Comparison of Southern Appalachian high-elevation outcrop plant communities with their Northern Appalachian counterparts

Abstract. Southern Appalachian high-elevation outcrops harbour six regionally rare Northern Appalachian taxa usually considered relicts of a Pleistocene alpine flora. For five of the six taxa, minimum elevation in the south was 367–1113 m higher than in the north. While habitats compared between the two regions share only 9% of their total flora, individual plots had up to 70% of their species occurring in the opposite region. The northern affinity of southern outcrops increased with elevation, slope steepness, soil Cu, B and SO₄ and decreased with potential solar radiation and soil Na. As a result, communities above 1600 m on felsic bedrock, and above 1350 m on mafic bedrock, were most northern in composition. Northern affinity of southern outcrops also increased with latitude, which may partly result from closer geographic proximity to past communities that provided progenitors for the current northern flora. Northern treeless habitats increased in southern affinity with increased slope steepness, perennial seepage, vegetation height, shade, soil pH, Al, Mn, Na and decreased elevation and organic matter. As a result, northern outcrop communities below treeline were most similar to those on southern outcrops. This suggests that southern outcrop vegetation may be more similar to Pleistocene outcrop vegetation than to Pleistocene alpine vegetation. Partial constrained ordination showed that while compositional differences between the Northern and Southern Appalachian habitats were largely explained by environmental differences, there was a significant component of residual variation explained by north or south position that was unrelated to environment. These residual compositional differences may result from historical influences on community structure involving stochastic extinction and colonization processes.     

High-elevation rock outcrop vegetation of the Southern Appalachian Mountains

Abstract. Species composition patterns and vegetation-environment relationships were quantified for high-elevation rock outcrops of the Southern Appalachian Mountains, an infrequent and insular habitat in a forested landscape. Outcrops occur over a wide geographic range encompassing extensive variation in both geology and climate. Geographic-scale factors interact with site-scale factors to produce variation in vegetation among outcrops. Similarly, site-scale factors interact with micro-scale factors to produce variation in vegetation within outcrops. To provide a quantitatively-based classification of outcrop vegetation we used a TWINSPAN analysis of 154 100-m² plots. We recognized nine communities that primarily correspond to different combinations of elevation, bedrock type, geography, and moisture. Within outcrops of a single bedrock type, vegetation composition of 100-m² plots was consistently correlated with elevation and solar radiation, but relationships to soil nutrients varied with bedrock type. Both site-scale (100 m²) factors (e.g. elevation, slope, aspect, and bedrock type) and plot-scale (1-m²) microsite factors (e.g. soil depth, vegetation height, soil nutrients) were strongly correlated with species composition at the 1-m² level. Environment can be used to predict composition more effectively for 100-m² plots on a single bedrock type than either across bedrock types or at a 1-m² scale. Composition-environment relationships resemble those described for outcrop systems from other regions with pronounced topographic relief more than they do those described for the nearby but flatter and lower-elevation outcrops of the Southeastern Piedmont. There is strong spatial autocorrelation in this community, perhaps owing to dispersal limitation. Consequently, a comprehensive conservation strategy must include reservation of both a range of geologic types and a range of geographic locations.     

Structure,function and floristic relationships of plant communities in stressful habitats marginal to the Brazilian Atlantic rainforest

The Brazilian Atlantic rainforest consists of a typical tropical rainforest on mountain slopes, and stands out as a biodiversity hotspot for its high species richness and high level of species endemism. This forest is bordered by plant communities with lower species diversity, due mostly to more extreme environmental conditions than those found in the mesic rainforest. Between the mountain slopes and the sea, the coastal plains have swamp forests, dry semi-deciduous forests and open thicket vegetation on marine sand deposits. At the other extreme, on top of the mountains (>2000 m a.s.l.), the rainforest is substituted by high altitude fields and open thicket vegetation on rocky outcrops. Thus, the plant communities that are marginal to the rainforest are subjected either to flooding, drought, oceanicity or cold winter temperatures. It was found that positive interactions among plants play an important role in the structuring and functioning of a swamp forest, a coastal sandy vegetation and a cold, high altitude vegetation in the state of Rio de Janeiro. Moreover, only a few species seem to adopt this positive role and, therefore, the functioning of these entire systems may rely on them. Curiously, these nurse plants are often epiphytes in the rainforest, and at the study sites are typically terrestrial. Many exhibit crassulacean acid metabolism. Conservation initiatives must treat the Atlantic coastal vegetation as a complex rather than a rainforest alone.     

Shrub vegetation on tropical granitic inselbergs in French Guiana

Abstract. In French Guiana, inselbergs in the form of granite outcrops rise abruptly from the surrounding rain forest. They constitute isolated islands of a special type of vegetation restricted to this peculiar substrate. Shrub granitic vegetation, organized in thickets on open exposed rocks of inselbergs, are described using the Braun‐Blanquet method combined with Correspondence Analysis. This phytosociological study revealed only one particular shrub community on each inselberg, including predominantly evergreen and sclerophyllous shrubs, especially microphanerophytes, belonging to the Clusiaceae, Myrtaceae and Bombacaceae. These outcrop communities exhibit species endemic to the Guianas region and also species rare in French Guiana. Affinities with flora of other inselbergs and vegetation types in South America are examined and discussed. Reasons for observed floristic and structural changes in each community are also discussed.     

Floristic relationships of the woody component in rocky outcrops savanna areas in Central Brazil

Rocky outcrops usually have a peculiar vegetation which differs from that of the surrounding vegetation matrix. In Brazil, a savanna type (cerrado rupestre) is particularly characterized by rocky outcrops and is one of eleven vegetation types which occur throughout Cerrado biome. It has been scarcely studied in detail. Therefore, the aim of this study was to investigate the phytogeographical patterns of the woody species in ten areas of the rocky outcrop savanna in the Brazilian Central Plateau. The difference in spatial distribution of the species was assessed through indicator species and CCA ordination analyses. Ten indicator species showed significantly different distribution. The first axis of the ordination showed association with the edaphic gradient, based on maximum temperature, pH and physical properties of the soils (eigenvalue 0.63). The second axis showed an eigenvalue 0.46, associated with poor rainfall and land declivity. It appears that the woody flora of the rocky outcrop savanna areas of the Brazilian Central Plateau is relatively homogeneous, with low β diversity. Apparently, the size of the woody species populations is what distinguishes the ten investigated areas. Therefore, initiatives and actions for the Cerrado biome conservation must consider not only the size of the protected areas but also that of the populations to be protected, especially the woody species on rocky outcrops.     

The ecology of granite outcrops at Wilson's Promontory,Victoria

Rock outcrops on spurs and summits on Wilson's Promontory tend to be more prevalent on hot dry slopes than cool moist ones. In this comparatively moist climate, the rock outcrop vegetation can be grouped into high altitude, low altitude and coastal types. The surrounding rock-heath is characterized below 400–500 m by Kunzea ambigua and above this altitude by Callistemon pallidus. Moss aprons of Campylopus bicolor on rock faces are often held in place by roots of surrounding shrubs on slopes up to 20°. Bryophyte and lichen communities vary with altitude, and annuals and geophytes are common. Fire may result in retrogression if forests and shrub zones are destroyed and humic top soil burnt or eroded away. Fossil root-grooves occur in granite on the summit of Mt Latrobe and are the legacy of a cloud forest of Nothofagus cunninghamii. The rock-heath species are more drought resistant than the‘climax’tree species and may develop very low water potentials. The rate of succession may be correlated with the highly variable rate of decay of each individual block of granite. The soils of the early successional stages are veneers of humic sands enriched with hill-wash, whilst the soils of the forest stages vary from duplex podzols to gradational podzolic soils of much greater moisture storage. Woody plant establishment is mostly initiated along rock joints: the rock surfaces and boulders so by-passed are colonized by more mesic species. Many zonations appear to be static since advances are cancelled by retreats. The rate of succession on bare granite is controlled by local climate, altitude, slope, fire and macro-and micro-jointing patterns. In general, the early stages bear strong similarities over a relatively wide range of climate. The so-called climaxes which have developed represent a continuum in response to the complex environments found on Wilson's Promontory.     

The role of outcrops in the diversity of Patagonian vegetation: Relicts of glacial palaeofloras?

Outcrops around the world enhance biodiversity as they provide heterogeneous environments and varied habitats for species with different requirements than those living in the surroundings, and in this way they increase alpha and beta diversity. We studied the floristic composition of the vegetation of rock outcrops in northwestern Patagonia, Argentina, and compared them to the surrounding matrix of shrubland and steppe. For this we sampled 50 outcrops and 50 matrix plots close to them, identified all the species present, and analyzed their floristic composition with Detrended Correspondence Analysis (DCA). We also analyzed the floras of the north and south faces of the outcrops. The resulting ordinations were related to geographic gradients of the region. The proportions of alien to native species, different life forms, and phylogenetic groups of the outcrop floras were compared to the matrix with G-tests. Our results showed similar dominant families, life form proportions and percentage of alien species in the outcrops and the surrounding matrix. However, species composition of the outcrops was markedly different. North Patagonian outcrops present several taxa not found in the matrix, especially in their southern, less insolated walls. The previously recorded distribution of most of these exclusive species is found in colder areas further south. Thus, we hypothesize that outcrops could be acting as relicts of cold-adapted glacial paleofloras that were probably more extended during glacial times and retreated south after the last glacial maximum.     

Managing rock outcrops to improve biodiversity conservation in Australian agricultural landscapes

Summary  Rocky outcrops are prominent geological features in agricultural landscapes worldwide. Reptiles are a major component of these habitats and some species are restricted to, and more abundant on, rocky outcrops than in remnant vegetation. Rock outcrops are important to reptiles because they provide resources that are often limited in the surrounding landscape (e.g. micro-gradients in climatic conditions, basking- and retreat-sites). However, there is a knowledge gap in the literature addressing the conservation value of small, rocky outcrops. Management may be necessary to reverse habitat degradation in these systems. We identify four key areas of management that need to be addressed to improve outcrop habitat values and enhance biodiversity conservation in agricultural landscapes. Specific actions involve: (i) protecting outcrops from processes that cause damage to rock microhabitat, (ii) monitoring and managing changes in vegetation structure to maintain thermally suitable environments, (iii) applying integrated pest animal control and (iv) improving matrix management to enhance inselberg function and landscape connectivity. Further research is required to evaluate the efficacy of different management regimes on outcrop biota. We hope this paper will provide the stimulus for land managers to incorporate rocky outcrops in future biodiversity conservation programmes.     

Plant communities on ironstone outcrops: a diverse and endangered Brazilian ecosystem

Mountain areas are recognized centres of endemism and diversity on account of their isolation and altitudinal diversity. In tropical regions, mountain tops usually stand as islands of xeric vegetation among mesophytic assemblages. Unlike the vegetation growing on other rock outcrops lithologies, such as inselbergs (granite/gneiss) or campos rupestres (quartz/arenite), ironstone outcrop plant communities still lack systematic studies in Brazil. These outcrops (locally known as canga) share most of the characteristics of other rock outcrops, such as isolation and edapho-climatic harshness, but differ in that they are the object of opencast mining, and thus subjected to irrecoverable degradation. In addition, they are expected to harbour metal-tolerant and hyperaccumulator plant species. A botanical survey of two ironstone outcrop locations in the most important mining region of southeastern Brazil, the Iron Quadrangle, revealed a high within-site (138 and 160 species per site), and between-site diversity (only 27% of common species), totaling 64 families and 234 species among basal families and eudicots (154 species), monocots (68 species), and ferns (12 species). Canga crusts are rich in dicots, several of which play an important role in community structuring, together with the more usual monocot aggregations. Distinct plant communities are found associated to different microhabitats within the iron crust, depending primarily on the amount of soil and moisture retention in the different microtopographies. The environmental uniqueness, high diversity, lack of studies and rapid destruction of these ecosystems pose an immediate challenge for their conservation.     

Post-pastoral changes in composition and guilds in a semi-arid conservation area, Central Otago, New Zealand

Changes in the vegetation of Flat Top Hill, a highly modified conservation area in semi;arid Central Otago, New Zealand, are described four years after the cessation of sheep and rabbit grazing. Unusually moist weather conditions coincide with the four-year period of change in response to the cessation of grazing. Between 1993 and 1997, the average richness and diversity (H') of species increased, and the average proportion of native species decreased significantly. The vegetation was significantly richer in exotic annual and perennial grass species, exotic perennial forbs, exotic woody species and native tussock grasses in 1997 than in 1993. Eight response guilds of species are identified. Most "remnant" native shrubs and forbs were stable, in that they remained restricted to local refugia and showed little change in local frequency. However, taller native grass species increased, some locally, and others over wide environmental ranges. Rare native annual forbs and several native perennial species from "induced" xeric communities decreased, and this may be a consequence of competition from exotic perennial grasses in the absence of grazing. The invasive exotic herb Sedum acre decreased in abundance between 1993 and 1997, but several other prominent exotic species increased substantially in range and local frequency over a wide range of sites. Exotic woody species, and dense, sward-forming grasses are identified as potential threats to native vegetation recovery.     

Exotic plant invasion in the highly degraded upland landscape of Hong Kong, China

Plant communities in the continental tropics have suffered less from exotic plant invasions than their oceanic island counterparts. Most studies have focused on near-pristine communities. By contrast, we examine the resistance of semi-natural continental plant communities in Hong Kong, which have been suffering from chronic and massive human impacts. We compiled a list of all naturalized non-native species recorded in Hong Kong and then sampled the plant communities for exotic species along roadsides, a stream through semi-natural vegetation, and in semi-natural vegetation away from both roads and streams on Tai Mo Shan, Hong Kong’s highest peak (957 m). Similar surveys were repeated in other areas of Hong Kong. More than 162 naturalized exotic plant species have been recorded in Hong Kong. On Tai Mo Shan, 29 exotic species were recorded in roadside vegetation, with the diversity but not percentage cover declining significantly with altitude. Fifteen exotic species were found along the stream, including two not found along the roadside. Only six exotic species were found away from roads and streams, all in unshaded areas disturbed by feral cattle. In all surveys, no exotics were found in closed woody vegetation or in open areas without feral cattle, except for one species. The shade-tolerant tropical Asian tree Syzygium jambos was found invading along some streamsides without anthropogenic disturbance. Despite centuries of massive human impacts, exotic plant invasions in Hong Kong are still largely confined to habitats that suffer from chronic human disturbance. Feral cattle promote invasion where people are absent, but this problem still seems potentially reversible. Only Syzygium jambos is of possible current conservation concern.     

Distribution and morphology of columnar cacti in tropical deciduous woodland,Jalisco, Mexico

The distribution and branching morphologies of four species of columnar cactus species in tropical deciduous woodland in Jalisco, western Mexico, are examined. Two species, Stenocereus marginatus and especially Pachycereus pecten-aboriginum, are taller and extend through the woodland canopy, and only these two species occupy habitats where the woodland vegetation is taller, on N- and NE-facing aspects where incident radiation levels are lowest. Two species, Stenocereus standleyi and Cephalocereus purpusii, are shorter, subcanopy species, and are more restricted to low-angle slopes and slopes with S- to W-aspects, where the woodland vegetation is shorter and more open, with a greater penetration of radiation into the vegetation. Branching morphology is affected by the vegetation cover in all species, but in different ways between the two taller and the two shorter species. Ecological segregation in these four species is achieved apparently by differences among the light regimes of subhabitats correlated to the morphology of the cacti. This contrasts to segregation in Sonoran Desert columnar cactus species, which segregate by different strategies of water uptake and utilization, and in columnar cacti in woodlands in Oaxaca further south in Mexico, where adult individuals are very similar in branching morphology, but the younger individuals differ in this respect and presumably become established in different sorts of light gaps.Nomenclature follows Gibson & Horak (1978).     

木论喀斯特峰丛洼地森林群落空间格局及环境解释

基于广西壮族自治区木论国家级自然保护区典型峰丛洼地景观尺度内不同微生境条件和植物群落类型50个样地 (20 m × 20 m)的系统取样调查, 用二元物种指示方法(TWINSPAN)对样地内胸径(DBH) ≥ 1 cm的木本植物进行分类, 选择10个土壤环境因子和5个空间因子, 利用除趋势典范对应分析(DCCA)研究了森林群落分布的土壤环境与空间格局, 并给予定量化的合理解释。结果如下: 1) TWINSPAN将森林群落划分为11组, 在三级水平上分为4类生态群落类型。2) DCCA第一排序轴集中了排序的大部分信息, 突出反映了各森林群落所在的坡向和土壤主要养分梯度, 沿第一轴从左到右, 坡向由阴转阳, 岩石裸露率越来越高, 土壤主要养分逐渐降低, 森林群落分别出现了由原生性和耐阴性强逐步向阳性先锋树种为主的次生林和人工林变化的格局。3)因子分离分析结果表明, 土壤环境因子对森林群落分布格局的解释能力为39.16%, 其中21.02%单纯由土壤环境因子所引起, 空间因子的解释能力为31.34%, 其中13.16%独立于土壤环境的变化, 18.15%是土壤环境和空间因子相互耦合作用的结果, 不可解释部分达47.66%, 表明喀斯特峰丛洼地森林群落的物种共存受生态位分化理论和中性理论双重控制。     

山西五台山高山林线的植被景观

 过草本植物群落的分类和排序,结合对乔木和灌木分布的分析,确定了五台山高山林线的几条植被界线以及五台山森林上限附近植被的性质。结果表明：1）阳坡林线的海拔范围为2 605～2 790 m,阴坡林线的海拔范围为2 810～3 015 m;2）草本植物群落随海拔高度的变化比较明显,阴坡和阳坡从郁闭林到山顶均依次分布林下草本层、林缘草甸、亚高山灌丛草甸、高山草甸,草本植物的分布很好地体现了林线内部景观的差异性;3）海拔高度是高山林线附近草本植物群落空间分异的决定性因素。     

Demographic variation of dwarf birch (<Emphasis Type="Italic">Betula nana</Emphasis>) in communities dominated by <Emphasis Type="Italic">Ledum palustre</Emphasis> and <Emphasis Type="Italic">Vaccinium uliginosum</Emphasis>

The structure and demographic processes were compared in shrub communities to test the effects of vegetation succession on population growth, fecundity and abundance of the dwarf birch (Betula nana L.), which is a rare and endangered plant species in Poland and a glacial relict in Central Europe. The effects of Ledum palustre L. and Vaccinium uliginosum L. were studied in the Linje nature reserve in Chełmińskie Lake District (northern Poland), in three permanent plots on a peat bog. Vegetative growth and reproduction of B. nana were lower in plant communities dominated by L. palustre and V. uliginosum, than in a reference site. Fecundity was also lower, despite the fact that the percentage share of potentially fertile age groups was similar in all study sites. Mortality of ramets was independent of vegetation, both for juvenile and mature stages. The results confirm that B. nana is intolerant of shade, and it is more abundant in vegetation without competitors. Light limitation can lead to its decline, primarily by a decrease in vegetative growth. Sexual reproduction may be negatively affected by shade, but it plays only small role in population growth. Butterfly larvae can destroy inflorescences, and thus contribute to low effectiveness of sexual reproduction. Increasing density of shrubs and trees in peat bogs can reduce the abundance of dwarf birch, and can lead to the extinction of its local populations.     

The vegetation of Flat Top Hill: An area of semi-arid grassland/shrubland in Central Otago, New Zealand

An account is given of the vegetation of Flat Top Hill, in the driest part of semi-arid lowland Central Otago, New Zealand. Although highly modified, the area was acquired for conservation in 1992, following almost 150 years of pastoral use. The vegetation was sampled in a composite scheme using permanent monitoring sites placed to include the majority of habitats and communities present. A number of environmental factors were measured in each sample. Native species comprise 53% of the vascular flora of the area (211 species). From multivariate analyses of the data collected over three seasons, fourteen 'communities' are recognised. Although there are few constant or faithful species, strong relationships are shown with certain environmental parameters. Moisture stress is the major environmental influence on the vegetation; soil depth and past disturbance are secondary determinants. The communities differ by a factor of 10 in vascular species richness; the richest communities, and those with the greatest native component, are those around rock tors. Many of the communities present have not been reported from other vegetation surveys in Central Otago Moisture stress at xeric sites in the dry core of the region has excluded some exotic species, and allowed the survival of the native component, including three tiny spring ephemerals. Near elimination of grazing, as a result of reservation, will probably lead to an increase in the cover of taller, palatable exotic grasses and Thymus vulgaris, which may threaten the survival of some native species. Optimum management, for recovery or persistence of native species, may comprise exclusion of grazers in some areas, but continuity of grazing in others.     

The influence of urban land use on seed dispersal and wetland invasibility

Urban habitats are generally considered highly invaded by exotic species due to the frequency and extent of disturbance caused by human activities and development. Our previous study had demonstrated that forested wetlands within residential areas are more extensively invaded than wetlands within industrial–commercial areas. In this study, we investigate whether the structure of the forest edge and seed dispersal can explain the differential in the invasion of wetlands surrounded by industrial and residential land use. Our results indicate that edges of industrially bordered wetlands are denser, with vegetation concentrated at the boundary of tree growth, whereas the edges of residentially bordered wetlands are more open and diffuse. This difference influences the number of seeds and species capable of dispersing into the wetland. Less dense edges resulted in a higher number of seeds entering residential wetlands; however, there were no differences in the numbers of exotic seeds or exotic species in the seed rain residential and industrial wetlands. Although seed dispersal could not directly explain differences in the current extent of invasion of these sites, seed dispersal did follow corridor pathways, including ditches and trails that breach the edges and extend through the wetlands. These disturbances act as corridors for seed dispersal into both types wetlands and may play a role in introducing new species to the interior of the wetland, an outcome supported by a higher number of exotic species in the seed banks of residential sites. Our results suggest that both the type of adjoining land use and the provision of access to people on trails in urban wetlands can affect the composition of these communities.     

Report of the Alpine Section

Background: Under climate-change scenarios, rock outcrops, by providing microclimatically diverse habitats, different from those of surrounding zonal vegetation matrix, may serve as climatic refugia and thereby facilitate the persistence of specialist species.

Aim: We tested whether rock outcrops of southern temperate latitudes could act as local refugia for a cold-adapted flora in the face of global warming.

Methods: We related species composition of 50 outcrops and that of their surrounding vegetation to climatic data at local and regional scales to establish whether species distributions reflected differences that could indicate the existence of climatic refugia.

Results: While at a regional scale, species composition of the outcrops across the study area was related to mean annual precipitation and mean annual maximum temperature, locally, southern faces of outcrops had different species and lower maximum temperatures than insolated north faces and surrounding vegetation plots.

Conclusions: South faces of outcrops by providing cool microhabitats, and currently harbouring species not found in the surrounding zonal vegetation matrix, could serve as local refugia for heat-intolerant plants and other microthermic organism. The degree of ‘stability’ of these refugia will depend on the degree of future change in climate. Therefore, it is important to plan protected area networks that maximise local environmental heterogeneity, including the protection of rock outcrops both as refugia for cold-adapted species and as potential stepping stones that would allow dispersal of these species between supportive environments through unfavourable ones.     

Facilitative interactions between an exotic mammal and native and exotic plants: hog deer (<Emphasis Type="Italic">Axis porcinus</Emphasis>) as seed dispersers in south-eastern Australia

Endozoochory by exotic mammalian herbivores could modify vegetation composition by facilitating the dispersal and establishment of exotic and native plant species. We examined the potential for endozoochoric dispersal of native and exotic plants by exotic hog deer (Axis porcinus) in south-eastern Australia. We quantified the germinable seed content of hog deer faecal pellets collected in five vegetation types within a 10,500-ha study area that was representative of their Australian range. Twenty exotic and 22 native species germinated from hog deer faecal pellets and significantly more native species germinated compared to exotic species. Seedlings of the encroaching native shrub Acacia longifolia var. sophorae emerged, but no native trees emerged and the percentage of grasses that germinated was low (11%). The species composition of germinants was similar among the five vegetation types. We estimated that the hog deer population in our study area could potentially disperse >130,000 viable seeds daily. Our study shows how an exotic mammal can disperse seeds from both native and invasive plants and highlights the need for endozoochory to be considered more widely in studies assessing the impacts of exotic mammals on plant communities.