Neotoma Paleoecology Database

Acknowledgements

This documentation would not be possible without the extrordinary work of Dr. Eric C. Grimm who has spent countless hours developing this manual. In addition, Neotoma rests on the work of a number of researchers who contributed to the original North American Pollen Database, and subsequent data contributors, including FAUNMAP contributors and the data contributions of Allan Ashworth. The Neotoma Database would not exist were it not for the ongoing contributions of authors, data analysts and funding agencies, in particular the National Sciences Foundation. This manual draws heavily from Eric Grimm’s original Neotoma manual (v2), published as:

Grimm, E.C., 2008. Neotoma: an ecosystem database for the Pliocene, Pleistocene, and Holocene. Illinois State Museum Scientific Papers E Series, 1.

The SQL Server snapshot is accessible from here: http://www.neotomadb.org/snapshots

Introduction

Neotoma is a public database containing fossil data from the Holocene, Pleistocene, and Pliocene, or approximately the last 5.3 million years. The database stores associated physical data from fossil bearing deposits, for example sediment loss-on-ignition and geochemical data. The database also stores data from modern samples that are used to interpret fossil data.

The initial development of Neotoma was funded by a grant from the U.S. National Science Foundation Geoinformatics program. This grant is collaborative between Penn State University [1] and the Illinois State Museum [2]. It has five Principle Investigators, Russell W. Graham, Eric C. Grimm, Stephen T. Jackson, Allan C. Ashworth, and John W. (Jack) Williams. The database is served from the Center for Environmental Informatics at Penn State University.

Initially, data are being merged from four existing databases: the Global Pollen Database, FAUNMAP (a database of mammalian fauna), the North American Plant Macrofossil Database, and a fossil beetle database assembled by Allan Ashworth. The design of this database is such that many other kinds of fossil data can easily be incorporated in the future, for example, the addition of ostracode, diatom, chironmid, and freshwater mussel datasets.

The existing databases were developed in the 1990’s and have not been updated structurally since. New data have been added, but the structures of these databases have not changed, despite significant advances in database and internet technology. Although structurally different, these databases contain similar kinds of data, and merging them was quite practical. The rationale for this merging was twofold: (1) to facilitate analyses of past biotic communities at the ecosystem level and (2) to reduce the overhead in maintaining and distributing several independent databases..

The new Neotoma database was initially designed by E. C. Grimm and implemented in Microsoft® Access®. This database will be ported to a higher end RDBMS for Internet distribution, but it will continue to be distributed as a standalone Access database for researchers who need access to the entire database.

Whence Neotoma

In the original NSF proposal, this database was called a “Late Neogene Terrestrial Ecosystem Database.” At the time this proposal was written, the Neogene Period included the Miocene, Pliocene, Pleistocene, and Holocene epochs. However, a proposal before the International Commission on Stratigraphy would elevate the Quaternary to a System or Period following the Neogene and terminate the Neogene at the end of Pliocene.

Because this proposal renders the Neogene description of this database obsolete, a new name was sought. Numerous names and companion acronyms were considered, but none engendered enthusiastic support. B. Brandon Curry proposed the name Neotoma, and this name struck a fancy. Neotoma is the genus for the packrat. Packrats are prodigious collectors of anything in their territory, and moreover they are collectors of fossil data. They collect plant macrofossils and bones, and pollen is preserved in their amberat—hardened, dried urine, which impregnates their middens and preserves them for millennia.

Rationale

Paleobiological data from the recent geological past have been invaluable for understanding ecological dynamics at timescales inaccessible to direct observation, including ecosystem evolution, contemporary patterns of biodiversity, principles of ecosystem organization, particularly the individualistic response of species to environmental gradients, and the biotic response to climatic change, both gradual and abrupt. Understanding the dynamics of ecological systems requires ecological time series, but many ecological processes operate too slowly to be amenable to experimentation or direct observation. In addition to having ecological significance, fossil data have tremendous importance for climatology and global change research. Fossil floral and faunal data are crucial for climate-model verification and are essential for elucidating climate-vegetation interactions that may partly control climate.

Basic paleobiological research is site based, and paleobiologists have devoted innumerable hours to identifying, counting, and cataloging fossils from cores, sections, and excavations. These data are typically published in papers describing single sites or small numbers of sites. Often, the data are published graphically, as in a pollen diagram, and the actual data reside on the investigator’s computer or in a file cabinet. These basic data are similar to museum collections, costly to replace, sometimes irreplaceable, and their value does not diminish with time. Also similar to museum collections, the data require cataloging and curation. Whereas physical specimens of large fossils, such as animal bones, are typically accessioned into museums, microfossils, such as pollen, are not accessioned, and the digital data are the primary objects, and their loss is equivalent to losing valuable museum specimens. The integrated database that we propose ensures safe, long-term archiving of these data.

Large independent databases exist for fossil pollen, plant macrofossils, and mammals: the Global Pollen Database (GPD), the North American Plant Macrofossil Database (NAPMD), and FAUNMAP. In addition, a database of fossil beetles (BEETLE) has been assembled, but it is not yet publicly available. These databases have become essential cyberinfrastructure. Nevertheless, they were developed as standalone databases in the early 1990’s with PC database software. GPD and NAPMD are in Paradox®; FAUNMAP is in Access. Since initial database development, emphasis has been placed on ingest of new and legacy data. However, database and Internet technology have advanced greatly in the past 15 years, and the current relational database software, ingest programs, data retrieval algorithms, output formats, and analysis tools are outdated and minimal. Moreover, the databases are not linked, so that integrated analyses are difficult.

Although GPD, NAPMD, and FAUNMAP were developed independently, they have much in common. The basic data of all three databases as well as BEETLE are essentially lists of taxa from cores, excavations, or sections, often with quantitative measures of abundance. The three databases include similar metadata. The objective of Neotoma is to build a unified data structure that will incorporate all of these databases. The database will initially incorporate pollen, plant macrofossil, mammal, and beetle data. However, the database designed facilitates the incorporation of all kinds of fossil data.

Various teams of investigators have developed databases for paleobiological data that have been project or discipline based, including the four databases to be integrated in this project. However, long-term maintenance and sustainability have been problematic because of the need to secure continuous funding. Nevertheless, these databases have become the established archives for their disciplines and, new data are continuously contributed. However, because of funding hiatuses, long spells may intervene between times of data contribution and their public availability. For example, the plant macrofossil database has not incorporated any new data since 1999. The number of different databases and disciplines exacerbates the problem, because each database requires a database manager. Consolidation of informatics technology helps address this overhead issue. However, specialists are still essential for management and supervision of data collection and quality control for their disciplines or organismal groups.

The purposes of Neotoma are (1) to facilitate studies of ecosystem development and response to climate change, (2) to provide the historical context for understanding biodiversity dynamics, including genetic diversity, (3) to provide the data for climate-model validation, (4) to provide a safe, long-term, low-cost archive for a wide variety of paleobiological data. Site-based studies are invaluable in their own right, and they are the generators of new data. However, much is gained by marshalling data from geographic arrays of sites for synoptic, broad-scale ecosystem studies. In order to carry out such studies efficiently, a queryable database is required. Thus, it is much more than an archive; it is essential cyberinfrastructure for paleoenvironmental research. The database facilitates integration, synthesis, and understanding, and it promotes information sharing and collaboration. The individual databases have been extensively used for scientific research, with several hundred scientific publications directly based upon data drawn from these databases. This project will enhance those databases and will continue their public access. By integrating these databases and by simplifying the contributor interface, we can reduce the number of people necessary for community-wide database maintenance, and thereby help ensure their long-term sustainability and existence.

History of the Constituent Databases
Global Pollen Database

In an early effort, the Cooperative Holocene Mapping Project (COHMAP Members 1988, Wright et al. 1993) assembled pollen data in the 1970s and 1980s to test climate models. Although data-model comparison was the principal objective of the COHMAP project, the synoptic analyses of the pollen data, particularly maps showing the constantly shifting ranges of species in response to climate change, were revelatory and led to much ecological insight (e.g. Webb 1981, 1987, 1988).

The COHMAP pollen “database” consisted of a multiplicity of flat files with prescribed formats for data and chronologies. FORTRAN programs were written to read these files and to assemble data for particular analyses. Thompson Webb III managed the COHMAP pollen database at , but as the quantity of data increased, data management became increasingly cumbersome. Clearly, the data needed to be migrated to a relational database management system. Discussions with E. C. Grimm led to the initiation of the North American Pollen Database (NAPD) at the in 1990.

At the same time in , the International Geological Correlation Project IGCP 158 was conducting a major collaborative synthesis of paleoecological data, primarily of pollen, and the need for a pollen database became painfully obvious. In the forward to the book resulting from this project (Berglund et al. 1996), J.L. de Beaulieu describes the role that this project had in launching the European Pollen Database. A workshop to develop a European Pollen Database (EPD) was held in in 1989. North American representatives also attended, and the organizers of NAPD and EPD commenced a long-standing collaboration to develop compatible databases. NAPD and EPD held several joint workshops and developed the same data structure. Nevertheless, the two databases were independently established, partly because Internet capabilities were not yet sufficient to easily manage a merged database. The pollen databases were developed in Paradox, which at the time was the most powerful RDBMS readily available for the PC platform. NAPD and EPD established two important protocols:

  1. the databases were relational and queryable
  2. they were publicly available.

As the success the NAPD-EPD partnership escalated, working groups initiated pollen databases for other regions, including the Latin American Pollen Database (LAPD) in 1994, the Pollen Database for and the Russian Far East (PDSRFE) in 1995, and the African Pollen Database (APD) in 1996. At its initial organizational workshop, LAPD opted to merge with NAPD, rather than develop a standalone database, and the Global Pollen Database was born. PDSRFE also followed this model. APD developed independently, but uses the exact table structure of GPD and EPD. Pollen database projects have also been initiated in other regions, and the GPD contains some of these data, including the Indo-Pacific Pollen Database and the Japanese Pollen Database.

The pollen databases contain data from the Holocene, Pleistocene, and Pliocene, although most data are from the last 20,000 years. Included are fossil data, mainly from cores and sections, and modern surface samples, which are essential for calibrating fossil data. NAPD data are not separate from the GPD, but rather NAPD is the North American subset of GPD. EPD has both public and restricted data—a concession that had to be made early on to assuage some contributors.

North American Plant Macrofossil Database

Plant macrofossils include plant organs generally visible to the naked eye, including seeds, fruits, leaves, needles, wood, bud scales, and megaspores. Synoptic-scale mapping of plant macrofossils from modern assemblages (Jackson et al. 1997) and fossil assemblages (Jackson et al. 1997, Jackson et al. 2000, Jackson and Booth 2002) have shown the utility of plant macrofossils in providing spatially and taxonomically precise reconstructions of past species ranges. Although plant macrofossil records are spatially precise, synoptic networks of high-quality sites can scale up to yield aggregate views of past distributions (Jackson et al. 1997). In addition, macrofossils, with their greater taxonomic resolution, augment the pollen data by providing information on which species might have been present, and can resolve issues of long-distance transport (Birks 2003).

The North American Plant Macrofossil Database (NAPMD) has been directed by S.T. Jackson at the . Highest priority has been placed on data from the last 30,000 years, although some earlier Pleistocene and late Pliocene data are included. The database originated as a research database for selected taxa from Late Quaternary sediments of eastern North America (Jackson et al. 1997). In 1994, an effort was initiated with NOAA funding to build on this foundation to develop a cooperative, relational database comprising all of , a longer time span, and all plant taxa.

The structure of NAPMD was adapted from the pollen database and is also in Paradox. The principal modifications made to the pollen database structure to accommodate plant macrofossils were those to cope with different organs from the same species and to deal with the various quantitative measures of abundance. The database also includes surface samples, which are useful for interpretation of fossil data.

FAUNMAP

R.W. Graham, E.L. Lundelius, Jr., and a group of Regional Collaborators organized a project to develop a database for late Quaternary faunal data from the , which the U.S. NSF funded in 1990. This project had a research agenda, and its seminal paper focused on the individualistic behavior displayed by animal species (FAUNMAP Working Group 1996).

Two FAUNMAP databases exist, FAUNMAP I and FAUNMAP II. Both databases were coordinated by R. W. Graham and E. L. Lundelius, Jr. and funded by NSF. Both are relational databases for fossil mammal sites. The data were extracted from peer-reviewed literature, selected theses and dissertations, and selected contract reports for both paleontology and archaeology. Unpublished collections were not included. Data were originally captured in Paradox but were later migrated to Access.

FAUNMAP I contains data from sites in the lower 48 states that date between 500 BP and ~40,000 BP. Funding for this project ended in 1994, with the production of two major publications by the FAUNMAP Working Group (1994, 1996), as well as numerous other publications by individual members and by many others who accessed the database on-line. Graham and Lundelius continued the FAUNMAP project, developing FAUNMAP II with funding from NSF beginning in 1998. FAUNMAP II shares the same structure as FAUNMAP I but expands the spatial coverage to include and and extends the temporal coverage to the Pliocene (5 Ma). In addition, sites published since 1994, when FAUNMAP I was completed, have been added for the contiguous 48 states. In all, FAUNMAP I and II contain more than 5000 fossil-mammal sites with more than 600 mammal species for all of North America north of Mexico that range in age from 0.5 ka to 5 Ma­.

The detailed structure of the FAUNMAP database is described in FAUNMAP Working Group (1994). Sites identified by name and location were subdivided into Analysis Units (AU’s), which varied from site to site depending upon the definitions used in the original publications (e.g., stratigraphic horizons, cultural horizons, excavation levels, biostratigraphic zones). All data (i.e. taxa identified and counts of individual specimens) and metadata (sediment types, depositional environments, facies, radiometric and other geochronological dates, modifications of bone) were captured by AU. This structure allows for the extraction of information at either the level of the site or the smallest subdivision (AU). The AU permits fine-scale temporal resolution and analysis. Similar to GPD and NAPMD, FAUNMAP contains archival and research tables. Similar to the plant macrofossil database, FAUNMAP contains a variety of quantitative measures of abundance, and presence data are more commonly used for analysis.

BEETLE

Many beetles have highly specific ecological and climatic requirements and are valuable indicators of past environments (Morgan et al. 1983, Ashworth 2001, 2004). They are one of the most diverse groups of organisms on earth, and of the insects, perhaps the most commonly preserved as fossils. Allan Ashworth has assembled a database of fossil beetles from . The data, which were recorded in Excel, contain 5523 individual records of 2567 taxa from 199 sites and 165 publications. Metadata include site name, latitude and longitude, lithology of sediment, absolute age, and geological age. The basic data are similar to plant and mammal databases—lists of taxa from sites. The metadata have not been recorded to the extent of the other databases, especially chronological data, but Ashworth has resolved the taxonomic issues and has assembled the publications, so that the additional metadata can be easily pulled together.

Who Will Use Neotoma?

The existing databases have been used widely for a variety of studies. Because the databases have been available on-line, precise determination of how many publications have made use of them is difficult. In addition, the databases are widely used for instructional purposes. Below are examples of the kinds of people who have used these databases and who we expect will find the new, integrated database even more useful.

  • Paleoecologists seeking to place a new record into a regional/continental/global context (e.g., Bell and Mead 1998, Czaplewski et al. 1999, Bell and Barnosky 2000, Newby et al. 2000, Futyma and Miller 2001, Gavin et al. 2001, Czaplewski et al. 2002, Schauffler and Jacobson 2002, Camill et al. 2003, Rosenberg et al. 2003, Willard et al. 2003, Pasenko and Schubert 2004, and many others).
  • Synoptic paleoecologists interested in mapping regional to sub-continental to global patterns of vegetation change (e.g., Jackson et al. 1997, Williams et al. 1998, Jackson et al. 2000, Prentice et al. 2000, Thompson and Anderson 2000, Williams et al. 2000, Williams et al. 2001, Williams 2003, Webb et al. 2004, Williams et al. 2004, Asselin and Payette 2005).
  • Synoptic paleoclimatologists building benchmark paleoclimatic reconstructions for GCM evaluation (e.g., Bartlein et al. 1998, Farrera et al. 1999, Guiot et al. 1999, Kohfeld and Harrison 2000, CAPE Project Members 2001, Kageyama et al. 2001, Kaplan et al. 2003).
  • Paleontologists trying to understand the timing, patterns, and causes of extinction events (e.g., Jackson and Weng 1999, Graham 2001, Barnosky et al. 2004, Martínez-Meyer et al. 2004, Wroe et al. 2004).
  • Evolutionary biologists mapping the genetic legacies of Quaternary climatic variations (e.g., Petit et al. 1997, Fedorov 1999, Tremblay and Schoen 1999, Hewitt 2000, Comps et al. 2001, Good and Sullivan 2001, Petit et al. 2002, Kropf et al. 2003, Lessa et al. 2003, Petit et al. 2003, Hewitt 2004, Lascoux et al. 2004, Petit et al. 2004, Whorley et al. 2004, Runck and Cook 2005).
  • Macroecologists interested in temporal records of species turnover and biodiversity and historical controls on modern patterns of floristic diversity (e.g., Silvertown 1985, Qian and Ricklefs 2000, Brown et al. 2001, Haskell 2001).
  • Archeologists who are studying human subsistence patterns and interactions with their environment (e.g., Grayson 2001, Grayson and Meltzer 2002, Cannon and Meltzer 2004, Grayson in press).
  • Natural resource managers who need to know historical ranges and abundances of plants and animals for designing conservation and management plans (e.g., Graham and Graham 1994, Cole et al. 1998, Noss et al. 2000, Owen et al. 2000, Committee on Ungulate Management in Yellowstone National Park 2002, Burns et al. 2003)
  • Scientists trying to understand the potential response of plants, animals, biomes, ecosystems, and biodiversity to global warming (e.g., Bartlein et al. 1997, Davis et al. 2000, Barnosky et al. 2003, Burns et al. 2003, Kaplan et al. 2003, Schmitz et al. 2003, Jackson and Williams 2004, Martínez-Meyer et al. 2004)
  • Teachers who use the databases for teaching purposes and class exercises.
[1]Grant number 0622349
[2]Grant number 0622289

Database Design Concepts

Sites, Collection Units, Analysis Units, Samples, and Datasets

Fossil data are site based. A Site has a name, latitude-longitude coordinates, altitude, and areal extent. In Neotoma, Sites are designated geographically as boxes with north and south latitude coordinates and east and west longitude coordinates. If the areal extent is not known, the box collapses to a point, with the north and south latitudes equal and the east and west longitudes equal. Most of the legacy sites in Neotoma currently have point coordinates. The lat-long box can circumscribe the site, for example a lake, or it may circumscribe a larger area in which the site lies either because the exact location of the site is not known or because the exact location is purposely kept vague. In the case of many legacy sites, the exact location is not know precisely; for example, it may have been described as «on a gravel bar 5 miles east of town». The exact locations of some sites have purposely been kept vague to prevent looting and vandalism.

A Collection Unit is a unit from a site from which a collection of fossils or other data have been made. Typical Collection Units are cores, sections, and excavation units. A site may have several Collection Units. A Collection Unit is located spatially within a site and may have precise GPS latitude-longitude coordinates. Its definition is quite flexible. For pollen data, a Collection Unit is typically a core, a section, or surface sample. A Collection Unit can also be a composite core comprised of two or more adjacent cores pieced together to form a continuous stratigraphic sequence. A Collection Unit can also be an excavation unit. For faunal data, a Collection Unit could be as precise as an excavation square, or it could be a group of squares from a particular feature within a site. For example, consider a pit cave with three sediment cones, each with several excavation squares. Collection Units could be defined as the individual squares, or as three composite Collection Units, one from each sediment cone. Another example is an archaeological site, from which the reported Collection Units are different structures, although each structure may have had several excavation squares. The precision in the database depends on how data were entered or reported.

For many published sites, the data are reported from composite Collection Units. If faunal data are reported from a site or locality without explicit Collection Units, then data are assigned to a single Collection Unit with the name «Locality». This is a «quote».

Different kinds of data may have been collected from a single Collection Unit, for example fauna and macrobotanicals from an excavation, or pollen and plant macrofossils from a lake-sediment core. A composite Collection Unit may include data from different milieus, which, nevertheless, are associated with each other, for example a diatom sample from surficial lake sediments and an associated lake-water sample for water-chemistry measurements.

The Collection Unit is equivalent to the Entity in the Global Pollen Database but was not defined in FAUNMAP. When the FAUNMAP data were imported into Neotoma, most localities were assigned a single «Locality» Collection Unit. However, for some localities, the data were assigned to different Collection Units that were clearly identifiable in FAUNMAP (see Figure 1).

An Analysis Unit is a stratigraphic unit within a Collection Unit and is typically defined in the vertical dimension. An Analysis Unit may be a natural stratigraphic unit with perhaps irregular depth and thickness or it may be an arbitrary unit defined by absolute depth and thickness. An excavation may have been dug in arbitrary units, for example 10 cm levels, or it may have followed natural stratagraphic boundaries, for example the «red zone» or a feature in an archaeological site. Although Analysis Units could be designated by an upper depth and lower depth, in Neotoma they are designated by their midpoint depth and thickness, which is more convenient for developing age models. Pollen and other microfossils are typically sampled at arbitrary depths, and although these samples have thicknesses corresponding to the thickness of the sampling device (usually 1 cm or less), these thicknesses are often not reported, just the depths. Different kinds of samples may have been taken from a single analysis unit, for example pollen, diatoms, and ostracodes. The Analysis Unit links these various samples together.

In larger excavations, natural stratigraphic Analysis Units may cut across excavation squares or Collection Units, and the data are reported by Analysis Unit rather than by Collection Unit. In this case, the fossil data are assigned to a generic composite Collection Unit named «Locality», which has the explicitly defined Analysis Units. If the Analysis Units are not described or reported, then the data are assigned to a single Analysis Unit with the name «Assemblage». Thus, for a locality published with only faunal list, the fauna are assigned to a Collection Unit named «Locality» and to an Analysis Unit named «Assemblage».

In FAUNMAP, Analysis Units are the primary sample units, and fauna are recorded by Analysis Unit. In the GPD, Analysis Units correspond to samples.

Samples are of a single data type from an Analysis Unit. For example, there may be a vertebrate faunal sample and a macrobotanical sample from the same Analysis Unit; or there may be a pollen sample and an ostracode sample from the same Analysis Unit. There can be multiple samples of the same data type from an Analysis Unit, for example two pollen samples counted by different analysts. Normally, vertebrate fossils from an Analysis Unit comprise a single sample; however, if the fossils are of mixed age, individually dated bones may be treated as separate samples, each with a precise age. In addition to fossils, samples may also be used for physical measurements, such as loss-on-ignition. Geochronologic measurements, such as radiocarbon dates, are made on geochronologic samples.

A Dataset is a set of Samples of a single data type from a Collection Unit. For example the pollen data from a core comprise a pollen Dataset. The geochronologic samples from a Collection Unit form a geochronologic Dataset. Every Sample is assigned to a Dataset, and every Dataset is assigned to a Collection Unit. Samples from different Collection Units cannot be assigned to the same Dataset (although they may be assigned to Aggregate Datasets).

image1

Figure 1. Diagram showing the relationships between tables in Neotoma, the Pollen Database, and FAUNMAP. Because the pollen database has only pollen, no need exists for Analysis Units, which may have multiple data types. FAUNMAP does not make a hierarchical distinction between Collection Units and Analysis Units, and the data for both Analysis Units and fauna are contained in the Faunal table, although within the Faunal table, implicit one-to-many relationships exist between Localities and Analysis Units and between Analysis Units and faunal data.

Taxa and Variables

In general, a sample in Neotoma has a list of taxa with some measure of abundances. The Data table in Neotoma has fields for SampleID, VariableID, and Value. Variables, which are listed in the Variables table, consist of a Taxon, referenced in the Taxa table, as well as the identified Element, measurement Units, Context, and Modification. A Taxon is generally a biological Taxon, but a Taxon may also be a physical attribute such as loss-on-ignition.

For biological taxa, the Element is the organ or skeletal element. Typical faunal Elements are bones, teeth, scales, and other hard body parts. Bone and tooth Elements may be specifically identified (e.g. «tibia» or even more precisely «tibia, distal, left», «M2, lower, left»). Some soft Elements also occur in the database (e.g. «hair» and «dung»). For mammals, an unspecified element is «bone/tooth». Elements for plant macrofossils are the organs identified (e.g. «seed», «needle», «cone bract»). Pollen and spores are treated simply as taxon Elements. Thus, Picea seeds, Picea needles, and Picea pollen are three different Variables. All three refer to a single entry in the Taxa table for Picea.

  • Variable Units are the measurement units. For faunal data, the most common are «present/absent», «number of individual specimens» (NISP), and «minimum number of individuals» (MNI). Plant macrofossils have many different quantitative and semi-quantitative measurement Units, including concentrations and relative abundance scales. Measurement Units for pollen are NISP (counts) and «percent». For pollen the preferred measurement Unit is NISP, but for some sites only percentage data are available. Picea pollen NISP and Picea pollen percent are two different Variables.
  • Variable Contexts for fauna include «articulated», «intrusive», and «redeposited». A context for pollen is «anachronic», which refers to a pollen type known to be too old for the contemporary sedimentary deposit. Most Variables do not have a specified context.
  • Variable Modifications include various modifications to fossils or modifiers to Variables, including human modifications to bones (e.g. «bone tool», «human butchering», «burned») and preservational and taphonomic modifications (e.g. «carnivore gnawed», «fragment»). Modifications for pollen include preservational classifications such as «corroded» and «degraded».

Taxonomy and Synonymy

Neotoma does not change or question identifications from original sources, although taxonomic names may be synonymized to currently accepted names. Thus, for example, the old (although still valid) non-standard plant family names such as Gramineae and Compositae are synonimized to their standard family names terminated with «-aceae», viz. Poaceae and Asteraceae. Neotoma has not attempted to establish complete or comprehensive synonymies. However, the Synonyms table lists commonly encountered synonyms. The descriptions of the SynonymTypes and Taxa tables contain fuller discussions of synonymiztions made in Neotoma.

An important feature of Neotoma is that the Taxa table is hierarchical. Each Taxon has a HigherTaxonID, which is the TaxonID of the next higher taxonomic rank. Thus, data are stored at the highest taxonomic resolution reported by the original investigators, but can be extracted at a higher taxonomic level.

Synonymy presents a challenge for any organismal database, particularly for one such as Neotoma, which archives data collected for over a century and which archives extinct taxa, often for which few and fragmentary specimens exist. Many changes are due to increased understanding of the diversity within taxonomic groups and of the phylogenetic relationships within and among groups. Other changes are due purely to taxonomic rules or conventions set by the International Code of Botanical Nomenclature (McNeill et al. 2006) and the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999). Working groups representing the different taxonomic groups included in Neotoma have established appropriate taxonomic authorities:

  • Plants – There is no worldwide authority. The International Plant Names Index [3] lists validly published names, but a listed name is not necessarily the accepted name for a given taxon. For families, Neotoma follows the Angiosperm Phylogeny Group II (2003) and Stevens (2007+), which follows and updates APG II. The APG is an international consortium of plant taxonomists, and the APG classification utilizes the great quantity of phylogenetic data generated in recent years. For lower taxonomic ranks, the various pollen database cooperatives follow appropriate regional floras:
  • North American Pollen Database/North American Plant Macrofossil Database: Insofar as possible, follows the Flora of North America (Flora of North America Editorial Committee 1993+); about half of the planned FNA volumes have been published. Otherwise, appropriate regional floras are followed.
  • European Pollen Database: The EPD has a Taxonomy Support Group. In general, nomenclature follows Flora Europaea (Tutin 1964-1993).
  • African Pollen Database: The APD has a Committee for Nomenclature, which has produced a list of pollen types with misspellings, synonymy, and nomenclature corrected [4]. APD nomenclature follows Enumération des plantes à fleurs d’Afrique Tropicale (Lebrun and Stork 1991-1997).
  • Latin American Pollen Database: has a tremendously rich and diverse flora and no comprensive flora is available. Various regional floras are followed.
  • Indo-Pacific Pollen Database: For Australia and adjacent areas follows the Australian Plant Name Index (Chapman 1991). For other regions, appropriate regional floras are followed.
  • Pollen Database for Siberia and the Russian Far East Follows Vascular Plants of Russia and Adjacent States (Czerepanov 1995).
  • Mammals – For extant taxa, the authority is Wilson and Reeder’s (2005) Mammal Species of the World . Original sources are followed for extinct species, and the database is considered authoritative.
  • Birds – For North America, the authority is the American Ornithologists’ Union Check-list of North American Birds (American Ornithologists’ Union 1983).
  • Fish – Follows the Catalog of Fishes (Eschmeyer 1998).
  • Mollusks – For North America, follows Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks (Turgeon et al. 1998).
  • Beetles – Comprehensive manuals do not exist. Original taxonomic authorities are cited, and the database is considered authoritative.

Taxa and Ecological Groups

In the Taxa table, each taxon is assigned a TaxaGroupID, which refers to the TaxaGroupTypes table. These are major taxonomic groups, such as «Vascular plants», «Diatoms», «Testate amoebae», «Mammals», «Reptiles and amphibians», «Fish», and «Molluscs». Also included are «Charcoal» and «Physical variables». Ecological Groups are groupings of taxa within Taxa Groups, which may be ecological or taxonomic. Ecological Groups are assigned in the EcolGroups table, in which taxa are assigned an EcolGroupID, which links to the EcolGroupTypes table, and an EcolSetID, which links to the EcolSetTypes table. Ecological Groups are commonly used to organize taxa lists and diagrams. For any taxonomic group, more than one Ecological Set may be assigned. For example, beetles may be assigned to a set of ecological groups, such as dung and bark beetles, and to second set based on taxonomy. Vascular plants are assigned to a «Default plant» set comprised of groups such as «Trees and Shrubs», «Upland Herbs», and «Terrestrial Vascular Cryptogams». Default pollen diagrams can then be generated based on a pollen sum of these three groups. Mammals are assigned to a «Vertebrate orders» set.

Chronology

Neotoma stores both the archival data used to reconstruct chronologies as well as interpreted chronologies derived from the archival data. The basic data used to reconstruct chronologies occurs in three tables: Geochronology, Tephrachronology, and RelativeChronology. The Geochronology table includes geophysical measurements such as radiocarbon, thermoluminescence, uranium series, and potassium-argon dates. This table also includes dendrochronological dates derived from tree-ring chronologies, for example logs in archaeological structures. The Tephrachronology table records tephras in Analysis Units. This table refers to the Tephras lookup table, which stores the ages for known tephras. The RelativeChronology table stores relative age information for Analysis Units. Relative age scales include the archaeological time scale, geologic time scale, geomagnetic polarity time scale, marine isotope stages, North American land mammal ages, and Quaternary event classification. For example, diagnostic artifacts from an archaeological site may have cultural associations with a known age ranges, which can be assigned to Analysis Units. The faunal assemblage from an Analysis Unit may be assignable to particular land mammal age, which places it within a broad time range. Sedimentary units may be assigned to particular geomagnetic chrons, marine isotope stages, or Quaternary events, such as a particular interglacial. Many of these relative ages have rather broad time spans, but do provide some chronologic control.

Actual Chronologies are constructed from the basic chronologic data in the Geochronology, Tephrachronology, and RelativeChronology tables. These chronologies are stored in the Chronologies table. A Chronology applies to a Collection Unit and consists of a number of Chron Controls, which are ages assigned to Analysis Units. A Chron Control may be an actual geochronologic measurement, such as a radiocarbon date, or it may be derived from the actual measurement, such as a radiocarbon date adjusted for an old carbon reservoir or calibrated to calendar years. A Chron Control may by an average of several radiocarbon dates from the same Analyis Unit. Different kinds of basic chronologic data may be used to assign an age to an Analysis Unit, for example radiocarbon dates and diagnostic archaeological artifacts. Some relative Chron Controls are not from one of the established relative time scales. Examples of these are local biostratigraphic controls, which may be based on dated horizons from nearby sites. A familiar example in is the Ambrosia-rise, which marks European settlement. The exact date varies regionally, depending on when settlement occurred locally. For a given site, the date assigned to the Ambrosia-rise may be based on historical information about when settlement occurred or possibly on geophysical dating (e.g. 210Pb) of a nearby site.

image2For continuous stratigraphic sequences, such as cores, not every Analysis Unit may have a direct date. Therefore, ages are commonly interpolated between dated Analysis Units. In this case, the ChronControls are the age-depth control points for an age model, which may be linear interpolation between Chron Controls or a fitted curve or spline.

Figure 2. Smoothed quick radiocarbon calibration curve. At the scale of this figure the difference is mostly less than the line thickness.

Age is measured in different time scales, the two most commn being radiocarbon years before present (14C yr BP) or presumed calendar years before present (cal yr BP). For a calibrated radiocarbon date, «cal yr BP» technically stands for «calibrated years before present», i.e. calibrated to calendar years. In Neotoma, «cal yr BP» is used for both calibrated radiocarbon years and for other ages scales presumed to be in calendar years, viz. dendrochronologic years and other geochronlogic ages believed to be in calendar years. The zero datum for any «BP» age is ad 1950, regardless of its derivation. Thus, BP ages younger than ad 1950 are negative—ad 2000 = ‑50 BP.

image3Ages may be reported in ad/bc age units, in which case bc years are stored as negative values. If ages are reported with a datum other than ad 1950 for BP years, the ages must be converted to an ad 1950 datum or to the ad/bc age scale before entry into Neotoma. For example, 210Pb dates are often reported relative to the year of analysis; these must be converted to either ad/bc or «cal yr BP» with an ad 1950 datum.

Figure 3. An enlarged portion of Figure 2 showing the monontonic smoothed curve

Radiocarbon years can be calibrated to calendar years with a calibration curve. The current calibration curve for ≤26,000 cal yr BP (=21,341 14C yr BP) is the INTCAL04 calibration curve (Reimer et al. 2004). Various programs, both online and standalone, are available for calibrating individual radiocarbon dates, two of the more popular are CALIB [5] (Stuiver and Reimer 1993) and OxCal [6] (Bronk Ramsey 1995, 2001), both available online for download. Calibration of radiocarbon years beyond the INTCAL04 curve is more controversial. However, the Fairbanks0107 curve is available for calibration of radiocarbon dates to 50,000 cal yr BP, the practial limit of radiocarabon dating (Fairbanks et al. 2005, Chiu et al. 2007), with an online application [7].

image4Figure 4. Sample ages calculated from the Neotoma quick calibraton curve vs. ages calculated from traditional age models.

Calibrated radiocarbon dates better represent the true time scale and the true errors and probability distributions of the age estimates. In addition, other important paleo records, notably the ice cores and tree-ring records, have calendar-year time-scales. Therefore, for comparison among proxies and records, it is clearly desirable to place all records on the same time-scale, viz. a calendar-year time-scale. Although this goal is laudable, most of the data ingested into Neotoma from other databases is on a radiocarbon time scale. The majority of assigned ages and almost all the ages from the pollen database are interpolated ages derived from age models. The proper method for deriving calibrated ages is to calibrate the radiocarbon dates and then reinterpolate new ages between these calibrated dates.

Virtually all age models are problematic. A key problem is that most age models linearly interpolate between age-depth points or fit functions or splines to points. However, radiocarbon ages are not points, but probability distributions. Moreover, the probability distributions of calibrated ages are non-Gaussian. Each calibrated age has a unique probability distribution, and many are bimodal or multimodal. Various investigators have used different points, including the intercepts of the radiocarbon age with the calibration curve and the midpoint of the 1σ or 2σ probability distributon. The former is particularly inappropriate (Telford et al. 2004b). The 50% median probability is probably the best single point; however, because of multimodality, this particular point may, in fact, be very unlikely. Nevertheless, if it falls between more-or-less equally probable modes, it may still be the best single point. Most age models for cores are based on relataively few radiocarbon dates, and the uncertainties of the interpolated ages are unknown and large (Telford et al. 2004a). Indeed, chronology is perhaps the greatest challenge for future research with this database.

image5

Figure 5. Anomalies (Sample ages from Neotoma default calendar-year age models minus ages calculated with the Neotoma quick calibration curve) vs. time.

Given the need for a common age scale and the enormity of the task to properly develop new age models, a RadiocarbonCalibration conversion table was developed to quickly convert sample ages in radiocarbon years to calendar years. These calibrated ages are for perusal and data exploration; however, the differences between these ages and those calculated with traditional age models are relatively small. The table contains radiocarbon ages from -100 to 45,000 in 1-year increments with corresponding calibrated values. The table was generated by smoothing the INTCAL04 calibration curve with an FFT filter so that the curve is monotonically increasing, i.e. so that there are no age reversals in calibrated age. The INTCAL04 curve is in 5-yr increments from -5 to 12,500 14C yr BP, 10-yr increments from 12,500 to 15,000 14C yr BP, and 20-yr increments from 15,000 to 26,000 14C yr BP. The FFT filter was 50 points (250 yr) for the first interval, 25 points (250 yr) for the second interval, and 10 points (200 yr) for the third interval. For the calibration beyond 26,000 14C yr BP, a calibrated age was determined with the Fairbanks0107 calibration curve every 100 years with a standard deviation of ±100 years from 20,000±100 14C yr BP to 46,700±100 14C yr BP. These were then smoothed with a 5-sample (500-yr) FFT filter. The curve kinks sharply after 45,000 14C yr BP, so the quick calibration curve was terminated at this date. The Fairbanks0107 curve diverges somewhat from the INTCAL04 curve for the portion they overlap in age. From 20,000 to 26,000 14C yr BP, the difference was prorated linearly from zero divergence from the INTCAL04 curve at 20,000 14C yr BP to zero divergence from the Fairbanks0107 curve at 26,000 14C yr BP. Figure 2 shows the smoothed curve, and Error! Reference source not found. shows an enlargement of part of the curve.

An analysis was made to assess the deviation between ages derived from traditionally calibrated age models and ages derived from the quick calibration curve. From the database, 57 default Chronologies in calibrated radiocarbon years were selected. The Chron Controls were all calibrated radiocarbon dates, except for top dates, European settlement dates, and 210Pb dates in the uppermost portions of the cores. A few Chronologies used the Zdanowicz et al. (1999) calendar-year age from the GISP2 ice core. Ages beyond the reliable age limit (Chronologies.AgeBoundOlder) were not used. These 57 Chronologies had a total of 1945 Sample Ages in calibrated radiocarbon years. Figure 4 shows graph of ages from the Neotoma age models vs. the ages calculated with the quick calibration curve. Error! Reference source not found. shows the anomalies vs. time and Figure 6 shows a histogram of the distribution of anomalies. Nearly half (47%) of the anomalies are <25 years, 86% are <100 years, 97% are <200 years, and 99.4% are <300 years. The average absolute anomaly is 49.2 years, and the median is 29 years. Thus, the quick calibration curve provides remarkably good results. The ages have no confidence limits, but neither do the interpolated ages of most age models.

image6

Figure 6. Binned distribution of anomalies between Neotoma default calendar-year age models and ages calculated with the Neotoma quick calibration curve.

Sediment and Depositional Environments

Several tables deal with depositional environments, depositional agents, and sediment descriptions. In Neotoma, the Depositional Environment refers to the Depositional Environment of the site today, for example, «», «Fen», «Cave», «Colluvial Fan». Depositional Environments may vary within a Site. For example, a lake with a marginal fen has lake and fen Depositional Environments. Thus, Depositional Environments are an attribute of Collection Units and are assigned in the CollectionUnits table. Depositional Environments are listed in the in the DepEnvtTypes lookup table, and they are hierarchical, for example:

Glacial Lacustrine

Any of these Depositional Environments may be assigned to a Collection Unit, but because they are hierarchical, Collection Units may be grouped at higher levels, for example, all Collection Units from natural lakes. The top level Depositional Environments, with some examples, are:

  • Archaeological burials, middens, mounds
  • Biological packrat middens, dung, moss polsters
  • Estuarine mangrove swamps, salt marshes
  • Lacustrine lakes and ponds
  • Marine deep sea benthic, coastal bars
  • Palustrine wetlands including fens, bogs, and marshes
  • Riverine river channels, point bars, natural levees
  • Sampler Tauber traps for modern pollen samples
  • Spring tufa deposits, spring conduits
  • Terrestrial caves, rock shelters, colluvium, volcanic deposits, soils

The Depositional Environment may change through time. For example, as a basin fills with sediment, it may convert from a lake to a fen and perhaps later to a bog. A colluvial slope may have alluvial sediments at depth. A modern playa lake may have a buried paleosol. Thus, a sediment section may have units with different facies and depositional agents. The Facies is the sum total of the characteristics that distinguish a sedimentary unit. Facies are listed in the FaciesTypes lookup table and are assigned to Analysis Units in the AnalysisUnits.FaciesID field. A sedimentary unit may have one or more agents of deposition. For example, a cave deposit may be partly owing to human habitation and partly to carnivore activity. Depositional Agents are listed in the DepAgentTypes lookup table and are assigned to Analysis Units in the DepAgents table.

Whereas Facies and Depositional Agents are both keyed to Analysis Units, the Lithology table is keyed to Collection Units. Analysis Units, especially from cores, may not be contiguous but be placed at discrete intervals down section. Lithologic units are defined by depth in the Collection Unit. Whereas Facies have short descriptions and are keyed to the FaciesTypes lookup table, the Lithology.Description field is a memo, and lithologic descriptions much more detailed than Facies descriptions. FAUNMAP, which was built around Analysis Units, stores Facies and Depositional Agent data; whereas the pollen database, which was centered on Collection Units, stores lithologic data.

Date Fields

Neotoma uses date fields in several tables. Dates are stored internally as a double precision floating point number, which facilitates calculations and functions involving dates. The disadvantage is that complete dates must be stored, i.e. year, month, and day; whereas in many cases only the year or month are known, for example the month a core was collected. Neotoma had adapted the convention that if only the month is known, the day is set to the first of the month; if only the year is known, the month and day are set to January 1. Thus, «June 1984» is set to «June 1, 1984»; and «1984» is set to «January 1, 1984». The drawback, of course, is that these imprecise dates cannot be distinguished from precise dates on the first of the month. However, it was determined that the advantages of the date fields outweighed this disadvantage.

[3]http://www.ipni.org
[4]http://medias.obs-mip.fr/apd/
[5]http://calib.qub.ac.uk/calib/
[6]http://c14.arch.ox.ac.uk/embed.php?File=oxcal.html
[7]http://radiocarbon.ldeo.columbia.edu/research/radcarbcal.htm

SQL Quickly

SQL (Sturctured Query Language) is a standard language for querying and modifying relational databases. It is an ANSI and ISO standard, although various vendors have added proprietary extensions. It is beyond the scope of this document to describe SQL or the differences between Microsoft Access SQL and ANSI SQL. However, examples of SQL queries are provided in this document as a tutorial. Most users of Access probably use the graphical design view for queries, but SQL queries are better suited for examples. These queries can by typed or copied and pasted into the Access query SQL view. The query can then be executed or opened in design view to show the graphical representation. One difference between Access SQL and other flavors is the wildcard; Access uses * rather than %.

SQL Example

The following SQL example lists the number of sites by GeoPoliticalID (the name of the country) for and GeoPoliticalID that is defined as a country.

 1
 2
 3
 4
 5
 6
 7
 8
 9
10
11
12
13
14
15
16
17
18
19
20
SELECT
        COUNT(sites.SiteName),
        gpu.GeoPoliticalName,
        gpu.GeoPoliticalUnit
FROM
        (
                SELECT
                        *
                FROM
                        GeoPoliticalUnits
                WHERE
                        geopoliticalunits.GeoPoliticalUnit = "country"
        ) AS gpu
INNER JOIN (
        Sites
        INNER JOIN SiteGeoPolitical ON Sites.SiteID = SiteGeoPolitical.SiteID
) ON gpu.GeoPoliticalID = SiteGeoPolitical.GeoPoliticalID
GROUP BY
        gpu.GeoPoliticalID,
        gpu.GeoPoliticalUnit;

Table Keys

Within tables there are often Keys. A Key may be a Primary Key (PK), which acts as a unique identifier for individual records within a table, or they may be a Foreign Key (FK) which refers to a unique identifier in another table. Primary Keys and Foreign Keys are critical to join tables in a SQL query. In the above example we can see that the

Data Types

In the table descriptions in the following section, the SQL Server data types are given for field descriptions. The equivalent Access data types are given in the following table.

SQL Server data type Access data type
bit Yes/No
datetime Date/Time
float Double
int Long Integer
nvarchar(n), where n = 1 to 4000 Text
nvarchar(MAX) Memo

Neotoma Tables

The Neotoma Database contains more than 100 tables and as new proxy types get added, or new metadata is stored, the number of tables may increase or decrease. As a result, this manual should not be considered the final authority, but it should provide nearly complete coverage of the database and its structure. In particular, do our best to divide tables into logical groupings: Chronology & Age related tables, Dataset related tables, Site related tables, Contact tables, Sample tables and so on.

References Cited

American Ornithologists’ Union. 1983. Check-list of North American birds, seventh edition. American Ornithologists’ Union, Washington, D.C., USA. [http://www.aou.org/checklist/index.php3]

Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141:399-436.

Ashworth, A. C. 2001. Perspectives on Quaternary beetles and climate change. in L. C. Gerhard, W. E. Harrison, and B. M. Hanson, editors. Geological perspectives of global climate change. AAPG Studies in Geology 47. American Association of Petroleum Geologists, Tulsa, Oklahoma, USA.

Ashworth, A. C. 2004. Quaternary Coleoptera of the United States and Canada. Pages 505-517 in A. R. Gillespie, S. C. Porter, and B. F. Atwater, editors. The Quaternary Period in the United States. Developments in Quaternary Science 1. Elsevier, Amsterdam, The Netherlands.

Asselin, H., and S. Payette. 2005. Late Holocene opening of the forest tundra landscape in northern Québec, Canada. Global Ecology and Biogeography 14:307-313.

Barnosky, A. D., E. A. Hadly, and C. J. Bell. 2003. Mammalian response to global warming on varied temporal scales. Journal of Mammalogy 84:354-368.

Barnosky, A. D., P. L. Koch, R. S. Feranec, S. L. Wing, and A. B. Shabel. 2004. Assessing the causes of late Pleistocene extinctions on the continents. Science 306:70-75.

Bartlein, P. J., K. H. Anderson, P. M. Anderson, M. E. Edwards, C. J. Mock, R. S. Thompson, R. S. Webb, T. Webb, III, and C. Whitlock. 1998. Paleoclimate simulations for North America over the past 21,000 years: features of the simulated climate and comparisons with paleoenvironmental data. Quaternary Science Reviews 17:549-585.

Bartlein, P. J., C. Whitlock, and S. L. Shafer. 1997. Future climate in the Yellowstone National Park region and its potential impact on vegetation. Conservation Biology 11:782-792.

Bell, C. J., and A. D. Barnosky. 2000. The microtine rodents from the Pit locality in Porcupine Cave, Park County, Colorado. Annals of Carnegie Museum 69:93-134.

Bell, C. J., and J. I. Mead. 1998. Late Pleistocene microtine rodents from Snake Creek Burial Cave, White Pine County, Nevada. Great Basin Naturalist 58:82-86.

Berglund, B. E., H. J. B. Birks, M. Ralska-Jasiewiczowa, and H. E. Wright, editors. 1996. Palaeoecological events during the last 15 000 years: regional syntheses of palaeoecological studies of lakes and mires in Europe, volumes. John Wiley & Sons, Chichester, England.

Birks, H. H. 2003. The importance of plant macrofossils in the reconstruction of lateglacial vegetation and climate: examples from Scotland, western Norway, and Minnesota, USA. Quaternary Science Reviews 22:453-473.

Bronk Ramsey, C. 1995. Radiocarbon calibration and analysis of stratigraphy: The OxCal program. Radiocarbon 37:425-430.

Bronk Ramsey, C. 2001. Development of the radiocarbon calibration program OxCal. Radiocarbon 43(2A):355-363.

Brown, J. H., S. K. M. Ernest, J. M. Parody, and J. P. Haskell. 2001. Regulation of diversity: maintenance of species richness in changing environments. Oecologia 126:321-332.

Burns, C. E., K. M. Johnson, and O. J. Schmitz. 2003. Global climate change and mammalian species diversity in U.S. national parks. Proceedings of the National Academy of Sciences of the United States of America 100:11474-11477.

Camill, P., C. E. Umbanhowar, Jr., R. Teed, C. E. Geiss, J. Aldinger, L. Dvorak, J. Kenning, J. Limmer, and K. Walkup. 2003. Late-glacial and Holocene climatic effects on fire and vegetation dynamics at the prairie–forest ecotone in south-central Minnesota. Journal of Ecology 91:822-836.

Cannon, M. D., and D. J. Meltzer. 2004. Early Paleoindian foraging: examining the faunal evidence for large mammal specialization and regional variability in prey choice. Quaternary Science Reviews 23:1955-1987.

CAPE Project Members. 2001. Holocene paleoclimate data from the Arctic: testing models of global climate change. Quaternary Science Reviews 20:1275-1287.

Chapman, A. D. 1991. Australian plant name index, 4 volumes. Australian flora and fauna series 12. Australian Government Publishing Service, Canberra.

Chiu, T.-C., R. G. Fairbanks, L. Cao, and R. A. Mortlock. 2007. Analysis of the atmospheric 14C record spanning the past 50,000 years derived from high-precision 230Th/234U/238U, 231Pa/235U and 14C dates on fossil corals. Quaternary Science Reviews 26:18-36.

COHMAP Members. 1988. Climatic changes of the last 18,000 years: observations and model simulations. Science 241:1043-1052.

Cole, K. L., M. B. Davis, F. Stearns, K. Walker, and G. Guntenspergen. 1998. Historical landcover changes in the Great Lakes region. Pages 43-50 in T. D. Sisk, editor. Perspectives on the land use history of North America: a context for understanding our changing environment. Biological Science Report USGS/BRD/BSR 1998-0003. United States Geological Survey, Biological Resources Division.

Committee on Ungulate Management in Yellowstone National Park, N. R. C. 2002. Ecological dynamics on Yellowstone’s northern range. National Academy Press, Washington, D.C., USA.

Comps, B., D. Gömöry, J. Letouzey, B. Thiébaut, and R. J. Petit. 2001. Diverging trends between heterozygosity and allelic richness during postglacial colonization in the European beech. Genetics 157:389-397.

Czaplewski, N. J., J. I. Mead, C. J. Bell, W. D. Peachey, and T.-L. Ku. 1999. Papago Springs Cave revisited, part II: vertebrate paleofauna. Occasional Papers of the Oklahoma Museum of Natural History 5:1-41.

Czaplewski, N. J., W. L. Puckette, and C. Russell. 2002. A Pleistocene tapir and associated mammals from the southwestern Ozark highland. Journal of Cave and Karst Studies 64:97-107.

Czerepanov, S. K. 1995. Vascular plants of Russia and adjacent states (the former USSR). Cambridge University Press, Cambridge, England.

Davis, M. B., C. Douglas, R. Calcote, K. Cole, M. Winkler, and R. Flakne. 2000. Holocene climate in the western Great Lakes National Parks and Lakeshores: implications for future climate change. Conservation Biology 14:968-983.

Eschmeyer, W. N., editor. 1998. Catalog of fishes, 3 volumes. California Academy of Sciences, San Francisco, California, USA. [http://research.calacademy.org/research/ichthyology/catalog/]

Fairbanks, R. G., R. A. Mortlock, T.-C. Chiu, L. Cao, A. Kaplan, T. P. Guilderson, T. W. Fairbanks, A. L. Bloom, P. M. Grootes, and M.-J. Nadeau. 2005. Radiocarbon calibration curve spanning 0 to 50,000 years BP based on paired 230Th/234U/238U and 14C dates on pristine corals. Quaternary Science Reviews 24:1781-1796.

Farrera, I., S. P. Harrison, I. C. Prentice, G. Ramstein, J. Guiot, P. J. Bartlein, R. Bonnefille, M. Bush, W. Cramer, U. von Grafenstein, K. Holmgren, H. Hooghiemstra, G. Hope, D. Jolly, S.-E. Lauritzen, Y. Ono, S. Pinot, M. Stute, and G. Yu. 1999. Tropical climates at the Last Glacial Maximum: a new synthesis of terrestrial palaeoclimate data. I. Vegetation, lake-levels and geochemistry. Climate Dynamics 15:823-856.

FAUNMAP Working Group. 1994. FAUNMAP: a database documenting late Quaternary distributions of mammal species in the United States. Illinois State Museum Scientific Papers 35(1-2):1-690.

FAUNMAP Working Group. 1996. Spatial response of mammals to late Quaternary environmental fluctuations. Science 272:1601-1606.

Fedorov, V. B. 1999. Contrasting mitochondrial DNA diversity estimates in two sympatric genera of Arctic lemmings (Discrostonyx: Lemmus) indicate different responses to Quaternary environmental fluctuations. Proceedings of the Royal Society of London B 266:621-626.

Flora of North America Editorial Committee. 1993+. Flora of North America north of Mexico, 14+ volumes. Oxford University Press, New York, New York, USA. [http://hua.huh.harvard.edu/FNA/]

Futyma, R. P., and N. G. Miller. 2001. Postglacial history of a marl fen: vegetational stability at Byron-Bergen Swamp, New York. Canadian Journal of Botany 79:1425-1438.

Gavin, D. G., J. S. McLachlan, L. B. Brubaker, and K. A. Young. 2001. Postglacial history of subalpine forests, Olympic Peninsula, Washington, USA. The Holocene 11:177-188.

Good, J. M., and J. Sullivan. 2001. Phylogeography of the red-tailed chipmunk (Tamias ruficaudus), a northern Rocky Mountain endemic. Molecular Ecology 2001:2683-2695.

Graham, R. W. 2001. Late Quaternary biogeography and extinction of Proboscideans in North America. Pages 707-709 in G. Cavarretta, P. Giola, M. Mussi, and M. R. Palombo, editors. The World of Elephants: Proceedings of the 1st International Congress, Rome, Italy, 16-20 October 2001. Consiglio Nazionale delle Ricerche, Rome, Italy.

Graham, R. W., and M. A. Graham. 1994. The late Quaternary distribution of Martes in North America. Pages 26-58 in S. W. Buskirk, A. S. Harestoel, M. G. Raphael, and R. A. Powell, editors. The biology and conservation of martens, sables, and fishers. Cornell University Press, Ithaca, New York, USA.

Grayson, D. K. 2001. The archaeological record of human impacts on animal populations. Journal of World Prehistory 15:1-68.

Grayson, D. K. in press. Early Americans and Pleistocene mammals in North America. in Handbook of North American Indians. Volume 3. Environment, origins, and population. Smithsonian Institution Press, Washington, D.C., USA.

Grayson, D. K., and D. J. Meltzer. 2002. Clovis hunting and large mammal extinction: a critical review of the evidence. Journal of World Prehistory 16:313-359.

Guiot, J., J. J. Boreux, P. Braconnot, F. Torre, and PMIP participants. 1999. Data-model comparison using fuzzy logic in paleoclimatology. Climate Dynamics 15:569-581.

Hallett, D. J., L. V. Hills, and J. J. Clague. 1997. New accelerator mass spectrometry radiocarbon ages for the Mazama tephra layer from Kootenay National Park, British Columbia, Canada. Canadian Journal of Earth Sciences 34:1202-1209.

Haskell, J. 2001. The latitudinal gradient of diversity through the Holocene as recorded by fossil pollen in Europe. Evolutionary Ecology Research 3:345-360.

Hewitt, G. M. 2000. The genetic legacy of the Quaternary ice ages. Nature 405:907-913.

Hewitt, G. M. 2004. Genetic consequences of climatic oscillations in the Quaternary. Philosophical Transactions of the Royal Society of London B 359:183-195.

International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, fourth edition. International Trust for Zoological Nomenclature, London, United Kingdom. [http://www.iczn.org/iczn/index.jsp]

Iversen, J., and J. Troels-Smith. 1950. Pollenmorfologiske definitioner og typer. Danmarks Geologiske Undersøgelse Series 4 3(8):1-54.

Jackson, S. T., and R. K. Booth. 2002. The role of late Holocene climate variability in the expansion of yellow birch in the western Great Lakes region. Diversity and Distributions 8:275-284.

Jackson, S. T., J. T. Overpeck, T. Webb, III, S. E. Keattch, and K. H. Anderson. 1997. Mapped plant-macrofossil and pollen records of late Quaternary vegetation change in eastern North America. Quaternary Science Reviews 16:1-70.

Jackson, S. T., R. S. Webb, K. H. Anderson, J. T. Overpeck, T. Webb, III, J. W. Williams, and B. C. S. Hansen. 2000. Vegetation and environment in eastern North America during the last glacial maximum. Quaternary Science Reviews 19:489-508.

Jackson, S. T., and C. Weng. 1999. Late Quaternary extinction of a tree species in eastern North America. Proceedings of the National Academy of Sciences of the United States of America 96:13847-13852.

Jackson, S. T., and J. W. Williams. 2004. Modern analogs in Quaternary paleoecology: here today, gone yesterday, gone tomorrow? Annual Review of Earth and Planetary Sciences 32:495-537.

Kageyama, M., O. Peyron, S. Pinot, P. Tarasov, J. Guiot, S. Joussaume, and G. Ramstein. 2001. The Last Glacial Maximum climate over Europe and western Siberia: a PMIP comparison between models and data. Climate Dynamics 17:23-43.

Kaplan, J. O., N. H. Bigelow, I. C. Prentice, S. P. Harrison, P. J. Bartlein, T. R. Christensen, W. Cramer, N. V. Matveyeva, A. D. McGuire, D. F. Murray, V. Y. Razzhivin, B. Smith, D. A. Walker, P. M. Anderson, A. A. Andreev, L. B. Brubaker, M. E. Edwards, and A. V. Lozhkin. 2003. Climate change and Arctic ecosystems: 2. Modeling, paleodata-model comparisons, and future projections. Journal of Geophysical Research 108(D19):8171. DOI:10.1029/2002JD002559.

Kohfeld, K. E., and S. P. Harrison. 2000. How well can we simulate past climates? Evaluating the models using global palaeoenvironmental datasets. Quaternary Science Reviews 19:321-346.

Kropf, M., J. W. Kadereit, and H. P. Comes. 2003. Differential cycles of range contraction and expansion in European high mountain plants during the Late Quaternary: insights from Pritzelago alpina (L.) O. Kuntze (Brassicaceae). Molecular Ecology 12:931-949.

Lascoux, M., A. E. Palmé, R. Cheddadi, and R. G. Latta. 2004. Impact of Ice Ages on the genetic structure of trees and shrubs. Proceedings of the Royal Society of London B 359:197-207.

Lebrun, J.-P., and A. L. Stork. 1991-1997. Enumération des plantes à fleurs d’Afrique Tropicale, 4 volumes. Conservatoire et Jardins Botaniques de la Ville de Genève, Geneva, Switzerland.

Lessa, E. P., J. A. Cook, and J. L. Patton. 2003. Genetic footprints of demographic expansion in North America, but not Amazonia during the late Quaternary. Proceedings of the National Academy of Sciences of the United States of America 100:10331-10334.

Martínez-Meyer, E., A. T. Peterson, and W. W. Hargrove. 2004. Ecological niches as stable distributional constraints on mammal species, with implications for Pleistocene extinctions and climate change projections for biodiversity. Global Ecology and Biogeography 13:305-314.

McNeill, J., F. R. Barrie, H. M. Burdet, V. Demoulin, D. J. Hawksworth, K. Marhold, D. H. Nicolson, J. Prado, P. C. Silva, J. E. Skog, J. H. Wiersema, and N. J. Turland. 2006. International code of Botanical Nomenclature (Vienna Code) adopted by the Seventh International Botanical Congress Vienna, Austria, July 2005. A.R.G. Gantner Velag, Ruggell, Liechtenstein. [http://ibot.sav.sk/icbn/main.htm]

Morgan, A. V., A. Morgan, A. C. Ashworth, and J. V. Matthews, Jr. 1983. Late Wisconsin fossil beetles in North America. Pages 354-363 in S. C. Porter, editor. Late-Quaternary environments of the United States. Volume 1. The late Pleistocene. University of Minnesota Press, Minneapolis, Minnesota, USA.

Newby, P. E., P. Killoran, M. R. Waldorf, B. N. Shuman, R. S. Webb, and T. Webb III. 2000. 14,000 years of sediment, vegetation, and water-level changes at the Makepeace Cedar Swamp, southeastern Massachusetts. Quaternary Research 53:352-368.

Noss, R. F., R. Graham, D. R. McCullough, F. L. Ramsey, J. Seavey, C. Whitlock, and M. P. Williams. 2000. Review of scientific material relevant to the cccurrence, ecosystem role, and tested management options for mountain goats in Olympic National Park. Fulfillment of contract #14-01-0001-99-C-05, U. S. Department of Interior. Conservation Biology Institute, Corvallis, Oregon, USA. http://www.consbio.org/cbi/professional_services/goat/goat_pdf.htm.

Owen, P. R., C. J. Bell, and E. M. Mead. 2000. Fossils, diet, and conservation of black-footed ferrets (Mustela nigripes). Journal of Mammalogy 81:422-433.

Pasenko, M. R., and B. W. Schubert. 2004. Mammuthus jeffersonii (Proboscidea, Mammalia) from northern Illinois. PaleoBios 24(3):19-24.

Petit, J. R., I. Aguinagalde, J.-L. de Beaulieu, C. Bittkau, S. Brewer, R. Cheddadi, R. Ennos, S. Fineschi, D. Grivet, M. Lascoux, A. Mohanty, G. Müller-Starck, B. Demesure-Musch, A. Palmé, J. P. Martin, S. Rendell, and G. G. Vendramin. 2003. Glacial refugia: hotspots but not melting pots of genetic diversity. Science 300:1563-1565.

Petit, R. J., R. Bialozyt, P. Garnier-Géré, and A. Hampe. 2004. Ecology and genetics of tree invasions: from recent introductions to Quaternary migrations. Forest Ecology and Management 197:117-137.

Petit, R. J., S. Brewer, S. Bordács, K. Burg, R. Cheddadi, E. Coart, J. Cottrell, U. M. Csaikl, B. van Dam, J. D. Deans, S. Espinel, S. Fineschi, R. Finkeldey, I. Glaz, P. G. Goicoechea, J. S. Jensen, A. O. König, A. J. Lowe, S. F. Madsen, G. Mátyás, R. C. Munro, F. Popescu, D. Slade, H. Tabbener, S. G. M. de Vries, B. Ziegenhagen, J.-L. de Beaulieu, and A. Kremer. 2002. Identification of refugia and post-glacial colonisation routes of European white oaks based on chloroplast DNA and fossil pollen evidence. Forest Ecology and Management 156:49-74.

Petit, R. J., E. Pineau, B. Demesure, R. Bacilieri, A. Ducousso, and A. Kremer. 1997. Chloroplast DNA footprints of postglacial recolonization by oaks. Proceedings of the National Academy of Sciences of the United States of America 94:9996-10001.

Prentice, I. C., D. Jolly, and BIOME 6000 participants. 2000. Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa. Journal of Biogeography 27:507-519.

Qian, H., and R. E. Ricklefs. 2000. Large-scale processes and the Asian bias in species diversity of temperate plants. Nature 407:180-182.

Reimer, P. J., M. G. L. Baillie, E. Bard, A. Bayliss, J. W. Beck, C. J. H. Bertrand, P. G. Blackwell, C. E. Buck, G. S. Burr, K. B. Cutler, P. E. Damon, R. L. Edwards, R. G. Fairbanks, M. Friedrich, T. P. Guilderson, A. G. Hogg, K. A. Hughen, B. Kromer, G. McCormac, S. Manning, C. B. Ramsey, R. W. Reimer, S. Remmele, J. R. Southon, M. Stuiver, S. Talamo, F. W. Taylor, J. van der Plicht, and C. E. Weyhenmeyer. 2004. INTCAL04 terrestrial radiocarbon age calibration, 0-26 cal kyr BP. Radiocarbon 46:1029-1058.

Rosenberg, S. M., I. R. Walker, and R. W. Mathewes. 2003. Postglacial spread of hemlock (Tsuga) and vegetation history in Mount Revelstoke National Park, British Columbia, Canada. Canadian Journal of Botany 81:139-151.

Runck, A. M., and J. A. Cook. 2005. Postglacial expansion of the southern red-backed vole (Clethrionomys gapperi) in North America. Molecular Ecology 14:1445-1456.

Schauffler, M., and G. L. Jacobson, Jr. 2002. Persistence of coastal spruce refugia during the Holocene in northern New England, USA, detected by stand-scale pollen stratigraphies. Journal of Ecology 90:235-250.

Schmitz, O. J., E. Post, C. E. Burns, and K. M. Johnston. 2003. Ecosystem responses to global climate change: moving beyond color mapping. Bioscience 53:1199-1205.

Silvertown, J. 1985. History of a latitudinal diversity gradient: woody plants in Europe 13,000–1,000 years. Journal of Biogeography 12:519-525.

Stevens, P. F. 2007+. Angiosperm Phylogeny Website. Version 8, June 2007. [http://www.mobot.org/MOBOT/research/APweb/]

Stuiver, M., and P. J. Reimer. 1993. Extended 14C database and revised CALIB radiocarbon calibration program. Radiocarbon 35:215-230.

Telford, R. J., E. Heegaard, and H. J. B. Birks. 2004a. All age-depth models are wrong: but how badly? Quaternary Science Reviews 23:1-5.

Telford, R. J., E. Heegaard, and H. J. B. Birks. 2004b. The intercept is a poor estimate of a calibrated radiocarbon age. The Holocene 14:296-298.

Thompson, R. S., and K. H. Anderson. 2000. Biomes of western North America at 18,000, 6000 and 0 14C yr BP reconstructed from pollen and packrat midden data. Journal of Biogeography 27:555-584.

Tremblay, N. O., and D. J. Schoen. 1999. Molecular phylogeography of Dryas integrifolia: glacial refugia and postglacial recolonization. Molecular Ecology 8:1187-1198.

Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. Coan, F. G. Hochberg, W. G. Lyons, P. M. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F. G. Thompson, M. Vecchione, and J. D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: mollusks, second edition. American Fisheries Society, Special Publication 26, Bethesda, Maryland, USA.

Tutin, T. G., et al., editor. 1964-1993. Flora Europaea, 5 volumes. Cambridge University Press, Cambridge, United Kingdom. [http://rbg-web2.rbge.org.uk/FE/fe.html]

Webb, T., III. 1981. The past 11,000 years of vegetational change in eastern North America. Bioscience 31:501-506.

Webb, T., III. 1987. The appearance and disappearance of major vegetational assemblages: long-term vegetational dynamics in eastern North America. Vegetatio 69:177-187.

Webb, T., III. 1988. Eastern North America. Pages 385-414 in B. Huntley and T. Webb, III, editors. Vegetation history. Handbook of vegetation science 7. Kluwer Academic Publishers, Dordrecht.

Webb, T., III, B. Shuman, and J. W. Williams. 2004. Climatically forced vegetation dynamics in eastern North America during the late Quaternary Period. Pages 459-478 in A. R. Gillespie, S. C. Porter, and B. F. Atwater, editors. The Quaternary Period in the United States. Developments in Quaternary science 1. Elsevier, Amsterdam, The Netherlands.

Whorley, J. R., S. T. Alvarez-Castañeda, and G. J. Kenagy. 2004. Genetic structure of desert ground squirrels over a 20-degree-latitude transect from Oregon through the Baja California peninsula. Molecular Ecology 13:2709-2720.

Willard, D. A., T. M. Cronin, and S. Verardo. 2003. Late-Holocene climate and ecosystem history from Chesapeake Bay sediment cores, USA. The Holocene 13:201-214.

Williams, J. W. 2003. Variations in tree cover in North America since the last glacial maximum. Global and Planetary Change 35:1-23.

Williams, J. W., B. N. Shuman, and T. Webb, III. 2001. Dissimilarity analyses of late-Quaternary vegetation and climate in eastern North America. Ecology 82:3346-3362.

Williams, J. W., B. N. Shuman, T. Webb, III, P. J. Bartlein, and P. L. Luduc. 2004. Late-Quaternary vegetation dynamics in North America: scaling from taxa to biomes. Ecological Monographs 74:309-334.

Williams, J. W., R. L. Summers, and T. Webb, III. 1998. Applying plant functional types to construct biome maps from eastern North American pollen data: comparisons with model results. Quaternary Science Reviews 17:607-627.

Williams, J. W., T. Webb, III, P. H. Richard, and P. Newby. 2000. Late Quaternary biomes of Canada and the eastern United States. Journal of Biogeography 27:585-607.

Wilson, D. E., and D. M. Reeder. 2005. Mammal species of the world: a taxonomic and geographic reference, third edition, 2 volumes. The Johns Hopkins University Press, Baltimore, Maryland, USA. [http://www.bucknell.edu/msw3/]

Wright, H. E., Jr., J. E. Kutzbach, T. Webb, III, W. F. Ruddiman, F. A. Street-Perrott, and P. J. Bartlein, editors. 1993. Global climates since the last glacial maximum, volumes. University of Minnesota Press, Minneapolis.

Wroe, S., J. Field, R. Fullagar, and L. S. Jermiin. 2004. Megafaunal extinction in the late Quaternary and the global overkill hypothesis. Alcheringa 28:291-331.

Zdanowicz, C. M., G. A. Zielinski, and M. S. Germani. 1999. Mount Mazama eruption: calendrical age verified and atmospheric impact assessed. Geology 27:621-624.