Environmental influences on the distribution of the rare Eucalyptus paliformis and the common E. fraxinoides

Abstract Eucalyptus paliformis is restricted to seven small populations within an area of <5 km2 on the Wadbilliga Plateau of south-eastern NSW. It occurs as a dominant tree in habitats superficially similar to those of the more widespread E. fraxinoides, which dominates forests immediately adjacent to the E. paliformis populations. This study examined factors which may influence the replacement of E. fraxinoides by E. paliformis, and similarly, factors which exclude E. paliformis from sites dominated by E. fraxinoides. Seedling growth and competition experiments indicated that E. fraxinoides outcompetes E. paliformis under unburnt field conditions, on field soils, under high and low moisture regimes, under high light and high nutrient conditions, and under a range of temperature regimes. Thus, it is likely that E. paliformis is excluded from E. fraxinoides sites by competition. However, further investigation of the competitive relationship between the two species under recently-burnt conditions is still required to test this hypothesis. The exclusion of E. fraxinoides from E. paliformis sites is unlikely to be due to soil types or moisture regimes. Further, temperature regimes do not appear to be important in reducing the competitive advantage of E. fraxinoides over E. paliformis, leaving survival at low temperature extremes as the most promising avenue for further investigation. Temperature measurements, and evidence for low temperature dormancy mechanisms in seeds of E. paliformis but not E. fraxinoides, support the hypothesis that E. fraxinoides is excluded from E. paliformis sites by low temperature extremes.     

Alpine and subalpine snow patch vegetation on the Bogong High Plains,SE Australia

Abstract. Snow patch vegetation in Australia is rare, being restricted to the relatively small area of alpine and subalpine country in the highlands of southeastern Australia. Snow patch vegetation occurs on steeper, sheltered southeastern slopes, where snow persists until well into the growing season (December/January). We surveyed the vegetation of 33 snow patch sites in the alpine and subalpine tracts of the Bogong High Plains, within the Alpine National Park, in Victoria. The vegetation was dominated by herbs and graminoids, with few shrubs and mosses. Major structural assemblages identified included closed herb‐fields dominated by Celmisia spp, and grasslands dominated by Poa fawcettiae or Poa costiniana. These assemblages occurred on mineral soils. Open herb‐fields dominated by Caltha introloba and several sedge species occurred on rocky and stony substrata. Vegetation‐environment relationships were explored by ordination and vector fitting. There was significant variation in the floristic composition of snow patch vegetation as a function of duration of snow cover, altitude, slope and site rockiness. Alpine sites were floristically distinct from subalpine sites, with a greater cover of Celmisia spp. and a lesser cover of low shrubs in the former. There was floristic variation within some snow patches as a function of slope position (upper, middle or lower slope) but this was not consistent across sites. The current condition of snow patch vegetation on the Bogong High Plains is degraded, with bare ground exceeding 20% cover at most sites. Snow patch vegetation is utilized preferentially by domestic cattle, which graze parts of the Bogong High Plains in summer. Such grazing is a potential threat to this rare vegetation type.     

The abundance of ground-dwelling invertebrates in a Victorian forest affected by ‘dieback’(Phytophthora cinnamomi) disease

Abstract Phytophthora cinnamomi (cinnamon fungus) is a pathogenic soil fungus that infects plant communities along the southeastern coast of Australia and the southwestern corner of Western Australia. Infection of native plant communities with P. cinnamomi regularly leads to dramatic changes in both the structural and the floristic characteristics of these communities. This study aimed to assess the effect of P. cinnamomi induced changes in plant community attributes upon the abundance and diversity of invertebrates in an area of open forest in the Brisbane Ranges, Victoria. Pitfall trapping was conducted continuously for 1 week within each season over a 3 year period. Invertebrates were sorted to order level, and abundances were compared between times (season and year) and infection status for taxa with normalized distributions (Coleoptera, Collembola, Dermaptera and ants). Non-parametric comparisons were made for other groups (Aranae, Blattodea, Diptera, Hemiptera, Orthoptera, ant ‘morpho-species’ and unidentified larvae) to assess differences on the basis of infection status. Significant differences on the basis of infection were uncommon and, where identified, elevated abundances were more commonly observed at sites infected by P. cinnamomi. Consistent temporal effects (season and year) were observed in normalized data sets. Abundances both within individual taxa and from pooled counts were generally weakly associated with ground-level habitat features. Overall, the impact of P. cinnamomi on vegetative structure and floristics was not reflected in different abundances of ground-dwelling invertebrates.     

A phylogenetic and allozyme approach to understanding rarity in three “green ash” eucalypts (Myrtaceae)

Phylogenetic relationships among the 12 species of the green ash group of eucalypts were examined using allozyme data, to investigate the causes of rarity in three localized endemics of the group. The relationships suggested by the allozyme data showed both similarities to and differences from those estimated from morphological data byLadiges and coworkers. The phylogenetic relationships suggest that rarity inEucalyptus burgessiana may be related to recent divergence, whileE. paliformis andE. rupicola are relatively old species, more likely to be relicts, and/or restricted to a rare habitat.     

Effects of fire on the structure and composition of open eucalypt forests

Abstract Fires are integral to the healthy functioning of most ecosystems and are often poorly understood in policy and management, however, the relationship between floristic composition and habitat structure is intrinsically linked, particularly after fire. The aim of this study was to test whether the variability of habitat structure or floristic composition and abundance in forests at a regional scale can be explained in terms of fire frequency using historical data and experimental prescribed burns. We tested this hypothesis in open eucalypt forests of Fraser Island off the east coast of Australia. Fraser Island dunes show progressive stages in plant succession as access to nutrients decreases across the Island. We found that fire frequency was not a good predictor of floristic composition or abundance across dune systems; rather, its affects were dune specific. In contrast, habitat structure was strongly influenced by fire frequency, independent of dune system. A dense understorey occurred in frequently burnt areas, whereas infrequently burnt areas had a more even distribution of plant heights. Plant communities returned to pre‐burn levels of composition and abundances within 6 months of a fire and frequently burnt areas were dominated by early successional species of plant. These ecosystems were characterized by low diversity and frequently burnt areas on the east coast were dominated by Pteridium. Greater midstorey canopy cover in low frequency areas reduces light penetration and allows other species to compete more effectively with Pteridium. Our results strongly indicate that frequent fires on the Island have resulted in a decrease in relative diversity through dominance of several species. Prescribed fire represents a powerful management tool to shape habitat structure and complexity of Fraser Island forests.     

Towards an explanation of the causes of the rarity of two Tasmanian Spyridium species

Spyridium microphyllum (F. Muell. ex Reissek) Druce and Spyridium obcordatum (Hook. f) W. M. Curtis are narrow endemic shrubs which are confined to small parts of Tasmania, Australia, and are rare within their present ranges. Analysis of quadrat data from transects through populations of both species showed that at most individual sites the rare taxa occur in floristically and environmentally distinct situations. However, sites supporting the same species differ floristically and environmentally, with the only communality being a lack of dense tall shrub or tree cover. Both species are absent from many areas with closely similar physical environments and vascular plant species composition to those in which they occur. Thus, their current distributions cannot be explained by limitation to a rare habitat. Spyridium microphyllum appears to be susceptible to local extinction because it is an obligate seeder, is killed by fire, does not survive in taller, dense vegetation, and is not adapted for dispersal over the distances between its potential habitats. While S. obcordatum recovers vegetatively after fire, it also does not survive in dense taller vegetation, has limited dispersal ability and highly disjunct potential habitat. Both species may have been more widespread and competitive during cooler and drier times.     

The impact of invasive cane toads on native wildlife in southern Australia

Commonly, invaders have different impacts in different places. The spread of cane toads (Rhinella marina: Bufonidae) has been devastating for native fauna in tropical Australia, but the toads' impact remains unstudied in temperate‐zone Australia. We surveyed habitat characteristics and fauna in campgrounds along the central eastern coast of Australia, in eight sites that have been colonized by cane toads and another eight that have not. The presence of cane toads was associated with lower faunal abundance and species richness, and a difference in species composition. Populations of three species of large lizards (land mullets Bellatorias major, eastern water dragons Intellagama lesueurii, and lace monitors Varanus varius) and a snake (red‐bellied blacksnake Pseudechis porphyriacus) were lower (by 84 to 100%) in areas with toads. The scarcity of scavenging lace monitors in toad‐invaded areas translated into a 52% decrease in rates of carrion removal (based on camera traps at bait stations) and an increase (by 61%) in numbers of brush turkeys (Alectura lathami). The invasion of cane toads through temperate‐zone Australia appears to have reduced populations of at least four anurophagous predators, facilitated other taxa, and decreased rates of scavenging. Our data identify a paradox: The impacts of cane toads are at least as devastating in southern Australia as in the tropics, yet we know far more about toad invasion in the sparsely populated wilderness areas of tropical Australia than in the densely populated southeastern seaboard.     

The large‐scale drivers of population declines in a long‐distance migratory shorebird

Migratory species can travel tens of thousands of kilometers each year, spending different parts of their annual cycle in geographically distinct locations. Understanding the drivers of population change is vital for conserving migratory species, yet the challenge of collecting data over entire geographic ranges has hindered attempts to identify the processes leading to observed population changes. Here, we use remotely sensed environmental data and bird count data to investigate the factors driving variability in abundance in two subspecies of a long‐distance migratory shorebird, the bar‐tailed godwit Limosa lapponica. We compiled a spatially and temporally explicit dataset of three environmental variables to identify the conditions experienced by each subspecies in each stage of their annual cycle (breeding, non‐breeding and staging). We used a Bayesian N‐mixture model to analyze 18 years of monthly count data from 21 sites across Australia and New Zealand in relation to the remote sensing data. We found that the abundance of one subspecies L. l. menzbieri in their non‐breeding range was related to climate conditions in breeding grounds, and detected sustained population declines between 1995 and 2012 in both subspecies (L. l. menzbieri, –6.7% and L. l. baueri, –2.1% year^–1). To investigate the possible causes of the declines, we quantified changes in habitat extent at 22 migratory staging sites in the Yellow Sea, East Asia, over a 25‐year period and found –1.7% and –1.2% year^–1 loss of habitat at staging sites used by L. l. menzbieri and L. l baueri, respectively. Our results highlight the need to identify environmental and anthropogenic drivers of population change across all stages of migration to allow the formulation of effective conservation strategies across entire migratory ranges.     

Spatial ecology of bluetongue lizards (Tiliqua spp.) in the Australian wet–dry tropics

New technologies for quantifying animal locations enable us to document habitat‐selection patterns of cryptic taxa in extraordinary detail. Northern bluetongues (Tiliqua scincoides intermedia) and centralian bluetongues (Tiliqua multifasciata) are large heavy‐bodied scincid lizards that are broadly sympatric in the wet–dry tropics of north‐western Australia. We used data from GPS‐based radiotelemetry (n = 49 lizards, tracked for 2–121 days, total n = 61 640 locations) to examine the size, internal structure and overlap of lizard home ranges. Despite substantial habitat differences at our two study sites (semi‐arid and relatively pristine habitat at Keep River National Park, Northern Territory, vs. highly disturbed and fragmented flood plain habitat in an agricultural area near Kununurra, Western Australia), home ranges were similar between the two areas, and between the two species. Our radio‐tracked lizards continued to disperse into previously unused areas throughout the duration of the study, so that the total areas used by lizards continued to increase. Based on the minimum convex polygon method, total home ranges averaged 4 ha (range 2–12 ha), but only about two‐thirds of each home range was used intensively. Each home range had multiple core areas, and overlap of core as well as peripheral areas (especially with same‐sex conspecifics) was high at the disturbed (Western Australia) site where lizard densities were high. The concentration of lizard activity within small core areas, often used by multiple individuals, suggests that these heavily used sites are critical to lizard conservation. However, the lizards' infrequent long‐distance displacements also make them vulnerable to changes in the wider landscape mosaic. Because GPS‐based radiotelemetry can quantify habitat use at finer spatial and temporal scales than earlier technologies, it can provide a robust base for management of at‐risk fauna.     

Fire severity influences the post-fire habitat structure and abundance of a cool climate lizard

In the spring and summer of 2019–2020, the ‘Black Summer’ bushfires burned more than 97 000 km² of predominantly Eucalyptus dominated forest habitat in eastern Australia. The Black Summer bushfires prompted great concern that many species had been imperilled by the fires. Here, we investigate the effects that fire severity had on the habitat and abundance of a cool climate lizard Eulamprus tympanum that was identified as a species of concern because 37% of its habitat was burnt in the Black Summer bushfires. We quantified habitat structure and the abundance of E. tympanum at sites which were unburnt, burnt at low severity and at high severity 10, 15 and 23 months after the fires. Our classification of fire severity based on scorch height and canopy status corresponded well with the Australian Government Google Earth Engine Burnt Area Map (AUS GEEBAM) fire severity layer. Ten months after the fires, sites burnt at high severity had less canopy cover, more bare ground and less fine fuel than sites burnt at low severity or unburnt sites. The abundance of E. tympanum varied with survey occasion and was greatest during the warmest sampling period and lowest during the coolest sampling period. The abundance of E. tympanum was consistently lower on sites burnt at high severity than sites burnt at low severity or unburnt sites. Our findings show that higher severity fires had a greater effect on E. tympanum than low severity fires. Our results suggest that E. tympanum were likely to have persisted in burnt sites, with populations in low severity and unburnt sites facilitating population recovery in areas burnt at high severity. Our results also suggest that wildfire impacts on E. tympanum populations will increase because the frequency and extent of severe fires are expected to increase due to climate change.     

Habitat preference in the critically endangered yellow‐tailed woolly monkey (Lagothrix flavicauda) at La Esperanza,Peru

Habitat loss is one of the main threats to wildlife. Therefore, knowledge of habitat use and preference is essential for the design of conservation strategies and identification of priority sites for the protection of endangered species. The yellow‐tailed woolly monkey (Lagothrix flavicauda Humboldt, 1812), categorized as Critically Endangered on the IUCN Red List, is endemic to montane forests in northern Peru where its habitat is greatly threatened. We assessed how habitat use and preference in L. flavicauda are linked to forest structure and composition. The study took place near La Esperanza, in the Amazonas region, Peru. Our objective was to identify characteristics of habitat most utilized by L. flavicauda to provide information that will be useful for the selection of priority sites for conservation measures. Using presence records collected from May 2013 to February 2014 for one group of L. flavicauda, we classified the study site into three different use zones: low‐use, medium‐use, and high‐use. We assessed forest structure and composition for all use zones using 0.1 ha Gentry vegetation transects. Results show high levels of variation in plant species composition across the three use zones. Plants used as food resources had considerably greater density, dominance, and ecological importance in high‐use zones. High‐use zones presented similar structure to medium‐ and low‐use zones; thus it remains difficult to assess the influence of forest structure on habitat preference. We recommend focusing conservation efforts on areas with a similar floristic composition to the high‐use zones recorded in this study and suggest utilizing key alimentation species for reforestation efforts.     

Concentrated Distribution of Wild Yam Patches: Historical Ecology and the Subsistence of African Rainforest Hunter-Gatherers

Availability of annually reproducing wild yam species (Dioscorea praehensilis and Dioscorea semperflorens) is considered to be a key factor for the viability of forest hunter-gatherer subsistence in the African rainforests. I here describe the habitat and distribution of wild yam patches, and discuss possible human impacts on the formation of wild yam patches, which currently support long-term forest expeditions by Baka hunter-gatherers in southeastern Cameroon. An analysis of canopy photographs shows that annual yams favor a habitat of canopy gaps which receive sufficient sunlight. However, the fact that annual yams patches are limited to specific locations even though gaps in the canopy occur throughout the forest, combined with the mostly vegetative reproductive system of the yams, suggests annual yams do not propagate extensively in the wild. Moreover, areas with abundant annual yam patches that are presently situated far from any villages contain several abandoned village sites. Since the Baka often transplant parts of yam tubers to encourage their spread, it seems that human activities may have contributed to the formation of locally concentrated annual yam patches.     

Environmental and vegetation variables have a greater influence than habitat fragmentation in structuring lizard communities in remnant urban bushland

Abstract The expansion of urban areas and adjacent farming land into natural landscapes modifies habitats and produces small isolated pockets of native vegetation. This fragmentation of the natural habitat subdivides animal communities, reduces population sizes and increases vulnerability to extinction. In this paper we investigate whether fragmentation decreases lizard species richness, composition, overall abundance and abundance at the species level. Urban remnants consisting of five small (< 10 ha) and four large (> 10 ha) fragments of natural bushland were paired with continuous bushland areas located near Hobart, Tasmania, Australia. These remnants were surveyed six times, using pitfall traps, from November 2001 to March 2002. Lizard species richness and abundance were not significantly influenced by habitat fragmentation or fragment size. Egernia whitii was the only lizard species significantly influenced by fragment size, and was only present in large fragments and continuous bush. Vegetation type and structure as well as environmental variables (geology and aspect) influenced the structure of reptile communities. Lizard species that were able to use a number of different habitat types were found to persist at most sites, irrespective of fragment size. Edge environment did not significantly influence lizard species richness or abundance in remnant areas. Lizard species richness was significantly lower in sites that had a high ratio of exotic to native plant species. Therefore, if remnants continue to be invaded by exotic plants, lizard species that require native plant communities will become increasingly vulnerable to local extinction. Our results suggest that lizard species requiring specialized habitats, such as E. whitii, may persist in large urban remnants rather than small urban remnants because large reserves are more likely to encompass rare habitats, such as rocky outcrops. Habitat heterogeneity, rather than size, may be the key to their persistence.     

Ocean currents influence the genetic structure of an intertidal mollusc in southeastern Australia – implications for predicting the movement of passive dispersers across a marine biogeographic barrier

Major disjunctions among marine communities in southeastern Australia have been well documented, although explanations for biogeographic structuring remain uncertain. Converging ocean currents, environmental gradients, and habitat discontinuities have been hypothesized as likely drivers of structuring in many species, although the extent to which species are affected appears largely dependent on specific life histories and ecologies. Understanding these relationships is critical to the management of native and invasive species, and the preservation of evolutionary processes that shape biodiversity in this region. In this study we test the direct influence of ocean currents on the genetic structure of a passive disperser across a major biogeographic barrier. Donax deltoides (Veneroida: Donacidae) is an intertidal, soft‐sediment mollusc and an ideal surrogate for testing this relationship, given its lack of habitat constraints in this region, and its immense dispersal potential driven by year‐long spawning and long‐lived planktonic larvae. We assessed allele frequencies at 10 polymorphic microsatellite loci across 11 sample locations spanning the barrier region and identified genetic structure consistent with the major ocean currents of southeastern Australia. Analysis of mitochondrial DNA sequence data indicated no evidence of genetic structuring, but signatures of a species range expansion corresponding with historical inundations of the Bassian Isthmus. Our results indicate that ocean currents are likely to be the most influential factor affecting the genetic structure of D. deltoides and a likely physical barrier for passive dispersing marine fauna generally in southeastern Australia.     

Spatio‐temporal gradients in food supply help explain the short‐term colonisation dynamics of the critically endangered central rock‐rat (Zyzomys pedunculatus)

Currently, the impact of introduced predators on small mammal population decline is a focal research direction in the Australian desert literature. In all likelihood though, single‐factor explanation of population dynamics is inadequate, leaving gaps in our knowledge of the multitude of potential influences on small mammal abundance and occupancy patterns in time and space. Here, we investigated floristic gradients across four potential refuge sites of the central rock‐rat, Zyzomys pedunculatus, a granivore rodent (50–120 g) that is endemic to central Australia and is categorised as critically endangered. The study took place in Tjoritja/West MacDonnell National Park in the MacDonnell Ranges bioregion. Floristic sampling was allocated across the four sites, the locations of which were predetermined by an established monitoring and management programme for the central rock‐rat. Our aim was to examine the relationship between environmental gradients and floristic composition across the four sites, and thereby test the extent to which the patterns of food type and food availability can inform central rock‐rat spatio‐temporal dynamics. We found high site‐scale floristic patterning that related foremost to elevation and then to antecedent rainfall and time‐since‐fire and fire‐severity effects. To interpret these results, we applied the principles of refuge theory and we described a gradient from core refuge habitat to intermittent and then marginal habitat within the current central rock‐rat stronghold area. Overall, our results implied a strong floristic basis to central rock‐rat site occurrence, and they thus compel us to take explicit account of spatial (elevation) and temporal (rainfall–productivity and fire‐disturbance) influences on the food axis of potential refuge sites of this critically endangered species.     

Early response of the platypus to climate warming

Combining a climatic envelope modelling technique with more than two centuries (1800–2009) of distribution records has revealed the effects of a changing climate on the egg‐laying monotreme, the platypus, Ornithorhynchus anatinus. We show that the main factor associated with platypus occurrence switched from aquatic habitat availability (estimated by rainfall) to thermal tolerances (estimated by annual maximum temperature) in the 1960s. This correlates directly with the change in the annual maximum temperature anomaly from cooler to warmer conditions in southeastern Australia. Modelling of platypus habitat under emission scenarios (A1B, A2, B1 and B2) revealed large decreases (>30%) in thermally suitable habitat by 2070. This reduction, compounded by increasing demands for water for agriculture and potable use, suggests that there is real cause for concern over the future status of this species, and highlights the need for restoration of thermal refugia within the platypus’ modelled range.     

Phylogenetic systematics of the gondwanan Empis macrorrhyncha group (Diptera,Empididae, Empidinae)

The Empis macrorrhyncha group (Diptera: Empididae) from cool to warm temperate areas of South America and Australia is diagnosed and cladistically analysed, and five new species, Empis animosa sp.n. , E. austera sp.n. , E. maculosa sp.n. , E. occidentalis sp.n. and E. pedivillosula sp.n. , are described. Cladistic analysis of 23 adult morphological characters for 14 species of the group generated a single tree of 28 steps (CI = 0.82; RI = 0.93). Monophyly was established on the basis of a single apomorphy, possession of a bilobed cercus of the male hypopygium. Three main clades were inferred: clade 1 included three Patagonian and a single southwestern Australian species; clade 2 included two species from southeastern Australia; clade 3 included a large Patagonian group of five species and a single southeastern Australian species. The E. fulvicollis complex (clade 1) is a sister‐group of the E. macrorrhyncha complex (clades 2 + 3). A provisional historical biogeographic hypothesis is advanced correlating the appearance of the South American and Australian sister lineages with the timing of the break‐up of Gondwana.     

Ecological Restoration and Large-Scale Ecological Disturbance: The Effects of Drought on the Response by Fish to a Habitat Restoration Experiment

Human‐induced erosion regularly delivers massive quantities of fine sediments into streams and rivers forming large static bodies of sediment known as sand slugs, which smother in‐stream habitat, alter community structure, and decrease biodiversity. Sand slugs are widespread in parts of southeastern Australia as well as in many other parts of the world, and there is now considerable interest in restoring such affected streams. The reintroduction of large timber is widely suggested as a strategy for restoring habitat complexity, but this has rarely been tested in sand slug–affected streams. We examined the response of fish populations to wood addition to two streams in southeastern Australia that have been impacted by sand slugs. Manipulated sites (three per treatment) had either one or four timber structures added, and these sites were compared with (three) unmanipulated (control) sites before and after the manipulation occurred. Despite a supraseasonal drought during the study, we observed short‐term increases in the abundance of Mountain galaxias (Galaxias olidus) at the four‐structure sites, while both the four‐structure and the one‐structure treatments appeared to buffer against drought‐induced declines in two other species, River blackfish (Gadopsis marmoratus) and Southern pygmy perch (Nannoperca australis), relative to controls. However, drought eventually caused the complete loss of surface water from these streams and the loss of fish from both manipulated and unmanipulated sites. Thus, although the study supports the use of timber structures as a means of increasing local fish abundances, these beneficial effects were, in these streams, contingent upon permanency of flow. Because sedimentation has depleted the number of permanent refuge pools in these creeks, recovery rates of the fauna (i.e., resilience) are likely to be slow. We therefore conclude that in streams subjected to frequent disturbance, restoring refugia may be as, if not more, important as restoring what we term residential habitat.     

Early herbaceous succession along a topographical gradient on forest clear-felling sites in mountainous terrain,central Japan

Early successional patterns of herbaceous communities in forest clear-felling sites were investigated along a topographical gradient, which included ridge, slope and valley habitat types, in warm-temperate evergreen forest regions of central Japan for 5 years. Three dominant species with wind-dispersed seeds played a major role in the succession: an annual,Crassocephalum crepidioides, a biennial,Erigeron canadensis, and a perennial,Miscanthus sinensis. Pioneer herbs that have a seed-bank strategy, which are common in old field succession, were not found in the mountainous sites. The persistence of the annual or biennial dominants during the very early stages of secondary succession was different in the three topographical habitat types. In the ridge habitat,M. sinensis dominated from the first year state, andC. crepidioides andE. canadensis were less prominent.Crassocephalum crepidioides andE. canadensis became dominant as one moved down the slope. In the valley habitat,C. crepidioides dominated in the first year stage, was succeeded byE. canadensis in the second year, and thenM. sinensis gradually replaced it in later years. As all three wind-dispersed dominants simultaneously invaded in all the habitat types after clear-felling, the different successional patterns along the topographical gradient might have resulted from differences in the establishment ability and the growth rate of the three dominants depending on the three habitat types.