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Home , Atlantic (period), Bootherium, Glyptodon, Irvingtonian, Pedocal, Rancholabrean

REPRINTED FROM: THE OF THE UNITED STATES

A ILEVIEW VOLUME FOR THE VII CONGRESS

OF THE INTERNATIONAL ASSOCIATION

FOR QUATERNARY RESEARCH H. E. Wright, Jr. and David G. Frey, Editors

Copyright @ 1965 by Princeton Univenity Preas

Introduction : Historical Perspectives nINT

PART I: GEOLOGY

Quaternary Geology of Northern Great Plains LEMKE,LAIRD, TIPTON, LINDVALL Glaciation of Minnesota and Iowa WRIGHT,RUHE Outline of Glacial Geology of Illinois and Wisconsin FRYE, WILLMAN, BLACK Geology of Indiana and WAYNE,ZUMBEROE Pleistocene Deposits of the Erie Lobe GOLDTHWAIT,DREIMANIS, FORSYTH, -BOW, WHPPE Quaternary Geology of New York MULLER The Quaternary of New England SCHAFER,HARTSHORN

The Coastal Plain and the Appalachian Highlands in the Quaternary RICHARDS, J~SON R8sumC of the Quaternary Geology of the Northwestern Gulf of Mexico Province BERNARD, LE BLANC The Pleistocene in Nebraska and Northern Kansas REED,DREESZEN,BAY~, SCHULTZ Quaternary of the Southern Great Plains FRYE,LEONARD

Glaciation of the Rocky Mountains RICH~TOND The Cordilleran Ice Sheet of the Northern Rocky Mountains and Related Quaternary History of the Columbia Plateau RICHMOXD, FRYXELL,NEFF, WEIS Nonglacial Quaternary Geology of the Southern and Middle Rocky Mountains SC~TT Snake River Plain MALDE Quaternary Geology of the Great Basin MORRISON Quaternary Geology of the Southwest KOTTMWSKI,COOLEY, RU~ The Quaternary of the Pacific Mountain System in Cahfornia WAHRHAFTIG,BIRMAN The Glacial History of Western Washington and Oregon H AND ELL The Quaternary Geology and Archaeology of PBwB, HOPKINS,GIDDIKGS

ZOOGEOGRAPHY AND EVOLUTION

QUATERNARY OF NORTH AiliIERICA

THE CONSPICUOUS remains of large extinct Simpson, 1941, 1945; Skinner and Kaisen, 1947; Stock, such as and ground have long held the 1925). public interest. of Quaternary mammals both large The history of mammalian paleontology in North Amer- and small are common in most parts of , ica is largely one of emphasis on comparative morphology, but they have not been studied so intensively as those of phylogeny, and origin of higher categories. This has led the earlier Cenozoic. The special advantages that study of generally to emphasis on the earlier Tertiary record and Quaternary mammals offers are a relatively full record, the to the lack of integration of related aspects of geology possibility of establishing detailed phyletic lineages, and and biology. We believe that the stucly of fossil mammals the opportunity to draw on a vast reservoir of knowledge in general, and particularly of Quaternary mammals, must of ecology, morphology, general biology, and geographic provide a synthesis if it is to be more than descriptive distribution of the living fauna. morphology and phylogenetic speculation. Hence we have The known fossil record of Pleistocene mammals is more only briefly summarized the basic data of geographic and nearly complete than that of any other group. Figure 1 stratigraphic occurrences (Fig. 1, Table 2), and we discuss shows the distribution of the major faunal localities in more fully the promising avenues for future research. Com- North America, herein defined to exclude the Neotropical mon names of all genera mentioned in the text are shown Region (compare with the much smaller number of Terti- on Table 2. ary localities sh0.cv-n by Savage, 1958, Fig. 2). The wide geographic occurrence of the sites and the short time range of many genera and make mammals the most useful means of correlation within the Quaternary over wide SCOPE OF PLEISTOCENE areas of North America. The abundant fossil record, coupled General agreement in defining the -Pleistocene uith the fact that so many genera are still living (Table boundary is lacking. Application of potassium-argon racllo- I), offers hopes for rich synthesis of data from fossll and genic dates and geomagnetic reversal sequences may bring llving mammals. a consensus in the next decade. Our placement of the lower Previous summaries of North American Quaternary limit of the Pleistocene is based upon the earliest paleonto- mammals by Hay (1923, 1924, 1927) and Hlbbard (in logic evidence of markedly cooler clirnate in mid-latitudes. Flint, 1957, p. 458-467; Hlbbard, 1958) show the progres- The few radiogenic dates now available (Evernden et al, sive acceleration in knowledge that has been accomplished 1964) imply that the base of the Pleistocene as v-e define in the last 25 by bulk screen-washing or sifting of it is probably 2-3 million years old. fossiliferous matrix. In the past 37 years, 49 of 155 genera Mammalian evidence for recognition of the end of the have been added to the known Quaternary record; 10 are Pleistocene in North America is based on of large mammals but 39 are small, being mostly insectivores, such large species as , mastodons, camels, large bats, and rodents. Despite this progress the knowledge of peccaries (Mylohyus and Platygonus) , Cervalces, the larger the North American record lags far behind that of Europe , Woodland (Symbos), Giant Beaver (Casto- (Flint, 1957, p. 451-455; Fejfar, 1961), even allowing for roides), ground sloths, and horses. The gradual extinction the Merent density of paleontologists. Comprehensive tax- of these species occurred through an interval of at least onomic works on North American Quaternary mammals 10,000 years (Hester, 1960) ; some probably lived as re- are rare (Frick, 1937; Klingener, 1963; Osborn, 1936-42; cently as 4000 years B.C. The most conspicuous changes in Museum of Paleontology, University of Michigan, Ann the North American mammalian fauna thus occurred after Arbor, Michigan. the Wisconsin glaciation. 'US. National Rluseum, Washington, D.C. Four chronologies are especially signfieant for correla- Department of Paleontology, University of , tion of Quaternary events. They arc provicIec1 by (a) the Berkeley, California. biostratigraphic record of land mammals; (b) glacial-inter- 'Museum of Zoology, University of Michigan, Ann iirbor, Michigan. glacial climatic fluctuations, inferred from many sources Carnegie Museum, Pittsburgh, Pennsylvania. besides physical-stratigraphic data; (c) geomagnetic-re- 509 Figure 1. Major faunal sitcs. Most of the wcll-ltnown sites for Pleistoccnc mammalian assemblages in North for those faunas not correlated with the glacial and intcrglacial stagcs; and (N) America are indicated by numbcrs, on Figure 1, that correspond with the follow- Nebraskan, (A) Aftonian, (LA) Latc Aftonian, (10 Iiansan, (EIO Early ICansan, ing list. Following the name of the locality is the age according to the following (LK) Late Kansan, (Y) Yarmouth, (I) , (LI) Late Illinoian, (8) Sanga- dcsignations: (LB) Late , (Ir) , and (R) mon, (LS) Late Sangamon, (W) Wisconsin, (EW) Early Wisconsin, (MW) Mid-

512 BIOGEOGRAPHY: ZOOGEGIGRAPHY AND EVOLUTION ica of more prinlitive members of phyletic series whose Late Cenozoic Genera of Non-marine Nearctic Mammals later members occur in Villafranchian faunas of Europe. (Counts in the two right-hand columns exclude a few marginal Such series are the follo~ving: genera that are dominantly Neotropical. The number of genera known as fossils is taken from Table 2) Ogmodontomys + Cosomys + Mimonzys (s.s.) Nek~olagus + O~yctolagusand Lepus

CORRELATION OF IfAblMALIAN FAUNAS TITH GLACIAL SEQUENCE The Great Plains region of central North America is the only area where faunas older than the limit of C14 dating have been related to glacial- changes. Most information comes from the intensively studied Meade Basin of southwestern Kansas (H~bbard,1963b; H~bbard and Taylor, 1960; Taylor, 1960). Correlation of this local faunal and stratigraphic sequence with glacial chronology Marsupialia is based on (a) inferred climatic changes correlated with Insectivora Chiroptera glacial-interglacial fluctuations, and (b) recognition of Primates lithologic units in southwestern Kansas that can be corre- Edentata lated with those of the glaciated region. Lagomorpha The sequence of diversified vertebrate and invertebrate Rodentia faunas from southwestern Kansas and the climatic shifts Carnivora inferred from them (Fig. 2) is the basis for recognizing Perissodactyla glacial and interglacial intervals in this area. The magni- Artiodactyla tude of the faunal shifts, in a vast region of little relief, Total favors the interpretation that the climatic changes affected most of North America and were probably associated versa1 sequences, which yield a relative chronology that is with changes in global circulation. In the southern Great becoming tied to dates in years (Cox et al., this volume) ; Plains there is evidence of four major episodes of cooler and (d) potassium-argon radiogenic dates (Evernden et al., climate separated by three intervals of warmer climate. 1964). The correlation of these chronologies- with each other These intervals are matched with the traditional four glaci- and with the record of marine sediments and sea-level ations and three interglaciations of the Mississippi Valley. changes will gain from and richly benefit the study of fossil The petrographically distinctive Pearlette ash (Powers mammals for the foreseeable future. et al., 1958; Swineford, 1949) occurs widely but discon- Most of the well-known sites of Pleistocene mammalian tinuously in the Great Plains and is associated at many assemblages in North America are indicated by number in places with vertebrate and molluscan fossils of the Cudahy Figure 1, the caption of which provides an entrance to the fauna, including representatives of many northern or even literature. The faunas we exclude as late Pliocene are species (Hibbard, 1949, 1960; Paulson, 1961). These Tehama, Tulare, and San Joaquin (California) ; Hagerman northern elements, and the qontrast between this fauna and and Sand Point (Idaho) ; Benson (Arizona) ; Rexroad and the immediately younger interglacial Borchers local fauna, Bender (Kansas) ; and Red Corral (). ind~catethat the Pearlette ash is of glacial age. In north- western Iowa the Pearlette ash occurs stratigraphically above till judged to be Kansan and below Loveland NORTH AMERICAN LAND- AGES (Condra et al., 1950, p. 22-24; Frye et al., 1948). Successive phases in evolution of the North American The Loveland loess in Iowa is younger than Illinoian till manlnlalian fauna are the basis of the land-manlmal ages. and older than Iowan till (Kay and Graham, 1943, p. 50). The most recent three of these are the Blancan, Irvingto- It has been traced from Iowa westward into Nebraska nian, and Rancholabrean (Hlbbard, 1958; Savage, 1951; (Condra et al., 1950; Lugn, 1962) and southward to Wood et al., 1941). Table 2 shows the combinations of southern Kansas (Frye and Leonard, 1952, p. 118), where restricted ranges, as based on the first and last appearances much of the Kingsdown Formation (Cragin, 1896) is cor- that characterize these ages. Outside of central North relative with it (Hrbbard, 1955a, p. 188; Bbbard et al., America, a given fauna is assignable to a land-mammal age 1944). In the Meade Basin the Kingsdown Formation has even though its place in glacial chronology can scarcely yielded the Mt. Scott local fauna of late lllinoian glacial ever be established. Recent potassium-argon radiogenic age (ECibbard, 1963b) and the Cragin Quarry local fauna dates (Evernden et al., 1964) can be correlated roughly of early Sangamon interglacial age (Hibbard and Taylor, mth the mammalian ages and thus indirectly E-ith glacial- 1960), and it is capped by a bed that is interpreted interglacial events. At present only the earliest Blancan as the Cca horizon of a pedocal that formed when the faunas (about 2.5 to 4 x lo6 years old) seem to be Pliocene, Sangamon soil developed in more humid areas to the north i.e. older than the earliest recognized continental glaciation and east. The Sangamon soil has been recognized widely that reached mid-latitudes. These faunas also are pre- on top of Loveland loess in Kansas and Nebraska (Frye Villafranchian according to the occurrence in North Amer- and Leonard, 1952, p. 119-123). QI;IITER~YARY.IIAJI.IIALS OF NORTH AMERICA TABLE2

Range of Late Pliocene and Pleistocene llammalian Genera Arranged According to First Appearance (Genera marked xith an asterisk (') are till living, but extinct in the Nearctic Region)

I III I Ardodus, giant short-faced 8eors Sorex, longtoiled shrews I1 Microsorex, shrews Bossoriscus, ringtoil cots I Eptesicus, brown bots Peromyscus, white-footed mice I I Mammut, American mastodons I I I IGuIo, woiverine Hypabgus, extinct rabbits Dipodomys, kangaroo rots Mochoirodus, sabre-tooth cots Gndotro. muskrots Scopanus, western moles ~henocimys,phenocomys Sciurus, tree squirrels Microtus, voles Citellus, ground squirrels Pedomys, prairie voles Conis, and wolfs Pifymys, .voles Nmfiber, woter rats Mortes, martens and fishers Erethizon, porcupines Mustelo, weosels and minks Sylvilogus, cotton- robbits Nonnippus, three-toed horses Le~us.hares Rhynchotherium, mastodons Nothrotherrum, small ground sloths Pliouchenio, extinct camels TI I-I'Chlomvtherium, aiont ~inobdstis,sob;eItooth cot , sabre-tooth cats Toxidm, badgers Hydrochoerus,' capyboros Mormoto, woodchucks and marmots Mommuthus, mommoths Costor, beavers Mylohyus, woodlond peccaries Perognothus, pocket mice Eucerotherium, shrub-oxen Onychomys, grasshopper mice Preptoceras, shrub-oxen Ochotono, pikas Tetromeryx, extinct pronghorns Prcdipodomys, extinct kangaroo rat: Equus (Equus),' modern horses Bensonomys, extinct mice Equus (Hemionus?),' hemionus-like horses Dipoides, extinct beavers Topirus,' topirs Buisnictis, extinct mustelid Stockoceros, extinct pronghorns Ogmodontomys, extinct voles Brochyprotomo, extinct skunk Notologus, extinct rabbits Glyptodon, glyptodons Boiomys, pigmy mice Gsmotherium, extinct skunk Odocoileus, Tremorctos,' spectacled bears Megalonyx, ground sloths Neotoma, woodrats Tonupolorno, extinct llomos Plesiothomomys, extinct gophers Poenemormoto, giont woodchuck Etodonomys, extinct kangoroo rat Blorino, shortoiled shrews Plofycerobos, extinct bovid Cryptotis, least shrews Scolopus, eostern moles Notiosorex, desert shrews Poroscolops, hairytail mole Losiurus, tree buts Antrozous, pallid bots Urocyon, gray foxes Plecotus, big-eared bots Procyon, raccoons Mephitis, striped skunks Spilogole, spotted skunks Tomios, eostern chipmunks Tamiosciurus, red squirrels Geomys, eostern pocket gophers Cle'thrionomys, redback voles Thomomys, western pocket gophers Bison, bison Sigmodon, cotton rots Desmodus, vompire bat Reithrodontomys, harvest mice Dosvoterus,. yellow. bat Zopus, jumping mice Dosypus, ormodillos , extinct mastodons Didelohis. oDossum Plotygonus, extinct peccaries r 1 r \+I M yotls,'. l~ttle'. ' brown bats Parohodomys, extinct woodrots ~idorido,freetoil bots Stegomostodon, mastodons Conepotus, hognose skunks Titonotylopus, giont camel Gloucomys, flying squirrels Pliopheno~om~s,extinct voles Bootherium, extinct bovid Piesippus, zebrine horses Cervolces, extinct moose Borophogus, -eating dogs Brochyostrocon, glyptodon Trigonictis, extinct grison Bormstrocon,- glyptodon lschyrosmilus, sabre-tooth cot Eremotherium, giont ground Chosmoporthetes, extinct hyaena Porodipaides, extinct beover Procostoroides, extinct beaver Oryzomys, rice rats Nebroskomys, extinct voles Neochoerus, extinct capybora Hesperoscolops, extinct moles Condylura, starnose mole Neterogeomys, extinct gophers Mormoops, leafchin bat Protilepus, extinct robbits Leptonyderis, longnose bat Porocryptotis, extinct shrew Pipistrellus, pipistrel bats ~rochyopsigole,extinct mustelid Eumops, mostiff bot Symmetrodontomys, extinct mouse Aplcdontia, sewellel Nekrologus, extinct rabbit Eutomios, western chipmunks Cerotomeryx, extinct pronghorn Crotogwmys, Mexican pocket gopher Cosomys, extinct vole Liomys, Mexican pocket mice Conimortes, extinct mustelid Dicrostonyx, collared lemmings Pliopotomys, extinct voles Nopomropus, woodland jumping mice S~no~tomys,bog lemmings Rongifer, coribou Antilocopro, pronghorn Comelops, extinct camels Greamnos, mountain gwt EquuS (Asinus?)*, ass-like horses aibos, musk-oxen Pliolemmus, extinct vole &is, bighorn sheep Glyptotheriurn, glyptodons Heterogeomys,* tropical gophers Soigo; osiotic ontelope Bos,' yok Songomono, caribou? symbos, woodland musk-ox Ca~romeryx, extinct pronghorns Alces, moose Costoroides, giont beovers Homo, man Simanycteris, extinct bat Hoyoceros, extinct pronghorn BIOGEOGRAPHY: ZOOGEOGRAPHY AND EVOLUTION

MAMMALIAN GLACIAL CLIMATES AND FAUNAL SHIFTS ! EPOCHS FAUNAS SIGNIFICANT ! AGES summer AGES GEOLOGIC DATA - Mild - winter elements elements / \ Mesothermal, semiarid, I RECENT Living Many formerly sympatric ' Sigmodon / / continental species now allopatric \ [ hispidus ' \ erosional unconformlty Ambystoma, neotenic I \ Microthermal, subhurnid, Sorex cinereus 0 Wisconsin Jones 'I C continental Citellus richardsoni \ 0 C Microtus pennsylvanicus / erosional unconformity - - - z Sorex cinereus / Microtus Ambystoma, metamorphosed a Mesothermal, humid / Jinglebob W maritime pennsylvanicus ,' Terrapene llanensis a ,--C A Oryzomys fossilis m Sangomon . caliche bed - Semiarid -- / a I Geochelone Crotaphytus collaris _I Cragin Mesothermal, subhumid, Phrynosoma modestum I Terropene llanehsis 0 Quarry I maritime \ Holbrookia texona NotiosoreA crowfordi 0 \r -- - Dasypterus golliheri z Sorex arcticus. a \ Terrapene llanensis Microthermal, subhumid, S. cinereus \ Blarina b. carolinensis a M~,Scott maritime 2 S. palustris Microtus penmyfvanicus DryZomys lllinoian '\,,-- W Perca flavescens \ z Butler Microthermal, subhumid, Ambystoma, neotenlc : W Spring continental? Sorex cinereus 1 0 Microtus pennsylvanicus 0 erosi~aluncmformity /---' ------/ caliche bed Semiarid Ambystoma, metomorphosed I- Z Mesothermal, subhumid, Geochelone UJ 5 Yormouth Borchers / Z maritime - 0 o I Sigmadon hilli w t- ' Spilogole cf S. ambarvalis -1 a '------z Pearlette ash bed Ambystorno, neotenic \ Q Microswex protensis \ a - Microthermal, subhumid, Sorex cinereus > ah Synoptomys (Mictomys) a- maritime S. lacustris Microtus paroperarius / c S. megapalustris / Kansan - 4 No climatically Se ger significant species erosional unconformity 1 Ambystoma, metamorphosed Mesothermal, subhumid, Sanders Some warm- I, Sigrnodon intermedius maritime temperate and Bensonomys meadensis subtropicol Pliolemmus ontiquus Aftonian caliche bed -Semiarid ! elements from I z Geochelone Mesothermal, Rexroad fauna a Deer Pork I maritime absent Pliopotomys meodensis u \ Pliolernmus antiquus Z \

Q No clirnatically Nebraskan Unnamed -I I significant species m 1 erosional unconfwmity f w Mesothermal, subhumid, I Mammalian f auna z Bender w moritime 9 not published 0 caliche bed -Semiarid 1 -0 I Geochelone rexroadensis Nerterogeorny s -I Mesothermal, subhumid, O.' Notiosorex jacksoni minor a Rexroad c I maritime I Baiornys spp. Bassariscus 1 Sigmodon intermedius casei

Fiwre 2. Late Cenozoic faunal shifts and inferred climatic changes in the southern Great Plains. QUATERNARY A1fAL11ML4LSOF NORTH AMERICA 515

Changes- in geographic--- distribution of species through geo- Earliest Xorth American Records of Eurasian Immigrants logic time are usually accompanied by morphological GESERA EARLIEST RECORDS changes. Hence biogeography is scarcely separable from UTSU Late Blancan (Loc. 46) e~olution.JVe consider three aspects of these topics: faunal illammu thus Late Kansan (Loc. 45) shifts. which are changes- in distribution associated with Preptoceras? Late Kansan (LOC. 44) ecological changes but with little taxonomic differentiation; is& Illinoian (Loc. 48) historical biogeography, involvmg changes in distribution Smilodon Irvingtonian (Loc. 100) Gulo Irvingtonian (Loc. 100) over much of Sorth America (including the Bering Straits Euceratheriunz Irvingtonian (Loc. 9) area), associated with some taxonomic differentiation; and Cervalces Rancholabrean (Loc. 99) evolution, in which geographic distribution is relatively Rangijer lT7isconsin (Hibbard, 1958) insignificant. Oreamnos JVisconsin (Hibbard, 1958) Ouibos lyisconsin (Hihbard, 1958) Ouis llTisconsin (Hihbard. 1958) FAUXAL SHIFTS Alms llTisconsin(~ibbard; 1958) Bos Wisconsin (Hibbard, 1958) The fossil record is so discontinuous that virtually no well- Saiga Wisconsin (Hibbard, 1958) documented evolutionary sequence of American Pleistocene Bootherium Wisconsin (Loc. 95) mammals have been described. Even at the generic level Symbos ll'isconsin (Loc. 95) the number of first appearances seems to be correlated with the number of known localities (Table 3). One of the mollusks). The climatic interpretations are drawn from principal reasons for the discontinuous record is the large- a variety of organisms, including mammals. These inter- scale faunal shifts that include such occurrences as pretations are the simplest possible based on the assump- in Pennsylvania (Simpson, 1945), ground sloths in Alaska tion that fossils imply past habitats like those of their (Stock, 1942), and boreal lemmings in the southern Great living morphological equivalents. A consistent application Plains (Hibbard, 1949). of this assumption suggests that present-day climates, with The most detailed studies of Pleistocene faunal sequence their seasonal extremes of temperature and aridity, are come from the Great Plains, mainly southwestern Kansas geologically atypical, even of the Pleistocene. Most of the and northwestern . A general hypothesis of major specific fossil associations would be inexplicable if the climatic shifts in this region has been developed by Hib- climate of the present were taken as the key to the past. bard a,nd Taylor (Hibbard, 1944, 1949, 1960, 196313; Taylor, But if different distributions of precipitation and seasonal 1960, 1965; Hibbard and Taylor, 1960). The inferred extremes are considered as separate variables, seemingly climatic sequence explains a wide variety of data, such as anomalous associations of species are rationalized. the former association of species that are now allopatric The first major southward faunal shift of ecological (living in widely separated areas), the repetitive strati- significance is shown by some of the mammals of the graphic occurrence of species, and the present-day relic Cudahy fauna (Hibbard, 1949, p. 1421) of late Kansan occurrence of others. It applies to (and is drawn from) such age. Some living species of mammals of the Boreal Sub- diverse groups as fishes, amphibians, reptiles, birds, mam- region extended their ranges southward during Illinoian mals, and mollusks. The same hypothetical climates are and/or Wisconsin time. Their remains have been recovered unlikely to have occurred outside central North America, in southern localities, e.g. Ovibos, Muskox (Fig. 3) ; Mi- but they unify so many facts and interpretations in this crotus xanthognathus (Fig. 4) ; Sorex arcticus (Fig. 5) ; and area that they will be a useful tool for future research. Dicrostonyx, Collared Lemming (Fig. 6). Remains of Sorex Climate can affect individual mammals directly by lethal cinereus, Masked Shrew, and Synaptomys cooperi, Bog extremes of temperature or by drought, but its effects on Lemming, are found as far south as San Josecito Cal-e, species or genera of mammals t,hrough geologic time are hfexico (Fig. 1, Loc. 26) ; and Sorex palustris, Northern probably indirect, acting through the and veget,ation Water Shrew, is known from Oklahoma, Kansas, and north- of the habitat. Smaller mammals, with a narrower range central Tennessee (Fig. 1, Loc. 48, 56, and 93). of habitat and individual home range than the larger The Yellow-cheeked Vole (Microtus xanthognathus) and carnivores or herbivores, are influenced more by the micro- the Rock Vole (M. chrotorrhinus) lived in the same area climate in which they live than by regional climatic change in Pennsylvania and (Fig. 4) during late Wiscon- (Burt, 1958). Hence it is reasonable and a justification of sin time; they now live in widely separated areas in Boreal uniformitarian interpretations that the ecological conclu- North America. Guilday (1963) recovered fossil remains sions drawn from rodents and insectivores are similar to of Dicrostonyx (iMisothermus) hudsonius, Labrador Col- those drawn from the associated lower vertebrates, mol- lared Lemming, in Pennsylvania (Fig. 6). He considered the lusks, and plants. The consistency of inferences from dif- present tundra population of this species to be a probable ferent elements of late Cenozoic biotas in the Great Plains relic of a former Holarctic pre-JVisconsin distribution. is decisive proof (and the only test possible) of the validity Southward and southwestward retractions of ranges of of the method. ecological significance during the later Pleistocene are Figure 2 summarizes the latest Cenozoic sequence of shon-n by the present distribution of Dusypterus, Yellow faunas and inferred climates in southwestern Kansas (see Bat (Fig. 4) ; Hydrochoerus, Capybara; Oryzomys, Rice Taylor, this volume, for a similar summary of associated Rat (Fig. 3) ; Neofiber, Water Rat (Fig. 1, Loc. 40, 61, and 516 BIOGEOGRAPHY: ZOOGEOGRAPHY AND EVOLUTION Loc. 93, 96, 97, 98, and 104); Taxidea, Badger (Fig. 1, Loc. 97 and 100); Conepatus, Hognose Skunk (Fig. 5); and Ochotona, Pika (Fig. 7).

HISTORICAL BIOGEOGRAPHY The Nearctic Region (Canada, continental United States, northern Mexico, including the central plateau but not the coastal lowlands on either side) is inhabited by 111 genera of mammals. These have various patterns of dis- tribution and are variably distinct from the mammals of the Palaearctic and Neotropical Regions. The most marked endemism is at the family level: the pronghorn (Antilo- capridae) and sewellel (Aplodontidae) are restricted to the Nearctic. At the other extreme 21 species may be common to North America and Eurasia (Rausch, 1963), and many more to the southern Nearctic and the Neotropical Re- gions. These different patterns of distribution and different degrees of endemism imply different histories. The Cenozoic (mainly pre-Quaternary) differentiation and spread of the North American familles and many genera of mammals have been summarized by Burt (1958), Savage (1958), and Simpson (1947). We comment on the Quaternary spread of mammals (a) between the Neotropical and Ne- Figure 3. Recent distribution (A) of the Regions, (b) between the Palaearctic and Nearctic Muskox, Ovibos moschatus, with fossil records (99, and unnum- Regions in the Bering Straits area, and (c) within the bered circles). Open circles indi- Nearctic. cate questionable identifications. Data from reference citations 28, 43, 49, 50, 52, 83. Northerly por- tion of the Recent distribution (B) of the Marsh Rice Rat, Ory- zomys palustris (including 0. couesi, fide E. R. Hall, 1960, Southw. Nat., v. 5, No. 3, p. 171- 173), with fossil records for 0. fossilis (43, 56, 58). Data from reference citations 20, 43, 60, 67. Oryzomys palustris is known also from several post-Pleistocene pre- historic localities, not mapped, east of the Mississippi River and north of its modern limits.

100) ; Conepatus, Hognose Skunk (Fig. 5) ; Felis pardalis, Ocelot (Fig. 7) ; Panthera onca, Jaguar (Fig. 8). The presence of maritime climates and extensive broad, marshy, undissected valleys seems to have provided the habitat for the northern ranges of the Capybara, Rice Rat, and Water Rat. Neofiber, the Water Rat, is a good example of a form whose range has become greatly restricted because of the lack of habitat or because of competition with Ondatra, the Muskrat, or for both reasons (Birkenholz, 1963). Neo- fiber is not closely related to Ondatra, and in middle Pleis- Figure 4. Recent distribution (A) of the tocene faunas (Port Kennedy Cave, Fig. 1, Loc. 100, and Rock Vole, ?Microtus chrotorrhi- nus, with fossil records (9,6, 98, Rezabek, Fig. 1, Loc. 61) these two forms were associated, 104), (B) of the Yellow-cheeked although at the present time their ranges do not overlap. Vole, M. xanthognathus, with fos- It is of interest to note that the range of the Jaguar during sil records (96, 98, 104), and the the Pleistocene has been approximately that of the peccaries. northerly portion of the Recent Westward shifts in ranges are shown by Mormoops, Leaf- distribution (C) of the yellow bat, Dasypterus, with fossil records chimed Bat (Fig. 6); Desmodus, Vampire Bat, (Fig. 1, (57, 82). Data from reference cita- Loc. 26, 78, 79, and 82) ; Citellus, Ground Squirrel (Fig. 1, tions 39,42-43,71. QUATERNARY JfAAIJrlALS OF NORTH AMERICA 517 Eurasia. Such immigrations are particularly noticeable at the family level ( and ) but range through lower taxonomic ranks. Table 3 summarizes the earliest known occurrences of genera that probably en- tered Worth America as the same or closely related genera during Quaternary or immediately pre-Quaternary t,imes. A recent study by KurtCn (1963) points the way ion-ard future refinements. Zoogeographical studies of living and fossil mammals (Burt, 1958; Rausch, 1963; Savage, 1958; Simpson, 1947; Stokes and Condie, 1961) have cast light on the approxi- mate t,imes and range of habitats in Quaternary faunal dispersal in the Bering Straits region. Direct paleontologic and stratigraphic evidence in the Seward Peninsula athwart the path of mammalian spread has become available only recently (Hopkins, 1959a, b) . Table 3 shows that half of the immigrant genera listed have an earliest known record in the Wisconsin. These genera are predominantly those characteristic or tolerant of arctic-alpine environments. The relative predominance of fossil deposits in Alaska of Wisconsin age is partly re- sponsible for the first records of such forms, but the relative severity of Wisconsin-age climates is probably a Figure 5. Recent distribution (A) of the significant cause also. Fossils of latest Pleistocene mam- Arctic Shrew, Sorex , arcticus, in moth, bison, and horse in a region then vegetated with North America, with fossil records (48, 56, 93, 96, 98, 104), and the northerly portion of the Recent distribution (B) of the Hognose Skunks, Conepatus (including the nominal species C. mesoleucus and C. leuconotus) with fossil rec- ords (27, 78, 81). Data from ref- erence citation 43 and from those cited under each locality in the explanation to Fig. 1.

hTeotropicd-Nearctic relationships. The fossil record in and L~Iexicois so little known that times of faunal interchange in Pliocene and Pleistocene times can scarcely be defined. Some groups of ultimate South Ameri- can ancestry (hystricomorph rodents and edent,ates) ex- tended their ranges northward into the Nearctic Region during this time. Quaternary events may have had little effect on these genera except in changing t,he limits of habitable areas during glaciations. Some genera that are now Neotropical (for example, the peccaries and , Tapirus [Tapirella]) occurred dur- ing Quaternary t,ilne far north of their present range. They arc most reasonably regarded not as tropical or subt,ropical element's in Quaternary faunas but as genera that were once Figure 6. Recent mainland distribution (A) widespread in the southern Nearctic Eegion and that have of the Collared Lemming, Di- been es-tinguishecl in much of their range by late Quater- crostonyx groenlandicus, (B) of nary climatic changes. the Labrador Collared Lemming D. hwlsonius, mith fossil record Polaearctic-hiearctic relatiu7zships. During late Pliocene (98), and the northerly portion of the Recent distribution (C) of and Quaternary times the spread of mammals beheen Peters' Leaf-chinned Bat, Mor- the Old and New Worlds could only have occurred in the moops ntegalophylla, with fossil Bering Straits region, although for the early Tertiary n-e record (84). Data from reference cannot categorically exclude faunal connections around the citations 40,43, 116. Rawch (1963) Korth Atlantic. The clearest paleontologic e\+dence is the regards the above forms of Di- crostonyx as conspecific mith D. appearance in North il~nericaof groups with ancestors in torquatus. 518 BIOGEOGRAPHY: ZOOGEOGRAPHY AND EVOLUTION tundra (Hopkins, 1959b) show that many of these mam- mals could have crossed the emergent Bering-Chukchi Platform during a time of lowered sea level in the . An earlier immigration that will be of significance both to regional stratigraphy in North America and to chronol- ogy of the Bering Straits region is that of Mammuthus. Two separate stocks of this genus seem to have entered North America. The later was M. primigenius, the Woolly , during the Rancholabrean. The earlier was the stock first seen as Mammuthus haroldcooki in Kansas and Oklahoma in late Kansan deposits. It may have given rise to all later American species except M. primigenius. These early were not boreal animals; they are rare in southwestern Kansas (Hibbard, 1953) and in Oklahoma (Meade, 1953), and are common in the Seymour For- mation of north-central Texas (Hibbard, in preparation). The passage of these animals across t'he Bering Strait dur- ing a glacial interval seems most unlikely, and yet the morphological grade of the first immigrants seems to pre- clude a Pliocene age. Perhaps the of our definition precedes the early Pleistocene of HopkinsJ (1959a) usage. Figure 8. Northerly portion of the Recent Intra-Neurctic relationships. The present patterns of dis- distribution of the Jaguar, Pan- tribution and the composition of the living fauna are the thera (Jaguarius) onca, with fossil records including P. (J.) onca au- gusta but excluding P. (J.) atrox. The' locality in Nebraska repre- sented by the unnumbered solid circle is the type locality ("Nio- brara RiverJJ) for P. (J.) onca augusta. Open circles indicate questionable identifications. Data from reference citations 43, 94, 105, 129, 131.

product of extinction and evolution as well as of consid- erable restrictions in distribution. Such changes can be studied most effectively by synthetic biogeographic studies of the type made by Blair (1958) and Neil1 (1957). The frequent disagreements about the chronology of range changes and about the age of a given pattern of distribu- tion indicate that biogeography is unlikely to be a fruitful area of research without an objective method of analyzing its data. Quaternary restrictions of range are known in many genera. The gopher, Geomys, and shrew, Blarina, which formerly occurred west of the continental divide but are now eastern (Hibbard, 1959; White and Doms, 1961) probably were affected by the regional disappearance of Figure 7. Recent distribution (A) of the t,heir habitat. Late Pleistocene aridity that deprived them Plka, Ochotona (Pika), in North of moist soil and humus is a plausible cause. In other cases America, with fossil record (971, the reasons remain obscure. and the northerly portion of the Recent distribution -(B) of the EVOLUTION Ocelot, Felis (Leopardus) pardalis, with fossil record (82).Data from The Pleistocene fossil record of mammals in North America reference citations 35, 43, 116. is more nearly complete than that of any other class of The fossil Ochotona belongs to organisms. As shown in Table 4, only 20% of t.he extant the subgenus Pika on the basis of partial separation of the incisive genera are unrepresented as fossils. Despite t,his relatively and palatine foramina. full record, the phg-letic development of practically no QUATERATARY JfAJlMdLS OF NORTH AiIIERICfA 519 TABLE 4 3. KurtCn (1963) has discussed Old JVorld-Sew World First and Last Appearances of Late Cenozoic llneages in the saber-toothed cats of the -Dino- Alammalian Genera Shorrn in Tables 1 and 2 bastis and Sfegantereon-Snzilodon lines and in the black (K = number of localities that are shox-n in Fig. 1. Late Pliocene bears of the lirsus thibetanus-U. americanus line. localities are listed in the text under Scope of Pleistocene) -1 possible example of apparent evolutionary change that First appearance Last appearance re~dtsonly from ecological changes has been described by Hibbard (196%). Stratigraphically distinct samples of the Per- Per- cent Number cent Number Short-tailed Shren., Blarina, represent different-sized ani- mals that ~vouldbe ascribed to an evolving lineage in con- Late Pliocene (N = 9) 52 37of72 ventional study. Similar differences can be found in the Early Pleistocene large northern and small southern subsoecies of Blarina ( = post-Pliocene brevicauda, and associated faunas support the interpreta- Blancan) (N = 14) Irvingtonian (A7 = 14) tion that the observed morphological differences are due to Rancholabrean (N = 81) shifts of a cline. TTisconsin (N = 59) Living

The progressive modernization of the Quaternary Kearctic species has been tlocumented. Evidently changes in habitat mammalian fauna has taken place through the evolution of and geographic range have added to the other gaps in the new genera and species from autochthonous stocks, through record caused by geologic and human factors. immigration from the Xeotropical and Palaearctic Regions, through elimination by absolute extinction, or through ex- PHYLETIC SEQUENCES tinction of widespread groups in the Nearctic Region but The better-documented phyletic sequences include species with survival elsewhere, e.g. camels and horses. of the Jumping Mouse, Zapzss, and of the Wh~te-footed The only extant species known from pre-Quaternary Mouse, Peromyscus. Klingener (1963) described a series deposits is the badger, Taxidea tazus, in the Rexroad local of four successional subspecies leading from the late fauna. The known fossils ,include only lower jaws, and if Pliocene Zapus sandersi rexroadensis to the Recent Zapus a is found it will probably reveal specific differences. hzsclsonius (Fig. 9). Hibbard (1955a, p. 212; Hibbard and Even if not, the generalization that mammalian species are Taylor, 1960, p. 175) found that the Sangamon Peromyscus of Quaternary origin is valid. (The modernization of genera progressus is closely related to the living P. leucopus but and species in the faunal sequence of southwestern Kansas has more primitive tooth characters. P. leucopus may there- fore have evolved during the latest Sangamon and Wis- consin. Less well documented but plausible lineages have been inferred for several other genera of mammals. These phy- letic lines are the simplest and shortest that are consistent nith the available fossil record and the data of comparative anatomy, and hence their errors will prove mainly due to oversimplification. Such synthesis of fossil interpretation with data from related fields, and the discovery of more fossils, will favor the inference of longer stratigraphic ranges and of slov-er rates of change and geographic spread. Students of evolution, biogeography, and Pleistocene stra- tigraphy should understand that the mammalian evolu- tionary sequences are only the simplest interpretations of a fragmentary record. Such phyletic lines have been pro- posed, mainly on dental characters, in many genera of I\-hich we list only a sample. 1. Hibbard (1963a) has interpreted evolution of some rabbit genera as taking place in the early Pleistocene- Sylvilagus, Caprolagus, Oryctolagus, and Lepus from a Figure 9. Evolution of the species and sub- "pro-Sylvilagus" stock. species of the jumping mouse, 2. The stratigraphic succession of fossil mammoths is Znnus.. . after Klinnener (19631.. The consistent with an interpretation of progressix-e increase Pleistocene fosils are from locali- in number of enamel plates in the teeth. Hibbard (1953) ties 48, 51, 53, 54, 56, and 58 (Fig. suggested that the oldest known American species, Mam- 1) in southwestern Kansas and ad- jacent Oklahoma, but Zapus lives muthus haro:dcooki from the late Kansan of southwestern only farther north and east in the Kansas and Oklahoma, is ancestral to M. imperator. Great Plains. 520 BIOGEOGRAPHY: ZOOGEOGRAPHY AND EVOLUTION has been contrasted by Taylor, this volume, with the elimination of old stocks rather than the development of record of mollusks.) new ones. Taking the stratigraphic ranges (Table 2) at face value PROBLEMS one might wrongly infer the evolution of many new genera during the pleistocene. ~h~ ecologic replacement of some The problems available for future research include a herit- genera by others, the inverse correlation of number of age of old, unsolved ones, along with newly raised questions known fossil localities with age, and the immigration of that result from an increase in knowledge in paleontology animals into areas where fossiliferous sediments have been as as in preserved are all more likely causes of the known range The problem is as as the study than taxonomic evolution. Quaternary mammals. Why did the most conspicuous ex- ~h~ following pairs of genera occur in successive strata, tinctions occur so late and after the last glaciation? Re- with no zone of overlap or with a relatively short interval cent studies (Hester, 1960; Martin, 1958; Martin et al., of joint occurrence. Their rnorphologic differences are 1961) and the application of C1* dates have ~rovided a great enough to preclude ancestral-descendant relation- data but no consensus. We suggest that any ship, but their similarities are such that they may have plausible explanation of late Pleistocene extinction cannot overlapped ecologically. In such cases the first appearance be ad but must shed light On previous of the younger genera in the fossil record may be due , contemporaneous non-mammalian extinctions, primarily to ecological replacement. and on contemporaneous range changes in organisms that did not become extinct. OLDERGENERA YOUNGERGENERA Newly apparent topics, and those that can be newly at- Equus s.s., large horse tacked with profit, have come from new techniques as well Plesippus (zebrine) Equus (Asinus?) and as the gradual accumulation of data. All problems of Nunnippus (three rep1aced by Equus (Hemionus?), Quaternary chronology and correlation, and all topics that toed) smaller horse-like forms involve these matters, will be influenced by the results of 11replaced by Mammuthus, mammoth C14 and K/Ar dating and by research on geomagnetic Capromeryx replaced by Antilocapra, pronghorn polarity. Study of rates of evolution, of geographic spread, Xylvilagus, rabbit and and of regional changes in habitat are particularly cle- Hypolagus by(Lepus, hare pendent upon a rehed chronology. Pliophenacomys replaced by Alicrotus, vole A particularly worthwhile study with worldwide ramifi- Arctodus replaced by Ursus (brown and grizzly cations would be paleoecological analysis of the biota asso- bear but not the black ciated with the Pearlette ash. This ash provides a unique, bear) virtually contemporaneous datum throughout much of the Great Plains for a time when a continental ice sheet cov- The correlation of first appearance of genera with num- ered part of northern North America and the climate and her of known fossil localities is evident in Table 4. This its zonation was radically different from that of today. shows that a very high percentage of Pleistocene first ap- Such a study would require the long-term collaboration of pearances could be due entirely to quality of the fossil diverse specialists; it thus reflects our belief that knotvl- record, without any evolutionary changes or immigration edge of Quaternary mammals will be most advanced from outside areas. through studies devoted to geology and entire biotas, and The fossil record is so scanty, particularly in Alaska and not solely to Quaternary mammals. adjacent areas, that we do not know when groups arrived Examples of a few recent studies that exemplify valuable or how much of their taxonomic differentiation took place data gained from a synthetic approach are given below. It after their arrival and before their earliest record as fos- is no coincidence that some of the most valuable work is sils. Study of these immigrants can be carried out most by those not trained as mammalian paleontologists. effectively by biogeographic studies such as those by Stokes The occurrence of beaver-cut wood in peat deposits of and Condie (1961) on Ovis or by Rausch (1963) on the New England led Kaye (1962) to suggest that the region mammals of the Bering Straits area. in early postglacial time was less swampy than today. If The genera and subgenera Sylvilagus, Lepus, Lagurus, beavers mere responsible for providing sites of accumula- Microtus, Dinobastis, Smilodon, Mammuthus, Equus, tion for pollen-bearing sediments perhaps they also account Mylohyus, Camelops, Rangifer, Alces, Antilocapra, Bison, for irregularities in pollen stratigraphy. Oreamnos, Symbos, Ovibos, and Ovis might have origi- Late Pleistocene wood rat (Neotoma) middens in south- nated as late as early Pleistocene times. If so, they are the ern Nevada include twigs and seeds of juniper not now highest-ranking groups that are geologically so young. living in the lonr desert mountains. The relatively large Although mammalian evolution might have been acceler- size of the plant remalns makes them more significant ated during the Pleistocene in some groups, this time-span indicators of local vegetation than n-indblown pollen; hence was so short that differentlation hardly ever went beyond Wells and Jorgensen (1964) T+-ereable to ~nferlate Pleis- the species-group level If Ondatra is derived from Plio- tocene changes in climate and in distribution of ~troodrat potamys this would be the latest orlgin of a genus that and ~uniper75-ooclland. can be documented in the Pleistocene of North America. In an area near Denver, Colorado, Scott (1962, 1963) In general, the Pleistocene record is conspicuous by the studied Quaternary history in a particularly thorough way. S OF NORTH AMERICA 5.21 The fossil mammals he recorded do not include great Recent land mammals of n-estern North America: in diversity of species, but the integration of fossil mammals Hubbs, C. L. (ed.), Zoogeography: Amer. Asoc. Adv. with geomorphology, buried soils, detailed physical stra- Sci. Publ. 51, p. 131-154 tigraphy, and the record of other fossil organi-ms points 14. Camp, C. L., et al., 1940-1961, Bibliography of fossil the way for future research of this kind. vertebrates, 1928-1953: Geol. Soc. Amer. Spec. Pap. 27, The most abundant and detailed information about 503 p.; Spec. Pap. 42, 663 p.; biem. 37, 371 p.; Mem. glacial-interglacial changes in climate and habitats comes 57,465 p.; Mem. 84,532 p. from the Illinoian and Sangamon faunas of southwestern 15. Carr, W. J., and Trimble, D. E., 1963, Geology of the Kansas and northwestern Oklahoma (see references to American Falls quadrangle, Idaho: U.S. Geol. Surv. faunas 47, 48, 55-58). Fossils representing angiosperms, Bull. 1121-G, 44 p. (17) mollusks, fishes, amphibians, reptiles, birds, and mammals 16. Colbert, E. H., 1950, The fossil vertebrates: in E. W. provide a wide variety of sources of ecological and strat- Haury and collaborators, The stratigraphy and ar- igraphic data. From these studies far more information chaeoloa of Ventana Cave, Arizona: Tucson, Univ. about environmental changes has been gained than the Arizona Press, p. 126-148 (21) study of any one group could yield, and far more informa- 17. Condra, G. E., Reed, E. C., and Gordon, E. D., 1950, tion about mammals than a strictly mammalogical study Correlation of the Pleistocene deposits of Nebraska: would produce. Nebraska Geol. Surv. Bull. 15A, 74 p. 18. Cox, AIlan, Doell, R. R., and Dalrymple, G. B., this volume, Quaternary paleomagnetic stratigraphy Numbers in parentheses refer to list of faunas in explana- 19. Cragin, F. W., 1896, Preliminary notice of three late tion of Figure 1. 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Only 22 out of 111 living Nearctic mammalian genera are unknown as fossils, so mammals are the best known Quaternary group of animals in North America. The 155 Quaternary genera have restricted and overlapping ranges that are the basis for the Blancan (only the latter part is Quaternary), Irvingtonian, and Rancholabrean mammalian ages. The chronology based on mammals can now be related to those of glacial-interglacial climatic changes, geo- magnetic-polarity epochs, and potassium-argon dates, from which it appears that the Quater- nary of our definition began about 2.5 million years ago. The basic data of stratigraphic ranges (177 genera) and geographic occurrences (104 localities) are summarized with 160 references that provide entrance to a much larger liter- ature. Most of the text is devoted to discussion of promising avenues for future research and examples of the kinds of synthetic study that have proved fruitful. Faunal shifts correlated with glacial-interglacial climatic changes have been documented thoroughly only in the southern Great Plains. Interpretations based on small mammals are consistent with those based on lower vertebrates, mollusks, and plants. Immigration from the Old World via the Bering Straits is most marked among hardy boreal forms that may have crossed when sea level was lower. In spite of the relatively full fossil record, well-dxumented phyletic sequences of species are unknown outside the rodents. Although many genera are first known from Quaternary sediments, in practically all cases they probably lived before then. Probably all Recent mam- malian species originated during the Quaternary, but generic change was by extinction rather than evolution into new genera. PART I1 : BIOGEOGRAPHY

Phytogeography and Palynology of Northeastern United States DAVIS Problems in the Quaternary Phytogeography of the Great Lakes Region CUSHING Palynology and Pleistocene Phytogeography of Unglaciated Eastern North America WHITE- HEAD Pleistocene Pollen Analysis and Biogeography of the Southwest MARTIN,MEHRINQER Plant Geography in the Southern Rocky Mountains WEBER A Pleistocene Phytogeographical Sketch of the Pacific Northwest and Alaska HEUSSER The Boreal Bryophyte Flora as Affected by STEERE Polyploidy, Distribution, and Environment JOHNSON,PACKER, REESE

ZOOGEOGRAPHYAND EVOLUTION Quaternary Mammals of North America HIBBARD,RAY, SAVAGE, TAYLOR, GUILDAY Avian Speciation in the Quaternary SELANDER Amphibian Speciation BLAIR Reptiles in the Quaternary of North America AUFFENBERG,MILSTEAD Quaternary Freshwater Fishes of North America MILLER Pleistocene Events and Insects ROSS The Study of Pleistocene Nonmarine Mollusks in North America TAYLOR Other Invertebrates-An Essay in Biogeography FREY Recent Adjustments in Ranges SMITH GENERAL Pleistocene Nonmarine Environments DEEVEY

PART 111 : ARCHAEOLOGY

Late Quaternary Prehistory in the Northeastern Woodlands GRIFFIN An Outline of Southeastern United States Prehistory with Particular Emphasis on the Paleo- Indiau Era WILLIAMS,STOLTMAN Quaternary Human Occupation of the Plains STEPHENSON Postglacial Climate and Archaeology in the Desert West BAUMHOFF,HEIZEB Pacific Coast Archaeology MEIGHAN PART IV : MISCELLANEOUS STUDIES Late Quaternary History, Continental Shelves of the United States CURRAY Isotope Geochemistry and the Pleistocene Climatic Record BROECKER Quaternary RUHE Geochemistry of Some Quaternary Lake Sediments of North America SWAIN Quaternary Paleohydrology SCHUMM Glaciers and Climate MELER Volcanic-Ash Chronology-. wucox Quaternary Paleomagnetic Stratigraphy cox, DOELL, DALRYMPLE Tectonics of Quaternary Time in Middle North America KING Dendrochronology FRITTS Theoretical Paleoclimatology MITCHELL