Biogeographic implications of a packrat midden sequence from the Sacramento Mountains, south-central New Mexico

Thirteen packrat (Neotoma spp.) and two porcupine (Erethizon dorsatum) middens from 1555 to 1690 m elevation from the Sacramento Mountains, New Mexico, provide an 18,000-yr vegetation record in the northern Chiuahuan Desert. The vegetation sequence is a mesic, Wisconsin fullglacial (18,000–16,000 yr B.P.) pinyon-juniper-oak woodland; a xeric, early Holocene (ca. 11,000–8000 yr B.P.) juniper-oak woodland; a middle Holocene (ca. 8000-4000 yr B.P.) desert-grassland; and a late Holocene (ca. 4000 yr B.P. to present) Chihuahuan desertscrub. The frequency of spring freezes and summer droughts in the late Wisconsin probably set the northern limits of Pinus edulis and Juniperus monosperma at about 34°N, or 6° south of today's limit. Rising summer tempratures in the early Holocene eliminated pinyon and other mesic woodland plants from the desert lowlands and allowed the woodland to move upslope. At this time pinyon-juniper woodland and pine forest dominated by Pinus ponderosa probably began their spectacular Holocene expansions to the north. Continued warming in the middle Holocene led to very warm summers with strong monsoons, relatively dry, cold winters, and widespread desert-grasslands. Desertscrub communities in the northern Chihuahuan Desert did not develop until the late Holocene when the biseasonal rainfall shifted slightly back toward the winter, catastrophic winter freezes decreased, and droughts in all seasons increased. The creosote bush desertscrub corridor across the Continental Divide between the Chihuahuan and Sonoran deserts was probably connected for the first time since the last interglaciation.     

A long‐term vegetation history of the Mojave–Colorado desert ecotone at Joshua Tree National Park

Thirty‐eight dated packrat middens were collected from upper desert (930–1357 m) elevations within Joshua Tree National Park near the ecotone between the Mojave Desert and Colorado Desert, providing a 30 ka record of vegetation change with remarkably even coverage for the last 15 ka. This record indicates that vegetation was relatively stable, which may reflect the lack of invasion by extralocal species during the late glacial and the early establishment and persistence of many desert scrub elements. Many of the species found in the modern vegetation assemblages were present by the early Holocene, as indicated by increasing Sørenson's Similarity Index values. C₄ grasses and summer‐flowering annuals arrived later at Joshua Tree National Park in the early Holocene, suggesting a delayed onset of warm‐season monsoonal precipitation compared to other Sonoran Desert and Chihuahuan Desert localities to the east, where summer rains and C₄ grasses persisted through the last glacial–interglacial cycle. This would suggest that contemporary flow of monsoonal moisture into eastern California is secondary to the core processes of the North American Monsoon, which remained intact throughout the late Quaternary. In the Holocene, northward displacement of the jet stream, in both summer and winter, allowed migration of the subtropical ridge as far north as southern Idaho and the advection of monsoonal moisture both westward into eastern California and northward into the southern Great Basin and Colorado Plateau. Copyright © 2009 John Wiley & Sons, Ltd.     

The lower Colorado River Valley: A Pleistocene desert

A chronological sequence of plant macrofossil assemblages from twenty-five pack rat middens provides a record of desert scrub vegetation for most of the last 13,380 yr B.P. from a hyperarid portion of the lower Colorado River Valley. At the end of the late Wisconsin, and probably during much of the Quaternary, the Picacho Peak area, Imperial County, California, supported a typical Mohave Desert association of Larrea divaricata (creosote bush), Coleogyne ramosissima (blackbrush), Yucca brevifolia (Joshua tree), and Y. whipplei (Whipple yucca). Recent arrivals of Sonoran Desert plants such as Olneya tesota (ironwood) and Fouquieria splendens (ocotillo) suggest that the area supported relatively modern Sonoran desert scrub species for relatively short periods during interglaciations.     

Late Pleistocene–Holocene environments in Valle Carbajal, Tierra del Fuego, Argentina

The authors discuss Late Pleistocene–Holocene depositional environments in one of the Fuegian Andes valleys on the basis of palynological, geomorphological, and sedimentological analyses from two sites located near the Beagle Channel. The results obtained at these localities reinforce and refine the Late Pleistocene–Holocene climatic pattern previously recorded there. A colder period, associated with the Younger Dryas stadial event, is suggested by low Nothofagus pollen frequency, and communities of grass, low scrub, and shrub heath expanded into the low/middle slopes (10,310 ¹⁴C yr BP). By ca. 9500 ¹⁴C yr BP, warmer and drier conditions occurred, as evidenced by the development of open-grown vegetation in the valley floors (pollen zone O-3), followed by the expansion of open Nothofagus woodland (pollen zone O-2) in the middle Holocene. The milder climate subsequently changed, as indicated by the spreading of the closed forest and mire (pollen zone O-1), to more humid and cooler conditions during the last ca. 5000 yr BP.     

Late Holocene vegetation changes in Greenwater Valley,Mojave Desert,California

Small-scale late Holocene vegetation changes were determined from a series of 13 modern and fossil packrat middens collected from a site in the Greenwater Valley, northern Mojave Desert, California. Although the site is above the modern lower limit of Coleogyne ramosissima (black-brush), macrofossils of this shrub are only present in samples younger than 270 yr B.P. In order to measure changes more subtle than presence vs absence, macrofossil concentrations were quantified, and principal components and factor analyses were used to distinguish midden plant assemblages. Both the presence/absence data and the statistical analyses suggest a downward shift of 50 to 100 m for Coleogyne (blackbrush) communities between 1435 and 1795 A.D.     

An 11 000-year record of vegetation and environment from Lago di Martignano,Latium, Italy

A pollen diagram from Lago di Martignano, a maar lake in central Italy, provides an 11000-year record of vegetation and environment change. The earliest pollen spectra are dominated by Artemisia and Gramineae, representing late glacial steppe vegetation typical of the Mediterranean region. Broad-leaved forests were established by ca. 11 000 yr BP. Although Quercus initially dominated their canopy, a wide range of other mesophyllous trees were also present. Pollen values for sclerophyllous tree and shrub taxa characteristic of Mediterranean woodlands and scrub are initially low (<10%). After ca. 7000 yr BP, however, they begin to increase and rise to a peak of >40% of total land pollen at ca. 6700 yr BP, with Olea europaea the single most abundant taxon. Human influence upon the vegetation only becomes significant somewhat after this peak, with progressive clearance of woodland and expansion of herbaceous communities. Castanea sativa and luglans regia pollen is recorded consistently from the beginning of the rise in pollen values for taxa characteristic of Mediterranean scrub communities. Pollen values for arable crops increase progressively after ca. 5500 yr BP, following the peak pollen values for taxa characteristic of Mediterranean scrub vegetation. Late glacial and Holocene climate changes have been complex in this region, with the present character of the climate developing only during the last millennium. Rates of change of pollen spectra peak during this period.     

Postglacial vegetation history of Mitkof Island, Alexander Archipelago, southeastern Alaska

An AMS radiocarbon-dated pollen record from a peat deposit on Mitkof Island, southeastern Alaska provides a vegetation history spanning ∼12,900 cal yr BP to the present. Late Wisconsin glaciers covered the entire island; deglaciation occurred > 15,400 cal yr BP. The earliest known vegetation to develop on the island (∼12,900 cal yr BP) was pine woodland (Pinus contorta) with alder (Alnus), sedges (Cyperaceae) and ferns (Polypodiaceae type). By ∼12,240 cal yr BP, Sitka spruce (Picea sitchensis) began to colonize the island while pine woodland declined. By ∼11,200 cal yr BP, mountain hemlock (Tsuga mertensiana) began to spread across the island. Sitka spruce-mountain hemlock forests dominated the lowland landscapes of the island until ∼10,180 cal yr BP, when western hemlock (Tsuga heterophylla) began to colonize, and soon became the dominant tree species. Rising percentages of pine, sedge, and sphagnum after ∼7100 cal yr BP may reflect an expansion of peat bog habitats as regional climate began to shift to cooler, wetter conditions. A decline in alders at that time suggests that coastal forests had spread into the island's uplands, replacing large areas of alder thickets. Cedars (Chamaecyparis nootkatensis, Thuja plicata) appeared on Mitkof Island during the late Holocene.     

Pollen diagram from the Nebraska Sandhills and the age of the dunes

Radiocarbon dates of organic alluvium beneath as much as 40 m of dune sand along the Dismal River have led to the suggestion that the Nebraska Sandhills date from the Holocene rather than the last glacial period. On the other hand, the basal layers of lake and marsh deposits in interdune depressions at three localities date in the range of 9000 to 12,000 yr B.P., implying a pre-Holocene age for the sand dunes. A pollen diagram for one of these sites, Swan Lake, indicates prairie vegetation throughout the last 9000 yr, with no suggestion that the landscape was barren enough to permit the shaping of the massive dunes characterizing the area. Sand was not transported across the site during the Holocene, either during the marsh phase, which lasted until 3700 yr B.P., or during the subsequent lake phase. The sand that buries the alluvium along the Dismal River may represent only local eolian activity, or it may indicate that the younger of the two main dune series identified by H. T. U. Smith (1965, Journal of Geology73, 557–578) is Holocene in age, and the older one Late Wisconsin in age.     

A record of late Quaternary vegetation from Davis Lake,southern Puget Lowland,Washington

Pollen and macrofossil analyses of a core spanning 26,000 yr from Davis Lake reveal late Pleistocene and Holocene vegetational patterns in the Puget Lowland. The core ranges lithologically from a basal inorganic clay to a detritus gyttja to an upper fibrous peat and includes eight tephra units. The late Pleistocene pollen sequence records two intervals of tundra-parkland vegetation. The earlier of these has high percentages of Picea, Gramineae, and Artemisia pollen and represents the vegetation during the Evans Creek Stade (Fraser Glaciation) (ca. 25,000–17,000 yr B.P.). The later parkland interval is dominated by Picea, Tsuga mertensiana, and Gramineae. It corresponds to the maximum ice advance in the Puget Lowland during the Vashon Stade (Fraser Glaciation) (ca. 14,000 yr B.P.). An increase in Pinus ontorta pollen between the two tundra-parkland intervals suggests a temporary rise in treeline during an unnamed interstade. After 13,500 yr B.P., a mixed woodland of subalpine and lowland conifers grew at Davis Lake during a period of rapid climatic amelioration. In the early Holocene, the prolonged expansion of Pseudotsuga and Alnus woodland suggests dry, temperate conditions similar to those of present rainshadow sites in the Puget Lowland. More-mesic forests of Tsuga eterophylla, Thuja plicata, and Pseudotsuga, similar to present lowland vegetation, appeared in the late Holocene (ca. 5500 yr B.P.).     

Vegetation history of the English chalklands: a mid‐Holocene pollen sequence from the Caburn,East Sussex

A pollen diagram has been produced from the base of the Caburn (East Sussex) that provides a temporally and spatially precise record of vegetation change on the English chalklands during the mid‐Holocene (ca. 7100 to ca. 3800 cal. yr BP). During this period the slopes above the site appear to have been well‐wooded, with vegetation analogous to modern Fraxinus–Acer–Mercurialis communities in which Tilia was also a prominent constituent. However, scrub and grassland taxa such as Juniperus communis, Cornus sanguinea and Plantago lanceolata are also regularly recorded along with, from ca. 6000 cal. yr BP onwards, species specific to Chalk grassland (e.g. Sanguisorba minor). This supports suggestions that elements of Chalk grassland persisted in lowland England through the Holocene. Such communities are most likely to have occupied the steepest slopes, although the processes that maintained them are unclear. Human interference with vegetation close to the site may have begun as early as ca. 6350 cal. yr BP and initially involved a woodland management practice such as coppicing. From the primary Ulmus decline (ca. 5700 cal. yr BP) onwards, phases of limited clearance accompanied by cereal cultivation occurred. Taxus baccata was an important component of the woodland which regenerated between these phases. Copyright © 2000 John Wiley & Sons, Ltd.     

History of vegetation during the Holocene in the Courel and Queixa Sierras,Galicia, northwest Iberian Peninsula

Palynological studies undertaken in the mountainous regions of the northwest of the Iberian Peninsula are few in number and have been concerned largely with the second half of the Holocene. New pollen data from two Galician sierras, the Courel and Queixa Sierras, provide a 10000‐yr record of vegetation and climate change. In the Courel Sierra before 9750 yr BP, Laguna Lucenza (1420 m a.s.l.) reflects a period of open landscape covered by Poaceae and heliophilous plants, which may be correlated with the Younger Dryas. The onset of the Holocene is characterised by the expansion of oak woodland, prior to 9300 yr BP, following a short phase of birch along with the gradual decline of pine. The oak values reach a peak at 8350 ± 80 yr BP. Towards 8800 yr BP Corylus begins to expand, followed by Alnus (7500 yr BP) and Ulmus. During this period, the Fraga pollen assemblage (Queixa Sierra, 1360 m a.s.l.) indicates Betula woodland surrounding the site, masking the regional predominance of oak. After 5000 yr BP there is a gradual decrease in arboreal pollen values in both Sierras. Castanea appears in Laguna Lucenza (Courel Sierra) at 4075 ± 75 yr BP. There is widespread deforestation during the last 4000 yr. During this period the presence of large quantities of microcharcoal particles points to the occurrence of fire. The reduction in forest is associated with the arrival of cultivation at 4000 yr BP at low altitudes in the Queixa Sierra. At higher altitudes the first agricultural activity is dated at later than 2000 yr BP. This coincides with the first record of cereal cultivation at high altitude in the Courel Sierra. Copyright © 2000 John Wiley & Sons, Ltd.     

Upper Quaternary vegetation dynamics and palaeoclimatology of the La Chonta bog area (Cordillera de Talamanca,Costa Rica)

Two pollen records from La Chonta bog (2310 m altitude) and one pollen record of a soil profile (2430 m altitude) at a short distance from the bog permit the reconstruction of the vegetational history and climatic sequence of probably the last ca. 80 000 yr of the Cordillera de Talamanca, Costa Rica. The three pollen records can be correlated on the basis of three radiocarbon-dated horizons (> 39 900 yr BP, 34 850 yr BP and 1390 yr BP) and palynostratigraphy. Pollen concentration data provide indirect time control between radiocarbon-dated horizons of the La Chonta 2 core. During the Early Glacial successive fluctuations of the upper forest line occurred between ca. 2300 m and 2800 m altitude and the bog is alternately situated close to the subalpine rainforest belt and in the uppermontane rainforest belt. During the Pleniglacial (ca. 50 000-13 000 yr BP) the forest line fell to ca. 2000 m altitude and paramo vegetation surrounded the La Chonta site. During the Late Glacial the forest line shifted rapidly in ca. 270 yr to 2700–2800 m altitude and the lake became surrounded by Quercus-dominated upper montane rainforest during a period of ca. 1655 yr. A distinct climate cooling (by 2–2.5°C) followed during ca. 580 yr and the upper forest line dropped to ca. 2400 m altitude. This temperature rebound is probably equivalent to the Younger Dryas event. In the depression extensive Alnus carr and swamp vegetation developed. Finally, during the Holocene, the forest line shifted to 3300–3500 m elevation and the lower montane rainforest reached close to ca. 2300 m altitude. In the later part of the Holocene subparamo scrub with Hypericum, Puya, Compositae, Escallonia and Ericaceae largely replaced Alnus carr and the regional and local vegetation closely resembles the presentday composition. During the Last Glacial to Holocene transition a shift of the upper forest line of ca. 1400 m is inferred, corresponding to a glacial temperature depression at 2300 m altitude of ca. 8°C.     

Pollen analysis of hyena coprolites and sediments from Equus Cave,Taung, southern Kalahari (South Africa)

Equus Cave, in Quaternary tufa near Taung in the semiarid woodland of the southern Kalahari, yielded 2.5 m of sediment in which a rich assemblage of bones and coprolites was preserved. The fossils were accumulated mainly by hyenas during the late Pleistocene and Holocene. Pollen from coprolites reflects diet as well as vegetation over relatively large areas visited by hyenas, while pollen from sediments represents more local sources. The pollen sequence derived from coprolites and sediments demonstrates how the vegetation evolved from open grassland with small shrubs and occasional trees during the late Pleistocene, to open savanna with more small shurbs, then, during the last 7500 yr, to modern savanna. Temperatures were not more than 4°C cooler and it was slightly moister than today during the late Pleistocene phase; it became gradually warmer but relatively dry before optimal temperature and moisture conditions developed around 7500 yr B.P. Climatic conditions slightly less favorable for woodland occurred during part of the late Holocene.     

Holocene environmental and climatic changes inferred from Wulungu Lake in northern Xinjiang, China

Sedimentological, geochemical and palynological data from Wulungu Lake in northern Xinjiang, China, are used to reconstruct environmental and climate changes since 9550 cal yr BP. High abundance of Sparganium and Poaceae, low Md (median diameter) and δ¹³C_organic values indicate aridity between 9550 and 6730 cal yr BP. High Md and δ¹³C_organic values, and the prevalence of desert-steppe and steppe vegetation between 4200 and 560 cal yr BP, indicate that effective moisture increased after 6730 cal yr BP, peaking at 4200 and 560 cal yr BP. Low Md values, a negative excursion of δ¹³C_org, and the transition from steppe to desert vegetation since 560 cal yr BP reflect a decrease in effective moisture during the latest Holocene. Late Holocene human activities were indicated by sharp increase in the abundance of Pediastrum then. Variations in carbonate contents indicate that temperature was generally high between 9550 and 7740 cal yr BP, low between 7740 and 6730 cal yr BP, intermediate between 6730 and 560 cal yr BP, and low during the last 560 yr. Regional comparison indicates that the Asian monsoon did not extend to Wulungu Lake and westerlies were the main factor in determining the moisture availability during the Holocene.     

Late Quaternary vegetational and climatic change in the Popayán region,southern Colombian Andes

Late Pleistocene and Holocene vegetational and climatic change have been studied palynologically at a site at 1750 m elevation in the subandean vegetation belt near Popayán, in the southern Colombian Andes. Time control on the pollen record is based on six AMS ¹⁴C ages, ranging from possibly Middle Pleniglacial time (around 50000 yr BP) to 1092 ± 44 yr BP. Because of the presence of two hiatuses only the Middle Pleniglacial and Late Holocene periods (the last 2300 yr BP) are represented. Pollen data indicate the presence of closed subandean forest during glacial time. Changes in the contribution of pollen originating from the uppermost and lowermost subandean forest belts, changes in the contribution of a number of other subandean forest taxa, and changes in species composition between the three pollen zones, suggest that the climate during the Middle Pleniglacial was markedly colder, and perhaps also wetter, than during the Late Holocene. Pollen assemblages from the Late Holocene indicate that the landscape has been affected by deforestation and agriculture since at least 2300 yr BP, but that human impact decreased in the last 780 yr BP. © 1998 John Wiley & Sons, Ltd.     

Late Quaternary vegetation and climate history of the central Bering land bridge from St. Michael Island, western Alaska

Pollen analysis of a sediment core from Zagoskin Lake on St. Michael Island, northeast Bering Sea, provides a history of vegetation and climate for the central Bering land bridge and adjacent western Alaska for the past ≥30,000 ¹⁴C yr B.P. During the late middle Wisconsin interstadial (≥30,000-26,000 ¹⁴C yr B.P.) vegetation was dominated by graminoid-herb tundra with willows (Salix) and minor dwarf birch (Betula nana) and Ericales. During the late Wisconsin glacial interval (26,000-15,000 ¹⁴C yr B.P.) vegetation was graminoid-herb tundra with willows, but with fewer dwarf birch and Ericales, and more herb types associated with dry habitats and disturbed soils. Grasses (Poaceae) dominated during the peak of this glacial interval. Graminoid-herb tundra suggests that central Beringia had a cold, arid climate from ≥30,000 to 15,000 ¹⁴C yr B.P. Between 15,000 and 13,000 ¹⁴C yr B.P., birch shrub-Ericales-sedge-moss tundra began to spread rapidly across the land bridge and Alaska. This major vegetation change suggests moister, warmer summer climates and deeper winter snows. A brief invasion of Populus (poplar, aspen) occurred ca.11,000-9500 ¹⁴C yr B.P., overlapping with the Younger Dryas interval of dry, cooler(?) climate. During the latest Wisconsin to middle Holocene the Bering land bridge was flooded by rising seas. Alder shrubs (Alnus crispa) colonized the St. Michael Island area ca. 8000 ¹⁴C yr B.P. Boreal forests dominated by spruce (Picea) spread from interior Alaska into the eastern Norton Sound area in middle Holocene time, but have not spread as far west as St. Michael Island.     

Multiproxy Devensian Late‐glacial and Holocene environmental records at an Atlantic coastal site in Shetland

Marine erosion at Clettnadal, West Burra island off the west coast of Shetland, caused the drainage of a small water body at Clettnadal, exposing deposits of Late Devensian and Holocene age. Pollen, diatom and invertebrate analyses have provided variable records of environmental change during stratigraphical event GI‐1. Event GS‐1 is revealed by the non‐pollen evidence, especially by Coleoptera, by sediment stratigraphy, and by radiocarbon dating. In contrast, the pollen evidence indicates that an arctic tundra flora, in which dwarf shrubs were prominent, persisted throughout the Late‐glacial. The Holocene brought colonisation by tree birch, but by ca. 9000 ¹⁴C yr BP the taxon had almost disappeared. This contrasts strongly with other Holocene pollen records for Shetland where both Betula and Corylus avellana‐type survived longer—at some sites, for example, until ca. 2900 yr BP. The extreme westerly and exposed coastal situation of Clettnadal appears to be responsible both for a muted Late‐glacial response in the pollen record of terrestrial vegetation and for the early replacement of woodland by a maritime grassland. The results provoke questions concerning biological stability at times of marked climatic change. Copyright © 2003 John Wiley & Sons, Ltd.     

Paleoecological and archaeological implications of a late Pleistocene/Early holocene record of vegetation and climate from the pacific coastal plain of panama

A phytolith record from Monte Oscuro, a crater lake located 10 m above sea level on the Pacific coastal plain of Panama, shows that during the Late Pleistocene the lake bed was dry and savanna-like vegetation expanded at the expense of tropical deciduous forest, the modern potential vegetation. A significant reduction of precipitation below current levels was almost certainly required to effect the changes observed. Core sediment characteristics indicate that permanent inundation of the Monte Oscuro basin with water occurred at about 10,500 ¹⁴C yr B.P. Pollen and phytolith records show that deciduous tropical forest expanded into the lake’s watershed during the early Holocene. Significant burning of the vegetation and increases of weedy plants at ca. 7500 to 7000 ¹⁴C yr B.P. indicate disturbance, which most likely resulted from early human occupation of the seasonal tropical forest near Monte Oscuro and the development of slash-and-burn methods of cultivation.     

Evidence for prolonged El Nino-like conditions in the Pacific during the Late Pleistocene: a 43 ka noble gas record from California groundwaters

Information on the ocean/atmosphere state over the period spanning the Last Glacial Maximum – from the Late Pleistocene to the Holocene – provides crucial constraints on the relationship between orbital forcing and global climate change. The Pacific Ocean is particularly important in this respect because of its dominant role in exporting heat and moisture from the tropics to higher latitudes. Through targeting groundwaters in the Mojave Desert, California, we show that noble gas derived temperatures in California averaged 4.2 ± 1.1 °C cooler in the Late Pleistocene (from ～43 to ～12 ka) compared to the Holocene (from ～10 to ～5 ka). Furthermore, the older groundwaters contain higher concentrations of excess air (entrained air bubbles) and have elevated oxygen-18/oxygen-16 ratios (δ¹⁸O) – indicators of vigorous aquifer recharge, and greater rainfall amounts and/or more intense precipitation events, respectively. Together, these paleoclimate indicators reveal that cooler and wetter conditions prevailed in the Mojave Desert from ～43 to ～12 ka. We suggest that during the Late Pleistocene, the Pacific ocean/atmosphere state was similar to present-day El Nino-like patterns, and was characterized by prolonged periods of weak trade winds, weak upwelling along the eastern Pacific margin, and increased precipitation in the southwestern U.S.     

An equable glaciopluvial in the West: Pleniglacial evidence of increased precipitation on a gradient from the Great Basin to the Sonoran and Chihuahuan Deserts

Dated macrofossil evidence documents the widespread occurrence of woodland in what are now desert lowlands of southwestern North America from the last pleniglacial (ca. 20,000 yr B.P.) to late glacial/Holocene transition (12,000–8000 yr B.P.). The composition of the Pleistocene woodlands indicates that they had already differentiated geographically in modern form, though immensely more extensive than today. The pinyon-juniper woodland (Pinus monophylla, Juniperus osteosperma) of the Mohave Desert province had not yet penetrated the central Great Basin, but extended from southern Nevada south through the vast lowlands of the Mohave and westernmost Sonoran Deserts to southeastern California and Baja California. The strongly xerophytic Mohavean woodland was characterized by a very well-marked altitudinal and latitudinal zonation with juniper-Joshua tree (Yucca brevifolia) sorting out below pinyon-juniper woodland, and with live oaks restricted to the upper level along the lower Colorado River drainage. Southeastward, the Sonoran Desert province was similarly zoned, but with the more slender-leaved Pinus edulis var. fallax as pinyon and with more live oaks in the upper zone. However, the pleniglacial woodland of the Chihuahuan Desert province was almost unzoned, inasmuch as the less xerophytic species of pinyon and live oaks prevailed over the entire span of available elevation; the pinyon was the very slender-leaved P. cembroides var. remota.The overall paleozonation indicates a strong northwest-to-southeast gradient of increasing summer rain with decreasing distance from the monsoonal source area over the Gulf of Mexico, as at present, but augmented pluvially along the same gradient. A key piece of evidence is the counterintuitive latitudinal-zonational anomaly between about 30 and 40° N in southwestern North America; the lower limits of modern vegetational zones are depressed with decreasing latitude (e.g., ca. 500 m lower at 34° than at 36° N). The axis of the gradient actually extends from northwest to southeast, paralleling the monsoonal gradient of increasing summer rain, which no doubt causes the apparent anomaly. During the Wisconsinan glacial, the latitudinal anomaly was greatly steepened, a fact requiring a pluvial increase in precipitation over the Southwest. The monsoonalpluvial pattern is supported by the Neotoma record of a northwest-to-southeast gradient of increasing diversity of evergreen oaks requiring summer rain, and by a parallel segregation of pinyon species. Equability of seasons during the last glacial is also suggested by the Neotoma macrofossil data.