Latitudinal decrease in folivory within Nothofagus pumilio forests: dual effect of climate on insect density and leaf traits?

Aim The strength of consumer–plant interactions may decrease with latitude. Our objectives were to assess the spatial variation in folivory on Nothofagus pumilio and understand the influence of climate on folivory patterns as mediated by changes in folivore density and leaf traits. Location Nothofagus pumilio forests, between 38 and 55°S (Argentina). Methods We studied the correlation of leaf damage with latitude on data from 47 sampling sites, and evaluated spatial patterns of autocorrelation on latitudinally detrended data with a principal coordinates of neighbour matrices method. Path analysis was used to test the association of temperature and precipitation with leaf damage, mediated by folivore density and leaf traits. We evaluated the adequacy of this ecological model by examining the spatial pattern of autocorrelation in the residuals, and combined spatial and environmental predictors of leaf damage into partial regression. Results Leaf damage decreased with latitude, which was the only significant spatial predictor. The latitudinal decrease in temperature and precipitation was correlated with a decrease in the density of folivores and leaf size, and diminished leaf damage. Our ecological model adequately explained the spatial autocorrelation in the data: 44% of the variation in leaf damage was explained by the latitudinally structured component of the environment, whereas local environmental effects accounted for another 22%. Main conclusions We conclude that N. pumilio forests show consistent latitudinal patterns of variation in folivory, folivore density and leaf traits. Our study suggests that the latitudinal variation in folivory rates is partly driven by the influence of climate on both plants and herbivores. This warns us about the potential susceptibility of folivory rates to climate warming. We emphasize the value of large‐scale analyses as complementary to local experimental approaches to understanding the regulation of herbivory.     

Regional climate oscillations and local topography shape genetic polymorphisms and distribution of the giant columnar cactus Echinopsis terscheckii in drylands of the tropical Andes

Aim

We sought to determine if the present fragmentary distribution of the giant columnar cactus Echinopsis terscheckii in tropical drylands is a relict of a previously more widespread range during cold and dry phases of the Last Glacial Maximum (LGM).

Location

Tropical and subtropical dry ecotonal areas of northern and central Andes of Argentina.

Methods

We combined ecological niche models (ENM) with molecular polymorphisms of isozymes and DNA sequences. We collected samples from 30 individuals at 24 locations for genetic analysis covering a wide range of environmental conditions. We sequenced the nuclear ITS and three non‐coding regions of the chloroplast DNA and we resolved 15 isozyme loci. Potential distribution was modelled using 88 E. terscheckii presence training records and a reduced set of 10 modern bioclimatic variables. LGM and the Mid‐Holocene distributions were derived by projecting bioclimatic data under present to past environmental conditions according to CCSM4 and MIROC‐ESM Global Climate Models.

Results

We detected high isozyme diversity towards the south. The multivariate cluster analysis yielded two groups of populations that were geographically concordant with the DNA haplotypes located north and south of a divide at 27°S. Distribution models show range expansion during the LGM in two north and south areas separated by a gap of low suitability at 27°S. Suitable areas in the south were close to current populations, while in the north, populations survived in more disjunct locations that probably suffered from founder effects. In contrast, Mid‐Holocene bioclimatic conditions were relatively unsuitable in the south.

Main conclusions

Our results suggest that the divergence of north and south groups of E. terscheckii populations reflect long‐lasting persistence through climatic cycles that were reinforced by the presence of an orogenic divide at mid‐latitudes. Latitudinally divergent groups of populations should be treated as distinct evolutionary significant units that deserve independent conservation actions. Increased genetic diversity and inbreeding towards the south may guide setting up priorities for the long‐term protection of a dominant element of drylands as E. terscheckii.     

Environmental and genetic control of insect abundance and herbivory along a forest elevational gradient

Environmental conditions and plant genotype may influence insect herbivory along elevational gradients. Plant damage would decrease with elevation as temperature declines to suboptimal levels for insects. However, host plants at higher elevations may exhibit traits that either reduce or enhance leaf quality to insects, with uncertain net effects on herbivory. We examined folivory, insect abundance and leaf traits along six replicated elevational ranges in Nothofagus pumilio forests of the northern Patagonian Andes, Argentina. We also conducted a reciprocal transplant experiment between low- and high-elevation sites to test the extent of environmental and plant genetic control on insect abundance and folivory. We found that insect abundance, leaf size and specific leaf area decreased, whereas foliar phosphorous content increased, from low-, through mid- to high-elevation sites. Path analysis indicated that changes in both insect abundance and leaf traits were important in reducing folivory with increasing elevation and decreasing mean temperature. At both planting sites, plants from a low-elevation origin experienced higher damage and supported greater insect loads than plants from a high-elevation origin. The differences in leaf damage between sites were twofold larger than those between plant origins, suggesting that local environment was more important than host genotype in explaining folivory patterns. Different folivore guilds exhibited qualitatively similar responses to elevation. Our results suggest an increase in insect folivory on high-elevation N. pumilio forests under future climate warming scenarios. However, in the short-term, folivory increases might be smaller than expected from insect abundance only because at high elevations herbivores would encounter more resistant tree genotypes.     

Environmental correlates of mammal species richness in South America: effects of spatial structure, taxonomy and geographic range

Although some consensus exists regarding the positive synergism between energy and heterogeneity in increasing species diversity, the role of environmental variability remains controversial. We examine how these factors interact to explain spatial variation in mammal species richness in South America. After taking into account the effects of spatial autocorrelation and area, elevation variability and energy mainly drive spatial variation in mammal species richness. The effect of environmental variability is less important. When different taxonomic groups of mammals are analyzed separately, three ways emerge whereby energy and heterogeneity interact to promote species richness. Heterogeneity may have no effect on species richness, habitat heterogeneity and energy availability contribute independently to species richness, or heterogeneity increases in importance with an increase in energy availability. The partition of species into range size quartiles shows that habitat heterogeneity and temporal instability in the resource supply account for the species richness pattern in the narrowest- ranging species. Habitat heterogeneity is significant also for intermediate ranging species but not for the widest-ranging species. Energy alone drives the species richness pattern in the latter species. The interplay between ecology and biogeographic history may ultimately explain these differences given that narrow- and wide-ranging species show distinct biogeographic patterns, and different taxonomic groups also unequally represent them.     

Patterns of use and damage by exotic deer on native plant communities in northwestern Patagonia

Ungulate overabundance can strongly alter composition and structure of forest communities through selective damage independently of the herbivory history of the recipient system. In the early 1900s, red and fallow deer (Cervus elaphus and Dama dama, respectively) were introduced to northwestern Patagonia. We studied patterns of relative habitat use, damage, and species selection by introduced deer populations in three major plant communities that have the longest history of invasion in the region. We also evaluated community structure and composition characteristics as possible explanatory factors for the observed patterns of browsing, fraying, and bark-stripping. At the species level, exotic deer browsed more than expected on two evergreen species (Austrocedrus chilensis and Schinus patagonicus) and two spiny species (Colletia hystrix and Dasyphyllum diacanthoides), while one dominant shrubland tree (Lomatia hirsuta) was selected for fraying and bark-stripping. Browsing was the more widespread damage in all plant communities, while fraying and bark-stripping occurred at low incidence. Furthermore, species identity was found as the main driver of deer damage over plant community type. Finally, variation in damage and in habitat use was explained by community structure and composition characteristics. Bark-stripping and fraying are best predicted by community composition, whereas browsing and habitat use are best predicted by structural properties including understory cover, cover of fallen logs, and tree density. The patterns of damage and the association with community structure characteristics reported here provide insight for developing management strategies.     

Landscape responses to a century of land use along the northern Patagonian forest-steppe transition

Land use history reconstructions in temperate regions of the Northern Hemisphere indicate that periods of deforestation are often followed by natural afforestation, so that the long-term outcome at the landscape level will be a balance of retractions and advances of plant communities associated with varying local land uses. During the last decades of the XIX century, large forest areas were cleared in Northwestern Patagonia to open farmland. In this article, we compared historical land use/land cover maps with land cover maps derived from Landsat images to analyze the factors that may have influenced the dynamics of land cover change of the forest-steppe ecotone during the last 100 years. Our results indicate that Patagonian forests underwent a rapid initial recovery after the extensive fires of last century, replacing mainly shrublands. More than 50% of the old burns are currently covered by forests, and modern fires affect areas characterized by fire-prone vegetation. Whereas natural afforestation is an ongoing process positively associated with moisture, the rate of forest losses has increased during the last three decades, concentrating on xeric aspects and the vicinity of roads. We conclude that the outcome of the dynamics between fire-intolerant forests and fire-prone plant communities will largely depend on human-related activities, modeled by structural features of the landscape (i.e., topography, dominant winds), and processes triggered by past land uses.     

Inter-hemispheric comparison of fire history: The Colorado Front Range,U.S.A., and the Northern Patagonian Andes,Argentina

Fire history was compared between the Colorado Front Range (U.S.A.) and northern Patagonia (Argentina) by dating fire-scars on 525 Pinus ponderosa and 418 Austrocedrus chilensis, respectively, and determining fire weather on the basis of instrumental and tree-ring proxy records of climatic variation. Years of above average moisture availability preceding fire years, rather than drought alone, is conducive to years of widespread fire in the Colorado Front Range and the northern Patagonian study areas. Above-average precipitation promotes fire by enhancing the growth of herbaceous plants which increases the quantity of fine fuels during the fire season a few years later. The short-term variability in moisture availability that is conducive to widespread burning is strongly related to El Niño Southern Oscillation (ENSO) activity. The warm (El Niño) phase of ENSO is associated with greater moisture availability during the spring in both regions which leads to peaks in fire occurrence several years after El Niño events. The warmer and drier springs associated with la Niña events exacerbate the drying of fuels so that fire years commonly coincide with La Niña events. In both regions, there was a dramatic decline in fire occurrence after the early 1900s due to a decline in intentionally set fires by Native Americans and European settlers, fuel reduction by livestock grazing, and increasingly effective organized fire suppression activities after the 1920s. In both regions there was a marked increase in fire frequency during the mid-and late-19th centuries which coincides with increased ignitions by Native Americans and/or European settlers. However, year-to-year variability in ring widths of Pinus ponderosa and Austrocedrus chilensis also increased from relatively low values in the late 1700s and early 1800s to peaks in the 1850s and 1860s. This implies frequent alternation of years of above and below average moisture availability during the mid-19th century when the frequencies of major fire years rise. The high correlation of tree-growth variability betweem the two regions implies a strong inter-hemispheric variation in climatic variability at a centennial time scale which closely parallels a variety of proxy records of ENSO activity. Based on the relationship of fire and ENSO events documented in the current study, this long-term trend in ENSO activity probably contributed to the mid- and late-19th century increase in fire spread in both regions. These similar trends in fire occurrence have contributed to similar patterns of forest structures, forest health, and current hazard of catastrophic wildfire in the Colorado Front Range and northern Patagonia.     

Interannual changes in folivory and bird insectivory along a natural productivity gradient in northern Patagonian forests

Trophic regulation models suggest that the magnitude of herbivory and predation (top-down forces) should vary predictably with habitat productivity. Theory also indicates that temporal abiotic variation and within-trophic level heterogeneity both affect trophic dynamics, but few studies addressed how these factors interact over broad-scale environmental gradients. Here we document herbivory from leaf-feeding insects along a natural rainfall/productivity gradient in Nothofagus pumilio forests of northern Patagonia, Argentina, and evaluate the impact of insectivorous birds on foliar damage experienced by tree saplings at each end of the gradient. The study ran over three years (1997–2000) comprising a severe drought (1998–1999), which allowed us to test how climatic events alter top-down forces. Foliar damage tended to increase towards the xeric, least productive forests. However, we found a predictable change of insect guild prevalence across the forest gradient. Leaf miners accounted for the greater damage recorded in xeric sites, whereas leaf chewers dominated in the more humid and productive forests. Interannual folivory patterns depended strongly on the feeding guild and forest site. Whereas leaf-miner damage decreased during the drought in xeric sites, chewer damage increased after the drought in the wettest site. Excluding birds did not affect leaf damage from miners, but generally increased chewer herbivory on hydric and xeric forest saplings. Indirect effects elicited by bird exclusion became most significant after the drought, when total folivory levels were higher. Thus, interannual abiotic heterogeneity markedly influenced the amount of folivory and strength of top-down control observed across the forest gradient. Moreover, our results suggest that spatial turnovers between major feeding guilds may need be considered to predict the dynamics of insect herbivory along environmental gradients.     

Climatic influences on fire regimes along a rain forest-to-xeric woodland gradient in northern Patagonia, Argentina

ABSTRACT. Influences of annual climatic variation on fire occurrence were examined along a rainfall gradient from temperate rainforest to xeric woodlands in northern Patagonia, Argentina. Fire chronologies were derived from fire scars on trees and related to tree-ring proxy records of climate over the period 1820–1974. Similarly, fire records of four Patagonian national parks for the period 1940–1988 were compared to instrumental weather data. Finally, the influences of broad-scale synoptic weather patterns on fire occurrence in northern Patagonia were explored.
Fire in Nothofagus rainforests is highly dependent on drought during the spring and summer of the same year in which fires occur and is less strongly favoured by drought during the spring of the previous year. The occurrence of fire in dry vegetation types near the steppe ecotone is less dependent on drought because even during years of normal weather fuels are thoroughly desiccated during the dry summer. In xeric Austrocedrus woodlands, fire occurrence and spread are promoted by droughts during the fire season and also appear to be favoured by above-average moisture conditions during the preceding 1 to 2 growing seasons which enhances fuel production. Thus, in the xeric woodlands fire is not simply dependent on drought but is favoured by greater climatic variability over time scales of several years.
Fire activity in northern Patagonia is greatly influenced by the intensity and latitudinal position of the subtropical high pressure cell of the southeast Pacific. Greater fire activity is associated with a more intense and more southerly located high pressure cell which blocks the influx of Pacific moisture into the continent. Although long-term changes in fire occurrence along the rainforest-to-xeric woodland gradient have been greatly influenced by human activities, annual variation in fire frequency and extent is also strongly influenced by annual climatic variation.