Climate,landscape and microenvironment interact to determine plant composition in naturally discrete gravel beach communities

Question: What is the relative importance of national‐, regional‐ and within‐beach‐scale influences on vegetation composition and floristic affinities of New Zealand gravel beaches? Location: Coastal New Zealand. Methods: We sampled vegetation composition at 61 gravel beaches, quantifying site factors and adjacent landscape characteristics. Site, climate and geographic relationships between gravel beaches and related ecosystems were inferred using GIS data layers. To simultaneously investigate influences at different spatial scales, we used ordination and variation partitioning to examine relationships between composition and environment, and hierarchical models to understand floristic affinities with related ecosystems. Results: At a national scale, compositional variation among beaches reflects mean annual temperature and spring vapour pressure deficit; within regions, proximity of native woody vegetation and coastal turfs are important; within‐beach variation is related to substrate stability and particle size distribution. The gravel beach flora is 50% exotic, reflecting the highly modified nearby landscapes; 30% of species are characteristic of coastal sands, 20% of braided riverbeds and 8% of coastal turfs. Affinities with coastal sand communities are unrelated to microsite sandiness or area of sand dunes within 50 km. Affinities with braided riverbeds are related to the bed area of those rivers draining within 200 km and proportion of gravel in the substrate. Affinities with coastal turfs are related to proximity to the nearest turf and the proportion of humus in the substrate. Conclusions: Examining multiple scales of influence in a landscape context is essential to understand composition of naturally discrete ecosystems that span wide geographic ranges and to underpin their conservation management.     

Impacts on biodiversity at Baltic Sea beaches

Sandy and shingle beaches were investigated in 2009 and 2010 along the coast of the Baltic Sea in northern Germany with the purpose of assessing biodiversity gradients from shingle to sandy beaches, from beach to primary dunes, and the impacts of tourism on biodiversity. On nine beach sites, ranging between 100% shingle and 99% sand without shingle, Carabidae, Staphylinidae, and Araneae were studied. Two of the six sandy beaches were open and four were closed to tourists. Additionally, trampling effects from tourists, species richness of plants, and plant cover were investigated on sixteen beaches. According to results, primary dunes showed higher species richness in carabids and spiders, but not in staphylinds. Shingle beaches exhibited lower species richness in Staphylinidae and Araneae, but not in Carabidae. As estimated by the Jackknife II method, shingle beaches were the lowest in total species richness. Trampling intensity ranged from 0 footprints m⁻² day⁻¹ on closed beaches, up to a maximum of 30 footprints m⁻² day⁻¹. On “intensively” used beaches (12 footprints m⁻² day⁻¹, on average), reduction of plant cover was more pronounced than on “extensively” used beaches (7 footprints m⁻² day⁻¹, on average). Both plant cover and plant species richness were lower on intensively and extensively used beaches than on closed beaches. In arthropods, only staphylind and spider species richness was significantly lower on open beaches than on closed beaches, but no differences were found in carabids. Referring to our results, trampling effects from tourists have high impact on species richness of sandy beaches, on both intensively and extensively used sites.     

Determinants of regional and local patterns in the floras of braided riverbeds in New Zealand

Aim To determine the composition and biogeographical origins of the native and naturalized flora of braided shingle riverbeds in New Zealand and whether the proportion of naturalized species is greater than in similar habitats elsewhere in the world. To test whether regional species pools, landscape matrix configuration, and local riverbed environment are all equally important in determining community structure in these systems. Location The braided reaches of four catchments on each of the eastern and western sides of North Island and South Island, New Zealand. Methods Plotless records of all native and naturalized seed plant and fern species on disturbed mobile surfaces were made over the length of four rivers. Altitude, climate variables, riverbed attributes, and surrounding land‐cover from maps were recorded for each site. For all species, the taxonomy, life form, origin and history, and presence within the province through which the rivers flowed, was obtained from published floras. Direct and indirect ordination and variance partitioning were employed to examine how native and naturalized species composition varies among and within rivers, and the degree to which this variation reflects climate, characteristic of riverbeds, and the surrounding land‐cover. Regression was used to determine how much introduction date and native geographical range influence the frequency of naturalized species. Results The total riverbed flora of 289 species comprises 40% native species and 60% naturalized species, both dominated by Asteraceae and Poaceae. The relative contributions of other plant families differ, and the two groups comprise different life‐form spectra. Native species occur across fewer rivers (mean 1.6 rivers) than naturalized species (mean 2.1 rivers). Species common in at least one river system tend to be widespread, occurring in at least three rivers. The rivers differ in their floras, with distinctions between North Island and South Island, and eastern and western rivers. The South Island rivers have more native species and a higher proportion of their regional native species pools than North Island rivers, whereas they have a lower proportion of their naturalized species pools. Introduction date and native geographical range are correlated with frequency of individual naturalized species. Geographical position and climate, riverbed variables of substrate size and seepage presence, and the type of adjacent land‐cover, differ between rivers and are significantly related to species composition. Land‐cover variables alone account for 46% of the explained variation in species composition, environmental variables alone 32% and components of the environment and land‐cover that co‐vary, 22%. Main conclusions New Zealand shingle riverbeds are among the riparian communities in the world most invaded by naturalized species but comparisons are difficult because studies of similar riverbeds are rare. New Zealand riverbeds are dominated by short‐lived terrestrial species from Eurasia. Native species are most frequent in South Island rivers draining extensive high mountains, and their abundances are determined to a greater extent by riverbed processes than are those of the naturalized species, which are more abundant when the surrounding landscape is modified. The distribution of the naturalized flora is not yet at equilibrium with the environment. Each river has a distinctive flora determined by ambient environmental factors, aspects of the riverbed environment itself, adjacent land‐cover reflecting the presence of native vegetation, the intensity of human modification or use, and invasion history.     

If you build it,will they come? Use of restored dunes by beach mice

Restoration of coastal habitat fragmented, degraded, or destroyed by development and climate‐related processes such as sea level rise and storm surge usually involves planting native plants to restore habitat structure, but whether and how restored areas benefit taxa other than plants is rarely reported. Installing restoration plantings is one method used to build habitat such as beach dunes where dunes have been lost, potentially creating habitat for dune‐dependent species. We compared use of natural vegetated dunes, open sand gaps, and restoration plantings (habitat treatment) by Perdido Key beach mice (Peromyscus polionotus trissyllepsis) over 3 years using tracking tubes to assess the value of restoration plantings for beach mice. Tubes were monitored in two seasons (early and mid‐summer), and under new and full moon conditions. Mice used restoration plantings less than natural vegetated dunes but more than open sand gaps, which suggests restoration plantings may facilitate movement of mice across fragmented areas. Both season and moon phase influenced the effect of habitat treatment, interactions which may be attributable to perceived risk associated with movement under a combination of different conditions of ambient light, vegetation cover, and habitat novelty. Our results show restoration plantings provide habitat for movement and foraging, and may ameliorate some consequences of sea level rise and storms for beach mice and potentially other dune‐dependent species into the future.     

Comparison of the breeding habitats of Little Ringed Plover Charadrius dubius and Kentish Plover Charadrius alexandrinus on a shingle bed

The breeding habitat preferences of Kentish Plover and Little Ringed Plover were studied on a shingle bed. The location of each nest was measured as the distance to the sea wall, nearest pond and nearest path. The habitat surrounding nests was measured at radii of 0.15 m and 15 m as the proportional cover of shingle, gravel, sand, live and dead vegetation. When all variables were considered simultaneously, the differences between the habitats used by the species were related to the cover of shingle within 15 m of the nest and the distance to the sea wall. Nest sites located close to the sea wall and with high shingle cover almost always belonged to Kentish Plover, while those more distant and with low shingle cover almost always belonged to Little Ringed Plover. Kentish Plover nested in areas of newly exposed shingle, while Little Ringed Plovers used areas which had been stable for longer and hence were colonized by vegetation. This study suggests that management which maintains a high proportion of exposed substrate on shingle beds will increase their suitability for Kentish Plover.     

Geographical differentiation inferred from flavonoid content between coastal and freshwater populations of the coastal plant Lathyrus japonicus (Fabaceae)

Lake Biwa, one of the few ancient lakes in the world, harbors many coastal species that commonly inhabit seashores. The beach pea (Lathyrus japonicus) is a typical coastal species of this freshwater lake, and morphological and genetic differentiation between inland and coastal populations of this species have been reported. Inland and coastal habitats inflict distinct environmental stresses to plants, the latter imposing salt stress and high-light intensity, which leads to physiological differentiation. These abiotic stresses affect phenolic compounds, which play an important role in the response of plants to the toxic by-products of stress metabolism. We investigated physiological differentiation of phenolic compounds of the beach pea between inland and coastal habitats using high-performance liquid chromatography (HPLC) analysis. Flavonoid composition analyses revealed that patterns of flavonoid composition of inland populations at Lake Biwa were differentiated from those of coastal populations. All Lake Biwa individuals were fixed in the same flavonols glycosylated at 3- and 7-positions. In contrast, most coastal individuals contained flavonols glycosylated at 3-position alone, and these populations exhibited higher variation in flavonoid composition compared to among/within inland populations. Variation was likely lower in inland populations because of a bottleneck during landlocked periods, which is consistent with previous phylogeographic studies. A qualitative HPLC survey of flavonoid content revealed substantial variation among individuals regardless of locality. These results suggest that changes in the habitat environment may have led to beach pea acclimation via alteration of the quantity and quality of flavonoids.     

Coastal fish otoliths from the early Pleistocene of Rhodes (eastern Mediterranean)

Coastal fish assemblages are especially vulnerable to environmental changes, but little is known about their evolution through time, mainly due to the scarcity of fossil material from such settings. The aim of this study is to characterize the early Pleistocene coastal fish assemblages of the eastern Mediterranean and to reconstruct the related paleobathymetric and paleoecologic conditions. Based on otolith findings, we identified thirty-seven teleost fish species from three sedimentary outcrops on the northeastern part of Rhodes Island (southeastern Aegean, Greece, Eastern Mediterranean), which have been placed within the Gelasian (early Pleistocene). The stratigraphic distribution of fifteen taxa is expanded for this interval, while five species are reported for the first time as fossils. The otolith assemblages provided paleodepth estimates indicative of shallow coastal environments. The ecosystem consisted mostly of substrates of sand and mud, with significant rocky micro-habitats and underwater vegetation; the climate was subtropical. These data complement existing knowledge on the study area, while providing new information on the composition of the ichthyofauna of the eastern Mediterranean during the Gelasian.     

Holocene sand beaches of southern California: ENSO forcing and coastal processes on millennial scales

A proxy record of sand beach accretion for the past 10,000 years has been assembled from radiocarbon dates on the Pismo clam, Tivela stultorum, in archaeological sites along the southern California coast. When this record is compared with numerous climate proxies, it appears that El Niño-Southern Oscillation (ENSO) controls on wave climate and sediment flux have acted upon regional geomorphology at different sea levels to either accrete or erode the Holocene beaches of southern California. Tivela dates from the Santa Maria coast indicate that perennial sand beaches built by 9000 years ago in response to abundant riverine sediment contained by the natural groin at Point Sal, wave sheltering by the massive headland of Point Buchon, and Early Holocene El Niño events. On the western Santa Barbara coast, sand beaches were forming by 7000 years ago in response to high sand fluxes from the Santa Ynez Mountains to the many small littoral catchments, possibly aided by high local rates of uplift. Decline of these sand beach habitats 5000-4000 years ago coincides with increased El Niño-driven wave energy. In accord with slowing in sea-level rise ca. 6000 years ago, sand beaches were most widespread in the period 6000-5000 years ago on Estero Bay, the western Santa Barbara coast, and west of Point Dume. However, Tivela dates first appear 5000 years ago in the Oceanside and Silver Strand littoral cells of the San Diego region. This lag coincides with the Middle Holocene shift to a more variable climate and modern periodicity in El Niño events that increased sediment supply to the southern coast. The ontogeny of the littoral cells provides timelines for modeling coastal evolution with implications for sand beach ecology, prehistoric human coastal adaptations, and coastal planning for future climate change.     

Recovery of sandy beach and maritime forest vegetation on Phuket Island (Thailand) after the major Indian Ocean tsunami of 2004

Question: How rapidly has the sandy beach and maritime forest vegetation on Phuket recovered and regenerated after the impact of the major Indian Ocean tsunami of 2004? What are the characteristics of sandy beach species for regenerating their populations and the invasion patterns of originally non-sandy beach species or other newcomers after the tsunami? Location: Phuket Island, southern Thailand. Methods: Species composition of beaches was studied on the same research plots 6 months before and 9 months after the tsunami. The changes in individual species cover before and after the tsunami were determined by χ² tests. Change in community composition was analysed by detrended correspondence analysis. The relationship between species and environmental factors was analysed by canonical correspondence analysis. Results: The sites disturbed by the tsunami were often invaded by annuals, especially grasses and asteraceous plants, rather than by perennials. In contrast, species with clonal growth by stolons decreased significantly. Factors determining the species habitat differences were soil hardness (penetration resistance of sandy soil), per cent silt content, soil water content and beach management. Habitat differences among originally non-sandy beach herbaceous species that expanded their population or moved to the coast after the disaster were defined by sand accretion or erosion caused by the tsunami. Many sandy beach herbaceous communities changed into Dactyloctenium aegyptium communities because of the tsunami were originally constituted by non-sandy beach D. aegyptium with Cenchrus echinatus. Although the forest floors of most maritime forests were invaded by originally non-sandy beach Tridax procumbens, Eleusine indica or D. aegyptium because of the tsunami, this did not result in a change in the vegetation unit, because species' loss was restricted to the understorey. In time, these forests will recover their previous community composition. Conclusions: Our results suggest that originally non-sandy beach native species invaded the disturbed beaches rapidly after the tsunami but their habitats differ. Sites where sand accumulated on a beach because of the tsunami were invaded by D. aegyptium and E. indica, whereas soil erosion permitted invasion by Digitania adscendens. Tridax procumbens establishes rapidly on wet sites with hard soil, high per cent silt content and low beach management pressure. Sandy beach species with subterranean long rhizomes are strongly tolerant of such disasters. We concluded that the species composition of the beaches disturbed by a temporary large disaster is determined by dormancy and growth forms, with radicoid form being influential.     

Factors responsible for Honckenya peploides (Caryophyllaceae) and Leymus mollis (Poaceae) spatial segregation on subarctic coastal dunes

Low water and nutrient availability and significant sand movement, salt spray, and soil salinity are typical of coastal dunes. These conditions are generally unfavorable for the various life stages of plants and especially for seedlings. However, the intensity of these stresses decreases landward, even over short distances, with significant effects on community composition. On coastal dunes in subarctic Québec, Canada, Honckenya peploides (Caryophyllaceae) colonizes the upper beach where it forms small mounds called embryo dunes. Leymus mollis (Poaceae) is mostly restricted to the foredune; however, a few individuals successfully establish on the upper beach, particularly on embryo dunes. We hypothesized that this differential distribution is associated with differences in the tolerance of the two species' seedlings to physical stresses. Honckenya peploides and L. mollis seedling tolerance to sand burial, salt spray, soil salinity, and nutrient and water availability was assessed in greenhouse experiments. Unexpectedly, our results showed that tolerance to sand burial, salt spray, and soil salinity was lower for H. peploides than for L. mollis. If seeds are available and seedlings tolerate the conditions prevailing on the upper beach well, why are mature L. mollis individuals rare in this habitat? We suggest that massive abrasion events (e.g., violent storm waves and ice thrust) restrict the presence of the species on the upper beach.     

Genetic polymorphism in the bladder campion,Silene maritima

A survey of protein variability has been made in wild populations of Silene maritima, a perennial outcrossing coastal plant. In the most extensively studied populations, at Beesands, six out of 21 loci were polymorphic, and the average heterozygosity was 0.153. Four polymorphic loci were studied on an extensive shingle beach at Dungeness. Three loci showed no frequency cline with distance from the sea; a fourth locus, that for adenylate kinase, showed a barely significant cline, which was not confirmed by a repeat study in the same place a year later. Five polymorphic loci were studied at Beesands along a strip of shingle beach and up a contiguous cliff. No frequency difference between the two habitats was observed. There are significant differences in allelic frequencies between localities. It is concluded that the data fail to show an association between habitat and gene frequency, although they were collected in such a way as to be capable of demonstrating such an association if it existed.     

Plant coexistence in coastal heaths: Floristic patterns and species attributes

Abstract An investigation was made of plant coexistence in coastal heaths. A 4 km stretch of Pleistocene beach sand was selected, and patterns of variation in heath vegetation on ridges and slopes in the beach system were studied. The sampling design covered variation with geographic distance among transects, plots within transects and quadrats within plots which were orthogonal to the alignment of ridges and slopes. Cover of vascular plant species was scored within each quadrat and both multivariate and univariate analyses tested the hypothesis that variation in the data was associated with environmental difference and geographic distance. There was a strong flohstic variation between the ridge and slope habitats. However, within these habitats there was no significant variation with distance at scales of kilometres, nor at hundreds of metres within transects nor at tens of metres within plots in floristic composition, in numbers of species, and in cover. Though differing floristically, the vegetation of the slopes and ridges had similar curves of importance values and fairly similar spectra of life forms and forms of post-fire regeneration in their constituent species, although the ridges had slightly more obligate-seeding species and fewer species of sedges and herbs than the slopes. The two habitats shared several genera, however, within such genera, species with similar attributes mostly occurred in different habitats. Hypotheses were suggested to explain segregation of species between the two habitats and the coexistence of species within habitats. Since patterns of variation among habitats were consistent over both smaller and larger scales of distance examined, experiments to test hypotheses may be designed so that their findings might be applied generally to the whole system studied.     

Green beach vegetation dynamics explained by embryo dune development

Sandy coastlines are dynamic environments with potential for biodiverse habitats, such as green beaches. Green beach vegetation can develop on nutrient-poor beaches landward from embryo dunes. It is characterised by low-dynamic coastal wetland habitat such as salt marshes and dune slacks. It has been hypothesised that the establishment of green beach vegetation is facilitated by the shelter provided by embryo dunes, however evidence is lacking.We explored the importance of geomorphology and soil conditions on the species richness and turnover of green beach vegetation over a time period of 10 years. We recorded 107 plots along 11 transects over a gradient from beach to dune on the island of Schiermonnikoog, the Netherlands. We characterised transect geomorphology at transect level and soil conditions and vegetation at plot level in 2006 and 2016.We found that the green beach vegetation was highly dynamic, total plant cover increased by 62% within 10 years. In 2006 beach width was an important factor in explaining species richness, with the highest number of species occurring on narrow beaches with a large volume of embryo dunes. In 2016, species richness was positively associated with the build-up of organic matter. Overall species richness declined relative to 2006 and was accompanied by an increase in elevation due to sand burial and the expansion of embryo dune volume.Our data suggests that geomorphology influenced the vegetation indirectly by affecting sand burial rate. Plant species richness declined less at sheltered conditions where sand burial was limited, allowing the build-up of organic matter. This indicates a time-dependent relationship between the development of embryo dunes and plant species richness: embryo dunes can be a source of shelter, thus increasing species richness, but can compete for space over time, lowering species richness again. Our results are relevant for engineering and management of biodiverse sandy shores.     

The coastal-disjunct mesic flora in the inland Pacific Northwest of USA and Canada: refugia, dispersal and disequilibrium

Aim Understanding the history of the mesic‐adapted plant species of eastern British Columbia and northern Idaho, disjunct from their main coastal distribution, may suggest how biotas reorganize in the face of climate change and dispersal barriers. For different species, current evidence supports establishment of the disjunction via an inland glacial refugium, via recent dispersal from the coast, or via a combination of both. In this study, the modern distributions of the coastal‐disjunct vascular plants are analysed with respect to modern climate to examine how refugia and/or dispersal limitation control regional patterns in species richness. Location North‐west North America. Methods The distributions of nine tree and 58 understorey species with a coastal‐disjunct pattern were compiled on a 50‐km grid. The relationship between species richness and an estimate of available moisture was calculated separately for formerly glaciated and unglaciated portions of the coastal and inland regions. Growth habit and dispersal mode were assessed as possible explanatory variables for species distributions. Results Species richness shows a strong relationship to climate in coastal‐unglaciated areas but no relationship to climate in inland‐glaciated areas. In inland‐glaciated areas, richness is c. 70% lower than that expected from climate. Species with animal‐dispersed seeds occupy a larger portion of coastal and inland regions than species with less dispersal potential. Main conclusions Modern patterns of diversity are consistent with both refugia and dispersal processes in establishing the coastal‐disjunct pattern. The inland glacial refugium is marked by locally high diversity and several co‐distributed endemics. In the inland‐glaciated area, dispersal limitation has constrained diversity despite the nearby refugia. Onset of mesic climate within only the last 3000 years and the low dispersal capacity of many species in the refugium may explain this pattern. This study suggests that vascular plant species will face significant challenges responding to climate change on fragmented landscapes.     

Determining vulnerability of stream communities to climate change at the landscape scale

1. As the climate changes, species are expected to shift to higher latitudes and altitudes where suitable habitat is available if dispersal is not constrained by geographic barriers. We analyse patterns of turnover in freshwater macroinvertebrate assemblages to identify which communities are most likely to be at risk from climate change, and the location of geographic barriers that could impede such adaptive range shifts. 2. We analysed macroinvertebrate data from standard biological assessments at the family level, from surveys of all coastal basins of New South Wales, Australia, covering a latitudinal gradient of more than 1000 km. We used variance partitioning to separate the variation in composition explained by climate, among‐site distance, human disturbance and other stream factors. 3. Montane stream assemblages showed high turnover in response to climatic variation. Turnover in coastal‐fringe streams was least affected by climate, but strongly correlated with distance and stream variables. Significant shifts in assemblage composition occurred between habitats within catchments and across catchment boundaries. 4. Montane stream assemblages are most vulnerable to climate change because their distribution is most responsive to climatic factors, and elevated sites are isolated from one another, reducing the scope for altitudinal migration. Dispersal limitations in coastal‐fringe assemblages will also increase their vulnerability to habitat loss from sea‐level rise. For all stream classes, the separation of many neighbouring catchment assemblages, owing to either limited dispersal or the lack of suitable habitat, is likely to constrain adaptive range shifts. This would lead to an overall reduction in beta diversity among reaches and subsequently to a reduction in landscape‐level gamma diversity.     

Sandy beaches at the brink

Sandy beaches line most of the world's oceans and are highly valued by society: more people use sandy beaches than any other type of shore. While the economic and social values of beaches are generally regarded as paramount, sandy shores also have special ecological features and contain a distinctive biodiversity that is generally not recognized. These unique ecosystems are facing escalating anthropogenic pressures, chiefly from rapacious coastal development, direct human uses — mainly associated with recreation — and rising sea levels. Beaches are increasingly becoming trapped in a 'coastal squeeze' between burgeoning human populations from the land and the effects of global climate change from the sea. Society's interventions (e.g. shoreline armouring, beach nourishment) to combat changes in beach environments, such as erosion and shoreline retreat, can result in severe ecological impacts and loss of biodiversity at local scales, but are predicted also to have cumulative large-scale consequences worldwide. Because of the scale of this problem, the continued existence of beaches as functional ecosystems is likely to depend on direct conservation efforts. Conservation, in turn, will have to increasingly draw on a consolidated body of ecological theory for these ecosystems. Although this body of theory has yet to be fully developed, we identify here a number of critical research directions that are required to progress coastal management and conservation of sandy beach ecosystems.     

Climate‐change impacts on sandy‐beach biota: crossing a line in the sand

Sandy ocean beaches are iconic assets that provide irreplaceable ecosystem services to society. Despite their great socioeconomic importance, beaches as ecosystems are severely under‐represented in the literature on climate‐change ecology. Here, we redress this imbalance by examining whether beach biota have been observed to respond to recent climate change in ways that are consistent with expectations under climate change. We base our assessments on evidence coming from case studies on beach invertebrates in South America and on sea turtles globally. Surprisingly, we find that observational evidence for climate‐change responses in beach biota is more convincing for invertebrates than for highly charismatic turtles. This asymmetry is paradoxical given the better theoretical understanding of the mechanisms by which turtles are likely to respond to changes in climate. Regardless of this disparity, knowledge of the unique attributes of beach systems can complement our detection of climate‐change impacts on sandy‐shore invertebrates to add rigor to studies of climate‐change ecology for sandy beaches. To this end, we combine theory from beach ecology and climate‐change ecology to put forward a suite of predictive hypotheses regarding climate impacts on beaches and to suggest ways that these can be tested. Addressing these hypotheses could significantly advance both beach and climate‐change ecology, thereby progressing understanding of how future climate change will impact coastal ecosystems more generally.     

江苏省海滩植被演替的研究

江苏省海滩植被可分为滨海盐土植被、盐沼植被及海滩沙生植被三个基本类型。本文论述了这些植被类型的演替规律。滨海盐土植被与盐沼植被的演替,外因于土壤盐分含量递减与有机质含量的递增;海滩沙生植被的演替,外因于土壤沙颗粒大小及其相应的土壤含水量的变化,所以海滩植被演替为外因动态演替。     

Waikanae River estuary: Changes to habitat and bird fauna evident from surveys thirty years apart

Abstract

A second ecological survey of the Waikanae estuary, in the North Island of New Zealand, was carried out 30 years after the first (1941–1943), and was of similar duration. The observer for the first survey participated in the second, and the same methods were used. Brief accounts are given of the prevailing climate and geology, and of physical changes at the estuary. The effect of periodic storms and of the various encroachments by man is recorded; the latter appear to be particularly important. The vegetation of the estuary is described. Records of beach patrols and of other ornithologists supplement the authors' observations on the bird life. The numbers of birds recorded in the two surveys are compared by analyses of variance. Changes in the flora deduced from aerial photographs spanning the two surveys and from notes and photographs are related to man-induced disturbance, natural physical changes, and natural succession. The total of 79 bird species recorded at the estuary and in its immediate vicinity compares favourably with the 60 species recorded during the first survey. The increase of nearly one-third is due to such factors as the appearance of new Australian bird species, the information provided by the beach patrols, and the extension of sand-flat habitat. In terms of numbers observed, 10 bird species have increased, 11 remain unchanged, and 8 have declined. The distribution of birds is considered in relation to nine areas of habitat: the beach and open sea; open dunes at the beach front; river and sand flats; unfixed and fixed dunes; saltmarsh; grassed fields; and lakelets. The most species (29) were recorded on sand flats and the least (7) on unfixed dunes. Periodic rechannelling of the river mouth has led to expansion of the sand flats as a habitat for waders and some other shore birds, but other man-induced developments, such as housing, are threatening the present ecological balance of the estuary. Use of the area for recreation and for wildlife study is increasing, and we consider that action by conservation authorities is urgently required.