Dynamic and static views of succession: Testing the descriptive power of the chronosequence approach

Chronosequence and permanent plot studies are the two most common methods for evaluating successional dynamics in plant communities. We combined these two approaches by re-sampling an old-field chronosequence at Cedar Creek Natural History Area (Minnesota, USA) to: (1) measure rates of secondary succession; and (2) to test the ability of the chronosequence approach to predict actual successional dynamics over a 14-year survey interval. For each of 19 chronosequence fields we calculated four complimentary indices of succession rate for community changes that actually occurred within each of these fields between 1983 and 1997. We found that measures of compositional dissimilarity, species turnover, and the change rates of perennial and native species cover over this 14-year period were all negatively correlated with field age, indicating that the rate of successional change in these old-fields generally declines over time. We also found that data collected from the initial static chronosequence survey (1983) accurately predicted many of the observed changes in species abundance that occurred between 1983 and 1997, but was a poor predictor of changes in species richness. In general, chronosequence re-sampling confirmed the validity of using the chronosequence approach to infer basic patterns of successional change.     

Recovery of Amphibian and Reptile Assemblages During Old‐Field Succession of Tropical Rain Forests

Conversion of tropical forests to agriculture affects vertebrate assemblages, but we do not know how fast or to what extent these assemblages recover after field abandonment. We addressed this question by examining amphibians and reptiles in secondary forests in southeastern Mexico. We used chronosequence data (12 secondary forests fallow for 1–23 yr and 3 old‐growth forest sites) to analyze successional trajectories and estimate recovery times of assemblage attributes for amphibians and reptiles. We conducted 6 surveys at each site over 14 mo (1200 person‐hours) and recorded 1552 individuals, including 25 species of amphibians and 36 of reptiles, representing 96 and 74 percent of the expected regional number of species, respectively. Abundance, species richness, and species diversity of amphibians increased rapidly with successional age, approaching old‐growth forest values in < 30 yr. Species richness and species diversity of reptiles reached old‐growth forest values in < 20 yr. By contrast, the abundance of reptiles and the assemblage composition of amphibians and reptiles recovered more slowly. Along the chronosequence, we observed more species replacement in reptile assemblages than in amphibian assemblages. Several species in the old‐growth forest were absent from secondary forests. Dispersal limitation and harsh conditions prevailing in open sites and early successional environments appear to preclude colonization by old‐growth forest species. Furthermore, short fallow periods and isolation of forest remnants lead to the formation of new assemblages dominated by species favored by human disturbances.     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

Litter dynamics recover faster than arthropod biodiversity during tropical forest succession

Litterfall and litter decomposition are key elements of nutrient cycling in tropical forests, a process in which decomposer communities such as macro-arthropods play a critical role. Understanding the rate and extent to which ecosystem function and biodiversity recover during succession is useful to managing the growing area of tropical successional forest globally. Using a replicated chronosequence of forest succession (5–15, 15–30, 30–45 years, and primary forest) on abandoned pastures in lowland tropical wet forest, we examined litterfall, litter chemistry, and effects of macro-arthropod exclusion on decomposition of two litter types (primary and 5- to 15-years-old secondary forest). Further, we assessed macro-arthropod diversity and community composition across the chronosequence. Overstory cover, litterfall, and litter nutrients reached levels similar to primary forest within 15–30 years. Young secondary forest litter (5–15 years) had lower initial N and P content, higher C:N, and decayed 60 percent faster than primary forest litter. The presence of macro-arthropods strongly mediated decomposition and nutrient release rates, increasing litter mass loss by 35–44 percent, N released by 53 percent, and P release by 84 percent. Forest age had no effect on soil nutrients, rates of litter decomposition, nutrient release, or macro-arthropod influence. In contrast, abundance and community composition of macro-arthropods remained significantly lower and distinct in all ages of secondary compared with primary forest. Order richness was lower in 5–15 years of secondary compared with primary forest. Our results suggest that in highly productive tropical wet forest, functional recovery of litter dynamics precedes recovery of decomposer community structure and biodiversity.     

Dispersal mode,shade tolerance,and phytogeographical affinity of tree species during secondary succession in tropical montane cloud forest

Secondary succession following land abandonment, represented by a chronosequence of 15 old fields (0–80 years old) and two old-growth forests, was studied in the tropical montane cloud forest region of Veracruz, Mexico. The objective was to determine successional trajectories in forest structure and species richness of trees ≥5 cm DBH, in terms of differences in seed dispersal mode, shade tolerance, and phytogeographical affinity. Data were analyzed using AIC model selection and logistic regressions. Mean and maximum canopy height reached values similar to old-growth forest at 35 and 80 years, respectively. Species richness and diversity values were reached earlier (15 and 25 years, respectively) while basal area and stem density tended to reach old-growth forest values within 80 years. Along the chronosequence, the proportion of species and individuals of wind-dispersed trees declined, that of bird dispersed small seeded trees remained constant, while that of gravity and animal dispersed large seeded trees increased; shade-intolerant species and individuals declined, while intermediate and shade-tolerant trees increased. Shade-tolerant canopy trees were rare during succession, even in the old-growth forest. Tropical tree species were more frequent than temperate ones throughout the chronosequence, but temperate tree individuals became canopy dominants at intermediate and old-growth forest stages.     

Patterns and Correlates of Tropical Dry Forest Structure and Composition in a Highly Replicated Chronosequence in Yucatan,Mexico

Research on tropical dry forest (TDF) succession i0s needed for effective conservation and management of this threatened and understudied ecosystem. We used a highly replicated chronosequence within a 37,242‐ha TDF landscape to investigate successional patterns by plant size class and to evaluate the influence of stand age, topographic position, soil properties and spatial autocorrelation on forest structure and composition. We used a SPOT5 satellite image to obtain a land‐cover thematic map, and sampled woody vegetation (adults: >5 cm diam; saplings: 1–5 cm) and soil properties in 168 plots distributed among four vegetation classes: VC1 (3–8‐yr‐old forest), VC2 (9–15‐yr‐old forest), VC3 (>15‐yr‐old forest on flat areas), VC4 (>15‐yr‐old forest on hills). Stem density decreased with stand age and was lowest in VC3, while height, basal area and species density increased with age and were higher in older than in younger forests. Topographic position also influenced forest structure and composition. Basal area and height were largely determined by stand age, whereas stem and species density, and composition were influenced mostly by soil variables associated with fertility, and by spatial autocorrelation. Adults and saplings showed contrasting patterns and correlates of community structure, but similar patterns and correlates of composition, possibly due to the prevalence of coppicing. Our results show that our sampling approach can overcome several limitations of chronosequence studies, and provide insights in the patterns and drivers of succession, as well as guidelines for forest management and conservation. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Removal of cattle accelerates tropical dry forest succession in Northwestern Mexico

Domestic livestock influence patterns of secondary succession across forest ecosystems. However, the effects of cattle on the regeneration of tropical dry forests (TDF) in Mexico are poorly understood, largely because it is difficult to locate forests that are not grazed by cattle or other livestock. We describe changes in forest composition and structure along a successional chronosequence of TDF stands with and without cattle (chronic grazing or exclusion from grazing for ~ 8 year). Forest stands were grouped into five successional stages, ranging from recently abandoned to mature forest, for a total of 2.7 ha of the sampled area. The absence of cattle increased woody plant (tree and shrub) density and species richness, particularly in mid-successional and mature forest stands. Species diversity and evenness were generally greater in sites where cattle were removed and cattle grazing in early successional stands reduced establishment and/or recruitment of new individuals and species. Removal of cattle from forest stands undergoing succession appears to facilitate a progressive and non-linear change of forest structure and compositional attributes associated with rapid recovery, while cattle browsing acts as a chronic disturbance factor that compromises the resilience and structural and functional integrity of the TDF in northwestern Mexico. These results are important for the conservation, management, and restoration of Neotropical dry forests.     

Rapid Liana Colonization along a Secondary Forest Chronosequence

Lianas (woody vines) can have profound effects on tree recruitment, growth, survival, and diversity in tropical forests. However, the dynamics of liana colonization soon after land abandonment are poorly understood, and thus it is unknown whether lianas alter tree regeneration early in succession. We examined the liana community in 43 forests that ranged from 1 to 31 yr old in central Panama to determine how fast lianas colonize young forests and how the liana community changes with forest succession. We found that lianas reached high densities early in succession, commonly exceeding 1000 stems/ha within the first 5 yr of forest regeneration. Lianas also increased rapidly during early succession in terms of basal area but did not show evidence of saturation within the 30 yr of our chronosequence. The relative contribution of lianas to total woody plant community in terms of basal area and density increased rapidly and reached a saturation point within 5 yr (basal area) to 15 yr (density) after land abandonment. Our data demonstrate that lianas recruit early and in high density in tropical forest regeneration, and thus lianas may have a large effect on the way in which secondary forests develop both early and throughout succession.     

Vegetation Structure,Composition, and Species Richness Across a 56‐year Chronosequence of Dry Tropical Forest on Providencia Island,Colombia1

We compared vegetation structure and species richness across a 56‐yr chronosequence of six replicated age classes of dry tropical forest on the island of Providencia, Colombia, in the Southwest Caribbean. Stand age classes were determined using sequential, orthorectified panchromatic aerial photos acquired between 1944 and 1996 and Landsat 7 ETM + satellite imagery from 2000. Along the chronosequence we established 59 plots of 2 × 50 m (0.01 ha) to document changes in species richness, basal area, tree height, stem density, and sprouting ability. All woody trees and shrubs >2.5 cm diameter at breast height (DBH) were censused and measured. Although woody species density reached a peak in stands from 32 to 56 yr old, rarefaction analysis showed that species richness increased linearly with stand age and was highest in stands 56 yr old or greater. Nonparametric, abundance‐based estimators of species richness also showed positive and linear associations with age. Basal area and mean tree height were positively associated with age since abandonment, while sprouting ability showed a negative relationship. Our results indicate rapid recovery of woody species richness and structural characteristics along this tropical dry forest chronosequence.     

Gall-inducing insect species richness as indicators of forest age and health

The changes in the plant community that occur during the process of succession affect the availability of resources for the community of herbivores. In this study, the richness of galling insects was evaluated in restored stands of Amazonian tropical rain forest of several ages (0-21 yr), as well as in areas of primary forest in Brazil. The richness of gallers increased with the age of the restored stands. Fifty-eight percent of the variation in the richness of galling insects was explained by forest stand age, but an increase in richness was observed at intermediate stages of succession. The greatest similarity among groups was found between the initial successional stages and intermediate ones. The results indicate a recovery of both host plants and insect community and that succession directly affects the richness and composition of these herbivores.     

Life History Traits of Lianas During Tropical Forest Succession

Tropical secondary forests form an important part of the landscape. Understanding functional traits of species that colonize at different points in succession can provide insight into community assembly. Although studies on functional traits during forest succession have focused on trees, lianas (woody vines) also contribute strongly to forest biomass, species richness, and dynamics. We examined life history traits of lianas in a forest chronosequence in Costa Rica to determine which traits vary consistently over succession. We conducted 0.1 ha vegetation inventories in 30 sites. To examine the establishment of young individuals, we only included small lianas (0.5–1.5 cm diameter at 1.3 m height). For each species, we identified seed size, dispersal mode, climbing mode, and whether or not the seedling is self‐supporting. We found a strong axis of variation determined by seed size and seedling growth habit, with early successional communities dominated by small‐seeded species with abiotic dispersal and climbing seedlings, while large‐seeded, animal‐dispersed species with free‐standing seedlings increased in abundance with stand age. Contrary to previous research and theory, we found a decrease in the abundance of stem twiners and no decrease in the abundance of tendril‐climbers during succession. Seed size appears to be a better indicator of liana successional stage than climbing mode. Liana life history traits change predictably over succession, particularly traits related to seedling establishment. Identifying whether these trait differences persist into the growth strategies of mature lianas is an important research goal, with potential ramifications for understanding the impact of lianas during tropical forest succession.     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.     

Posthurricane Seedling Structure in a Multi‐aged Tropical Dry Forest: Implications for Community Succession

Hurricane‐caused tree mortality in tropical dry forests occurs predominantly in early successional species. Consequently, hurricanes may accelerate succession in these forests. Forest regeneration, however, must be measured over an extended posthurricane time period to demonstrate this pattern. In this study, we recorded tree seedlings in 19 Florida Keys forests during May–August 1995, 3 years after Hurricane Andrew. For these forests—spanning a chronosequence from 14 to over 100 years since the most recent clearing—we used weighted averaging regression on relative abundances of pre‐hurricane trees to calculate a successional age optimum for each species; and used weighted averaging calibration to calculate inferred successional ages for stands based on pre‐hurricane trees and on posthurricane seedlings. To test the hypothesis that successional stage of seedlings exceeded successional stage of pre‐hurricane trees, we compared inferred stand ages based on posthurricane seedlings with those based on pre‐hurricane trees. Across the study area, inferred stand ages based on posthurricane seedlings were greater than those based on pre‐hurricane trees (P < 0.005); however, more seedlings in the youngest stands were early successional than in older stands. Of 29 species present both as pre‐hurricane trees and posthurricane seedlings, 23 had animal‐dispersed seeds. These results provide evidence that: (1) hurricanes do not ‘reset’ succession, and may accelerate succession; and (2) a strong legacy of stand successional age influences seedling assemblages in these forests.     

Successional dynamics of the bee community in a tropical dry forest: Insights from taxonomy and functional ecology

Despite the recent rapid growth of tropical dry forest succession ecology, most studies on this topic have focused on plant community attribute recovery, whereas animal community successional dynamics has been largely overlooked, and the few existing studies have used taxonomic approaches. Here, we analyze the successional changes in the bee community in a Mexican tropical dry forest, by integrating taxonomic (species, genus, and family diversity) and functional (sociability, nesting strategy, and body size) information for bees. Over one year, in a successional chronosequence (2–67 years after abandonment) we collected 469 individual bees, representing five families, 36 genera, and 69 species. Linear modeling showed decreases in taxonomic diversity with succession, more strongly so for species. Bee species turnover along succession ranged from moderate to high, decreasing slightly at intermediate stages. An RLQ analysis (ordination method that allows relating environmental variables with functional attributes) revealed clear relations between bee functional traits and the plant community. RLQ axis 1 was positively related to vegetation structural and diversity variables, and to eusociality, while solitary, parasociality, and ground nesting was negatively associated with it. Early successional fallows attract mostly solitary and parasocial bees; older fallows tend to attract eusocial bees with aerial nesting. The continuous taxonomic turnover observed by us and the functional analysis suggest that the disappearance of old fallows from agricultural landscapes would likely result in significant reductions and even local extinctions of particular bee guilds. Considering the low viability of preserving large mature tropical dry forest tracts, the conservation of older successional stands emerges as a crucial component of landscape management.Abstract in Spanish is available with online material.     

Changes in Mediterranean plant succession: old-fields revisited

Abstract. Old-field plots used for a study of succession in Mediterranean France were revisited after 12–14 yr. Our aims were: (1) to verify if predicted patterns of species richness, turnover and composition are confirmed; (2) to compare the development in disturbed plots with that in undisturbed ones; (3) to discuss the impact of management changes. In undisturbed plots species richness and turnover decreased with successional age. Floristic composition changed in a way consistent with the predicted successional development in most plots. Therophytes decreased and phanerophytes increased; anemochorous species decreased and endozoochorous species increased, as expected. In plots disturbed since the first analysis richness decreased with successional age, but generally remained higher than in undisturbed plots. Floristic composition, species turnover and an increase in therophytes indicated changes towards younger successional stages. Thus, disturbance changed succession but not much. This is probably linked with the regeneration abilities typical of mediterranean species, e.g. resprouting. At the landscape scale, richness did not change and species turnover was low. The plots studied were situated in two distinct locations. One had not been disturbed between the two observation periods, while the other is a mosaic of undisturbed and disturbed sites. Observations fitted predictions much more closely at the undisturbed location. We conclude that permanent plot studies are powerful in identifying successional trends and can also provide additional insights into the effects of disturbance some of the mechanisms underlying the dynamics of diversity.     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Vegetation succession on reclaimed coal wastes in Spain: the influence of soil and environmental factors

Question: How is vegetation succession on coal mine wastes under a Mediterranean climate affected by the restoration method used (topsoil addition or not)? How are plant successional processes influenced by local landscape and soil factors? Location: Reclaimed coal mines in the north of Palencia province, northern Spain (42°47′‐42°50′ N, 4°32′‐4°53′ W). Methods: In Jun–Jul 2008, vascular plant species cover was monitored in 31 coal mines. The mines, which had been restored using two restoration methods (topsoil addition or not), comprised a chronosequence of different ages from 1 to 40 yr since restoration started. Soil and environmental factors at each mine were monitored and related to species cover using a combination of ordination methods and Huisman–Olff–Fresco modeling. Results: Plant succession was affected by restoration method . Where topsoil was added, succession was influenced by age since restoration and soil pH. Where no topsoil was added, soil factors seem to arrest succession. Vegetation composition on topsoiled sites showed a gradient with age, from the youngest, with early colonizing species, to oldest, with an increase in woody species. Vegetation on non‐topsoiled sites comprised mainly early‐successional species. Response to age and pH of 37 species found on topsoiled mines is described. Conclusions: Restoration of coal mines under this Mediterranean climate can be relatively fast if topsoil is added, with a native shrub community developing after 15 yr. However, if topsoil is not used, it takes more than 40 yr. For topsoiled mines, the species found in the different successional stages were identified, and their tolerance to soil pH was derived. This information will assist future restoration projects in the area.     

Pathways, mechanisms and predictability of vegetation change during tropical dry forest succession

The development of forest succession theory has been based on studies in temperate and tropical wet forests. As rates and pathways of succession vary with the environment, advances in successional theory and study approaches are challenged by controversies derived from such variation and by the scarcity of studies in other ecosystems. During five years, we studied development pathways and dynamics in a chronosequence spanning from very early to late successional stages (ca. 1–60 years) in a tropical dry forest of Mexico. We (1) contrasted dynamic pathways of change in structure, diversity, and species composition with static, chronosequence-based trends, (2) examined how structure and successional dynamics of guilds of trees shape community change, and (3) assessed the predictability of succession in this system. Forest diversity and structure increased with time but tree density stabilized early in succession. Dynamic pathways matched chronosequence trends. Succession consisted of two tree-dominated phases characterized by the development and dynamics of a pioneer and a mature forest species guild, respectively. Pioneer species dominated early recruitment (until ca. 10 years after abandonment), and declined before slower growing mature-forest species became dominant or reached maximum development rates (after 40–45 years). Pioneers promoted their replacement early in succession, while mature-forest species recruited and grew constantly throughout the process, with their lowest mortality coinciding with the peak of pioneer abundance. In contrast to prevailing stochastic views, we observed an orderly, community driven series of changes in this dry forest secondary succession. Chronosequences thus represent a valuable approach for revealing system-specific successional pathways, formulating hypotheses on causes and mechanisms and, in combination with repeated sampling, evaluating the effects of vegetation dynamics in pathway variation.