Seedling recruitment, survival and facilitation in alpine Pinus uncinata tree line ecotones. Implications and potential responses to climate warming

Aims Alpine tree line ecotones are harsh environments where low temperatures constrain tree regeneration and growth. However, the expected upward shift of tree line ecotones in response to climate warming has not been ubiquitous. The lack of coupling between tree line dynamics and climate warming might be explained by factors other than climate variation that determine seedling recruitment in these ecotones. We want to assess how the availability of suitable habitat for establishment and the effects of facilitation on seedling survival and growth affect tree recruitment within tree line ecotones and modulate their responses to climate. Location We evaluate the relevance of these factors for Pinus uncinata tree line ecotones in the Catalan Pyrenees (north‐east Spain) and Andorra. Methods We analysed the microhabitat of naturally established seedlings in rectangular plots at the tree line ecotone, assessing the habitat type and the proximity to potentially protective elements that may improve microsite conditions. We tested whether krummholz individuals influence regeneration at the tree line by performing a transplantation field experiment to evaluate the extent of facilitation on seedling survival and growth in height. A total of 820 seedlings were transplanted at different distances and orientations (resulting in 12 positions) from krummholz mats and monitored over 2 years. Results Safe sites for P. uncinata recruits consisted of sparse vegetation covering bare soil, gravel or litter, and close to protective elements that may ameliorate microsite conditions. The field experiment showed that directional positive interactions enhance seedling survival and growth, altering the spatial patterns of recruit survivorship, especially during harsh winter conditions (shallow and irregular snowpack). Main conclusions Our results suggest that scarce availability of safe sites and uneven facilitation by krummholz control seedling recruitment patterns within alpine tree line ecotones. Such constraints may distort or counter the response of tree line ecotones to climate warming at local and regional scales.     

Patterns and dynamics of tree-line response to climate change in the eastern Qilian Mountains,northwestern China

Tree-line ecotones are strongly climatically limited and serve as potential monitors of climate change. We employed annual growth increment from tree-rings, and tree density and age structure data derived from two Juniperus przewalskii tree-line sites in the eastern part of the Qilian Mountains, northeastern Tibetan Plateau, to detect the responses of tree growth and population dynamics to climate change. High temperature favors tree growth and is associated with increased tree density at tree-line, and an advance in tree-line position. Significantly positive correlations were found between ring-width and mean monthly air temperatures in current and previous June, July and August. Tree recruitment began to increase rapidly at the two sites after the Little Ice Age, but then decreased starting in the 1970s. The number of trees established coincides with temperature changes. The warming trend after the Little Ice Age favors increases of tree density and an advance of tree-line. The majority of trees established during the period of 1931–1970, which coincides well with the rapid radial growth of the trees.     

Tree-line dynamics in Ungava peninsula,northern Quebec

The characteristics of the present tree-line in continental Northern Quebec appear to be related to modern and past ecological conditions. ¹⁴C dating of fossil trees and charcoal, and age structure of tree populations are used as evidence of tree-line dynamics over the last 400 yr. As inferred from the reproductive strategies of the two tree species, black spruce Picea mariana (Mill.) BSP, and Tamarack Larix laricina (DuRoi) K. Koch, forming the present tree-line populations, major changes during postglacial time in Northern Quebec are not only concerned with tree-line position, but also with tree-line composition. The present tree-line is made by larch (or tamarack) and corresponds to its natural seed regeneration limit. Black spruce probably reached its northernmost holocene latitudes during a warmer period and it then formed the tree-line. During the Little Ice Age, fires destroyed large tracks of forest and black spruce krummholz (400, and 250–200 yr ago); afterwards, tree species reestablished only in protected sites, and exposed sites were not reseeded. At some places, fires did not burn completely the forest cover, and preserved isolated trees and small groves probably became locus of reafforestation. Differential forest regeneration and tree age structure suggest that fire and climate are intimately associated in controlling tree population dynamics. Favourable and less favourable forest regeneration periods are inferred from these data and it is suggested that tree-line displacements in Northern Quebec during the last 400 yr were less important than in the Northwest Territories.     

A simulation test of the resource‐averaging hypothesis of ecotone formation

Abstract. The pattern at an ecotone may indicate the processes that created that ecotone. Such patterns may in turn affect the responses of ecotones to environmental change. The resource averaging hypothesis suggests a process for the development of tree lines that should produce patterns that are modifications of patterns in soil resources. A computer simulation model that embodies the resource averaging hypothesis is used to generate tree‐line patterns. Different spatial patterns in the variation of soil resources are represented in the model. The patterns of tree line computed by the simulation closely correspond to the patterns of soil resources that were input. These patterns are compared to patterns recorded in the field and by aerial photography. For the patterns of soil resources observed at some alpine tree lines, the model cannot produce the kinds of patterns of vegetation observed. Resource averaging alone cannot be an explanation of such tree lines.     

Tree-line changes along the Andes: implications of spatial patterns and dynamics

The Andes provide an extensive latitudinal and topographical framework for studying the factors that control the spatial patterns of forests (timberlines) and their species components expressed through the presence of tree growth forms (tree lines). Despite consistent overall similarities in landscape patterns, many processes must be unique, given the dramatic differences in species richness and biophysical constraints along the Andes. In all cases evaluated to date, morphological plasticity is a common trait of plant species that dominate at tree lines. In fact, many changes observed can be related to species-specific traits. Physiological limitations on tree growth form only explain species limits, while disturbances and cyclical climate fluctuations interact to affect many landscape patterns. Over long periods of time, tree lines provide unique habitats and perhaps opportunities for speciation. Understanding the spatial organization of tree-line dynamics is one viable research approach for evaluating the likely past fluxes and possible future changes.     

Plant species distribution across two contrasting treeline ecotones in the Spanish Pyrenees

We describe the structure of two contrasting (elevation, topography,climate, vegetation, soil) alpine forest–pasture ecotones located in theCentral Pyrenees (sites Ordesa, O, and Tessó, T). We define ecotonestructure as the spatial distribution of trees of different size classes andgrowth-forms and the relationship between these aspects and the spatialdistribution of understory vegetation and substrate. The studied ecotones aredominated by Pinus uncinata Ram. and have been littleaffected by anthropogenic disturbances (logging, grazing) during this century.One rectangular plot (30 × 140 m) was located within eachsite with its longest side parallel to the slope and encompassing treeline andtimberline. The distribution of size and growth-form classes at site O followeda clear sequence of increasing size downslope from shrubby multistemmedkrummholz individuals to bigger arborescent trees. At site O, regeneration wasconcentrated near the krummholz area and over rocky substrates. This suggeststhat krummholz may modify microenvironment conditions and increase seedlingsurvival. At site T, regeneration was abundant above the treeline where thecover of the dominant understory shrub (Rhododendronferrugineum) decreased. In both ecotones the diversity of plants washigher above the treeline than in the forest and decreased going downslopecoinciding with the increase of P. uncinata cover. Thereduction of plant diversity appeared above the current timberline. At site O,the decrease was steep and spatially heterogeneous what may be due in part tothe edaphic heterogeneity. At site T the change was abrupt though smaller. Therelationships between the plant community and tree regeneration should be takeninto account in future ecological studies of treeline pattern.     

Current regeneration patterns at the tree line in the Pyrenees indicate similar recruitment processes irrespective of the past disturbance regime

Aim Impacts of global change, such as land‐use and climate changes, could produce significant alterations in the elevational patterns of alpine tree line ecotones and their adjacent vegetation zones. Because the responses of the tree line to environmental variations are directly related to successful tree regeneration, understanding recruitment dynamics is an indispensable step in tree line research. We aimed to compare potential ecological limitations on recent tree line regeneration in undisturbed and disturbed sites by analysing the demographic structure and spatiotemporal patterns of recruits and large trees. Location Alpine tree line ecotones comprising Pinus uncinata in the Catalan Pyrenees (north‐east Spain) and Andorra. Methods We assessed the demographic structure and spatial pattern of recent recruitment using techniques of point‐pattern and autocorrelation analyses. A total of 3639 P. uncinata individuals were mapped, measured and aged at 12 sites. To evaluate the effects of past disturbances on recent tree line response we compared tree lines that had either been recently affected by human‐induced disturbances or had remained undisturbed for many years. Results The age structure of the tree lines, together with the lack of an age gap between seedlings and saplings, did not indicate recent episodes of high seedling mortality and suggest that recruitment has been frequent under current climate conditions. Seedlings appeared highly aggregated at short distances (up to 3 m), irrespective of disturbance history, and were spatially segregated with respect to large trees. However, we found no evidence of patches of even‐aged seedlings, and our results suggest that dispersal events at intermediate distances (10–17 m) may be frequent. Autocorrelation analyses revealed different patterns of density and age of recruits between disturbed and undisturbed tree lines, but the strength and small‐scale clustering of seedlings and saplings were very similar between sites. Main conclusions We found no recruitment limitation on recent tree line dynamics in the Pyrenees. Furthermore, processes affecting tree recruitment seem to be similar among populations regardless of their past disturbance regime. Our results suggest that constraints on tree line dynamics causing differential responses between sites may operate on older life stages and not upon recruits, and that such constraints may be more contingent on local site conditions than on disturbance history.     

Relating hybrid advantage and genome replacement in unisexual salamanders

Unisexual vertebrates are model systems for understanding the evolution of sex. Many predominantly clonal lineages allow occasional genetic recombination, which may be sufficient to avoid the accumulation of deleterious mutations and parasites. Introgression of paternal DNA into an all-female lineage represents a one-way flow of genetic material. Over many generations, this could result in complete replacement of the unisexual genomes by those of the donor species. The process of genome replacement may be counteracted by contemporary dispersal or by positive selection on hybrid nuclear genomes in ecotones. I present a conceptual model that relates nuclear genome replacement, positive selection on hybrids and biogeography in unisexual systems. I execute an individual-based simulation of the fate of hybrid genotypes in contact with a single host species. I parameterize these models for unisexual salamanders in the Ambystoma genus, for which the frequency of genome replacement has been a source of ongoing debate. I find that, if genome replacement occurs at a rate greater than 1/10,000 in Ambystoma, then there must be compensating positive selection in order to maintain observed levels of hybrid nuclei. Future researchers studying unisexual systems may use this framework as a guide to evaluating the hybrid superiority hypothesis.     

Problems with edges: tree lines as indicators of climate change (or not)

Elevational tree lines are commonly seen as particularly important ecological ‘ecotones’ that should, according to models, respond sensitively to climate change. In this issue, Mathisen et al. report empirical analyses of two tree lines that fail to show expected upward movement. This study points up both methodological and conceptual challenges in understanding ecological edges and in connecting general expectations to specific cases.     

Ecohydrological gradients and their restoration on the periphery of extracted peatlands

The moss layer transfer technique is effective at restoring extracted peatland surfaces. However, remnant peatlands persist on the periphery of extracted surfaces. These remnant peatlands drop steeply to extracted surfaces, producing artificial ecotones that are more challenging to restore. We asked to what degree natural ecotones at undisturbed reference fens can act as models for the restoration of artificial ecotones around an extracted peatland, and whether management actions can ameliorate conditions in artificial ecotones. We compared changes in elevation, water table, peat, and multiple vegetation characteristics between natural ecotones and unmanaged artificial ecotones. We then clear‐cut peripheral strips, completely filled perimeter canals, and smoothed peripheral slopes around sections of the extracted surfaces to assess whether hydrological conditions improved. Without management, artificial ecotones are not good models of natural ecotones. The elevation gradient is steep, and water tables drop steeply within 8 m of blocked perimeter canals, with possible effects at 25 m. The consequent vegetation had denser tree saplings, faster tree growth, almost no moss cover, and low moss species richness. After these management actions, water tables increased to within approximately 5 cm of those along natural ecotones. Future study is required to assess the extent of vegetation recovery, but these results hold promise for a more holistic rehabilitation of ecotones on the periphery of extracted peatland surfaces. We present recommendations to optimize the management actions on the periphery of extracted peatlands.     

Functional responses in the habitat selection of a generalist mega‐herbivore,the African savannah elephant

Resource selection function (RSF) models are commonly used to quantify species/habitat associations and predict species occurrence on the landscape. However, these models are sensitive to changes in resource availability and can result in a functional response to resource abundance, where preferences change as a function of availability. For generalist species, which utilize a wide range of habitats and resources, quantifying habitat selection is particularly challenging. Spatial and temporal changes in resource abundance can result in changes in selection preference affecting the robustness of habitat selection models. We examined selection preference across a wide range of ecological conditions for a generalist mega‐herbivore, the African savanna elephant Loxodonta africana, to quantify general patterns in selection and to illustrate the importance of functional responses in elephant habitat selection. We found a functional response in habitat selection across both space and time for tree cover, with tree cover being unimportant to habitat selection in the mesic, eastern populations during the wet season. A temporal functional response for water was also evident, with greater variability in selection during the wet season. Selection for low slopes, high tree cover, and far distance from people was consistent across populations; however, variability in selection coefficients changed as a function of the abundance of a given resource within the home range. This variability of selection coefficients could be used to improve confidence estimations for inferences drawn from habitat selection models. Quantifying functional responses in habitat selection is one way to better predict how wildlife will respond to an ever‐changing environment, and they provide promising insights into the habitat selection of generalist species.     

Little change in the fir tree-line position on the southeastern Tibetan Plateau after 200 years of warming

? As one of the world's highest natural tree lines, the Smith fir (Abies georgei var. smithii) tree line on the southeastern Tibetan Plateau is expected to vary as a function of climate warming. However, the spatial patterns and dynamics of the Smith fir tree line are not yet well understood. ? Three rectangular plots (30 m × 150 m) were established in the natural alpine tree-line ecotone on two north-facing (Plot N1, 4390 m asl; Plot N2, 4380 m asl) and one east-facing (Plot E1, 4360 m asl) slope. Dendroecological methods were used to monitor the tree-line patterns and dynamics over a 50-yr interval. ? The three study plots showed a similar pattern of regeneration dynamics, characterized by increased recruitment after the 1950s and an abrupt increase in the 1970s. Smith fir recruitment was significantly positively correlated with both summer and winter temperatures. However, Smith fir tree lines do not show a significant upward movement, despite warming on the Tibetan Plateau. ? The warming in the past 200 yr is already having a significant impact on the population density of the trees, but not on the position of the Smith fir tree line.     

Simulating natural selection in landscape genetics

Linking landscape effects to key evolutionary processes through individual organism movement and natural selection is essential to provide a foundation for evolutionary landscape genetics. Of particular importance is determining how spatially-explicit, individual-based models differ from classic population genetics and evolutionary ecology models based on ideal panmictic populations in an allopatric setting in their predictions of population structure and frequency of fixation of adaptive alleles. We explore initial applications of a spatially-explicit, individual-based evolutionary landscape genetics program that incorporates all factors--mutation, gene flow, genetic drift and selection--that affect the frequency of an allele in a population. We incorporate natural selection by imposing differential survival rates defined by local relative fitness values on a landscape. Selection coefficients thus can vary not only for genotypes, but also in space as functions of local environmental variability. This simulator enables coupling of gene flow (governed by resistance surfaces), with natural selection (governed by selection surfaces). We validate the individual-based simulations under Wright-Fisher assumptions. We show that under isolation-by-distance processes, there are deviations in the rate of change and equilibrium values of allele frequency. The program provides a valuable tool (cdpop v1.0; http://cel.dbs.umt.edu/software/CDPOP/) for the study of evolutionary landscape genetics that allows explicit evaluation of the interactions between gene flow and selection in complex landscapes.     

Feeding modes in stream salmonid population models: is drift feeding the whole story?

Drift-feeding models are essential components of broader models that link stream habitat to salmonid populations and community dynamics. But is an additional feeding mode needed for understanding and predicting salmonid population responses to streamflow and other environmental factors? We addressed this question by applying two versions of the individual-based model inSTREAM to a field experiment in which streamflow was varied in experimental units that each contained a stream pool and the adjacent upstream riffle. The two model versions differed only in the feeding options available to fish. Both versions of inSTREAM included drift feeding; one also included a search feeding mode to represent feeding in which food availability is largely independent of streamflow, such as feeding from the benthos, or feeding from the water column or the water’s surface in low water velocities. We compared the abilities of the two model versions to fit the observed distributions of growth by individual rainbow trout (Oncorhynchus mykiss) in the field experiment. The version giving fish the daily choice between drift or search feeding better fit observations than the version in which fish fed only on drift. Values for drift and search food availability from calibration to the individual mass changes of fish in experimental units with unaltered streamflow yielded realistic distributions of individual growth when applied to experimental units in which streamflow was reduced by 80 %. These results correspond with empirical studies that show search feeding can be an important alternative to drift feeding for salmonids in some settings, and indicate that relatively simple formulations of both processes in individual-based population models can be useful in predicting the effects of environmental alterations on fish populations.     

Regional and local effects of disturbance and climate on altitudinal treelines in northern Patagonia

Abstract. In this field study we analysed the regional and local scale effects of disturbance and climate on altitudinal treelines dominated by Nothofagus pumilio in northern Patagonia. We compared two regions west and east of the Andes at 40° S, slopes with warm vs cool aspects and undisturbed vs locally disturbed treelines. This spatial framework allowed us to test (1) for differences among treelines affected by different types of local disturbance and (2) the traditional hypothesis that low temperature limits treeline. Contingency tables and ANOVA showed that local disturbance occurred more frequently than expected on slopes with cool aspects, steep slope angles and concave slope configuration. Disturbed treelines were locally lowered with longer ecotones and lower krummholz growth rates and vegetation cover than undisturbed treelines. Three‐way ANOVA showed the significant influences of study area (regional climate) and aspect (local climate) on treeline elevation, krummholz growth rates and density, tree density and vegetation cover, while accounting for local disturbance. Treeline elevations were higher east of the Andes reflecting the more continental climate in Argentina than in Chile, plus regional impacts of volcanic eruptions. Tree density and vegetation cover were greater west of the Andes reflecting greater precipitation in Chile. Within study areas, local climate had different influences on treeline elevations and krummholz growth rates west and east of the Andes. We predict that increased tree growth and upslope advance of treeline in response to global warming is more likely in Chile than in Argentina near 40° S, unless precipitation increases east of the Andes. To test these predictions, we recommend research be stratified to account for the influences of local disturbance, which may confound climatic impacts. In northern Patagonia, suitable control (undisturbed) study sites will most likely be found at upper slope positions with low slope angles, simple microtopography and straight topographic configuration.     

Temperate tree expansion into adjacent boreal forest patches facilitated by warmer temperatures

Temperate and boreal forests are forecast to change in composition and shift spatially in response to climate change. Local‐scale expansions and contractions are most likely observable near species range limits, and as trees are long‐lived, initial shifts are likely to be detected in the understory regeneration layers. We examined understory relative abundance patterns of naturally regenerated temperate and boreal tree species in two size classes, seedlings and saplings, and across two spatial scales, local stand‐scale ecotones (tens of meters) and the regional temperate–boreal transition zone (?250 km) in central North America, to explore indications of climate‐mediated shifts in regeneration performance. We also tested for the presence of strong environmental gradients across local ecotones that might inhibit species expansion. Results showed that tree regeneration patterns across ecotones varied by species and size class, and varied across the regional summer temperature gradient. Temperate tree species regeneration has established across local ecotones into boreal forest patches and this process was facilitated by warmer temperatures. Conversely, boreal conifer regeneration exhibited negative responses to the regional temperature gradient and only displayed high abundance at the boreal end of local ecotones at cool northern sites. The filtering effects of temperature also increased with individual size for both boreal and temperate understory stems. Observed regeneration patterns and the minor environmental gradients measured across local ecotones failed to support the idea that there were strong barriers to potential temperate tree expansion into boreal forest patches. Detectable responses, consistently in the directions predicted for both temperate and boreal species, indicate that summer temperature is likely an important driver of natural tree regeneration in forests across the temperate–boreal transition zone. Regeneration patterns point toward temperate expansion and reduced but continued boreal presence in the near‐future, resulting in local and regional expansions of mixed temperate‐boreal forests.     

Tall shrub and tree expansion in Siberian tundra ecotones since the 1960s

Circumpolar expansion of tall shrubs and trees into Arctic tundra is widely thought to be occurring as a result of recent climate warming, but little quantitative evidence exists for northern Siberia, which encompasses the world's largest forest‐tundra ecotonal belt. We quantified changes in tall shrub and tree canopy cover in 11, widely distributed Siberian ecotonal landscapes by comparing very high‐resolution photography from the Cold War‐era ‘Gambit’ and ‘Corona’ satellite surveillance systems (1965–1969) with modern imagery. We also analyzed within‐landscape patterns of vegetation change to evaluate the susceptibility of different landscape components to tall shrub and tree increase. The total cover of tall shrubs and trees increased in nine of 11 ecotones. In northwest Siberia, alder (Alnus) shrubland cover increased 5.3–25.9% in five ecotones. In Taymyr and Yakutia, larch (Larix) cover increased 3.0–6.7% within three ecotones, but declined 16.8% at a fourth ecotone due to thaw of ice‐rich permafrost. In Chukotka, the total cover of alder and dwarf pine (Pinus) increased 6.1% within one ecotone and was little changed at a second ecotone. Within most landscapes, shrub and tree increase was linked to specific geomorphic settings, especially those with active disturbance regimes such as permafrost patterned‐ground, floodplains, and colluvial hillslopes. Mean summer temperatures increased at most ecotones since the mid‐1960s, but rates of shrub and tree canopy cover expansion were not strongly correlated with temperature trends and were better correlated with mean annual precipitation. We conclude that shrub and tree cover is increasing in tundra ecotones across most of northern Siberia, but rates of increase vary widely regionally and at the landscape scale. Our results indicate that extensive changes can occur within decades in moist, shrub‐dominated ecotones, as in northwest Siberia, while changes are likely to occur much more slowly in the highly continental, larch‐dominated ecotones of central and eastern Siberia.     

Using GIS to model tree population parameters in the Rocky Mountain National Park forest–tundra ecotone

Climatic change may alter vegetation composition and structure, but the response to climatic change can be expected to be spatially heterogeneous. Tree populations in the alpine forest–tundra ecotone, for example, may find only certain locations to be favourable for regeneration and growth. If monitoring and detection of vegetation responses to climatic change is to be most successful, the monitoring system must be tuned to the locations where a response is most likely. We used the grass geographical information system ( gis ) to map population parameters indicating potential change throughout the forest–tundra ecotone (FTE) of Rocky Mountain National Park (RMNP). Seedling density in patch forest and krummholz openings, as well as annual krummholz height growth, were measured in the field. These parameters were then modelled over the heterogeneity of the FTE environment, using principle components regression analysis. The grass gis was used to extrapolate the resulting predictive equations to the entire RMNP FTE. Potential FTE responses to climate change were evaluated in the context of species-specific differences in how tree seedling density and krummholz height growth are associated with the present environment. For example, climate change leading towards moister conditions, causing currently xeric environments to become more mesic, might increase the spatial extent of existing tree invasion into patch forest openings. This would increase the potential for widespread conversion of patch forest to closed forest. Present population parameters extrapolated spatially may provide a useful guide to where future change is likely.     

Drought limitation on tree growth at the Northern Hemisphere’s highest tree line

The alpine tree line is generally assumed to be controlled by low temperatures, and thus to be experiencing an upward shift under global warming. As global temperatures rise, tree growth at the tree line could either increase if temperature is the limiting factor or decrease if a warming-induced loss of moisture limits growth. Here, we use dendrochronological techniques to understand the abiotic drivers of the Northern Hemisphere’s highest tree line ecotones on the southern Tibetan Plateau (TP). Ring-width measurements from three juniper sites between 4680 and 4900 m asl were significantly and negatively correlated with May-June-July evapotranspiration (ET₀), and positively correlated with relative humidity and other moisture-related meteorological variables. At the same time, ring widths were negatively correlated with temperature means and sunshine rates. Our results highlight the common sensitivity of tree growth to moisture variations despite the differential growth trends occurring since 1850 (end of the Industrial Revolution) at the three tree line ecotones. These findings indicate that low temperatures may not be the sole driving force behind tree growth and the range dynamics of alpine tree lines. Tree lines in the dry parts of the TP and possibly also beyond are likely to retreat rather than to advance in a warmer world due to water limitations.