Climate-driven range extension of a sea urchin: inferring future trends by analysis of recent population dynamics

Patterns of climate-forced range shift in the marine environment are informed by investigating the population dynamics of an ecologically important sea urchin ( Centrostephanus rodgersii – Diadematidae) across its newly extended range in Tasmania (southeastern Australia). A growth model of C. rodgersii is developed allowing estimation of a sea urchin age profile and, in combination with abundance data, we correlate the sea urchin population dynamic with respect to environmental signals across the range extension region. Growth patterns did not vary across the extension region; however, there was a strong pattern of decreasing sea urchin age with increasing distance from the historic range. The sequential poleward discovery of the sea urchin, a pattern of declining age and a general poleward reduction in abundance along the eastern Tasmanian coastline are consistent with a model of range extension driven by recent change in patterns of larval dispersal. We explore this hypothesis by correlating C. rodgersii population characteristics with respect to the East Australian Current (EAC), i.e. the chief vector for poleward larval dispersal, and reveal patterns of declining sea urchin age and abundance with increasing distance from this oceanic feature. Furthermore, C. rodgersii is generally limited to sites where average winter temperatures are warmer than the cold threshold for its larval development. Potential dispersal and physiological mechanisms defining the range extension appear to be strongly coupled to the EAC which has undergone recent poleward advance and resulted in coastal warming in eastern Tasmania. Predicted climate change conditions for this region will favour continued population expansion of C. rodgersii not only via atmospheric-forced ocean warming, but also via ongoing intensification of the EAC driving continued poleward supply of larvae and heat.     

Building Resilience Against Climate-Driven Shifts in a Temperate Reef System: Staying Away from Context-Dependent Ecological Thresholds

As climate-driven environmental changes and anthropogenic perturbations increasingly affect ecological systems, the number of abrupt phase shifts in ecosystem dynamics is rising, with far-reaching ecological, economic and social effects. These shifts are notoriously difficult to study, anticipate and manage. Although indicators of impending phase shifts in ecosystems have been described theoretically, they have only been observed empirically either after the fact or under controlled experiments. Here we demonstrate the usefulness of case-specific simulation models to estimate tipping points in the dynamics of real ecological systems, characterise how these thresholds may vary depending on local conditions and derive safe management targets associated with low risk of undesirable phase shifts. Under the combined effects of ocean changes and fishing, inshore rocky reefs in eastern Tasmania can transition from dense seaweed beds to sea urchin ‘barrens’ habitat, realising severe local loss of habitat, productivity and valuable fisheries. Using Monte-Carlo simulations with a validated model that realistically captures reef dynamics, we characterise the hysteresis in community dynamics and the variability in ecological thresholds along the gradient of environmental conditions. Simulation suggests that prevention of ongoing sea urchin destructive grazing of macroalgal beds is achievable but the yet-to-be-observed restoration of seaweed beds from extensive sea urchin barrens is highly unlikely. To guide management against undesirable phase shifts, we define target points associated with low risk of widespread barrens formation and show that, along with sea urchin culling, recognising the role of lobsters in mitigating sea urchin destructive grazing through predation is key to maintain reef productivity.     

Reproductive potential of a marine ecosystem engineer at the edge of a newly expanded range

Global climate change is leading to redistribution of marine species and altering ecosystem dynamics. Given recent poleward range extension of the barrens‐forming sea urchin Centrostephanus rodgersii (Diadematidae) from mainland Australia to Tasmania, there is a need to understand the population dynamics of this ecologically important species in the Tasmanian environment. This paper informs possible population dynamics of C. rodgersii in Tasmania by examining its reproductive ecology in this new environment. Reproductive periodicity (gonad index and propensity to spawn) was assessed bimonthly over 18 months at four sites in eastern Tasmania spanning ～2° in latitude. At all sites, C. rodgersii displayed a strong seasonal cycle in gonad production with major spawning occurring in winter (～August) at minimum annual water temperature. Gametes from Tasmanian C. rodgersii were viable as determined by fertilization and early development trials. However, development to the two‐arm stage at ～3 weeks was strongly dependent on water temperature across the 8–20 °C temperature range, with poor development occurring below 12 °C. The range of temperatures tolerated by Tasmanian C. rodgersii larvae was similar to that of larvae from its native New South Wales range, indicating that this species has not undergone an adaptive shift to the cooler Tasmanian environment. There was also no evidence for an adaptive shift in reproductive phenology. Importantly, coastal water temperatures in eastern Tasmania during the peak spawning in August fluctuate about the 12 °C larval development threshold. Recent warming of the eastern Tasmanian coast and further warming predicted by global climate change will result in an environment increasingly favourable for the reproduction and development of C. rodgersii.     

The effects of an invasive habitat modifier on the biotic interactions between two native herbivorous species and benthic habitat in a subtidal rocky reef ecosystem

Range expanding species can have major impacts on marine ecosystems but experimental field based studies are often lacking. The urchin Centrostephanus rodgersii has recently undergone a southerly range expansion to the east coast of Tasmania, Australia. We manipulated densities of C. rodgersii and algal regrowth in urchin barrens habitat to test effects of the urchin on biotic interactions between two native herbivores, black-lip abalone (Haliotis rubra) and another urchin (Heliocidaris erythrogramma), and their benthic habitat. After 13 months, removals of only C. rodgersii resulted in overgrowth of barrens habitat by algae and sessile invertebrates. Densities of abalone increased (+92 %) only in patches from which C. rodgersii was removed and algal regrowth allowed. In contrast, densities of H. erythrogramma increased in all treatments (+45, +28, +25 %) in which C. rodgersii was removed, irrespective of the algal regrowth manipulations. These results suggest that C. rodgersii has a negative influence on the densities of abalone through competition for food and on densities of H. erythrogramma through competition for preferred habitat. Densities of abalone (+65 %) but not H. erythrogramma (+25 %), were lower in the patches from which C. rodgersii and canopy algae regrowth were removed relative to patches from which only C. rodgersii was removed (+92 and +28 %, respectively). These results suggest that C. rodgersii overgrazing of canopy-algae results in loss of structural complexity which could increase abalone susceptibility to predation, cause abalone to seek shelter in cryptic microhabitats and/or prevent their return to patches where canopy algae are absent. The ongoing spread of C. rodgersii and expansion of barrens habitat in eastern Tasmania will continue to negatively affect populations of these two native herbivores and their associated fisheries at a range of spatial scales. This example shows that habitat modifying species which become highly invasive can have disproportionate negative impacts on the structure and dynamics of the recipient community.     

Effects of algal canopy clearance on plant, fish and macroinvertebrate communities on eastern Tasmanian reefs

Changes in assemblages of plants, macroinvertebrates and fishes on three eastern Tasmanian reefs were monitored over 12 months in replicated control blocks and adjacent 10×12-m blocks cleared of fucoid, laminarian and dictyotalean algae. Removal of canopy-forming plants produced less change to biotic assemblages than reported in studies elsewhere, with the magnitude of change for fish and invertebrate taxa lower than variation between sites and comparable to variation between months.The introduced annual kelp Undaria pinnatifida exhibited the only pronounced response to canopy removal amongst algal taxa, with a fivefold increase in cleared blocks compared to control blocks. Marine reserves are suggested to assist reef communities resist invasion by U. pinnatifida, through an indirect mechanism involving increased predation pressure on sea urchins and reduced formation of urchin barrens that are amenable to U. pinnatifida propagation.Large invertebrates were more associated with turfing algae or the reef substratum than the macroalgal canopy. The herbivorous sea urchin Heliocidaris erythrogramma and abalone Haliotis ruber showed the strongest response to clearing amongst common macroinvertebrate species, with a halving of population numbers. Observed densities of the common monacanthid fish Acanthaluteres vittiger also declined by about 50%. The relatively high level of resistance shown by eastern Tasmanian reef biota to patch disturbance was attributed largely to high diversity and biomass of turfing macroalgae damping effects of canopy clearance.     

Genetic structure of a recent climate change‐driven range extension

The life‐history strategies of some species make them strong candidates for rapid exploitation of novel habitat under new climate regimes. Some early‐responding species may be considered invasive, and negatively impact on ‘naïve’ ecosystems. The barrens‐forming sea urchin Centrostephanus rodgersii is one such species, having a high dispersal capability and a high‐latitude range margin limited only by a developmental temperature threshold. Within this species’ range in eastern Australian waters, sea temperatures have increased at greater than double the global average rate. The coinciding poleward range extension of C. rodgersii has caused major ecological changes, threatening reef biodiversity and fisheries productivity. We investigated microsatellite diversity and population structure associated with range expansion by this species. Generalized linear model analyses revealed no reduction in genetic diversity in the newly colonized region. A ‘seascape genetics’ analysis of genetic distances found no spatial genetic structure associated with the range extension. The distinctive genetic characteristic of the extension zone populations was reduced population‐specific F_ST, consistent with very rapid population expansion. Demographic and genetic simulations support our inference of high connectivity between pre‐ and post‐extension zones. Thus, the range shift appears to be a poleward extension of the highly‐connected rangewide population of C. rodgersii. This is consistent with advection of larvae by the intensified warm water East Australian current, which has also increased Tasmanian Sea temperatures above the species’ lower developmental threshold. Thus, ocean circulation changes have improved the climatic suitability of novel habitat for C. rodgersii and provided the supply of recruits necessary for colonization.     

Do sulphuric acid and the brown alga <Emphasis Type="Italic">Desmarestia viridis</Emphasis> support community structure in Arctic kelp patches by altering grazing impact,distribution patterns,and behaviour of sea urchins?

Macrobenthic community structure and the distribution of the green sea urchin (Strongylocentrotus droebachiensis) were recorded inside and outside (=barrens) of kelp patches (Alaria esculenta) at Kongsfjordneset, Svalbard between August 2002 and October 2006. In manipulative field experiments, conducted at Kongsfjordneset, Svalbard in August 2002, the effect of the presence of the brown seaweed Desmarestia viridis on sea urchin distribution and kelp grazing was determined. Additionally, we studied the effect of sulphuric acid, which is produced and stored by D. viridis, on sea urchin movements in the laboratory at Ny-Ålesund, Svalbard, in May 2006. Sea urchin densities were two- to threefold lower in kelp patches than on barrens. The macrobenthic community inside kelp patches hosted 39% more species and was of different species composition than on barrens. Anchored pieces of the kelp A. esculenta were less consumed when surrounded by D. viridis than non-surrounded conspecifics. Changes in pH affected the behaviour of sea urchins. Exposing sea urchins to 500 μl seawater at pH 7.5 caused them to stop, while the exposure of as little as 25 μl at pH 1 caused sea urchins to move in the opposite direction. Acid-mediated escape responses in sea urchin behaviour suggest chemical protection by D. viridis as an additional mechanism to mechanical protection in the generation of kelp refuges. These results improve our understanding of how isolated kelp beds can persist over a wide range of environmental conditions, like wave-sheltered sites, and suggest that changes in community structure may be in part attributable to altered trophic interactions.     

Changing Community States in the Gulf of Maine: Synergism Between Invaders, Overfishing and Climate Change

Human activities, including overfishing and species introductions, have had a dramatic impact on benthic communities in the Gulf of Maine within the past two decades. Prior to the 1970s, the climax community in the shallow subtidal was composed of Laminaria spp. kelp beds with an understory of arborescent red algae. In the 1980s, a population explosion of the green sea urchin, Strongylocentrotus droebachiensis, created an alternate community state, urchin barrens. Recently, a new community has been observed in former urchin barrens and kelp beds. This assemblage is principally composed of the introduced species: Codium fragile subsp. tomentosoides (green alga), Membranipora membranacea (bryozoan), Diplosoma listerianum (tunicate), Bonnemaisonia hamifera (red alga) and the opportunistic species Mytilus edulis (mussel) and Desmarestia aculeata (brown alga). In addition to changes in relative abundance, many of these species have greatly expanded their distribution and habitat selection. A model detailing mechanisms for the transition of the traditional kelp bed and urchin barren communities to others is presented and implications for this new community are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Global regime shift dynamics of catastrophic sea urchin overgrazing

A pronounced, widespread and persistent regime shift among marine ecosystems is observable on temperate rocky reefs as a result of sea urchin overgrazing. Here, we empirically define regime-shift dynamics for this grazing system which transitions between productive macroalgal beds and impoverished urchin barrens. Catastrophic in nature, urchin overgrazing in a well-studied Australian system demonstrates a discontinuous regime shift, which is of particular management concern as recovery of desirable macroalgal beds requires reducing grazers to well below the initial threshold of overgrazing. Generality of this regime-shift dynamic is explored across 13 rocky reef systems (spanning 11 different regions from both hemispheres) by compiling available survey data (totalling 10 901 quadrats surveyed in situ) plus experimental regime-shift responses (observed during a total of 57 in situ manipulations). The emergent and globally coherent pattern shows urchin grazing to cause a discontinuous ‘catastrophic’ regime shift, with hysteresis effect of approximately one order of magnitude in urchin biomass between critical thresholds of overgrazing and recovery. Different life-history traits appear to create asymmetry in the pace of overgrazing versus recovery. Once shifted, strong feedback mechanisms provide resilience for each alternative state thus defining the catastrophic nature of this regime shift. Importantly, human-derived stressors can act to erode resilience of desirable macroalgal beds while strengthening resilience of urchin barrens, thus exacerbating the risk, spatial extent and irreversibility of an unwanted regime shift for marine ecosystems.     

Survival of kelp adjacent to areas grazed by sea urchins in New South Wales,Australia

Abstract The sea urchin Centrostephanus rodgersii grazes areas of reef free of large brown algae (the barrens habitat). Survival of the kelp Ecklonia radiata adjacent to patches of barrens habitat was compared to that in the centre of kelp forests (centre) and edges of forests not adjacent to patches of barrens habitat (ungrazed). Estimates of rates of instantaneous mortality for tagged plants, as described by the slope of a negative exponential model (± 95% CI) were: barrens, 0.078 ± 0.004; centre, 0.051 ± 0.004 and ungrazed edge, 0.065 ± 0.007. Survival of plants was greatest in the middle of forests and least on the margins of patches of barrens habitat. A significant proportion of mortality in the barrens and centre positions was caused by herbivorous fish. When these plants were excluded from analysis there were no significant differences in survival between the barrens and ungrazed positions. It is concluded that C. rodgersii has little impact on the abundance of Ecklonia outside sharply defined boundaries.     

Herbivore control in connected seascapes: habitat determines when population regulation occurs in the life history of a key herbivore

Herbivore outbreaks often trigger catastrophic overgrazing events in marine macrophyte ecosystems. The sea urchin Paracentrotus lividus, the dominant herbivore of shallow Mediterranean seascapes, is capable of precipitating shifts to barrens when its populations explode. Paracentrotus lividus is found ubiquitously in rocky macroalgal communities and in sandy seagrass meadows of Posidonia oceanica, two of the most important subtidal habitats in the Mediterranean. We explored if habitat‐specific regulation across the principal stages of the urchin life cycle could help explain the persistence of these populations in connected mosaics. We measured each of three relevant ecological processes (i.e. settlement, post‐settlement survival and predation) across a wide stretch of the Mediterranean coast (ca 600 km). Our results show that habitat‐specific regulation is critical in determining urchin populations: each habitat limited urchin sub‐populations at different life stages. Settlement was never limiting; urchins settled at similar rates in both habitats across the coast. Post‐settlement survival was a clear bottleneck, particularly in seagrass meadows where no juvenile urchins were recorded. Despite this bottleneck in seagrasses, adult urchin populations were very similar in both seagrass and macroalgal habitats indicating that other processes (potentially migration) could be key in determining adult distributions across the mosaic. The fact that population regulation is clearly habitat‐specific suggests that sea urchin populations may be significantly buffered from bottlenecks in mixed seascapes where both habitats co‐occur. Sea urchin populations can therefore persist across the seascape despite strong habitat‐specific regulation either by maintaining reproductive output in one habitat or by migrating between them. By affording these regulatory escapes to habitat‐modifying species, patchy mosaics may be much more prone to herbivore outbreaks and a host of cascading effects that come in their wake.     

A COMPARISON OF DRAGON KELP,EUALARIA FISTULOSA, (PHAEOPHYCEAE) FECUNDITY IN URCHIN BARRENS AND NEARBY KELP BEDS THROUGHOUT THE ALEUTIAN ARCHIPELAGO1

The Aleutian Archipelago coastal ecosystem has undergone a dramatic change in community composition during the past two decades. Following the removal of ～99% of the sea otters, Enhydra lutris, from the ecosystem, changes to the benthic communities resulted in widespread losses to most of the region’s kelp beds and corresponding increases in the prevalence of urchin barrens. Within the urchin barrens, the few kelps that have remained are exposed to elevated light, nutrients and currents, all of which may enhance their physiological condition and thus result in greater fecundity. To explore this further, we examined patterns of sporophyte fecundity in the dominant canopy‐forming kelp, Eualaria fistulosa, in both urchin barrens and in nearby kelp beds at seven Aleutian Islands spanning a range of 800 km. We found that the average weight of E. fistulosa sporophyll bundles was significantly greater on sporophytes occurring in the urchin barrens than in the nearby kelp beds. Furthermore, the average number of zoospores released per cm² of sporophyll area was also significantly greater in individuals from the urchin barrens than the nearby kelp beds. When these two metrics were combined, our results suggest that individual E. fistulosa sporophytes occurring in the urchin barrens may produce as many as three times more zoospores than individual E. fistulosa sporophytes occurring in the nearby kelp beds, and thus they may contribute disproportionately to the following year’s sporophyte recruitment in both urchin barrens and the adjacent kelp beds.     

Long Term Variability in the Structure of Kelp Communities in Northern Chile and the 1997–98 ENSO

This is the first study on the south eastern Pacific coast of South America which details long term, interannual variability in the structure of subtidal rocky-bottom kelp-dominated communities before, during, and after the El Niño Southern Oscillation (ENSO) event of 1997–1998 in northern Chile (23^°S). The temporal patterns of the main components of these ecosystems, which included Macrocystis integrifolia, Lessonia trabeculata, echinoids and asteroids, were evaluated seasonally between 1996 and 2004. M. integrifolia demonstrated high interannual variability in temporal patterns of abundance. The 1997–1998 ENSO did not significantly modify the temporal patterns of Macrocystis, although local extinction of M. integrifolia beds occurred during negative thermal anomalies in 1999–2000 (La Niña event), facilitating the establishment of urchin dominated “barren grounds”. The abundance of Lessonia trabeculata showed little temporal variability, and this species dominated the deeper regions of the kelp assemblage (8–13 m depth). The structure of the kelp communities in the study area is regulated by a trophic cascade which modulates alternation between kelp dominated areas and sea urchin barrens. In this context, frequent and intense upwelling of cold water high in nutrients favors the establishment and persistence of kelp assemblages. During ENSO, coastal upwellings can mitigate superficial warming of coastal water and increase the nutrient concentration in the water column. Superficial warming during the 1997–1998 ENSO induced spawning by different species of echinoderms, which resulted in major recruitment of these species during 1999. Top-down events, such as the decrease in densities of the asteroids after the 1997–1998 ENSO event, favored increases in densities of benthic grazers, which caused significant decreases in abundance of M. integrifolia. The re-establishment of the adult fraction of the carnivore (starfish) guild coincided with a decrease in the density of sea urchins and thus re-establishment of the kelp. In the temperate south eastern Pacific, oceanographic events, which act on different spatial-temporal scales, trigger trophic cascades that act at local levels, producing interannual variability in the structure of kelp communities. On the other hand, considering the high macroinvertebrate diversity associated with kelp assemblages, the transitions between kelp-dominated areas and sea urchin barrens do not appear to significantly affect the biodiversity of these assemblages of benthic invertebrates.     

Alien marine fishes deplete algal biomass in the Eastern Mediterranean

One of the most degraded states of the Mediterranean rocky infralittoral ecosystem is a barren composed solely of bare rock and patches of crustose coralline algae. Barrens are typically created by the grazing action of large sea urchin populations. In 2008 we observed extensive areas almost devoid of erect algae, where sea urchins were rare, on the Mediterranean coast of Turkey. To determine the origin of those urchin-less 'barrens', we conducted a fish exclusion experiment. We found that, in the absence of fish grazing, a well-developed algal assemblage grew within three months. Underwater fish censuses and observations suggest that two alien herbivorous fish from the Red Sea (Siganus luridus and S. rivulatus) are responsible for the creation and maintenance of these benthic communities with extremely low biomass. The shift from well-developed native algal assemblages to 'barrens' implies a dramatic decline in biogenic habitat complexity, biodiversity and biomass. A targeted Siganus fishery could help restore the macroalgal beds of the rocky infralittoral on the Turkish coast.     

Increased macroalgal abundance following mass mortalities of sea urchins (Strongylocentrotus droebachiensis) along the Atlantic coast of Nova Scotia

Summary Recurrent outbreaks of disease between 1980 and 1983 caused catastrophic mortality of sea urchins (>260,000 t fresh weight) along 280 km (straight line distance) of the Atlantic coast of Nova Scotia. The complete elimination of sea urchins and concomitant development of fleshy macroalgal communities have occurred along different parts of this coast in different years. Macroalgal communities in areas where sea urchins died off 1, 3 and 4 years previously are compared to existing sea urchin-dominated barren grounds and to a mature kelp bed without sea urchins. Changes in macroalgal cover and species composition, and increases in biomass, density and size of kelp (Laminaria) species, characterize the succession from barren grounds to 3- and 4-year-old kelp beds. The greatest change occurred between one and three years following sea urchin mass mortality. Within 3 years, kelp beds attained a level of biomass (7.6 kg m^-2) comparable to that of mature beds. Recovery of sea urchin populations via recruitment of planktonic larvae has been slow and spatially variable. Large-scale reciprocal fluctuations in kelp and sea urchin biomass may characterize the trajectory of a dynamic system which cycles between two alternate community states: kelp beds and sea urchin-dominated barren grounds. Periodic decimation of sea urchin populations by disease may be an important mechanism underlying this cyclicity.     

The distribution of macroinvertebrates and fishes in Tasmanian estuaries

The distributions of 390 taxa of benthic macroinvertebrates collected in forty-eight estuaries and 101 fish species collected in seventy-five Tasmanian estuaries were related to geographical and environmental variables. Distribution patterns for the two taxonomic groups were largely congruent at both between and within-estuary scales. Faunal composition and the number of species collected at a site were primarily related to site salinity, the biomass of seagrass and tidal range. At the broader estuary scale, the distributions of macroinvertebrate and fish assemblages were primarily correlated with the presence of an entrance bar. Species richness varied with geographical location for both macrofauna and fishes, with highest numbers of species occurring in the Furneaux Group, north-eastern Tasmania and south-eastern Tasmania. These patterns primarily reflected differences in estuary type between regions rather than concentrations of locally endemic species. Although the majority of species collected during the study were marine vagrants, they constituted a very low proportion of total animal densities within estuaries. Only four species considered exotic to Tasmania were identifed. Nearly all species recorded from Tasmanian estuaries occurred widely within the state and have also been recorded in south-eastern Australia. Only 1% of estuarine fish species and < 5% of invertebrate species were considered endemic to the state. The generally wide ranges of species around Tasmania were complicated by (i) the absence of most species from the west coast (ii) a small (< 10%) component of species that occurred only in the north-east and Furneaux Group (eastern Bass Strait), and (iii) a few species (< 5%) restricted to other regions. The low number of species recorded from estuaries along the western Tasmanian coast reflected extremely low faunal biomass in that area. This depression in biomass on the west coast was attributed to unusually low concentrations of dissolved nutrients in rivers and dark tannin-stained waters which greatly restricted algal photosynthesis and primary production.     

Spatial analysis of Tasmania’s native vegetation cover and potential implications for biodiversity conservation

Summary The landscape modification model proposed by McIntyre and Hobbs (1999) was used to assess the modification of Tasmania’s native vegetation and its potential implications for biodiversity conservation. The inclusion of new ‘substates’ in the model allowed the varying degrees of landscape variegation and fragmentation observed in Tasmania to be quantified. The mapped extent of Tasmania’s native vegetation is approximately 5.06 million ha or 74% of the land area. The extent of native vegetation varies across bioregions from a low of around 36% in the Tasmanian Northern Midlands bioregion to a high of 94% in the Tasmanian West bioregion. Overall, the Tasmanian landscape can be described as medium variegated as the State retains 76% cover of native vegetation, by area. Two of Tasmania’s nine bioregions are in an intact state, four are variegated, and three are fragmented. Seven of the State’s 48 catchments are in an intact state, 24 catchments are variegated, and 17 are fragmented. Tasmania was estimated to support 33 760 patches of native vegetation. Fewer than 3% of these patches exceed 50 ha in area. Small and medium patches occur predominantly on freehold land with grazing as a major land use, whereas large patches occur predominantly on crown land with conservation and production forestry as the major land uses. One feature of the State’s native vegetation is the large tract of native vegetation ecosystems in western Tasmania. Opportunities arise to sustain the resilience of these native ecosystems both by consolidating the formal protection of vegetation within catchments such as the Arthur and Pieman and by strengthening environmental management in adjacent areas. Bioregions and catchments where climate change may be of particular concern for biodiversity conservation and management include the Tasmanian Northern Midlands bioregion and Cam catchment in north‐western Tasmania. The maintenance and enhancement of patches of remnant vegetation in these areas will be challenging and appears likely to require strategic, multiscale and coordinated natural resource management over decades. Limiting the loss of native vegetation across the entire range of landscape states in Tasmania appears essential to mitigate the further decline of biodiversity.     

Marine reserves demonstrate top-down control of community structure on temperate reefs

Replicated ecological studies in marine reserves and associated unprotected areas are valuable in examining top-down impacts on communities and the ecosystem-level effects of fishing. We carried out experimental studies in two temperate marine reserves to examine these top-down influences on shallow subtidal reef communities in northeastern New Zealand. Both reserves examined are known to support high densities of predators and tethering experiments showed that the chance of predation on the dominant sea urchin, Evechinus chloroticus, within both reserves was approximately 7 times higher relative to outside. Predation was most intense on the smallest size class (30-40 mm) of tethered urchins, the size at which urchins cease to exhibit cryptic behaviour. A high proportion of predation on large urchins could be attributed to the spiny lobster, Jasus edwardsii. Predation on the smaller classes was probably by both lobsters and predatory fish, predominantly the sparid Pagrus auratus. The density of adult Evechinus actively grazing the substratum in the urchin barrens habitat was found to be significantly lower at marine reserve sites (2.2ǂ.3 m^-2) relative to non-reserve sites (5.5ǂ.4 m^-2). There was no difference in the density of cryptic juveniles between reserve and non-reserve sites. Reserve populations were more bimodal, with urchins between 40 and 55 mm occurring at very low numbers. Experimental removal of Evechinus from the urchin barrens habitat over 12 months lead to a change from a crustose coralline algal habitat to a macroalgal dominated habitat. Such macroalgal habitats were found to be more extensive in both reserves, where urchin densities were lower, relative to the adjacent unprotected areas that were dominated by urchin barrens. The patterns observed provide evidence for a top-down role of predators in structuring shallow reef communities in northeastern New Zealand and demonstrate how marine reserves can reverse the indirect effects of fishing and re-establish community-level trophic cascades.     

Response of a southern temperate marsupial, the Tasmanian pademelon (Thylogale billardierii), to historical and contemporary forest fragmentation

Despite only limited Pleistocene glacial activity in the southern hemisphere, temperate forest species experienced complex distributional changes resulting from the combined effects of glaciation, sea level change and increased aridity. The effects of these historical processes on population genetic structure are now overlain by the effects of contemporary habitat modification. In this study, 10 microsatellites and 629 bp of the mitochondrial control region were used to assess the effects of historical forest fragmentation and recent anthropogenic habitat change on the broad-scale population genetic structuring of a southern temperate marsupial, the Tasmanian pademelon. A total of 200 individuals were sampled from seven sites across Tasmania and two islands in Bass Strait. High mitochondrial and nuclear genetic diversity indicated the maintenance of large historical population sizes. There was weak phylogeographical structuring of haplotypes, although all King Island haplotypes and three Tasmanian haplotypes formed a divergent clade implying the mid-Pleistocene isolation of a far northwestern population. Both the mitochondrial and nuclear data indicated a division of Tasmanian populations into eastern and western regions. This was consistent with a historical barrier resulting from increased aridity in the lowland 'midlands' region during glacial periods, and with a contemporary barrier resulting from recent habitat modification in that region. In Tasmania, gene flow appears to have been relatively unrestricted during glacial maxima in the west, while in the east there was evidence for historical expansion from at least one large glacial refuge and recolonization of Flinders Island.     

Predation of the sea urchin Heliocidaris erythrogramma by rock lobsters (Jasus edwardsii) in no-take marine reserves

The formation of sea urchin ‘barrens’ on shallow temperate rocky reefs is well documented. However there has been much conjecture about the underlying mechanisms leading to sea urchin barrens, and relatively little experimentation to test these ideas critically. We conducted a series of manipulative experiments to determine whether predation mortality is an important mechanism structuring populations of the sea urchin Heliocidaris erythrogramma in Tasmania. Tethered juvenile and adult sea urchins experienced much higher rates of mortality inside no-take marine reserves where sea urchin predators were abundant compared to adjacent fished areas where predators were fewer. Mortality of tagged (but not tethered) sea urchins was also notably higher in marine reserves than in adjacent areas open to fishing. When a range of sizes of sea urchins was exposed to three sizes of rock lobsters in a caging experiment, juvenile sea urchins were eaten more frequently than larger sea urchins by all sizes of rock lobster, but only the largest rock lobsters (> 120 mm CL) were able to consume large adult sea urchins. Tagging (but not tethering) juvenile and adult sea urchins in two separate marine reserves indicated that adult sea urchins experience higher predation mortality than juveniles, probably because juveniles can shelter in cryptic microhabitat more effectively. In a field experiment in which exposure of sea urchins to rock lobster (Jasus edwardsii) and demersal reef fish predators was manipulated, rock lobsters were shown to be more important than fish as predators of adult sea urchins in a marine reserve. We conclude that predators, and particularly rock lobsters, exert significant predation mortality on H. erythrogramma in Tasmanian marine reserves, and that adult sea urchins are more vulnerable than smaller cryptic individuals. Fishing of rock lobsters is likely to reduce an important component of mortality in H. erythrogramma populations.