Albert J. Parker

THE TOPOGRAPHIC RELATIVE MOISTURE INDEX: AN APPROACH TO SOIL-MOISTURE ASSESSMENT IN MOUNTAIN TERRAIN

The Topographic Relative Moisture Index (TRMI), designed to indicate the relative soil moisture availability among sites in mountain terrain, is described. The TRMI is a summed scalar index of four slope parameters: topographic position, slope aspect, steepness, and slope configuration. A review of other methods of characterizing site moisture relations, including direct monitoring, water-balance climatology, site index, and other inferential topographic/edaphic indices, reveals that the simple, straightforward application of the TRMI is rapid, inexpensive, and effective in field research. The ability of the TRMI to portray significant patterns of plant response to environment is demonstrated from empirical studies at both the species-level (the ecology of Arizona cypress) and community-level (forest patterns in Yosemite National Park).

Fine-scale genetic structure in Pinus clausa (Pinaceae) populations: effects of disturbance history.

Spatial autocorrelation analyses of 12 allozyme loci were used to compare genetic structure within populations of two varieties of Pinus clausa. P. clausa var. immuginata populations tend to be uneven-aged, with continuous recruitment in small gaps created by wind damage, whereas P. clausa var. clausa populations are more even-aged, with recruitment postdating periodic canopy fires. Three var. immuginata populations and three matched pairs of var. clausa populations, including both a mature and a nearby recently burned population, were examined. Aggregation of multilocus genotypes at small distances was evident in all young var. clausa populations. Little inbreeding was apparent among juveniles or adults in these populations; their genetic structure is likely to have resulted from limited seed dispersal. Genotypes were not significantly spatially structured in nearby matched mature populations. Genetic structure was less evident in var. immuginata populations. Aggregated genotypes were only apparent in the population where patches included juveniles of similar ages; dense juvenile clumps in the other two var. immuginata populations comprised a variety of ages. Interannual variability in allele frequencies of surviving seedlings may account for the absence of genetic structure in these populations.

Climate remains an important driver of post‐European vegetation change in the eastern United States

Department of Geography, University of Wisconsin-Madison, 550 North ParkStreet, Madison, WI 53706, USAThe influence of climate on forest change during thepast century in the eastern United States was evalu-ated in a recent paper (Nowacki & Abrams, 2014)that centers on an increase in ‘highly competitivemesophytic hardwoods’ (Nowacki & Abrams, 2008)and a concomitant decrease in the more xerophyticQuercus species. Nowacki & Abrams (2014) con-cluded that climate change has not contributed sig-nificantly to observed changes in forest composition.However, the authors restrict their focus to a singleelement of climate: increasing temperature since theend of the Little Ice Age ca. 150 years ago. In theirstudy, species were binned into four classifications(e.g., Acer saccharum – ‘cool-adapted’, Acer rubrum –‘warm-adapted’) based on average annual tempera-ture within each species range in the United States,reducing the multifaceted character of climate into asingle, categorical measure. The broad temperatureclasses not only veil the many biologically relevantaspects of temperature (e.g., seasonal and extremetemperatures) but they may also mask other influ-ences, both climatic (e.g., moisture sensitivity) andnonclimatic (e.g., competition).Understanding the primary drivers of forest changeis critically important. However, using annual tem-perature reduces the broad spectrum of climaticinfluence on forests (e.g., Jackson & Overpeck, 2000;Jackson et al., 2009) to a single variable. Tsuga canad-ensis illustrates one example of the complex interac-tion between trees and temperature. In the southernpart of its range, Tsuga canadensis growth is weakly,but positively correlated with early growing-seasontemperature. However, this relationship becomesstronger and shifts to later in the season toward thenorthern part of its range (Cook & Cole, 1991). More-over, Tsuga canadensis growth is significantly andnegatively correlated with just May temperaturesduring the current growing season in the northeast-ern United States (Cook, 1991; Cook & Cole, 1991;Vaganov et al., 2011), while in the southeastern Uni-ted States it is strongly and negatively correlatedwith summer (June–August) temperatures (Hart et al.,2010). Trees can also be sensitive to diverse and ofteninteracting climate variables at various stages of theirlife cycles (Jackson et al., 2009). Interactions betweenprecipitation and temperature are clearly important(Harsch & Hille Ris Lambers, 2014; Martin-Benito &Pederson, accepted), and often lead to counterintui-tive responses. For example, some plant species thatwould have been expected to move north and ups-

Climate, Lightning, and Wildfire in the National Forests of the Southeastern United States: 1989-1998

We compiled the record of wildfires from each national forest unit in the southeastern United States for the period 1989 to 1998, then regressed annual fire numbers against lightning flash densities (as recorded by the National Lightning Detection Network) and measures of climatic drought (seasonal precipitation and Palmer Drought Severity Index [PDSI] values). Lightning-caused fires were most common in Florida, where seasonally-dry winters and high lightning-flash density elevate fire risk. Overall regression models consistently identified fire-season PDSI as the most influential explanatory variable in accounting for the spatial and temporal pattern of fires, whereas the contribution of lightning-flash density to regional patterns of wildfires is muted. Lightning density only enters the overall model when a single outlier forest unit from Florida is included in the regression analysis. With respect to physiographically defined regional regression models, lightning density was only a significant influence in the Coastal Plain-East, which included Florida. Some measure of climatic drought was uniformly present in all other regions, except in the driest portion of the study region, the Ozark-Ouachita Uplands. Elsewhere, geographic variation in effective moisture is highlighted. Drought for periods of up to 12 months is necessary to promote burning in the wetter forests of the Appalachian Highlands, but periods of antecedent drought necessary to trigger widespread wildfires are shorter in the warmer and drier Piedmont and Coastal Plain.

Spatial components of biotic diversity in landscapes of Georgia, USA

Traditional measures of species diversity and spatially-explicit measures of landscape diversity (derived from Romme 1982) are used to compare biotic diversity in six landscapes across Georgia, USA; two each from the Appalachian Highlands, Piedmont, and Coastal Plain. Species richness and evenness of plots generally increased from the Coastal Plain to the Appalachian Highlands. Community richness, landscape contrast, and landscape heterogeneity increased from the Appalachian Highlands to the Coastal Plain, opposite the gradient of topographic complexity. Coastal Plain landscapes possessed greater contrast and heterogeneity than landscapes in the other two physiographic provinces. This high level of landscape diversity is interpreted as a response to two factors: the increased role of human activity in shaping landscape structure, and the increased range of soil moisture regimes encountered in the sand-rich substrates of the Coastal Plain (from permanently flooded hydric communities to well drained xeric uplands only a few meters higher in elevation).

Disturbance-Mediated Variation in Stand Structure between Varieties of Pinus clausa (Sand Pine)

Geographically mediated differences in exposure to disturbance agents that operate at different scales impose demographic contrasts between plant populations. We examined such effects on the stand-scale population structure of the two varieties of Pinus clausa (sand pine). Pinus clausa var. immuginata stands of the Florida panhandle exhibited reverse-J diameter and age structures, with ongoing stem recruitment and relatively high seedling and sapling densities in canopy gaps. Seedling radial growth rates were relatively slow in P. c. var. immuginata stands, reflecting the partially closed character of the forest canopy. Radial growth-release events among trees were recorded in 35 – 65% of trees in P. c. var. immuginata stands. Timing of releases matched exposure to hurricanes and extratropical storms in these coastal settings. Wind damage provides opportunities for recruitment and growth of stems, so that stands of P. c. var. immuginata do not require stand-destroying disturbances to persist in the landscape. Mature stands of Pinus clausa var. clausa on the Florida peninsula were broadly unimodal in diameter form and narrowly even-aged, with virtually all trees established within a decade following stand-initiating disturbances in the 1920s and 1930s; seedlings were absent. Regeneration of P. c. var. clausa in our study was limited to stands that have experienced crown fire since 1970. Seedlings in the recently burned P. c. var. clausa stands exhibited rapid growth rates. Growth-release events were uncommon in P. c. var. clausa stands, except for an Atlantic coastal population that experienced repeated exposure to hurricanes from 1947 – 1950. Crown fires appear essential to long-term maintenance of naturally seeded populations of Pinus clausa var. clausa on most sites.

Allozyme diversity in Pinus virginiana (Pinaceae): intraspecific and interspecific comparisons

Two of the four members of subsection Contortae of the genus Pinus occur in the southeastern United States: Pinus virginiana, which ranges throughout the southern and central Appalachian Mountains, and P. clausa, which is restricted to Florida and southern Alabama. We examined allozyme variation within P. virginiana and genetic relationships between this species and the two varieties of P. clausa (var. clausa and var. immuginata). P. virginiana maintains more genetic diversity at both the species (Hes = 0.139) and population (Hep = 0.128) levels than the other three species in the subsection, which may reflect the combination of its widespread distribution and the absence of cone serotiny. Genetic differentiation among populations in P. virginiana was relatively low (GST = 0.053), but significant contrasts in allozyme frequencies and genetic diversity were apparent between populations to the northwest vs. outheast of the Appalachian Mountains. These regional differences likely resulted initially from historical processes that occurred during the Pleistocene and early Holocene, and have been reinforced by modern selective pressures and barriers to gene flow. The mean genetic distance between populations of P. virginiana and P. clausa (D = 0.071) was greater than that between populations of the two varieties of P. clausa (D = 0.012), which suggests that the two varieties diverged at some point after the separation of the two species.

Inferring ancient Agave cultivation practices from contemporary genetic patterns

Several Agave species have played an important ethnobotanical role since prehistory in Mesoamerica and semiarid areas to the north, including central Arizona. We examined genetic variation in relict Agave parryi populations northeast of the Mogollon Rim in Arizona, remnants from anthropogenic manipulation over 600 years ago. We used both allozymes and microsatellites to compare genetic variability and structure in anthropogenically manipulated populations with putative wild populations, to assess whether they were actively cultivated or the result of inadvertent manipulation, and to determine probable source locations for anthropogenic populations. Wild populations were more genetically diverse than anthropogenic populations, with greater expected heterozygosity, polymorphic loci, effective number of alleles and allelic richness. Anthropogenic populations exhibited many traits indicative of past active cultivation: fixed heterozygosity for several loci in all populations (nonexistent in wild populations); fewer multilocus genotypes, which differed by fewer alleles; and greater differentiation among populations than was characteristic of wild populations. Furthermore, manipulated populations date from a period when changes in the cultural context may have favoured active cultivation near dwellings. Patterns of genetic similarity among populations suggest a complex anthropogenic history. Anthropogenic populations were not simply derived from the closest wild A. parryi stock; instead they evidently came from more distant, often more diverse, wild populations, perhaps obtained through trade networks in existence at the time of cultivation.

Genetic consequences of pre-Columbian cultivation for Agave murpheyi and A. delamateri (Agavaceae)

Pre-Columbian farmers cultivated several species of agave in central Arizona from ca. A.D. 600-1350. Because of the longevity and primarily asexual reproduction of these species, relict agave clones remain in the landscape and provide insights into pre-Columbian agricultural practices. We analyzed variation in allozyme allele frequencies to infer genetic effects of prehistoric cultivation on Agave murpheyi and A. delamateri, specifically to estimate genetic diversity and structure, to determine whether cultivated populations descended from a single clone, and to examine regional-scale genetic variation. Agave murpheyi maintained more genetic diversity at the species and population levels than A. delamateri, and A. murpheyi populations typically included more multilocus genotypes. Relict plants from prehistoric fields reflect a more complex history than descent from a single clone; A. murpheyi populations may have included more diversity initially because bulbils (produced routinely in A. murpheyi but not A. delamateri) and possibly seed would have facilitated transport of genetically diverse planting stock. Genetic variation in both cultigens was lower than in most contemporary commercial crops but similar to that observed in modern traditional agricultural systems.

Evergreen understory dynamics in Coweeta forest, North Carolina

A number of studies have elucidated the distributional patterns of various components of Southern Appalachian forests. The evergreen understory here is composed largely of a dominant ericaceous shrub, Rhododendron maximum L., which is believed to be expanding and inhibiting the development of other species with consequent impacts on overall forest structure and composition. We use a GIS and logistic regression to examine this less-studied forest element in the Coweeta Hydrological Laboratory, North Carolina, over a 17-year period to determine whether expansion is occurring and, if so, whether it can be predicted based on terrain characteristics. We examine two adjacent, physically similar basins with differing degrees of experimental manipulation in the 20th century in an attempt to examine the role of environmental and historical factors in determining spatial patterns of persistence, expansion, and decline of the evergreen understory. Results indicate that significant expansion of the evergreen understory occurred in both basins during the period 1976-1993, and that stream proximity, topographic setting, and elevation are related to patterns of evergreen-understory dynamics. Patterns differ between the two basins, suggesting that disturbance and differing land-use histories are also influential. Predictive power of models based on terrain factors alone ranges from <10% to >50%.