenté ailleursdans le Pacifique Nord, de la présence d’eaux tempéréesau milieu dumioche. Cependant, le refroidissementde la rkgion aucours de la fin dunéogtne fit fuir des latitudes dlevtes ce groupe appartenant en grande partieà la faune tropicale, ce qui explique sa faible representation dans la faune boréale de I’holoc&ne. Mots clés: paldontologie, biogéographiehisbrique, palddcologie, fossiles gastkropodes,mioche, patelles gasteropodes, Patelloidu, patelles Traduit pour lejournal par Ndsida Loyer.

PE3QME. Onncanw nsa KOBWX snna narennoracspononnuxonmneqex unouetta-nnnouena Anucre. nepaun - &- FëïXTdagradatus n. ~p.11s ynrwcxoro ~onrnouepat~~~oronnacTa ~~P~~RKCKORCBHTU na nuce AnUKCHHe, HP AJIUCKHCXOU nonyocrpoae B mro3anonnonAnrcrte, H eropo~- Niveotecturamyrakeenae n. sp. HS urararnn- CKOA CBHTU na cesepo~ocro~n~ndepery AnucKnncroro sanuna. TperxA ann, HS H3ppOYK3IICKOti CBHTY na oc- rpose KonuKe, noze~6hlTb orneceti K Patelloida sookensisClark 8 Arnold,1923. Ao lwHe TOT ann scspe- VanCr TOnbKO na KanancKon ocrpoBe BaHNyBepe. 3rn TPH anna, BHOCT~c npyrnnw annann Patelloidinae, rocnoncrBoBann B +aynenarennoracrponon ceBeponocroqnoro THxoro oKeana a Teqeane 90 nnn. ner.npncy- cTBne OTXX rennosonnux~nnos dnloz~euer B Anrcxenoxer durb carsano CO cpennewnouenosuuTennosonnun co- 8b1THen. KOI~DO~secbua xopomo noKynentnposanno B npyrwxnynrrax Cenepworo Tnxoro oKeana. Onnaxo, pe- rnonanbnoenoxonoxnenne BO Bpenunoonnero neorexa Burecnnno STY npenwymecraexaorponnqecKym rpynny c BYCOKHX mnpor, OCT~B~RUeecnadoe npeacraanrenbcrso B ronouenonoR6opeanbnoR aaywe Ceaepnoro Tnxoro olceaaa.

INTRODUCTION Cap-shaped or limpet shells are common components of some excellent condition and are readily apparent in cross section Tertiary fossil assemblages. However, because of the simple (Fig. 1). shell morphology and the reliance on soft part characters for mostlimpet classification schemes, fossil limpets areoften tentatively assigned to a Holocene regional familyor genus and are not further studied. This practice has contributed to a fossil record for limpets that lacks a historical biogeographical perspective. Patellogastropod limpets, whichtypically occur in high- energy, rocky, nearshore environments, are usually represented in fossil assemblages by few individuals and often reflect the vagaries of post-mortem transport into more benign environ- ments, where they were preserved. However, if some of the original material of the patellogastropod shell is preserved, identification of the specimen need notbe uncertain. The use of shell microstructure in patellogastropodsystematics has already been demonstrated(MacClintock, 1963,1967; Lindberg, 1976, 1978,1979,1981a; Lindberg and McLean, 1981; Lindberg and Hickman, 1986), and its use in recognizing distributions of patellogastropod taxa inspace and timeis invaluable for under- standing the evolution of members of the order. For example, although manyof the specimens discussed herein are Miocene mG. I. Cross section through the shell of Niveotectura myrukeenue Lindberg and in age (24-5 Ma), their shell microstructure characters are in Marincovich, new species (holotype, USNM 414579). Scale bar = 2 mm.

‘Museum of Paleontology, Universityof California, Berkeley, California94720, U.S.A. ’U.S. Geological Survey, 345 Middlefield Road, Menlo Park, California94025, U.S.A. @TheArctic Instituteof North America 168 / D.R. LINDBERG and L. MARINCOVICH, JR.

The three limpets from the Neogene of Alaska discussed here Genus Patelloida Quoy and Gaimard, 1834 include two new species and are the first fossil members of the Patelloida Quoy and Gaimard, 1834:349 patellogastropod subfamily Patelloidinae known from Alaska. Shell oflow to high profile; typically stout, withthick Members of this predominantly tropical clade were common in intermediate area composedof concentric crossed-lamellar layer. Tertiary limpet faunas along the west coast of North America Sculpture variable, but radial components usually present and (Vancouver to San Diego) for over 62 m.y. (Lindberg and concentric growth line sculpture typically threadlike and pro- Hickman, 1986). With the onset of regional cooling during the nounced in complex prismatic layer. Lateral teeth variable in Neogene (Durham, 1950; Addicott, 1969), most Patelloida taxa size and shape, cusps complex; marginal teethin two pairs were excluded from higher latitudes, and only tworelict genera, (radular tooth formula: 2-3-0-3-2). Radular ventral plates com- Erginus Jefferys, 1877, and Niveotectura Habe, 1944, remain plex; sutures, anterior processes and lateral extensions present in the boreal Pacific Ocean today (Lindberg, 1983). Patelloida in most taxa. Gut looping complex; two excretory organs, one gradatus new species from the Alaska Peninsula and ?Patelloida on either side of the rectum; single gill present on left side of sookensis (Clark and Arnold, 1923) from nearby Kodiak Island nuchal cavity. Pericardial sac triangular, positioned on anterior are members of a genus that was excluded from the region, left side of visceral mass. Oral lappets present in some species. while Niveotectura myrakeenae new species from the Bering Mantle edge often thickened with large tentacle/gland com- Glacier region of southeast Alaska is the first fossil occurrence plexes. Broadcast spawning. of this relict genus in Alaska. Age. Cretaceous to Holocene. The presence in Alaska of these three warm-water limpet Distribution. Past andpresent tropical and warmtemperature species may be related tothe middle Miocene warm-water event seas in both hemispheres. that is well documented elsewhere in the North Pacific. Evi- Remarks. Modem species of the monophyletic taxon (or dence for this warm-water event in Alaska has recently been clade) Patelloida s.1. (sensu lato) live in tropical and subtropi- documented by the presence in the Bear Lake Formationof the cal regions. As now defined, the group consists of at least two warm-water gastropod Turritella(Hataiella) sagai Kotaka, subclades (Lindberg and Vermeij, 1985). One subclade includes 1951, previously known only from earliest middle Miocene those species with low to medium shell profiles, strong radial (16-15 Ma old) faunas of Honshu, Japan (Marincovich and ribs or many fine riblets, reduced third lateral radular teeth, and Kase, 1986). Patelloida gradatus new species, known from a that occur on various substrata. The type species of the genus, Bear Lake outcrop inferred herein to be of middle Miocene age, Patelloida rugosaQuoy andGaimard, 1834, is a member of this is further evidence of a warm-water event that influenced the subclade; hencemembers of this subclade are assigned to Bear Lake fauna. The less certain dating of the Miocene or Patelloida S.S. (sensu stricto). The other subclade, which has Pliocene occurrence of Niveotectura myrakeenaenew species in been called the Patelloida profundagroup by Christiaens (1 975) the Yakataga Formation makes it more debatable to ascribe the and Lindberg andVermeij (1989, includes species with medium presence of this species to the earliest middle Miocene warm- tohigh shell profiles, many ribs, equal-sized lateral radular water event. The same may be saidfor the presence of ?Patelloida teeth, and typicallylimited to calcareous substrata. Both groups sookensis in the Narrow Cape Formation of Kodiak Island. have Tethyan distributions and are recognizable in the Creta- However, this formation on Kodiak Island is considered to be of ceous; the Patelloida S.S. group dates from the Late Cretaceous early and middle Miocene age and contains several other (Campanian, about 75 Ma) of California (MacClintock, 1967), warm-water molluscan genera (Allison, 1978), so the presence and the Patelloida profunda group from the Early Cretaceous of ?I? sookensis there is possibly related to the same warm- (Albian, about 100 Ma) of England (Lindberg, in press). These water event that introduced T.(H.) sagai and P. gradatus into clades are defined by, and can be recognized in the fossil record the Bear Lake fauna. by, shell morphology (Lindberg and Vermeij, 1985), shell The higher classification used here follows Lindberg (1983, structure (MacClintock, 1967; Lindberg, in press), and pre- 1986a,b). The time scale is fqom Palmer (1983). Abbreviations served features of soft part anatomy such as gill morphology in the text are as follows: C&(S) - Stanford University Type (Hickman andLindberg, 1985) and radular morphology (Akpan Collection (now on permanent loan to Department of Inverte- et al., 1982). brates, California Academy of Sciences, San Francisco, Cali- Patelloida gradafus Lindberg and Marincovich, fornia); LACM- MalacologySection, Natural History Museum new species (Fig. 2a) of Los Angeles County, Los Angeles, California; UCMP - Shell medium sized (aperture diameter about 20 mm). Apex Museum of Paleontology, University of California, Berkeley, subcentral, positioned in anterior third of shell; profile high California; USGS - U.S. Geological Survey, Menlo Park, (lengtwheight ratio approximately 1.5). All slopes convex, with California; USNM - U.S. National Museum of Natural His- strong concentric shell sculpture. Concentric sculpture consists tory, Washington, D.C. of alternating opaque and translucent growth increments that are grouped into distinct, approximately 1-mm-wide bands around SYSTEMATICS the shell. Aperture oval. Order Patellogastropoda Lindberg, 1986a Holotypedimensions. Length 19.9 mm, width 16.7 mm, Family Lottiidae Gray, 1840 height 12.4 mm. Subfamily Patelloidinae Chapman and Gabriel, 1923 Type locality. ALASKA: USGS Cenozoic locality M8028, sea cliff on northwest shore of Cape Aliaksin, Alaska Peninsula; Shell patelliform and composed of four layers. Outer surface Port Moller (B-3) quadrangle; 55"30'10N, 16O05O'07'W (Fig. of shell and interior margin complex prismatic. Next inner layer 3). Unga Conglomerate Member of the Bear Lake Formation. concentric crossed-lamellar, followed by myostracum, and radial Collected by Louie Marincovich, Jr., August 1982. crossed-lamellar layers. Type material. USNM 414578, holotype. LIMPETS FROM THE NEOGENE OF ALASKA I 169

FIG. 2. Fossil and Holocene Patellogastropoda of the boreal Pacific Ocean. Scale bars = 5 mm. a) Putelloidu grdurus Lindberg and Marincovich, new species (holotype, USNM 414578). Unga Conglomerate Memberof the BearLake Formation, Alaska Peninsula, Alaska.b) Putelloidu sookensis (Clark and Arnold, 1923) (holotype, CAS(S) 67). Sooke Formation, Vancouver Island, British Columbia, Canada. c) ?Purelloidu sookensis (hypotype, UYNM 414580). Narrow Cape Formation, Narrow Cape, Kodiak Island, Alaska. d) Putelloidu sookensis? (hypotype, UCMP 35251). Quimper Sandstone, Discovery Bay, Jefferson County, Washington. e) Niveorecruru myrukeenueLindberg and Marincovich, new species (holotype,USNM 414579). Yakataga Formation, northeastern Gulfof Alaska, Alaska. f) Niveotecrurupullidu (Gould, 1859) (LACM Loc. No. 82-12). MutsuBay, Aomori Prefecture,Japan. g) Niveorecrurufuniculuru(Carpenter, 1864) (LACM Loc. No. 60-24). Carmel, Monterey County,California. h) Acmuea rnitra Rathke, 1833 (UCMP Loc. No. A-6601). Moss Beach, San Mateo County,California.

Distribution. Known only from the type locality. Scurriu orbignyi (Dall, 1909). Discurriu insessu ranges from Age. Middle Miocene. The Cape Aliaksin strataare overlain Wrangell, southeastern Alaska, to southern Baja California, by a lava flow that yields a potassium-argonof age 10.4 Ma (F. Mexico, and lives on the stipes of the brown algaEgregiu spp. Wilson, USGS, pers. comm.1986) and contain molluscan (Lindberg, 1981b). The curved rather than parallel lateral shell species whoseoldest inferred age inthe Bear Lake Formationis margins of P. grudutus argue against a similar habitat for the earliest middle Miocene or 16-15 Ma (Marincovich andKase, Miocene species. Scurriuorbignyi occurs on rockyshores 1986; L. Marincovich, Jr., unpub. data). between Paita, Peru, and Talcahuano, Chile (Marincovich, Etymology. The specific name is derived fromthe Latin noun 1973). The apertures of both S. orbignyi and P. grudutus are grudus, or step, and refers to the stepped arrangement of the only slightly irregular and are not similar to the apertures of growth phases of this species. patellogastropods that hometo specific sites on the substratum Discussion. The distinct bands of growth lines that encircle after foraging. the shell distinguishP. grdurus from all other known Holocene The wavy, grouped growth increments of P. grudutus are also and fossil species of Purelloidu. The gross shell morphology of present inPutelfoidu sookensis from the late Oligocene or early P. gradatus is similar to Discurriu insessu (Hinds, 1842) and Miocene Sooke Formation of Vancouver Island, Canada (Fig. 170 / D.R. LINDBERG and L. MARINCOVICH. JR.

their greatest diversity in the boreal Pacific Ocean; Patelloida species occur worldwide in tropical and warm-temperate seas. Members of the genusErginus differ from Niveotectura species by having markedly different anatomical organizations and by brooding their young (Lindberg, 1983). Like the Patelloidaprofunda groupdiscussed above, Niveotectura is primarily subtidal and restricted to calcareous substrata. However, these similarities appear to result from convergence rather than from commonancestry. One difference between these two groups is found the in relative thicknesses of the outer two shell layers. In the genus Niveotectura the outer complex prismatic layer is equal to or thicker than the inner concentric crossed-lamellar layer (Fig. 1); in members of the Patelloidaprofunda group the concentric crossed-lamellar layer is substantially thicker than thecomplex prismatic layer. Almost every known patellogastropodclade has subtidal taxa that have convergedon common shell and radular morphologies (McLean, 1966;LindbergandMcLean, 1981; Lindberg,1983). The genusNiveotectura probably represents this expansion into the subtidal habitat by early Neogene North Pacific Patelloida S.S. species. Erginus was either subsequently derivedfrom Niveotectura or represents an earlier or later second invasion of the North Pacific Ocean subtidal habitat by a Patelloida S.S. 182O 158. I species (Fig. 4).

FIG. 3. Location of Alaskan fossil localities.

2b). However, unlike P. gradatus, P. sookensis has weak radial P ribs. Patelloida sookensis may be represented in Alaskanstrata 3 of approximately late early to early middle Miocene age by a 0, single, deformedlimpet from the Narrow Cape Formation, w Kodiak Island, Alaska (Fig. 2c). Remarks. Patelloidagradatus, P. sookensis and a third speci- men from the Quimper Sandstone of Washington (Durham, 1944) (Fig. 2d), which is probably notreferrable to P. sookensis, Q all share similar gross shell morphology. They all have medium- highshell profiles, lackstrong radial ribbing, haveapexes Q positioned at or in the anterior third of the shell and, when .4 .4 preserved, have characteristically coarse, wavy growth lines. s v) s 0 5 0 Although the gross shell morphology of these species is not P C P aa .4 aa known in Holocene species of Patelloida, it was present in Ir Ir Oligocene and Miocene species of the northeastern Pacific. P Whether theseextinct species are all members of a third subclade8 ru B within the Putelloida cannot be determined from the available data. Genus Niveotectura Habe, 1944 Niveotectura Habe, 1944: 185 Shell of high profile, large (to 50 mm long), white; sculpture of strong radial ribs. Pericardial sac triangular, positioned on anterior left side of visceral mass;single gill presenton left side of nuchal cavity. Radular tooth formula: 0-3-0-3-0. Remarks. Niveotectura appears to be intermediate in shell form and anatomy betweenPatelloida and Erginus. Niveotectura and Patelloida species have similar anatomical organizations (radulae excepted) and exterior shell sculpture. Niveofectura and Erginus species typically have whiteshells with mediumto high profiles, lack marginal radular teeth, have similar radular basal plate morphologies, equal-sized lateral teeth, and occuron calcareous substrates in subtidal habitats; all are characters FIG.4. Hypothesizedrelationships of subcladeswithin the patellogastropod shared by subtidal Patellogastropoda. Niveotectura and Erginus subfamily Patelloidinae Chapman and Gabriel,1923. Each branch representsa species are restricted to the Northern Hemisphere and reach monophyletic taxon. LIMPETS FROM THE NEOGENE OF ALASKA / 171

Niveotectura myrakeenae Lindberg and Marincovich, Based on present correlations, N. myrakeenae and N. pallida new species (Fig. 2e) were contemporary species on either side of the North Pacific Shell medium sized (aperture diameter about 25 mm). Apex Ocean during the early Neogene.The oldest specimens referrable central; profile high(lengtwheight ratio approximately 1.O). All to N. pallidaare reported from the "Shirado beds" of the Joban slopes straight or slightly convex, with both radial andconcen- coal fieldof northeastern Honshu, Japan,by Yokoyama (1925b). tric shell sculpture. Radial sculpture consists of approximately These strata are now referred to the Nakayama Formationof the 28 primary and secondary radialriblets; concentric sculpture of Shirado Group (Kamada, 1962) andof earliestare middle Miocene weaker concentriccords that cross the radialriblets and produce age (equivalent to planktonic foraminiferal zones N8 or N9; S. pustules at the intersections. Interspaces between ribs about Ohara, pers. comm. 1986). A Late Miocene specimen is known three times as wide as ribs. Concentric growth lines distinct. fromSakae(Yokoyama, 1925a) (OgawaFormation [Makiyama, Most of theexterior of the shell is coveredby a crustose coralline 1958]), and Pliocene specimens have been reported from the alga that forms excrescences. Shigarami Formation (Yokoyama, 1925a), the Omma Forma- Holotypedimensions. Length 21.3 mm, width 25.7 mm, tion(Kaseno and Matsuura, 1965), the SawaneFormation height 24.5 mm. Shell has been secondarily deformed by com- (Yokoyama, 1926), and the Imaizumi Formation(Shikama and pression in the anterio-posterior axis. Masujima, 1969). Pleistocene specimens are known from the Type locality.ALASKA:USGS Cenozoic locality M7540, on Koshiba Formation of Yokohama (Yokoyama, 1920). slope southof and below 1 m139 (3737 ft) benchmark of unnamed Niveotectura myrakeenae could beancestral to N. funiculata, peak that is immediatelyeast of White RiverGlacier; northeast- which first appears inthe lower Pleistocene Lomita Marl Mem- em Gulf of Alaska; Bering Glacier (A-3) quadrangle; about ber of the San Pedro Formation in SanPedro, southernCalifor- 60"04'43"N, 142"OO'OO"W (Fig. 3). 1669 26 m(5475 r20 ft) nia (Woodring et al., 1946). above base of Yakataga Formation. Coll. John Armentrout, 1974. Type material. USNM 414579, holotype. ACKNOWLEDGEMENTS Distribution. Known only from type locality. Age. Mioicene or Pliocene. We thank S. Ohara, of Chiba University, Chiba,and T. Kase, of the Etymology. We are pleased to name this species in honor of National Science Museum, Tokyo, Japan, for discussions of the mod- em ages of the older literature records of Japanese fossil limpets; R. the late Dr. A.Myra Keen, of Stanford University, an exempla- Allison,University of Alaska, T. Kase,and S. Stone(USGS) for ry paleontologist and malacologist, who taught the importance reviewingthe manuscript; G. Shkurkin(UCMP) for preparing the of interfacing the present with thepast. Russian abstract;H. Schom (UCMP) for photographing the specimens Discussion. Niveotectura myrakeenaediffers from the Holoceneillustrated in Figure 2; and M. Krup (UCMP) for preparing Figures 1 northwestern Pacific Niveotectura pallida (Gould, 1859) (Fig. and 3. 2f) by having fewer and narrower radial ribs and much wider inter- spaces betweenribs. In N. pallidathe ribs are broad, almost flat topped in profile, and the width of the interspaces between ribs REFERENCES is equalto or slightly less than the widthof the ribs. The Holocene northeastern Pacificspecies Niveotecturafuniculata (Carpenter, ADDICOTT, W.O. 1969. Tertiary climatic change in the marginal northeastern 1864) (Fig. 2g) differs from N. myrakeenae by having more Pacific Ocean. Science 165583-586. AKPAN, E.B., FARROW, G.E., and MORRIS, N. 1982. Limpet grazing on numerous ribs and narrower interspaces betweenribs. The indi- Cretaceous algal-bored ammonites. Palaeontology25(2):361-367. vidual ribs on theshell of N. funiculataare not as broad as those ALLISON, R.C. 1978.Late Oligocene through Pleistocene molluscanfaunas in on N. pallida, but they aresubstantially broader than those N.on the Gulf of Alaska region. Veliger21(2):171-188. myrakeenae. Like N. pallida and unlike N. myrakeenae,N. CARPENTER, P.P. 1864. 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Informations de la Socittk Patelloidinae discussed hereinby having moderately strong belge de Malacologie 4:91-116. radial ribbing and by lacking the distinctly convex shell slopes CLARK, B.L., andARNOLD, R. 1923. Fauna of the SookeFormation, that characterize the other species. Instead, the slopes of the VancouverIsland, with description of anew coral by T.W. Vaughan. University of California Publications, Bulletinof the Departmentof Geologi- shell arestraight or only slightly convex, which givesthe shell a cal Sciences 14(5):123-234. classical conical shape. DALL, W.H. 1909.Report on a collection of shellsfrom Peru, with a summary Remarks. The presence of encrusting calcareous algae with of the littoral marineMollusca of the Peruvian zoological province.Prweed- elaborate excrescences onthe holotype suggests minimal aerial ings of the United States National Museum 37( 1704):147-294. exposure for this species. Holocene subtidal species are com- DURHAM, J.W. 1944. 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