Contrast in the importance of arrow squid as prey of male New Zealand sea lions and New Zealand fur seals at The Snares,subantarctic New Zealand

New Zealand sea lions (Phocarctos hookeri) are threatened by incidental bycatch in the trawl fishery for southern arrow squid (Nototodarus sloanii). An overlap between the fishery and foraging sea lions has previously been interpreted as one piece of evidence supporting resource competition for squid. However, there is currently no consensus about the importance of squid in the diet of New Zealand sea lions. Therefore, we investigated this importance independently of spatial and temporal differences in squid availability through a simultaneous study with sympatric New Zealand fur seals (Arctocephalus forsteri), a species known to target arrow squid. Diet sampling at The Snares (48°01′S 166°32′E), subantarctic New Zealand, in February 2012 coincided with peak annual catch in the nearby squid fishery. Diets were deduced by analyses of diagnostic prey remains from scats (faeces) and casts (regurgitations). The contribution of each prey species to the diet was quantified using the per cent index of relative importance (% IRI) that combined frequency of occurrence, mass and number of prey items. Arrow squid was a minor component in sea lion scats (2 % IRI), and none was found in their casts. In contrast, arrow squid was the major component in fur seal scats and casts (93 and 99 % IRI, respectively). This study found that New Zealand sea lions ate minimal squid at a time when squid was clearly available as evidenced by the diet of New Zealand fur seals; hence, there was no indication of resource competition between sea lions and the squid fishery.     

A demographic study of a subtidal population of the New Zealand articulate brachiopod Terebratella inconspicua

Size-frequency distributions from subtidal populations of Terebratella inconspicua (Sowerby, 1846) consistently display prominent modes at 18 to 22 mm shell length, representing many years accumulation of non-growing adults. In addition, these distributions contain seasonally variable modes made up of new recruits which are present in very high numbers towards the end of the breeding season, but are reduced to very low numbers by the end of their first year of life. Growth of recruits follows a sigmoid curve with considerable variability between individuals. Some adults survive at least 14 years. Females are 2 years old before they breed. Larvae are brooded internally by the parent and spawned in autumn and spring. The density of recruitment is positively correlated with the density of residents. Larvae commonly settle on the shells of large brachiopods; these recruits can survive to breed, but their survivorship as adults is lower than that of conspecifics attached to rock. Total mortality in the population is size (=age)-dependent; the mean life expectancy of a new recruit is 0.54 years compared with 2.57 years for a 1 year-old. Sources of mortality are considered.     

Shading facilitates sessile invertebrate dominance in the rocky subtidal Gulf of Maine

Dramatic shifts in community composition occur between vertical and horizontal rocky surfaces in subtidal environments worldwide, yet the forces mediating this transition are poorly understood. Vertical rock walls are often covered by lush, diverse communities of sessile suspension-feeding invertebrates, while adjacent horizontal substrates are dominated by algae, or corals in the tropics. Multiple factors, including light, sedimentation, water flow, and predation have been proposed to explain this pattern, but experimental tests of these hypotheses are lacking. We manipulated light level and predation to test whether variation in these mechanisms could be responsible for the shift in composition of sessile communities between vertical and horizontal surfaces in the rocky subtidal Gulf of Maine. Shaded horizontally oriented granite plots were dominated by invertebrates (e.g., ascidians, barnacles, bryozoans) after 25 months. Unshaded plots were dominated by macroalgae, which was virtually absent in shaded plots. Exclusion of grazers with cages had no effect on percent cover of invertebrates or algae. Preferential settlement of invertebrate larvae to shaded plots, due to larval behaviors such as negative phototaxis, did not seem to play a role. Shading likely affects post-settlement mortality of invertebrates by alleviating competition for space with algae, although greater abundance of micropredators in algal-dominated communities could also be important. Communities on shaded plots lacked many taxa present on natural wall communities, likely due to greater disturbance on horizontal substrates and/or lack of sufficient time for colonization of these taxa. These results suggest that light plays a key role in controlling the structure, composition, and function of shallow subtidal communities.     

The effects of catchment land use on water quality and macroinvertebrate assemblages in Otara Creek,New Zealand

The effect of catchment land use on water quality and macroinvertebrate communities was examined by using data gathered during a 2004 reconnaissance of nine sites in the Otara Creek, New Zealand. Data collected included macroinvertebrate, water chemistry and sediments characteristics. Macroinvertebrate data were used in metric and index calculations. A total of 61 macroinvertebrate taxa, with 3032 total individuals, were identified from the macroinvertebrates samples collected from nine sites in Otara Creek. The greatest number of macroinvertebrate taxa was recorded within bush sites (mean>25), while the urban sites had the least number of taxa (mean=10). Pasture sites were intermediate with the mean>17. Taxa number differed significantly across land use. Mean macroinvertebrates abundance varied across the sites and land uses. The highest macroinvertebrates mean abundance was recorded in urban and pasture sites, while bush sites had significantly lower mean abundance. Physico-chemical parameters decreased from bush toward urban streams. Biotic indices were sensitive to changes in macroinvertebrates community structure across land uses with mean scores decreasing from bush to urban and pasture streams. Ordination of biological data showed a clear separation of bush from urban and pastures streams. Analysis of similarities revealed significant differences in macroinvertebrates between both stream groups and land-use groups. The observed macroinvertebrate assemblage pattern was best correlated with a single variable, conductivity, temperature, turbidity, nitrate and dissolved oxygen. The combination of these environmental variables best explained the changes in the macroinvertebrate assemblages between sites. This study demonstrates that catchment land use may significantly affect the water quality and macroinvertebrate communities in an ecosystem.     

Differential responses of vertebrate and invertebrate herbivores to traits of New Zealand subalpine shrubs

Plant traits are influenced by herbivore diet selection, but little is known about how traits are affected by different types of herbivores. We related eight traits of 27 subalpine shrub species in South Island, New Zealand, to damage of these shrubs by introduced red deer (Cervus elaphus) and native invertebrate herbivores using phylogenetically explicit modeling. Deer preferentially consumed species that grew quickly, were low in foliar tannins, or had high leaf area per unit mass. However, these traits did not trade off against each other; rather, they could be related to different multivariate defense strategies. Although the proportion of leaves damaged by leaf-chewing invertebrates also increased with stem growth, invertebrates did not damage the same fast growing species as those preferred by deer. Other traits may also be important in determining herbivore preferences, as suggested by the high proportion of variation in herbivory explained by phylogeny. Last, we found that the composition of invertebrate herbivore communities was more similar among closely related shrubs, and consequently, the range of invertebrate-plant associations may change if introduced deer shift plant composition toward slow-growing species. Overall, our results demonstrate the importance of herbivore type and coevolved interactions for the adaptive significance of plant traits.     

The invasibility of marine algal assemblages: role of functional diversity and identity

The emergence of the biodiversity-ecosystem functioning debate in the last decade has renewed interest in understanding why some communities are more easily invaded than others and how the impact of invasion on recipient communities and ecosystems varies. To date most of the research on invasibility has focused on taxonomic diversity, i.e., species richness. However, functional diversity of the communities should be more relevant for the resistance of the community to invasions, as the extent of functional differences among the species in an assemblage is a major determinant of ecosystem processes. Although coastal marine habitats are among the most heavily invaded ecosystems, studies on community invasibility and vulnerability in these habitats are scarce. We carried out a manipulative field experiment in tide pools of the rocky intertidal to test the hypothesis that increasing functional richness reduces the susceptibility of macroalgal communities to invasion. We selected a priori four functional groups on the basis of previous knowledge of local species characteristics: encrusting, turf, subcanopy, and canopy species. Synthetic assemblages containing one, two, three, or four different functional groups of seaweeds were created, and invasion by native species was monitored over an eight-month period. Cover and resource availability in the assemblages with only one functional group showed different patterns in the use of space and light, confirming true functional differences among our groups. Experimental results showed that the identity of functional groups was more important than functional richness in determining the ability of macroalgal communities to resist invasion and that resistance to invasion was resource-mediated.     

Spatial variability of infaunal nematode and polychaete assemblages in two muddy subtidal habitats

Spatial variation of populations and their assemblages is an important component of many aspects of ecology, including the maintenance of species diversity. In nature, organisms are generally aggregated in patches or form gradients or other spatially related patterns. This investigation quantified the degree of spatial structure in benthic invertebrate assemblages. It examined the distribution of infaunal assemblages of different size, mobility and contrasting life-history strategies (i.e. meiofaunal nematodes and macrofaunal polychaetes) in two offshore muddy habitats in the Celtic Deep and the NW Irish Sea off the west coast of the United Kingdom. The more heterogeneous habitat in the NW Irish Sea was characterised by higher tidal stress and bottom temperature while greater water depth, mean particle diameter and organic carbon content was typical for the comparatively homogeneous environment in the Celtic Deep. In both habitats, the environmental conditions became increasingly dissimilar with separation. A total of 125 nematode and 88 polychaete species were recorded with 69% of the nematode and 49% of polychaete species present at both study sites. Occurrence frequency of species, species diversity and average similarity of assemblage composition was higher in the Celtic Deep than in the NW Irish Sea. Results from correlation analyses revealed statistically robust relationships of community similarity and sample distance. Given their small size and low mobility, nematodes were more susceptible to within-habitat physical variability than larger-sized, more mobile polychaetes. This, coupled with limits to long-distance dispersal and likely restrictions in gene flow, resulted in a significant decrease in community similarity with distance at the spatial scales sampled (i.e. within 0.1–23 km). Polychaetes, in contrast, combined a higher dispersal potential (>23 km) with a relatively high tolerance to within-habitat environmental variability and these were the most likely causes for non-significant relationships between the similarity of their assemblages and sample distance. The potential mechanisms causing the observed variation and implications of results for environmental management strategies are discussed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Vertical stratification of mobile epiphytal arthropod assemblages between the canopy and understorey of subtidal macroalgae

This study examined patterns of faunal stratification between vertical strata by partitioning subtidal macroalgal communities into vertically contiguous (and hence paired) canopy and understorey habitats. Such floristic strata are often clearly defined in temperate subtidal communities and have been the focus of a range of other, primarily botanical, studies. Previous studies that have touched upon this subject have used statistical tests that require spatially independent samples, yet have collected samples that are often spatially contiguous and therefore not statistically independent. Here an analytical approach for examining faunal stratification in subtidal macroalgal assemblages using split-plot ANOVA for univariate data and a novel multivariate test suitable for analysing assemblage-level paired-sampled data is proposed. Differences in faunal assemblage structure between habitats were primarily attributed to the distribution of two functional feeding groups: filter-feeding and scavenging arthropods. Filter-feeders were more abundant amongst the canopy, whereas scavengers were more abundant in the understorey. No pattern was detected for herbivorous arthropods. These patterns are consistent with the feeding ecology of these functional groups and reflect expected differences in the relative availability of trophic resources between habitats, commensurate with the physical structure of the environments. Contrary to expectation, understorey algal habitats supported faunal assemblages that were as diverse, or more diverse than the canopy habitats, despite comprising smaller habitable spaces (in terms of algal biomass) and generally supporting fewer faunal individuals. Trophic stratification of epifaunal assemblages is suggested as a mechanism whereby faunal diversity is enhanced in these environments.     

Drivers of Seabird Population Recovery on New Zealand Islands after Predator Eradication

Eradication of introduced mammalian predators from islands has become increasingly common, with over 800 successful projects around the world. Historically, introduced predators extirpated or reduced the size of many seabird populations, changing the dynamics of entire island ecosystems. Although the primary outcome of many eradication projects is the restoration of affected seabird populations, natural population responses are rarely documented and mechanisms are poorly understood. We used a generic model of seabird colony growth to identify key predictor variables relevant to recovery or recolonization. We used generalized linear mixed models to test the importance of these variables in driving seabird population responses after predator eradication on islands around New Zealand. The most influential variable affecting recolonization of seabirds around New Zealand was the distance to a source population, with few cases of recolonization without a source population ≤25 km away. Colony growth was most affected by metapopulation status; there was little colony growth in species with a declining status. These characteristics may facilitate the prioritization of newly predator‐free islands for active management. Although we found some evidence documenting natural recovery, generally this topic was understudied. Our results suggest that in order to guide management strategies, more effort should be allocated to monitoring wildlife response after eradication. Conductores de la Recuperación de Poblaciones de Aves Marinas en Islas de Nueva Zelanda después de la Erradicación de Depredadores     

Habitat and reef-fish assemblages of banks in the Northwestern Hawaiian Islands

The moderately deep terraces and banks of the Northwestern Hawaiian Islands (NWHI) were surveyed to describe their habitat and reef-fish assemblages. These tracts of bottom at 30–40 m comprise more than 4,500 km² of the regions reef area. The habitat was found to be dominated by algal meadows (65% cover of exposed bottom), with infrequent relief features. Annual monitoring of select stations for 4 years at Necker Bank indicated that the relative difference in algal abundance between stations persisted from year to year (at least in summer). Temperature records from year-long deployments of archival thermistors in high-cover (>70%) and low-cover (<30%) algal biotopes were indistinguishable, providing no explanation of the algal differences between stations. At all banks, Microdictyon was the primary alga, averaging 1.22 kg/m². In spite of the extensive standing primary production, and a historical lack of fishing, bank reef-fish populations were impoverished. Mean densities, sizes, and biomass of trophic groups were considerably less than values reported for NWHI reef shallows. An overall mean biomass was estimated at 22.5 g/m², which is a fifth of that reported for shallow reefs of the region. Fish biomass of all trophic groups was associated with the few sources of relief available on the banks. Apex predators (sharks, jacks, and snappers), common on all surveys (with a mean of five per station), were proposed to constrain fish populations to sparse sources of relief resulting in a skewed size structure of the two primary fish trophic components. Sizes of lower-level carnivores were tightly correlated with sources of relief whereas the size of herbivores were not, indicating that herbivores more often venture out and risk the exposed algal meadows. These bank summits are a rare example of a near pristine reef system with high benthic primary productivity and low fish biomass, and are a stark contrast to shallower coral-reef ecosystems of the NWHI.Communicated by P.W. Sammarco, Chauvin     

Foliose algal assemblages and deforested barren areas: phlorotannin content,sea urchin grazing and holdfast community structure in the Aleutian dragon kelp,Eualaria fistulosa

In the Aleutian Archipelago, two distinct organizational states of kelp forest communities exist, foliose algal assemblages and deforested barren areas. The canopy-forming kelp Eualaria fistulosa can be found in both states, although it is much less abundant in the deforested state. In contrast, sea urchins also occur in both states, but they are considerably more abundant in the deforested state. This study determined whether the protective phlorotannin content in E. fistulosa sporophylls that originated in foliose algal assemblages would differ from those that originated in deforested areas. We also examined sea urchin grazing rates on these sporophylls to determine whether there were any differences in grazing rates related to the origin of the sporophylls. Next, we determined whether taxon richness and abundance of E. fistulosa holdfast communities would differ depending on the origin of the holdfast. Our results showed that sporophylls that were collected in barren areas generally had higher phlorotannin content than sporophylls found in foliose algal assemblages, although phlorotannin content varied across the study area. Grazing rates on sporophyll tissue and holdfast community structure did not differ between foliose algal assemblages and barren areas. These results show that phlorotannin content is greatest in areas under high grazing pressure and that phlorotannins may possibly protect E. fistulosa and the holdfast communities from being grazed in barren areas.     

Genetic structure of the subtidal red alga Delisea pulchra

We used random amplified polymorphic DNA (RAPDs) to examine small-scale spatial genetic structure in the red alga Delisea pulchra (Greville) Montagne at two locations near Sydney, Australia. We examined genetic structure among plants at four spatial scales ranging from 2 km apart down to <50 cm apart between locations, among sites within locations, among quadrats within sites, and among plants within quadrats. Haploid stages of D. pulchra were absent from the populations studied, suggesting that they are maintained through asexual reproduction of diploid plants. Consistent with this, we found that 19 RAPD phenotypes scored in this study had multiple individuals, indicating the presence of clones in these populations. However, there were no RAPD phenotypes common to two locations separated by only 2 km. Analysis of molecular variance revealed that strong genetic differences occurred between plants from these two locations, with 46.3% of the total genetic variation occurring at this scale, most probably reflecting limited gene flow. Within each location, <25% of the genetic variation was attributable to differences among sites or quadrats, indicating gene flow at those smaller scales. Most of the variation within each location occurred at the smallest spatial scale, among plants within 0.25 m² quadrats. Nonetheless, some pairwise genetic distances (φST) between sites or quadrats within locations were large, indicating some genetic divergence on smaller scales. Genetic distance was independent of spatial distance within both locations, suggesting that fine-scale differences within locations were most probably caused by variation in fine-scale patterns of water movement or fine-scale natural selection. We assessed the impact of one potential selective agent, grazing sea urchins, on the fine-scale genetic structure of D. pulchra. There was no evidence that grazing by sea urchins affected the genetic structure of D. pulchra. In combination with demographic data, our results indicated that local populations of D. pulchra within locations were relatively open and that fine-scale genetic structure was probably constrained by gene flow. At the larger scale however, strong genetic differentiation indicated little gene flow between locations and restricted dispersal of spores. Received: 22 April 1999 / Accepted: 29 November 1999     

Effects of long-term physical disturbance by commercial scallop fishing on subtidal epifaunal assemblages and habitats

 This paper examines spatial differences in the distribution of by-catch assemblages from the scallop [Pecten maximus (L.) and Aequipecten opercularis (L.)] fishing grounds in the North Irish Sea, during 1995. The sites examined have been exposed to differing known levels of fishing disturbance by scallop dredging, based on unusually high-resolution data extracted from fishermens' logbooks. Uni- and multi-variate techniques have been used on a production dataset (a value which incorporates both abundance and biomass figures), as well as abundance and biomass data individually. The original species list was reduced to higher taxonomic groupings in line with the theory that the latter is more appropriate for detecting anthropogenic change. Species diversity and richness, total number of species, and total number of individuals all decrease significantly with increasing fishing effort. Species dominance increases with effort. Total abundance, biomass and production, and the production of most of the major individual taxa investigated decrease significantly with increasing effort. Multivariate analysis reveals a significant relationship between fishing effort and by-catch assemblage structure. The taxa most responsible for the differences are the echinoids and cnidarians, but prosobranch molluscs and crustaceans also contribute to the differences. By-catch assemblage structure is more closely related to fishing effort than any other environmental parameter investigated, including depth and sediment type. We observed an approximately linear decrease in diversity with increasing fishing disturbance, and suggest this is primarily due to selective removal of sensitive species and, more importantly, habitat homogenisation. These results were interpreted in the light of ecological theories relating disturbance to community structure. The argument that invertebrate scavenger populations benefit from prolonged exposure to fishing disturbance was also examined, but no supporting evidence was found. Received: 29 January 2000 / Accepted: 31 May 2000     

Long-term variability in the structure of subtidal benthic communities in Puget Sound,Washington, USA

Data on benthic infauna from 4 permanent stations in Puget Sound off Seattle, USA, collected during 1963–1964, 1967, and 1969, revealed considerable stability in numbers of species and specimens and in diversity within stations among sampling dates. The species composition of the faunal assemblages also remained rather constant during the period of investigation, but the relative dominance among the numerically important species varied somewhat. Biomass data did not differ significantly in 1964 and 1969, but the 1967 data were considerably lower at all stations.     

Spatio-temporal structure of pelagic larval and juvenile fish assemblages in coastal areas of New Caledonia,southwest Pacific

This study aims at describing the diversity and composition of larval and juvenile fish assemblages in coastal areas of New Caledonia, southwest Pacific, and identifying the environmental factors that influence the seasonal and spatial patterns of these assemblages. A total of 97 taxa belonging to 7 orders and 26 familis were captured in three bays near Nouméa by light trapping every month between January 2002 and June 2003. The assemblages were dominated by Clupeiform larvae and juveniles (96.4% of total abundance) and followed by Perciform larvae (3%). The number of taxa per sample varied from less than five in July–August to more than ten in October–November and abundances followed the same seasonal pattern. Analyses of similarity showed significant differences in the assemblages caught in the three bays and analyses of contribution to the dissimilarity revealed that these differences were due to the most abundant families. The constant part of the relationship between environmental variables and the composition of assemblages was assessed by the partial triadic analysis STATICO, a statistical approach that takes into account the strong seasonality of the data. Rainfall, wind direction and thermal stratification of the water column were found to play a major role in the structure of the assemblages, although tidal amplitude and wind speed became important when Clupeidae and Engraulididae were excluded from the analyses. The richness, relative abundances and seasonal variations of the assemblages caught in three bays under study are close to what has been observed elsewhere in the tropics. This study shows the efficiency of the STATICO analysis for identifying the environmental factors that have a permanent effect on assemblages and sorting them out from those which act temporally or on specific locations. The high abundances and diversity of coral-reef fish larvae observed in coastal zones of New Caledonia suggest that further studies are needed to fully explore the role of the coastal zones of New Caledonia as nurseries.     

Limited change in the diversity and structure of subtidal communities over four decades

A unique archive of photographs from 1969 to 1974 permitted a test of the hypothesis that the diversity and composition of contemporary epilithic communities on subtidal rock walls in the San Juan Islands, WA, USA, has changed over 40 years. Notably, the richness and diversity of sessile taxa was significantly higher in 2008–2011. Furthermore, the multivariate community structure of sessile and mobile taxa differed between the historic and modern eras. Historic communities were characterized by a high percent cover of bare rock and non-calcified algal crusts, consistent with the effects of grazing by chitons and urchins. The rate of sessile community turnover, an index less susceptible to spatial sampling artifacts than richness or diversity, was not significantly different between the two eras. Together with the naturally high spatial variability in these epilithic communities, and the limited replication of historic quadrats, we interpret cautiously the data as evidence of limited change despite a clear shift in temperature and predator (fish) guild composition. The lack of substantial change in rock wall communities may be due in part to their vertical topography, which limits physical disturbance and the preemption of space by weedy algae, two processes that are often associated with “phase-shifts” in other marine ecosystems.     

Wave exposure,swimming performance,and the structure of tropical and temperate reef fish assemblages

We examined the relationship between swimming performance, wave exposure, and the distribution patterns of labrids on temperate rocky reefs, in comparison with previous functional analyses of a tropical assemblage. Visual censuses of the distribution and abundance of labrids across two major gradients of wave exposure (depth and aspect to prevailing winds) were made at two offshore islands near Port Stephens, New South Wales, Australia. Distinct shifts in species composition and abundance were evident between high and low wave exposure habitats on temperate rocky reefs, particularly between deep and shallow habitats on exposed reef fronts. The swimming performances of temperate labrids were assessed through examination of pectoral fin shape (aspect ratio) and in situ swimming speeds. A diversity of pectoral fin morphologies was exhibited within this temperate assemblage, ranging from rounded to tapered fins (aspect ratios of 0.52 and 1.43, respectively). Fin shape was strongly correlated (Pearsons correlation 0.884, P<0.001) with swimming speed (ranging from 1.05 and 3.06 body lengths s^–1), in a relationship comparable to that observed in tropical labrids. Inter-specific differences in swimming ability provided some explanation for differences in the distribution and abundance of temperate labrids in relation to wave exposure. However, our findings suggest that although coral reef labrids appear to predominantly use high aspect-ratio fins to successfully occupy wave-exposed habitats, temperate labrids appear to be using an enhanced swimming ability through increased body size.Communicated by G.F. Humphrey, Sydney