The natural exclusion of red deer from large boulder grazing refugia and the consequences for saxicolous bryophyte and lichen ecology

Large boulder grazing refugia permitted comparison of saxicolous bryophyte and lichen assemblages with those boulder tops accessible to red deer (Cervus elaphus) on a sporting estate in northwest Scotland. Plant succession was predicted to occur unchecked by grazing on the tops of these large boulders with cascading effects on bryophytes and lichens—assuming boulders had been in place over the same time period. Fifty pairs of boulders (one ≥2 m and the other accessible to red deer) were selected at random from various locations below north-facing crags. Percentage cover of each bryophyte and lichen species was estimated from three randomly placed quadrats on each boulder top. Due consideration was given to the influence of island biogeography theory in subsequent model simplification. Mean shrub cover and height, leaf-litter, bryophyte cover and bryophyte species richness were significantly higher within quadrats on large boulder tops that naturally excluded red deer. Lichen cover and lichen species richness were significantly higher on boulder tops accessible to red deer. Lichen cover was in a significant negative relationship with bryophyte cover, shrub cover and litter cover. Bryophyte cover showed a significant positive relationship with shrub height but there was an optimum shrub cover. Natural exclusion of red deer from the tops of large boulders has facilitated plant succession. The results suggest that grazing arrests the lithosere on boulder tops accessible to red deer at an early plagioclimax favouring saxicolous lichens. The results are relevant to situations where red deer might be excluded from boulder fields that hold lichen assemblages of conservation value.     

Patterns of distribution and abundance of chitons of the genus Ischnochiton in intertidal boulder fields

Abstract Chitons of the genus Ischnochiton are found predominantly on the undersurfaces of boulders, compared with other intertidal or subtidal habitats. They therefore appear to be habitat‐specialists at this scale. This, combined with the fact that boulder fields are relatively sparse compared with other intertidal habitats, makes these animals vulnerable to natural and anthropogenic disturbances. In addition, many species of Ischnochiton are relatively rare and appear to have very patchy abundances, making them likely to have very specific requirements for habitat. We need to understand the habitat requirements in order to manage, conserve and restore disturbed habitats. The present study was carried out at three intertidal boulder fields separated along approximately 200 km of the coast of New South Wales, Australia, centred around Sydney. The boulder fields were representative of those found in this region. The boulders were made of different materials: shale in the north, sandstone in Sydney and quartzitic sandstone in the south. Some boulders in each boulder field were covered by up to 0.4 m of water during low tide. The study showed that the seven species examined were overdispersed among boulders in each of these three intertidal boulder fields. Most boulders did not have associated chitons, but there were very large abundances on a very small number of boulders. Chitons were also overdispersed among boulders that they occupied. These patterns were consistent among shores and among species, even though patterns of abundance were extremely different among different species. These species appear therefore to show specific requirements for habitat at a small spatial scale, using only a small proportion of potential patches of habitat (i.e. boulders) in any place. Extremely patchy patterns of dispersion can be caused by variation in patterns of recruitment, mortality or behavioural responses to habitat or other species. Before performing experiments to investigate such processes, it is useful to test hypotheses of association with habitat using mensurative experiments to identify environmental correlates that might explain the observed patterns. In the present study, sizes of boulders and the associated sessile and mobile assemblages were proposed as mechanisms that could affect dispersion of chitons among boulders. None of these factors, however, showed strong associations with abundances of chitons. The lack of support of these models rules out some features of habitat to which species of Ischnochiton might respond, thereby precluding manipulative experiments involving these features, which are unlikely to be involved in the very patchy patterns of dispersion of species of Ischnochiton.     

A preliminary investigation of diversity,abundance, and distributional patterns of chitons in intertidal boulder fields of differing rock type in South Australia

The rock type of hard substrata marine habitats can affect numerous benthic invertebrates, but little is known of the effects on molluscan assemblages, for example, the chitons often found under intertidal boulders. We compared chiton assemblage composition, abundance, species richness, and patterns of frequency distribution in 10 boulder fields containing either hard metamorphic/igneous boulders or soft limestone boulders in two geographical areas in South Australia. Similar species richness occurred in both types of boulder fields, but hard rock boulder fields had greater overall abundances, because of particularly large abundances of some common species. Differences in abundances of common species also resulted in significantly different assemblages occurring between the boulder field types. Some species appeared aggregated among boulders, but this pattern was variable between boulders in differing areas and of differing rock type. In one area, a common species had variable aggregation that caused frequency distributions to differ significantly between boulders of different rock types. These results indicate that rock type needs to be considered when investigating ecological patterns and processes involving specialist molluscs such as under-boulder chitons and to ensure comprehensive marine reserve planning for protecting rare invertebrates in rocky intertidal reefs.     

Spatial and genetic investigation of aggregation in Ischnochiton (Polyplacophora; Neoloricata; Ischnochitonina; Ischnochitonidae; Ischnochitoninae) species with different larval development

Many species are aggregated at some spatial scale but for some species, due to habitat or behaviour, aggregation can be difficult to assess quantitatively. Here, aggregation of chiton species (Polyplacophora; Neoloricata; Ischnochitonina; Ischnochitonidae; Ischnochitoninae; Ischnochiton) living under boulders in intertidal boulder fields was investigated at several relatively small spatial scales. Closely related species were found to be aggregated at the level of individual boulders, but evidence for grouping at the larger scale of patches of boulders varied. When organisms are mobile, dispersal can have an important influence on spatial patterns. Some organisms, such as marine invertebrates, have a highly dispersive larval phase that can influence spatial patterns of adults, particularly where the adult is sessile or relatively sedentary. Aggregation was compared in two species to test whether different modes of larval development influence spatial pattern. There was some evidence that species with a planktonic larva were even more aggregated than expected, in contrast to predictions based on this mode of development. Both types of larval development (planktonic and non‐planktonic) produce larvae with short development times in these species, so one possible explanation for the grouping habit of these chitons is that they do not disperse, at all, from their natal boulder. The complexity of the boulder field habitat and the cryptic behaviour of these chitons may also contribute to a lack of dispersal. A simple application of a genetic method indicated, however, that philopatry at this scale is unlikely.     

Neither connectivity nor genetic diversity matter in the conservation of a rare fern and a moss on insular erratic boulders

Erratic boulders provide habitat for rock-dwelling species and contribute to the biodiversity of landscapes. In the calcareous Swiss lowlands, siliceous erratic boulders are exclusive habitat islands for the regionally critically endangered fern Asplenium septentrionale, about 20 bryophyte species and numerous lichens. Focusing on island biogeographical processes, we analysed the conservation genomics of A. septentrionale and the moss Hedwigia ciliata on insular erratic boulders in the Swiss lowlands and the adjacent “mainland” in siliceous mountains. We genotyped both species using double digest restriction associated DNA sequencing (ddRAD). For the tetraploid A. septentrionale, abundant identical multilocus genotypes within populations suggested prevalent intragametophytic selfing, and six out of eight boulder populations consisting of a single multilocus genotype each indicated single spore founder events. The genetic structure of A. septentrionale mainland populations coincided with Pleistocene glacial refugia. Four genetic lineages of H. ciliata were identified, and populations consisting of a single multilocus genotype were less common than in A. septentrionale. For both taxa, multilocus genotype diversity on boulders was lower than in mainland populations. The absence of common genetic groups among boulder populations, and the absence of isolation by distance patterns, suggested colonisation of boulders through independent long-distance dispersal events. Successful boulder colonisation of A. septentrionale seems to be rare, while colonisation by H. ciliata appears to be more frequent. We conclude that pivotal principles of conservation biology, such as connectivity and genetic diversity, are of less importance for the studied cryptogams on insular erratic boulders because of long-distance dispersal, intragametophytic selfing and polyploidy.

     

Early colonization of shallow subtidal boulders in two habitats

Intertidal and shallow subtidal boulder fields are relatively uncommon along the coast of New South Wales, Australia. Nevertheless, they provide habitat for a diverse suite of species, many of which are seldom found in other habitats. The types and abundances of animals found on or under boulders can be influenced by features of the boulders themselves, or by features of the substratum on which a boulder lies. Boulders can be colonized by larvae, or by adult or juvenile animals drifting in the water column or crawling up from the substratum. This experiment investigated the effect of an algal or sandy substratum on early stages of colonization of new boulders in two boulder fields by a suite of invertebrates, including gastropods, bivalves, chitons, polychaetes and insect larvae. Replicate times of three different periods (5, 18 and 38 days) tested for consistency of patterns of colonization at different times. Although the experiment was completed in a single season (within 2 months), there was considerable variation in patterns of abundances and diversity between replicate times. Thus, for each time in each period of a particular length, a different assemblage developed on the algal or sandy substratum in each boulder field. Despite this temporal variation, a few taxa showed some consistency of colonization among habitats, although most showed unpredictable patterns. The implications of these patterns of colonization for the creation or restoration of new intertidal and shallow subtidal boulder fields are discussed.     

Fish assemblages of Cais do Carvão Bay (Madeira Island) determined by the visual census technique

Fish assemblages in Cais do Carvão Bay, Madeira Island, a proposed marine protected area (MPA), were determined from a diver visual census. A total of 32 transect counts were performed. Habitats sampled included sandy bottom, rocky boulders, vertical walls and rocky outcrops. Species richness, diversity, density, trophic structure, size and spatial organization were documented for the fish assemblages. Forty‐four species from 23 families were encountered; 32% belonged to Sparidae (10) and Labridae (four). The greatest species richness (25) was observed in rocky boulder habitat at 10–15 m depth, while the lowest (five) occurred over a deeper sand habitat. The greatest density (760·5 individuals per 100 m²) was recorded over rocky outcropping (20–25 m deep), and the lowest of 11·6 individuals per 100 m² was over a sand bottom at 10–15 m depth. Thalassoma pavo , Abudefduf luridus and Chromis limbata had higher densities on hard bottoms, while Heteroconger longissimus was the most abundant species in sand bottom habitats. No significant differences were detected for all indices calculated among depth intervals for sand and rocky boulder stations. Sand and rock boulder substratum, however, differed significantly for the 10–15 m depth stratum.     

Restoring Intertidal Boulder‐Fields as Habitat for “Specialist” and “Generalist” Animals

Restoration is important in urban areas where habitat destruction is greatest. It incorporates many levels of intervention, with creation of new habitat the most extreme form. Most research on habitat creation has been terrestrial, or in marine habitats dominated by large structuring biota, such as mangroves. Intertidal boulder‐fields in urban areas are vulnerable to disturbances and habitat loss, which adversely affect numerous habitat specialists. This study describes experiments in which quarried stones were used to create new habitat outside natural boulder‐fields as a practical approach to restoring habitat. Colonization by specialist fauna and by common algae and invertebrates was measured for a year after deployment. Despite sessile assemblages on new boulders differing from those on natural boulders, common and rare animals rapidly colonized the new habitat. There was no clear succession, but colonization was variable and patchy at all scales examined, although diversities and abundances of some species in this novel habitat matched those of natural boulders within a few months. Rare and common animals generally colonized the new habitat as adults moving in from surrounding areas. Creating new boulder‐fields using quarried rocks is a successful approach to restoration and conservation of fauna where natural boulder‐fields are threatened.     

Borings in Quartzite Surf Boulders from the Upper Cambrian Basal Deadwood Formation,Black Hills of South Dakota

Shallow, semi-spherical borings occur in clusters with densities of 1–3.5/cm² in quartzite boulders and in vein quartz from the localized basal conglomerate of the Cambrian-Ordovician Deadwood Formation in the east-central Black Hills of South Dakota. Some borings are superimposed on primary but enigmatic semi-circular structures, 2.5–5 cm in diameter, which are soft-sediment trace fossils formed prior to lithification. The macroborings are the first to be recorded from quartzite and vein quartz. Host boulders were eroded from near vertically dipping Paleoproterozoic quartzites from several different stratigraphic units. The thin boulder conglomerate grades were abruptly changed to sandstone through a layer ≤2 m thick in the Marjuman transgression (regionally correlated to the Cedarina dakotaensis trilobite zone). This transgression occurred prior to the start of the globally recognized Upper Cambrian Paibian Stage. Physically similar rocky-shore settings are widely known from quartzite islands of Cambrian age in Wisconsin, Middle Ordovician age on Ontario's Manitoulin Island, Ordovician-Silurian age in Manitoba, and Devonian age in western Australia. Erosion of quartzite surf boulders of equal or larger size occurred in all those regions, but the Black Hills of South Dakota is the only region where borings in quartzite are documented.     

Evaluation of physical fish habitat quality enhancement designs in urban streams using a 2D hydrodynamic model

Creating a habitat for a variety of forms of life, such as riparian plants and various fish, is a necessity for stressed river ecosystems. In this study, the hydraulic characteristics of a fish habitat in an urban channel were analyzed using River2D, which is a two-dimensional (2D) depth-averaged finite element hydrodynamic model, to improve the habitat of two target fish in the Daejeon Stream, Korea. These species are Pseudopungtungia nigra, which is an endangered species in the Daejeon Stream, and Zacco platypus, which is a dominant species. In addition, changes in the weighted usable area (WUA) were compared and reviewed as boulders were placed in the stream. The best method for improving the P. nigra's habitat is proposed. A simulation analysis was performed on urban rivers for fish habitats. As a result, a straight and monotonous urban river flow was found to be an appropriate habitat environment for Z. platypus. The WUA for Z. platypus was about 20 times greater than that for P. nigra. Three different fish habitat enhancement methods were evaluated by calculating the WUA for the target fish in the study channel. By calculating the WUA to create fish habitats, the V-type riffle method was found to increase the usable area of the habitat environment for P. nigra by 360%, and the step stone method and single boulder method did so by 60% and 8%, respectively. For the single boulder method, boulders were placed in the channel bed at 3.5-m intervals, which significantly increased habitat availability. Moreover, centralizing the flow pattern in the channel among several types of boulder placements greatly expanded the habitat for P. nigra. Thus, an appropriate placement interval and boulder location that considers the characteristics of the riverbed and target fish species should be researched and implemented.     

Multiple-scale environmental modulation of lichen reproduction

It is necessary to understand how environmental changes affect plant fitness to predict survival of a species, but this knowledge is scarce for lichens and complicated by their formation of sexual and asexual reproductive structures. Are the presence and number of reproductive structures in Lobaria pulmonaria, a threatened lichen, dependent on thallus size, and is their formation sequential? Does any size-dependence and sequential formation vary along a climate gradient? Generalized linear mixed models were used to explore the effect of environmental predictors on the size and presence/abundance of each reproductive structure and to determine the probability of a given-sized thallus to develop any reproductive structure. The largest individuals are more likely to develop reproductive structures, and the lichen uses a mixed strategy of early asexual reproduction and late sexual. Macro and microclimatic variables also influenced reproductive capacity. Relationships among climate conditions and lichen size and reproductive capacity can compromise the future viability of the species in the most southern populations of Europe.     

Demography, competitive interactions and grazing effects of intertidal limpets in southern New Zealand

Population dynamics and the effects of intraspecific competition on limpet growth and maintenance of bare patches were investigated for the intertidal limpet Cellana ornata (Dillwyn) at a boulder-dominated site and on a rocky platform near Kaikoura (South Island), New Zealand. Distribution and abundance patterns of C. ornata were described in relation to other biota and tidal height. C. ornata occurred almost exclusively in patches devoid of macroalgae, particularly in the mid-tidal zone. Both adult and juvenile limpets were most abundant on the tops of boulders, where their numbers were positively correlated with barnacle cover, which averaged 77%. The size structure and growth patterns of C. ornata were different between populations. Mark-recapture studies showed that the slopes of annual growth increments regressed on initial sizes were similar at both sites but that the annual increments on the platform were about 6 mm greater than on boulders. Growth virtually ceased at 27 mm for limpets at the barnacle-dominated boulder site and at 40 mm at the platform site. Recruiting cohorts had 20% survival on boulders and 37% on the platform during their first year. The largest size classes at both sites had around 57% annual survival. To test the effects of varying limpet densities on the growth and mortality of limpets and the maintenance of bare patches, densities of C. ornata were experimentally increased at both sites. Beyond a density of 4 per 0.25 m², sizes and survival of limpets were reduced at both sites, but the effect was more pronounced at the boulder site. Limpets at the boulder site were more effective at maintaining bare space than those on the reef platform. Enclosing limpets in plots with and without barnacles showed that C. ornata and a co-occurring species (Cellana denticulata (Martyn)) grazed more effectively and had greater growth in cleared plots. Overall, there was considerable variation in the demographics of C. ornata between populations driven by site-specific characteristics.     

Macrofaunal diversity and habitat structure in intertidal boulder fields

Low-midshore boulder fields in Europe are increasingly subject to degradation related to beach fishing for crabs and abalones. The aim of the study was to better understand the structure and species diversity of macrofaunal assemblages living in a low-midshore boulder field in order to define conservation strategies for this biotope. Sampling strategy involved different spatial scales (macro- and microstrata) relative to the complexity and heterogeneity of the habitat. Assemblages of species and the corresponding habitats were identified by multifactorial analysis and compared by ANOVAs. The results show a hierarchical organization of the macrofauna within the boulder field, corresponding to three spatial levels of habitat structure: (i) eight microhabitats at the lowest level of spatial organization, each defined by a specific assemblage (subcommunity); (ii) three habitats at a middle level combining these microhabitats, each associated with a specific community (open rock, protected rock and sediment); and (iii) three macrohabitats at the uppermost level (corresponding to the scale of the entire boulder field and including the main geomorphological features of the beach), each defined by a specific assemblage of species (boulders on boulders, boulders on bedrock, and boulders on sediment). Two microhabitats with particularly high species diversity were regarded as the most important ecological units of the field in terms of conservation of specific and functional biodiversity. Comparison of habitat/community parameters showed (i) that habitat heterogeneity was not an accurate indicator of faunal diversity, and (ii) that habitat complexity enhanced the species richness of the fauna, but only above a threshold value. This enhancement was due to semi-sheltered microhabitats, which were found only in the most complex areas of the boulder field. It is likely that this complexity affects species richness qualitatively more than by the diversity of microhabitats. In other words, a cross-scale effect is apparent in which high habitat complexity on the middle spatial scale creates microhabitats on the lowest spatial scale that are characterized by stable semi-sheltered environmental conditions conducive to a maximum of species.     

A quantitative evaluation of the effects ofAscophyllum harvesting on the littoral ecosystem

Little is known of the ecological effects of harvesting littoral algae although this is a worldwide commercial activity. In 1976 an attempt to establish harvesting in Strangford Lough, Northern Ireland, was opposed on mainly theoretical conservation grounds. The attempt began and stopped within a single small bay leaving a sharp boundary between cut and uncut areas. A subjective survey apparently confirmed the predicted loss of cryptic fauna, decline through predation and the resorting of interboulder sediment. In April 1979 the cut and uncut areas were examined in detail to determine whether any of these effects had persisted and were demonstrable scientifically. Beach and boulder transects and various other studies showed some increases in the cut area. There was significantly moreFucus, Enteromorpha andUlva; Cirratulus (inhabitingRhodochorton-bound sediment on boulder surfaces) had a greater biomass. Some changes inLittorina colour morphs were apparent. Sediment in the cut area was coarser and had significantly more crustacean meiofauna.Ascophyllum internodal length and lateral branching were increased but it still provided 20% less shore cover than in the uncut area. There were significant decreases in the cover ofCladophora on the sides of boulders and ofHalichondria, Hymeniacodon andBalanus on undersurfaces. Indeed on the habitable underside of boulders total animal cover had been reduced by nearly two-thirds and the average number of species per boulder by one-third. It is concluded thatAscophyllum harvesting has a significant and persistent effect on shore ecology. Littoral algae are a valuable commercial asset but it is important that some fairly large intertidal areas should be left unharvested for general conservation purposes.     

Topographic and forest-stand variables determining epiphytic lichen diversity in the primeval beech forest in the Ukrainian Carpathians

The Uholka-Shyrokyi Luh area of the Carpathian Biosphere Reserve is considered the largest and the most valuable primeval beech forest in Europe for biodiversity conservation. To study the impact of different topographic and forest-stand variables on epiphytic lichen diversity a total of 294 systematically distributed sampling plots were surveyed and 198 epiphytic lichen species recorded in this forest landscape, which has an uneven-aged structure. The obtained data were analysed using a non-metric multidimensional ordination and a generalized linear model. The epiphytic lichen species density at the plot level was mainly influenced by altitude and forest-stand variables. These variables are related to both the light availability i.e. canopy closure, and the habitat diversity, i.e. the developmental stage of the forest stands and the mean stem diameter. We found that lichen species density on plots with a relatively open canopy was significantly higher than on plots with a fairly loose or closed canopy structure. The late developmental stage of forest stands, which is characterized by a large number of old trees with rough and creviced bark, had a strong positive effect on lichen species density. In the Uholka-Shyrokyi Luh primeval forest the mean stem diameter of beech trees significantly correlated with lichen species density per plot. Similar trends in the species diversity of nationally red-listed lichens were revealed. Epiphytic lichens with a high conservation value nationally and internationally were found to be rather abundant in the Uholka-Shyrokyi Luh area, which shows its international importance for the conservation of forest-bound lichens.     

Factors influencing the distribution of aquatic plant communities in Irish canals

Terrestrial tardigrades are often found in the lichens and mosses growing on trees and rocks. The assertion that tardigrades in these habitats are very patchy in their distribution has rarely been backed by quantitative sampling. This study assesses spatial variability in tardigrade populations inhabiting small patches (0.1 cm² to over 5 cm²) of moss and lichen on trees and rocks at three sites in the United States of America. Tardigrades were collected from four replicate rocks in the Ouachita Mountains of Arkansas, with 30 lichen patches collected on two adjacent boulders and 20 moss patches on a second pair of boulders. In Fort Myers and in Citrus Springs, Florida, 30 lichen patches per tree were collected from two pairs of trees. The tardigrades in each sample were extracted, mounted, identified, and counted. The variation in tardigrade abundance among lichen or moss patches within rocks or trees was very high; the only consistent pattern was that very small patches usually lacked tardigrades. Tardigrade diversity and abundance also varied greatly within sites when lichens and mosses of the same species from different rocks and trees were compared (in the most extreme case one tree had numerous individuals of two tardigrade species present while the other had almost no tardigrades). The results of this quantitative sampling support the assertion that tardigrades are very patchy in distribution. Given the considerable time investment required for the quantitative processing of tardigrade samples, this high spatial variability in tardigrade diversity and abundance requires that researches testing ecological hypotheses about tardigrade abundance check variability before deciding how many samples to take.     

浙江金华北山石生藓类分布与环境关系的排序分析

采用样方法调查了浙江金华北山石生环境30个样点167个样方,共记录86种,其中,丛藓科(Pottiaceae)、青藓科(B rachythec iaceae)、羽藓科(Thu id iaceae)和灰藓科(Hypnaceae)为4个大科,含的种数分别为11、11、10和8种。含种数较多的属为真藓属(Bryum)、凤尾藓属(F issidens)和青藓属(Brachythecium),分别含有7、7和6种。对86种藓类植物分布与环境关系的典范对应分析表明,海拔高度、水分条件和光照强度是影响金华北山石生藓类分布的主要环境因素。海拔决定藓类植被的类型,进而影响光照、岩表温度和水分条件。石面光洁度对石生藓类分布也有一定的影响,而石面坡度与其分布没有显著的相关性。     

Variation in Habitat use by Juvenile Acadian Redfish, Sebastes fasciatus

A basic paradigm in behavioral ecology is that organisms expand their distribution as preferred sites become saturated with individuals that reduce the availability of resources (e.g., shelter, prey) on a per capita basis. Previous fish community studies at Stellwagen Bank National Marine Sanctuary have shown that juvenile Acadian redfish Sebastes fasciatus (<20cm total length; TL) were primarily associated with boulder reefs that have deep interstices amongst the boulders; and that redfish expanded their distribution to adjacent gravel habitats when local abundance on reefs was high. Multibeam and sidescan sonar surveys in Stellwagen Basin (primarily a cohesive mud seafloor) have shown that discrete small areas of the basin floor are composed of mud draped gravel and partially buried boulders. Linear video transects using remotely operated vehicles and a video/photographic equipped grab sampler across five of these sites in 1997 showed that exposed boulders do not have crevices along their lower margins and are surrounded by dense patches of cerianthid anemones, Cerianthus borealis. These anemone patches are not present on the surrounding mud seafloor. Video image data showed that late juvenile redfish (11–20cm TL) occurred on boulder reefs as well as in the dense cerianthid patches but not on unstructured mud habitat (without cerianthid anemones). Comparisons of boulder reef and cerianthid habitats in 1998 showed that early demersal phase (0-year) redfish (0–10cm TL) occurred only on reefs but late juveniles occurred both on the reefs and in dense cerianthid habitats. Adult size classes (>20cm TL) also occurred in dense cerianthid habitats. Two explanations for these distributions can be advanced. The simplest is that redfish use both boulder and cerianthid habitats on an encounter basis, regardless of habitat saturation or predation pressure. Alternatively, boulder reefs serve as recruitment habitats and cerianthid habitats serve as a conduit for redfish moving away from saturated boulder reef sites, essentially serving as elements of a 'redfish pump'.     

内蒙古草原放牧恢复过程地衣生物量分布及其影响因素的研究

1　引　　言在干旱半干旱草原生态系统中,地衣是植物群落中的一个重要组成部分.地衣是真菌和具有光合功能的藻类的共生体,共生光合生物通过光合作用为自身和共生真菌提供碳水化合物,共生真菌为共生光合生物提供物理保护、水和矿物元素[9] .地衣芽枝可通过截获风蚀物质而增加土壤表面粗糙度,有利于积蓄降水和养分[1] ,地衣的共生真菌菌丝可以通过分泌粘质物而改善土壤团粒结构[3 ] .另外,地衣还可有效地分解矿物元素,同时还有较强的固氮功能[9,17] ,地衣不仅影响土壤的理化性状而且对维管植物也有潜在的影响.因此,地衣在草原生态系统中可促进…     

Analysis of growth directions of columnar stromatolites from Walker Lake, western Nevada

Samples of digitate, branching, columnar stromatolites were collected from the steep sides and near horizontal top of four in situ boulders located on the southwestern side of Walker Lake, Nevada, to test the widely held assumption that stromatolite column formation represents a phototropic response. We would predict that the columns on the steeply dipping sides of the boulder would bend upwards toward the light during growth if phototropism was significant during stromatolite morphogenesis. Angle of growth measurements on >300 stromatolites demonstrate that the stromatolites grew nearly normal to their growth surface, regardless of the inclination of their growth surface. No significant differences in the distribution of growth angles between north-, south-, east-, or west-facing samples were observed, and stromatolite lamina thickness did not systematically vary with position on the boulder. The lack of a strong phototropic response does not rule out a biological origin for the Walker Lake structures, but it does suggest that phototropic growth was not a dominant factor controlling stromatolite morphogenesis in these stromatolites and that column formation cannot be uniquely attributed as a phototropic response in stromatolites. It is interesting to note that the morphology of the stromatolites on the top of the boulder is identical to stromatolites on the steep sides. Stromatolite morphogenetic models that predict branching typically require a vertically directed sedimentary component, a feature that would have likely affected the stromatolites on the tops of the boulders, but not the sides, suggesting that other factors may be important in stromatolite morphogenesis.