Using space‐for‐time substitution and time sequence approaches in invasion ecology

1. Invasion biologists use two main approaches to evaluate the effects of non‐native species (NNS) on diversity of native species (DNS), namely space‐for‐time and time approaches. These approaches have pitfalls related to lack of controls: the former lacks pre‐invasion data, while the latter often lacks data from non‐invaded sites. 2. We propose a framework that combines space‐for‐time and time approaches and which should result in more focused mechanistic hypotheses and experiments to test the causes of invasibility and the effects of NNS on DNS. We illustrate the usefulness of our framework using two case studies: one with the submersed macrophyte, Hydrilla verticillata, in reservoir and the other with the fish, Geophagus proximus, in a large river–floodplain system. 3. Hydrilla verticillata invaded sites with DNS similar to that found in non‐invaded sites, indicating that biotic and/or abiotic factors did not influence invasion success; however, DNS increased over time in invaded sites compared with non‐invaded sites, suggesting that H. verticillata facilitated natives. In contrast, G. proximus invaded sites with higher DNS than non‐invaded sites, suggesting that biotic and/or abiotic factors favouring natives were important for invasion success, but DNS increased in invaded and non‐invaded sites over time, indicating that an independent factor contributed to DNS increases. 4. Conclusions from both studies would have been inaccurate or incomplete if the space‐for‐time and time approaches had not been used in combination as proposed in our framework.     

Butterfly phenology in Mediterranean mountains using space‐for‐time substitution

Inferring species' responses to climate change in the absence of long‐term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14‐day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23‐day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8‐day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space‐for‐time substitutions, understanding species‐specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.     

Effects of Top‐Soil Preparation and Broad‐Leaved Tree Mixture on Carabid Beetles in Afforested Fallow Plots

A 2‐year study to evaluate the effect of four soil preparation techniques and four species compositions in a pine plantation with an admixture of broad‐leaved trees on the species diversity and structure of ground beetle assemblages (Carabidae) was conducted. The soil preparation techniques were full shallow agricultural plowing (25 cm deep with three variants—plowing, plowing with ridge formation, and plowing with subsoiling) and full deep plowing (50 cm deep). The plantation species compositions were pine 7000/hectare, pine 5500/hectare + broad‐leaved species 1500/hectare, pine 4000/hectare + broad‐leaved species 3000/hectare, and pine 2000/hectare + broad‐leaved species 5000/hectare. Carabidae were sampled monthly with pitfall traps in the plantation from May to November in 2003 and 2004—the fourth and fifth years after afforestation. Only the soil preparation technique affected species diversity. The mean number of carabid species and the Shannon–Wiener index were higher in the case of the full agriculture plowing with subsoiling treatment and the full agriculture plowing with ridge formation than in the full shallow agriculture plowing. The structure of the assemblages was dependent on the soil preparation technique, but not on tree species composition. In general, deep plowing and shallow plowing with subsoiling were conducive to an increase in the proportion of individuals representing late succession species, whereas shallow plowing favored early succession species. The results suggest that for effective plantation of a managed stand, deep plowing or shallow plowing with subsoiling is more beneficial for afforestation of fallow fields than typical shallow agricultural plowing.     

Increasing representation of localized dung beetles across a chronosequence of regenerating vegetation and natural dune forest in South Africa

Aim Species assemblages with high proportions of localized taxa occur in regional islands with a history of strong eco‐climatic separation from adjacent systems. Current disturbance in such islands of relictualism or endemism disrupts the distinctive local character in favour of regionally distributed taxa with a wider range of tolerances. However, rehabilitation of the system should restore the localized biota. Thus, we used biogeographical composition to assess progress towards restoration of the dung beetle fauna associated with such an island of endemism following dredge‐mining. Location The study was conducted in natural coastal dune forest and a 23‐year chronosequence of regenerating dune vegetation in the Maputaland centre of endemism, KwaZulu‐Natal, South Africa. Methods Dung beetles were trapped in eight stands of regenerating vegetation of different ages (< 1 year to ~21 years) and in four stands of natural dune forest with differing ecological characteristics defined by measurements of vegetative physiognomy and microclimate. Species groups defined from multivariate analysis of biogeographical distribution patterns and vegetation associations were used to demonstrate quantitative compositional changes in the dung beetle assemblages across the chronosequence to natural forest. Results Three biogeographical groups were defined. One group comprised species widespread in southern Africa or both southern and east Africa. The other two groups were endemic, one to the east coast and the other to Maputaland. There was a general trend from dominance by regionally distributed dung beetle taxa to dominance by locally distributed taxa across the chronosequence of regenerating vegetation from grassland, to open Acacia karroo thicket, to dense A. karroo‐dominated woodland. However, this trend was linked closely to the relative physiognomic and microclimatic similarity between the regenerating vegetation and natural forest. Thus, proportions of locally distributed taxa were lower in older chronosequence woodland (~18–~21 years) with its low canopy cover and open understorey than in dense early chronosequence woodland (~9–~12 years), which is physiognomically and microclimatically closer to species‐diverse natural forest with its dense canopy and understorey. Overall, the present dung beetle community comprises five species groups. Single widespread (21 spp.) and endemic groups (14 spp.) showed similar patterns of association with early chronosequence grassland and open thicket stands. A single widespread (3 spp.) and two endemic shade‐associated groups (3 and 11 spp.) showed differing patterns of association centred, respectively, in late chronosequence woodland, natural forest, or all shaded stands. Main conclusions At 23 years, vegetative regeneration is still at an early stage, but abundant activity of most, although not all species recorded in natural forest, is recovered with the closure of the woodland canopy at ~9 years. Compositional differences with respect to natural forest vary closely with vegetative physiognomy and its effect on the microclimate. Therefore, full compositional recovery is dependent on the re‐establishment of natural forest physiognomy and microclimate.     

A phylogenetic and trait‐based analysis of community assembly in a subtropical forest in central China

Despite several decades of study in community ecology, the relative importance of the ecological processes that determine species co‐occurrence across spatial scales remains uncertain. Some of this uncertainty may be reduced by studying the scale dependency of community assembly in the light of environmental variation. Phylogenetic information and functional trait information are often used to provide potentially valuable insights into the drivers of community assembly. Here, we combined phylogenetic and trait‐based tests to gain insights into community processes at four spatial scales in a large stem‐mapped subtropical forest dynamics plot in central China. We found that all of the six leaf economic traits measured in this study had weak, but significant, phylogenetic signal. Nonrandom phylogenetic and trait‐based patterns associated with topographic variables indicate that deterministic processes tend to dominate community assembly in this plot. Specifically, we found that, on average, co‐occurring species were more phylogenetically and functionally similar than expected throughout the plot at most spatial scales and assemblages of less similar than expected species could only be found on finer spatial scales. In sum, our results suggest that the trait‐based effects on community assembly change with spatial scale in a predictable manner and the association of these patterns with topographic variables, indicates the importance of deterministic processes in community assembly relatively to random processes.     

南亚热带混交人工林树种丰富度与土壤微生物多样性和群落组成的关系

森林植被与土壤微生物作为森林生态系统的重要组成部分,它们之间的相互作用对维持森林生态系统功能和稳定性起着重要作用。以往多在天然草地和森林生态系统开展植物多样性与土壤微生物多样性关系的研究,但人工构建的多树种混交林生态系统中树种多样性对土壤微生物群落组成的影响及其机制尚不完全清楚。因此,以南亚热带人工块状造林后自然恢复形成的多树种混交森林生态系统为研究对象,利用高通量测序技术研究了随树种丰富度(1—10种)变化土壤细菌和真菌多样性的变化规律及主要影响因子。结果表明,随树种丰富度增加,土壤真菌α多样性显著提高,但土壤细菌α多样性差异不显著;不同树种丰富度梯度间土壤细菌和真菌的群落结构组成均差异显著;Pearson相关分析表明土壤细菌α多样性主要受土壤pH和土壤铵态氮影响,而土壤pH和有效磷是土壤真菌α多样性的主要影响因子。距离冗余分析(db-RDA)表明,对土壤细菌群落组成产生显著影响的环境因子分别为土壤pH、硝态氮和芳香碳组分,而土壤有机碳、硝态氮、细根生物量和氧烷基碳组分是影响土壤真菌群落组成的主要因子。本研究的结果说明了南亚热带人工林不同树种混交后形成多树种混交林生态系统的过程中,树...     

Quantifying climate–growth relationships at the stand level in a mature mixed‐species conifer forest

A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year‐to‐year variability in growth. Numerous dendrochronological (tree‐ring) studies have identified climate factors that influence year‐to‐year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand‐level (as opposed to species‐level) growth. We argue that stand‐level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed‐species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand‐level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year‐to‐year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand‐level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species‐ and canopy‐position level. Our climate models were better fit to stand‐level biomass increment than to species‐level or canopy‐position summaries. The relative growth responses (i.e., percent change) predicted from the most influential climate variables indicate stand‐level growth varies less from to year‐to‐year than species‐level or canopy‐position growth responses. By assessing stand‐level growth response to climate, we provide an alternative perspective on climate–growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate.     

Long‐term seed bank dynamics in a temperate forest under conversion from coppice‐with‐standards to high forest management

Questions: How do changes in forest management, i.e. in disturbance type and frequency, influence species diversity, abundance and composition of the seed bank? How does the relationship between seed bank and vegetation change? What are the implications for seed bank dynamics? Location: An ancient Quercus petraea — Carpinus betulus forest in conversion from coppice‐with‐standards to regular Quercus high forest near Montargis, France. Methods: Seed bank and vegetation were sampled in six replicated stand types, forming a chronosequence along the conversion pathway. The stand types represented mid‐successional stages of stands in transition from coppice‐with‐standards (to high forest (16 plots) and early‐ and mid‐successional high forest stands (32 plots). Results: Seed bank density and species richness decreased with time since last disturbance. Adjusting for seed density effects obscured species richness differences between stand types, but species of later seres were nested subsets of earlier seres, implying concomitant shifts in species richness and composition with time since disturbance. Later seres were characterized by species with low seed weight and high seed longevity. Seed banks of early seres were more similar to vegetation than to later seres. Conclusions: Abandonment of the coppice‐with‐standards regime altered the seed bank characteristics, as well as its relationship with vegetation. Longer management cycles under high forest yield impoverished seed banks. For their persistence, seed bank species will increasingly rely on management of permanently open areas in the forest landscape. Thus, revegetation at the beginning of new high‐forest cycles may increasingly depend on inflow from seed sources.     

Consequences of multiple mating‐system shifts for population and range‐wide genetic structure in a coastal dune plant

Evolutionary transitions from outcrossing to selfing can strongly affect the genetic diversity and structure of species at multiple spatial scales. We investigated the genetic consequences of mating‐system shifts in the North American, Pacific coast dune endemic plant Camissoniopsis cheiranthifolia (Onagraceae) by assaying variation at 13 nuclear (n) and six chloroplast (cp) microsatellite (SSR) loci for 38 populations across the species range. As predicted from the expected reduction in effective population size (N_e) caused by selfing, small‐flowered, predominantly selfing (SF) populations had much lower nSSR diversity (but not cpSSR) than large‐flowered, predominantly outcrossing (LF) populations. The reduction in nSSR diversity was greater than expected from the effects of selfing on N_e alone, but could not be accounted for by indirect effects of selfing on population density. Although selfing should reduce gene flow, SF populations were not more genetically differentiated than LF populations. We detected five clusters of nSSR genotypes and three groups of cpSSR haplotypes across the species range consisting of parapatric groups of populations that usually (but not always) differed in mating system, suggesting that selfing may often initiate ecogeographic isolation. However, lineage‐wide genetic variation was not lower for selfing clusters, failing to support the hypothesis that selection for reproductive assurance spurred the evolution of selfing in this species. Within three populations where LF and SF plants coexist, we detected genetic differentiation among diverged floral phenotypes suggesting that reproductive isolation (probably postzygotic) may help maintain the striking mating‐system differentiation observed across the range of this species.