Diurnal, horizontal and vertical dispersal of kelp-associated fauna

The kelp Laminaria hyperborea is host for a rich fauna of mobile invertebrates. Dispersal patterns of these animals may be crucial for their availability to visual predators like fish, which are known to search for food in these productive habitats. Diurnal, horizontal and vertical dispersal within and out of the kelp forest were studied by analysing colonization of artificial substrata placed among kelps. The species composition of the fauna was different on three parts of the kelp: lamina, stipes (with epiphytes) and holdfast. The artificial substrata were colonized mainly by species associated with kelp. More species and individuals colonized the artificial substrata at night than during the day. The most abundant faunal groups on the kelps and the artificial substrata were amphipods and gastropods. Both groups dispersed at a significantly higher rate at night than during the day. Rapid horizontal dispersal out of the kelp forest was found. The artificial substrata were also colonized quickly by kelp fauna at all vertical levels inside and above the kelp forest. However, species associated with the kelp holdfast tended to disperse close to the bottom, while stipe-associated fauna moved through all parts of the kelp forest and even above the canopy layer. A high dispersal rate appears common amongst the mobile species living on kelp and seems to be advantageous, despite the risks involved in emigrating from the habitat and being exposed to predators. Higher activity at night may reduce availability to predators.     

Artificial substrata in a shallow sublittoral habitat: do they adequately represent natural habitats or the local species pool?

Artificial substrata have been advocated as tools which have considerable potential for monitoring both natural and anthropogenic effects on invertebrate communities of shallow coastal environments. In this experiment, community structure was compared between two dominant natural algal habitats (kelp holdfasts and algal turf) and artificial substratum units (ASUs; nests of pan scourers) deployed in close contact with, and 20 cm above the substratum. Univariate and multivariate statistical analyses were applied to the data to determine the similarity of community structure between the four different habitats. In addition, recently developed measures of taxonomic distinctness were applied to the data from both sets of artificial substrata to determine if they provided a representative sample of the local epifaunal species pool and thus have the potential to be used as surrogate samples for this important faunal group. There were marked differences between community structure in each of the habitats. Both sets of artificial substrata were dominated by tubicolous polychaetes with abundances that were more than an order of magnitude greater than in the holdfast and turf samples. The fauna recruiting to the artificial substrata deployed above the substratum showed the lowest values in the univariate summaries of diversity and evenness and were unrepresentative of the local species pool. Artificial samples deployed in contact with the substratum showed greater diversity and evenness but were still mostly unrepresentative of the local species pool. The tendency for both sets of artificial substrata to under-sample amphipods and to be dominated by suspension-feeding polychaetes suggests that methods using these units may be relatively insensitive to the effects of anthropogenic impacts (e.g. sewage outfalls) where shifts in community structure including increased dominance of suspension-feeders and polychaetes and a reduced dominance of amphipods have been observed. Further studies, including the evaluation of temporal variation in community structure related to the time at which the ASUs are deployed and duration of deployment, are needed to test the wider utility of artificial substrata as tools for monitoring shallow, sublittoral, epifaunal communities.     

Patterns of abundance and assemblage structure of epifauna inhabiting two morphologically different kelp holdfasts

The mobile fauna associated with two sympatric kelp species with different holdfast morphology (Saccorhiza polyschides and Laminaria hyperborea) was compared to test for differences in the assemblage structure of holdfast-associated mobile epifauna. A total of 24,140 epifaunal individuals were counted from 30 holdfasts of each kelp species. Overall epifaunal abundances exceeded faunal abundances previously reported from holdfasts of other kelps. Three taxonomic groups, Amphipoda, Mollusca, and Polychaeta, accounted for ca. 85% of all individuals. Total abundances increased with the amount of habitat available, quantified either as the volume or the area provided by the holdfasts. The multivariate structure of the epifaunal assemblage did not differ between holdfasts of the two kelp species. However, epifaunal assemblages responded differentially to the habitat attributes provided by each type of kelp holdfast: multivariate variation in the assemblage structure of epifauna was mostly explained by holdfast area and volume for L. hyperborea, and by the surface-to-volume ratio for S. polyschides holdfasts. Therefore, the physical attributes of biogenic habitats, here kelp holdfasts that better predict patterns in the assemblage structure of associated fauna can differ according to their different physical morphology, even though the overall assemblage structure of associated fauna was similar.     

Kelp rafts in the Southern Ocean

Kelp rafts were surveyed during three summer traverses between Hobart and Macquarie Island (subAntarctic) to determine the potential for dispersal of kelp‐associated macroinvertebrates across the latitudes of 46–53°S. Rafts of Durvillaea antarctica dominated the sightings (94% of rafts) and extrapolations to the full Southern Ocean between these latitudes indicate a figure of over 70 million rafts afloat at any one time, 20 million of which support a holdfast, the habitat supporting the highest faunal diversity in attached kelp plants. In contrast, few, small rafts of Macrocystis pyrifera were observed. The potential for dispersal of fauna is presumed to be related to the species of kelp with which they are associated. Empirical studies of survival of animals while drifting at sea, and also on making land‐fall, are required to allow fuller interpretation of the significance of these findings.     

Changes in the fish fauna associated with a sub-Antarctic <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis> kelp forest in response to canopy removal

The fish fauna associated with a Macrocystis pyrifera forest and the effects of the canopy removal on this fauna were studied in the Beagle Channel, Tierra del Fuego. Seasonal changes and differences in the fish communities that inhabit the canopy and the holdfast were also studied. Two patches of kelp forest separated by 200 m were selected. In one, the canopy was removed periodically while the other was used as a control. Samples were collected seasonally, from autumn 1999 to 2001. Fish fauna in the water column was sampled using trammel nets and holdfast fish fauna was sampled by removing the complete holdfast. Different assemblages of fish species were captured in the water column (surface and bottom) and in the holdfast. In the former there were mainly pelagic and benthopelagic species and the latter predominantly demersal species. The principal effect of the canopy removal was a drop in the abundance (and total weight) of Paranotothenia magellanica (doradito), the species with the strongest relationship with the canopy, principally at the surface of the treated patch probably due to a reduction in the availability of refuges. A high degree of seasonality was observed for fish species’ total weight, abundance and diversity, with higher values in summer and autumn. Taking into account our data and available data on kelp growth in the same locality, we suggest of cutting the kelp forest (preferably those farther away from the coast) once a year in winter or early spring, in order to minimize impact on the fish community.     

The kelp fauna of northeast Britain. I. Introduction and the physical environment

As part of a multidisciplinary investigation of pollution in northeast Britain a study of the sublittoral kelp fauna is described. Sampling strategy is discussed and a programme adopted involving the investigation of an instantaneous faunal pattern over a wide area and its environmental correlation. The kelp holdfast as a sample unit is briefly evaluated. The total faunal content of the survey samples is described in gross terms. A review of the physical parameters which impinge upon the holdfast fauna is made and on this basis significant variables are resolved into turbidity, pollution, water movement, and holdfast morphology. Detailed treatment of these factors follows and their distribution over the northeast coast transect is described.Seasonal studies on suspended solids enable an approximate area north of the River Coquet in Northumberland to be considered as continually clear, whilst the area to the south is considered turbid. The region of turbid water is considered to be the consequence of erosion of superficial coastal deposits under the action of marine and atmospheric forces. The rôle of discharges in contributing to suspended solids is considered.Nutrient data are given; the amounts were spatially similar over the whole coastline. An area of ‘pollution’ is provisionally designated on the basis of published figures.The problems of assessment of wave exposure and present inadequate knowledge of local inshore current systems preclude other than crude separation of one site on considerations of water movement. Holdfast parameters are generally interdependent: however the degree of holdfast branching is shown to be independent of age and to influence significantly (along with water movement external to the holdfast) the degree of holdfast silting.     

Evaluating stress in rocky shore and shallow reef habitats using the macrofauna of kelp holdfasts

Kelp often dominates hard rocky substrata in the coastal watersof temperate, boreal and subantarctic regions of the world's oceans. The habitat providedby kelp holdfasts supports high species richness and high abundance of a range of invertebratetaxa. Studies conducted on the effects of stress on these communities have indicated that theyare sensitive to stressors and that the response is often predictable. With the growing body ofliterature on natural variation in holdfast community structure, studies of the impacts of stressare becoming more readily interpretable making the macrofauna of this habitat a potentiallyimportant tool for evaluating anthropogenic effects on hard substrata in coastal environments. Limitations of the use of holdfast fauna in studies of stress include the fact that brownseaweeds may disappear from polluted areas of reef, and where they persist, holdfaststructure may be altered directly by the stressor. An additional problem is that manipulative,experimental studies are difficult. Artificial holdfasts may provide a means by which thedemonstrated sensitivity of the holdfast community can be combined with a manipulative approach to testspecific hypotheses related to effects of stress in shallow coastal environments.     

Survey of the terrestrial and freshwater molluscan fauna of the Pilliga forest area in northern inland New South Wales,Australia

The Pilliga forest in northern inland New South Wales, Australia, is one of the largest surviving remnants of native forest on the western slopes of the Great Dividing Range. The Pilliga landscape is a challenging environment for molluscs, dominated by dry sclerophyll forest and with limited and largely ephemeral aquatic habitats. A field survey of the area in 2006–2012 identified a surprisingly rich and relatively intact aquatic native molluscan fauna with five species of bivalves in three families and nine species of freshwater gastropods (four families), including some rare species and range extensions. The native land snail fauna comprised 18 species (six families), including an unusually rich pupillid fauna with nine species. Some range extensions are recorded and some species are narrow-range endemics. The distributions of many aquatic and terrestrial species were correlated with geology or soil type. Introduced molluscs were predominantly found in anthropogenic habitats and include two freshwater gastropods (two families) and nine terrestrial snails and slugs (eight families). This study provides insight into the original molluscan fauna of the western slopes prior to landscape-scale agricultural development and provides a benchmark for future reference.     

A Reverse Taxonomic Approach to Assess Macrofaunal Distribution Patterns in Abyssal Pacific Polymetallic Nodule Fields

Heightened interest in the exploitation of deep seafloor minerals is raising questions on the consequences for the resident fauna. Assessing species ranges and determination of processes underlying current species distributions are prerequisites to conservation planning and predicting faunal responses to changing environmental conditions. The abyssal central Pacific nodule belt, located between the Clarion and Clipperton Fracture Zones (CCZ), is an area prospected for mining of polymetallic nodules. We examined variations in genetic diversity and broad-scale connectivity of isopods and polychaetes across the CCZ. Faunal assemblages were studied from two mining claims (the eastern German and French license areas) located 1300 km apart and influenced by different productivity regimes. Using a reverse taxonomy approach based on DNA barcoding, we tested to what extent distance and large-scale changes in environmental parameters lead to differentiation in two macrofaunal taxa exhibiting different functions and life-history patterns. A fragment of the mitochondrial gene Cytochrome Oxidase Subunit 1 (COI) was analyzed. At a 97% threshold the molecular operational taxonomic units (MOTUs) corresponded well to morphological species. Molecular analyses indicated high local and regional diversity mostly because of large numbers of singletons in the samples. Consequently, variation in composition of genotypic clusters between sites was exceedingly large partly due to paucity of deep-sea sampling and faunal patchiness. A higher proportion of wide-ranging species in polychaetes was contrasted with mostly restricted distributions in isopods. Remarkably, several cryptic lineages appeared to be sympatric and occurred in taxa with putatively good dispersal abilities, whereas some brooding lineages revealed broad distributions across the CCZ. Geographic distance could explain variation in faunal connectivity between regions and sites to some extent, while assumed dispersal capabilities were not as important.     

Studying exotics in their native range: Can introduced fouling amphipods expand beyond artificial habitats?

Knowledge of the habitat use patterns of introduced species in their native or naturalized range can provide unique insights into processes of secondary dispersal and colonization of natural habitats. Caprellid amphipods are small mobile marine epibionts with limited natural dispersal. The global distribution of some caprellid species is mostly the result of anthropogenic transport; however, their subsequent spread beyond artificial habitats is poorly understood. A biogeographic approach, mainly focused on the native-range ecology of introduced and common fouling caprellid amphipods of southern Europe, was used to understand the implications of habitat use patterns for predicting their spread in the introduced regions. Specifically, abundance and composition of caprellid populations were compared among different primary habitats including artificial (floating pontoons), sheltered and wave-exposed rocky shores along the southern and southeastern coasts of Brazil. The findings indicated that artificial habitats act as reservoirs for globally distributed species in both their native and introduced ranges, while endemic species are more scarcely represented. Environmental conditions provided by primary habitats appear important in structuring caprellid assemblages on secondary substrata (basibiont species). Most wide-ranging caprellids were negatively correlated with the level of wave exposure, being more abundant in sheltered (artificial or natural) than in exposed habitats. In this context, Caprella scaura and Paracaprella pusilla, the two introduced caprellids recorded in the Mediterranean, where they are virtually restricted to artificial habitats, may become established in sheltered and even highly polluted natural habitats but hardly colonize wave-exposed rocky shores.     

Environmental factors structuring polychaete communities in shallow rocky habitats: role of physical stress versus habitat complexity

Polychaetes inhabiting 12 different hard bottom habitats were studied. A total of 157 species belonging to 32 families were identified. Differences among habitats in polychaete density, species richness, and diversity were analysed, as well as the relationships between these ecological indices and depth range, slope and in-bay/out-bay gradient. A high faunal homogeneity was found: all biotopes were dominated by a low number of eurytopic species. Intertidal habitats and subtidal ones with scarce algal cover were typified by vagile polychaetes (syllids, nereids), while sessile polychaetes (serpulids, sabellids) appeared typically among subtidal large macrophytes, habitats with a calcareous substrate and shaded habitats. Multivariate analyses showed that habitat complexity, determined by physical disturbance, is the main structuring factor for polychaete populations. Biotopes with the highest structural complexity displayed a high number of companion species increasing ecological indices and denoting a well-structured habitat. On the other hand, communities such as those in the upper intertidal, mainly controlled by physical environmental variables, showed a poorer polychaete fauna, dominated by ubiquitous species and a few well-adapted specialists.     

Intertidal colonization rates. A matched latitude, north v. south, remote v. near shore island experiment

Colonization of artificial substrata was monitored on sheltered rocky shores at Husvik, South Georgia (54° 11'S; 36° 40'W) and Cumbrae, Scotland (55° 46'N; 4° 55'W) from mid summer to early autumn. South Georgia is a remote island (1,330 km from other land); Cumbrae is only 2 km from nearby coasts. Both islands were heavily glaciated for a period up to about 10,000 ybp, so the intertidal fauna is of geologically recent origin. The South Georgian fauna is depauperate and consists mainly of direct developers; that of Cumbrae is rich and largely larviparous. Colonization rates at Cumbrae were one to four orders of magnitude greater than at Husvik. It is suggested that the fauna of South Georgian shores stems from colonization by rafting from remote sources, while Cumbrae has been supplied predominantly by short-range pelagic larval dispersal. The measured differences in colonization rates reflect the substantial local advantage of larval dispersal over direct development in established communities.     

The kelp fauna of northeast Britain. II. Multivariate classification: Turbidity as an ecological factor

Detailed faunal data relating to a sublittoral survey of kelp holdfasts (Laminaria hyperborea (Gunn.) Fosl.) in northeast Britain are presented. These data have been ordered by multivariate statistical techniques. Analytical strategy-is outlined and results pertaining to the classificatory techniques of association analysis (ASOC), predictive attribute analysis (PRED) and divisive information analysis (DINF) using qualitative data are given. Normal and inverse hierarchies wree computed using ASOC and DINF and used to construct two-way tables. Recognition of environmentally definie species assemblages is possible from a consideration of particular analyses. Taking into account certain methodological and sampling limitations, a definition of groups of ubiquitous, clear-water, turbidity indifferent and turbid water species may be recognized; the significant species for which purposes are only one-third of the total number considered.A primary correlation between the local composition of the holdfast fauna and turbidity is indicated. Subsidiary pollution effects cannot be ruled out but their definition becomes complicated by the intervention at lower levels of heterogeneity of other correlating factors, e.g., holdfast morphology. The danger of attributing causal relationships to statistical correlations is stressed throughout.     

Gradients in the number of species at reef-seagrass ecotones explained by gradients in abundance

Gradients in the composition and diversity (e.g. number of species) of faunal assemblages are common at ecotones between juxtaposed habitats. Patterns in the number of species, however, can be confounded by patterns in abundance of individuals, because more species tend to be found wherever there are more individuals. We tested whether proximity to reefs influenced patterns in the composition and diversity ('species density' = number of species per area and 'species richness' = number of species per number of individuals) of prosobranch gastropods in meadows of two seagrasses with different physiognomy: Posidonia and Amphibolis. A change in the species composition was observed from reef-seagrass edges towards the interiors of Amphibolis, but not in Posidonia meadows. Similarly, the abundance of gastropods and species density was higher at edges relative to interiors of Amphibolis meadows, but not in Posidonia meadows. However, species richness was not affected by proximity to reefs in either type of seagrass meadow. The higher number of species at the reef-Amphibolis edge was therefore a consequence of higher abundance, rather than species richness per se. These results suggest that patterns in the composition and diversity of fauna with proximity to adjacent habitats, and the underlying processes that they reflect, likely depend on the physiognomy of the habitat.     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Chance and necessity: land-snail faunas of Sao Miguel, Azores, compared with those of Madeira

The land-snail faunas of 23 sites from forest, disturbed andopen habitats were sampled on São Miguel, Azores archipelago.Collectively, the samples contained nearly all the known faunaof the island. Forest sites were the richest; while the numberof non-endemic species differed little between habitats, forestsheld more endemic species, many of which were widely distributed.Open habitats have no unique endemic species. These resultsare compared with similar data from Madeira. Although the twoislands are of the same size, and share the same Laurisilvaforest cover, their faunas differ in important aspects of diversity.While many non-endemic species are common to both islands, andoccur in similar numbers, endemic species are more numerouson Madeira, even though the numbers at individual forest sitesare, on average, fewer than on São Miguel. Many endemicsare confined to open, coastal habitats. These differences arediscussed in terms of the islands' histories, the range of habitatsavailable and the ease of dispersal from continental sources.Age, accidents of colonization and the range of natural habitatsavailable must have major effects on the faunas, but presentendemic faunal diversity is more a product of speciation andextinction than of immigration. (Received 13 May 2006; accepted 7 July 2006)     

Long‐term colonization ecology of forest‐dwelling species in a fragmented rural landscape – dispersal versus establishment

Species colonization in a new habitat patch is an efficiency indicator of biodiversity conservation. Colonization is a two‐step process of dispersal and establishment, characterized by the compatibility of plant traits with landscape structure and habitat conditions. Therefore, ecological trait profiling of specialist species is initially required to estimate the relative importance of colonization filters. Old planted parks best satisfy the criteria of a newly created and structurally matured habitat for forest‐dwelling plant species. We sampled species in 230 ancient deciduous forests (source habitat), 74 closed‐canopy manor parks (target habitats), 151 linear wooded habitats (landscape corridors), and 97 open habitats (isolating matrix) in Estonia. We defined two species groups of interest: forest (107 species) and corridor specialists (53 species). An extra group of open habitat specialists was extracted for trait scaling. Differing from expectations, forest specialists have high plasticity in reproduction mechanisms: smaller seeds, larger dispersules, complementary selfing ability, and diversity of dispersal vectors. Forest specialists are shorter, less nutrient‐demanding and mycorrhizal‐dependent, stress‐tolerant disturbance‐sensitive competitors, while corridor specialists are large‐seeded disturbance‐tolerant competitors. About 40% of species from local species pools have immigrated into parks. The historic forest area, establishment‐related traits, and stand quality enhance the colonization of forest specialists. The openness of landscape and mowing in the park facilitate corridor specialists. Species traits in parks vary between a forest and corridor specialist, except for earlier flowering and larger propagules. Forest species are not dispersal limited, but they continue to be limited by habitat properties even in the long term. Therefore, the shady parts of historic parks should be appreciated as important forest biodiversity‐enhancing landscape structures. The habitat quality of secondary stands can be improved by nurturing a heterogeneous shrub and tree layer, and modest herb layer management.     

广州地区7种菊科入侵植物丛枝菌根侵染和根际土壤孢子密度的生境差异性分析

为了解广州地区7种菊科(Compositae)入侵植物与丛枝菌根真菌(Arbuscular mycorrhizal fungi,AMF)的互作共生关系,调查了这7种菊科入侵植物在4种生境中的AMF侵染和根际土壤孢子密度,并分析土壤因子对AMF的影响。结果表明,7种入侵植物根内均受到AMF侵染,根际土壤中均检测到AMF孢子;AMF侵染在宿主间差异显著,生境间的差异不显著;孢子密度在生境和宿主间的差异均显著,人工绿地、农田果园的AMF孢子密度均显著高于森林周边和滨海地带。相关性分析表明,农田果园生境的根际土壤孢子密度与土壤有机质含量呈显著负相关关系;森林周边生境的AMF总侵染率与土壤全氮呈极显著正相关关系;人工绿地的AMF总侵染率与土壤速效氮含量呈显著负相关关系;滨海地带的AMF总侵染率与土壤有效磷含量呈显著负相关关系。这些对理解菊科植物入侵机理具有非常重要的作用。