Method of estimating the standing stock of Trochus niloticus incorporating Landsat satellite data,with application to the trochus resources of the Bourke Isles,Torres Strait,Australia

The habitat of Trochus niloticus in the Bourke Isles, Torres Strait, was classified into areas of a Landsat image, using high-ratio values of green (Band 2) to red (Band 3) light, along the windward reef margins. These shallow-water (< 15 m) areas have a coral and rubble/algal pavement cover, which constitutes the optimal habitat for this gastropod. The habitat was sampled to estimate the abundance of T. niloticus. The proportion of commercial-sized individuals was estimated by measuring the basal width of all individuals in a sample. A multistage sample design incorporating three spatial scales -100 m² (transect), 1500 m² (site) and 1 km² (reef) — was used to provide variance estimates for sample-design optimisation and to provide data on the spatial variation of abundance. Most variation (68%) in abundance was within reefs and was attributable to differences in reef cover. Variations in abundance and time costs for sampling 2 and 4 m transects were compared; the 2 m transect was more efficient than the 4 m transect. The abundance estimates were combined with habitat-area estimates and the proportion of commerical-sized individuals was estimated at a standing stock of 186000 (24% precision), or 14 t of commerical-sized T. niloticus.     

Density and distribution patterns of the temperate marine fish Cheilodactylus spectabilis (Cheilodactylidae) in a reef environment

Cheilodactylus spectabilis (Hutton) is common over shallow reefs in north-eastern New Zealand. Replicated transect between-area differences in density and in size frequency. C. spectabilis is not nocturnally active. The between-area differences in density and size frequency remained constant over daylight hours, demonstrating that C. spectabilis does not undergo any systematic feeding migrations. Topographic complexity was shown to have a significant positive influence on fish density. Mean size was related to water depth at 5 localities investigated, with small (<200 mm standard length, SL) individuals being restricted to shallow water. All sizes of C. spectabilis examined had fed on small invertebrates, with gammarid amphipods predominating. No evidence of size-related differentiation in feeding patterns was observed, although small individuals spent significantly more time feeding than large ones. Small (<200 mm SL) individuals occupied feeding areas and shelter sites from which they excluded other small C. spectabilis. Large individuals showed no site-associated aggression, covered a greater area during daily movements and overlapped with other large and small C. spectabilis while feeding. No evidence of dawn or dusk peaks of feeding activity or movement was collected.     

Latitudinal variation in abundance of herbivorous fishes: a comparison of temperate and tropical reefs

The aim of the study was to provide comparable estimates of abundance of herbivorous reef fishes at temperate and tropical localities using a standardized methodology. Faunas of herbivorous fish were sampled on the rocky reefs of temperate northern New Zealand and on the coral reefs of the northern Great Barrier Reef (GBR), Australia, and the San Blas Archipelago in the Caribbean. A pilot study established the most appropriate habitat setting and the scale and magnitude of replication for the sampling program in temperate waters. Herbivorous fishes, including members of families endemic to the southern hemisphere (Odacidae and Aplodactylidae), were most abundant in turbulent, shallow water (0 to 6 m) and had patchy distributions within this habitat. A hierarchical sampling program using 10-min transect counts within the 0 to 6 m depth stratum examined abundance patterns at a range of spatial scales including mainland and island coasts, localities separated by up to 100 km and sites separated by up to 10 km. This program identified a characteristic fauna of seven species of herbivorous fishes with mean total abundances ranging from 23 to 30 individuals per 10-min transect. Species composition of the fauna varied between islands and coasts. A similar methodology was used to sample the major families of herbivorous fish in a number of sites in each of the tropical regions. These sampling programs revealed a fauna dominated by acanthurids and scarids in both the GBR and Caribbean localities. Estimates of abundance from these regions were similar, with a mean of 108 individuals recorded on the GBR and 129 per 10-min transect in the Caribbean. Species richness varied between each region, with 44 taxa recorded from the GBR and 11 from the Caribbean. Abundances of temperate water herbivores in New Zealand were found to be 75 to 80% lower than those recorded from shallow water habitats sampled on coral reefs. This was not related to species richness, since both New Zealand and the Caribbean locality had patterns of low richness. We suggest that the differences in abundance found by our study between temperate and tropical regions are not restricted to herbivorous fishes, but are representative of general latitudinal trends in reef fish faunas. Received: 4 November 1996 / Accepted: 15 December 1996     

Habitat harshness and morphodynamics: life history traits of the mole crab Emerita brasiliensis in Uruguayan sandy beaches

The habitat harshness hypothesis (HHH) postulates that in reflective beaches the harsh environment forces organisms to divert more energy towards maintenance and they therefore have lower abundance, fecundity, growth and survival rates than in dissipative beaches. Recent investigations have tested this hypothesis through single comparisons of only two beaches, and thus the observed trends in population level variables cannot be attributed incontestably to the beach state, but only to location. Here, abundance, reproduction, recruitment, population structure and body size of the intertidal mole crab Emerita brasiliensis were compared between populations from eight microtidal exposed sandy beaches with contrasting morphodynamics, sampled bimonthly during 22 months throughout the 180 km Uruguayan Atlantic coast. Physical variables and compound indices of the beach state were used to categorize sandy beaches. The results of this bi-annual large-scale analysis were fully consistent with the predictions of the HHH: abundance (total and population components), duration of the reproduction and recruitment seasons and the individual size of megalops and females of the mole crab E. brasiliensis decreased from dissipative to reflective beaches. This was reflected by linear or, mostly, nonlinear relationships between biological and both physical variables and compound indices of beach state. In conclusion, this multi-beach sampling provides compelling evidence of a consistent response of demographic and life history traits of an intertidal beach species to morphodynamic characteristics.     

Determining Optimal Population Monitoring for Rare Butterflies

Abstract: Determining population viability of rare insects depends on precise, unbiased estimates of population size and other demographic parameters. We used data on the endangered St. Francis' satyr butterfly (Neonympha mitchellii francisci) to evaluate 2 approaches (mark–recapture and transect counts) for population analysis of rare butterflies. Mark–recapture analysis provided by far the greatest amount of demographic information, including estimates (and standard errors) of population size, detection, survival, and recruitment probabilities. Mark–recapture analysis can also be used to estimate dispersal and temporal variation in rates, although we did not do this here. Models of seasonal flight phenologies derived from transect counts (Insect Count Analyzer) provided an index of population size and estimates of survival and statistical uncertainty. Pollard–Yates population indices derived from transect counts did not provide estimates of demographic parameters. This index may be highly biased if detection and survival probabilities vary spatially and temporally. In terms of statistical performance, mark–recapture and Pollard–Yates indices were least variable. Mark–recapture estimates were less likely to fail than Insect Count Analyzer, but mark–recapture estimates became less precise as sampling intensity decreased. In general, count‐based approaches are less costly and less likely to cause harm to rare insects than mark–recapture. The optimal monitoring approach must reconcile these trade‐offs. Thus, mark–recapture should be favored when demographic estimates are needed, when financial resources enable frequent sampling, and when marking does not harm the insect populations. The optimal sampling strategy may use 2 sampling methods together in 1 overall sampling plan: limited mark–recapture sampling to estimate survival and detection probabilities and frequent but less expensive transect counts.     

Seals at sea: modelling seal distribution in the German bight based on aerial survey data

The Wadden Sea is an important habitat for harbour seals and grey seals. They regularly haul-out on sandbanks and islands along the coast. Comparably little is known about the time seals spend at sea and how they use the remainder of the North Sea. Yet, human activity in offshore waters is increasing and information on seal distribution in the North Sea is crucial for conservation and management. Aerial line transect surveys were conducted in the German bight from 2002 to 2007 to investigate the distribution and abundance of marine mammals. Distance sampling methodology was combined with density surface modelling for a spatially explicit analysis of seal distribution in the German North Sea. Depth and distance to coast were found to be relevant predictor variables for seal density. Density surface modelling allowed for a depiction of seal distribution in the study area as well as an abundance estimate. This is the first study to use aerial survey data to develop a density surface model (DSM) for a spatially explicit distribution estimate of seals at sea.

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Migration and stock studies on the Australian school prawn Metapenaeus macleayi

The results of tagging studies conducted to determine the relationship between estuarine (juvenile) populations and adults at sea suggest that maturing Metapenaeus macleayi leave the estuaries and move along the coast in a northerly direction; the longest migration recorded was 120 km, but most prawns appear to disperse in shallow water (<40 m) within about 70 km of their juvenile habitat. There are approximately 60 estuarine populations of M. macleayi in south-east Australia; the geographical range of prawns emigrating from most estuarine populations overlap (at sea) to some degree with that of adjacent or nearby populations, but there is little or no overlap between any two of the 6 major populations. The mixing of individuals from a major population and from nearby smaller populations is negligible for the purposes of fisheries management, hence the prawns in each of the major estuarine populations and the adults in the coastal area north for about 70 km may be regarded as a unit stock.     

Chilean Blue Whales as a Case Study to Illustrate Methods to Estimate Abundance and Evaluate Conservation Status of Rare Species

Abstract: Often abundance of rare species cannot be estimated with conventional design‐based methods, so we illustrate with a population of blue whales (Balaenoptera musculus) a spatial model‐based method to estimate abundance. We analyzed data from line‐transect surveys of blue whales off the coast of Chile, where the population was hunted to low levels. Field protocols allowed deviation from planned track lines to collect identification photographs and tissue samples for genetic analyses, which resulted in an ad hoc sampling design with increased effort in areas of higher densities. Thus, we used spatial modeling methods to estimate abundance. Spatial models are increasingly being used to analyze data from surveys of marine, aquatic, and terrestrial species, but estimation of uncertainty from such models is often problematic. We developed a new, broadly applicable variance estimator that showed there were likely 303 whales (95% CI 176–625) in the study area. The survey did not span the whales' entire range, so this is a minimum estimate. We estimated current minimum abundance relative to pre‐exploitation abundance (i.e., status) with a population dynamics model that incorporated our minimum abundance estimate, likely population growth rates from a meta‐analysis of rates of increase in large baleen whales, and two alternative assumptions about historic catches. From this model, we estimated that the population was at a minimum of 9.5% (95% CI 4.9–18.0%) of pre‐exploitation levels in 1998 under one catch assumption and 7.2% (CI 3.7–13.7%) of pre‐exploitation levels under the other. Thus, although Chilean blue whales are probably still at a small fraction of pre‐exploitation abundance, even these minimum abundance estimates demonstrate that their status is better than that of Antarctic blue whales, which are still <1% of pre‐exploitation population size. We anticipate our methods will be broadly applicable in aquatic and terrestrial surveys for rarely encountered species, especially when the surveys are intended to maximize encounter rates and estimate abundance.     

Community structure,density and standing crop of fishes in a subtropical Australian mangrove area

The fishes occurring in a subtropical mangrove (Avicennia marina) area in Moreton Bay, Australia, were studied for one year (November 1987 to November 1988, inclusive). Fishes within the mangroves were sampled using a block net, whilst those in adjacent waters were sampled using seine and gill nets. Forty six percent of the species, 75% of the number of fishes and 94% of the biomass taken during the study (all methods combined) were of direct importance to regional fisheries. The fish community utilising the habitat within the mangrove forest differed from that occurring in adjacent waters in terms of density, standing crop, species composition and diversity-index values. Standing-crop estimates for the fishes occurring within the mangroves (study period mean ± SD = 25.3 ± 20.4 g m^–2) were amongst the highest recorded values for estuarine areas whilst those for adjacent waters (2.9±2.3 g m^–2) were comparable to those of other estuarine studies.     

Preliminary patterns of distribution and abundance of loggerhead sea turtles,<Emphasis Type="Italic"> Caretta caretta</Emphasis>, around Columbretes Islands Marine Reserve,Spanish Mediterranean

Aerial surveys were conducted to estimate the abundance and distribution of loggerhead turtles (Caretta caretta) in the Columbretes Islands Marine Reserve and surrounding waters (western Mediterranean). Four surveys were carried out during 2000 and 2001, following the line transect methodology. Loggerheads appeared to be present at high densities in the area throughout the whole year, although density varied between seasons, being more abundant during the spring. Mean density in the study area was 0.322 turtles/km² (range 0.200–0.516) and the mean abundance was 1,324 turtles (range 825–2,124). The turtles were distributed homogeneously throughout the study area, we found no difference in loggerhead density between the water around the reserve and that in the rest of the study area. Current conservation measures planned by the local authorities, which include increasing the area of the reserve, would be very positive for the conservation of this stock.Communicated by S.A. Poulet, Roscoff     

Distribution of diatoms in the surficial sediments of the Mangyung-Dongjin tidal flat,west coast of Korea (Eastern Yellow Sea)

The relationship between the distribution of benthic diatoms and sediment characteristics of the Mangyung-Dongjin tidal flat on the west coast of Korea was investigated during June and July 1988. Diatoms were collected from the upper 5 mm of sediments at 60 sites along eight transect lines running perpendicular to the shore line. Of the 371 taxa encountered in the study area, 88% were pennate diatoms. Genera represented by the greatest number of species were Navicula, Nitzschia, Amphora, Cocconeis, Fragilaria and Achnanthes. The most abundant species were Paralia sulcata, Navicula sp. #1, N. arenaria and Cymatosira belgica; all were broadly distributed across the tidal flats. The 60 sites could be assigned to eight clusters with respect to similarity in species composition. Discriminant analysis showed that separation cluster was primarily related to the mean grain size of the sediment. The species could not be separated into groups based on similarities in occurrence; a high degree of spatial overlap was observed. The preferences of the more abundant species for grain size were, therefore, analysed by plotting numerical abundance against mean grain size. There were at least four patterns: species groups could be associated with finegrained sediments, those of intermediate size and coarser sediments and the last group showed no discernible pattern.     

Day-night differences in the emergence of demersal zooplankton from a sand substrate in a kelp forest

Day-night differences in abundance and biomass of demersal zooplankton in the water column were determined by trapping these animals as they emerged from the sand substrate in a kelp forest (Macrocystis pyrifera) ecosystem off Santa Catalina Island, California, USA. The day and night sampling periods of the 24 June 1979 new moon each lasted 12 h. Abundance and biomass of total demersal zooplankton were significantly higher in night samples. A mean of 2,425±1,168 demersal zooplankton m^-2 24 h^-1 migrated over a diel cycle; 97% of these animals were crustaceans. The mean biomass of demersal zooplankton was 94.2±27.6 mg ash-free dry wt m^-2 24 h^-1. No significant differences were found in either the abundance or biomass of demersal zooplanktion collected in low and high traps, suggesting that most animals collected 25 cm off the bottom can sustain swimming to at least 75 cm and that both traps give comparable estimates of the amount of demersal zooplankton available to planktivorous predators.     

The importance of incorporating functional habitats into conservation planning for highly mobile species in dynamic systems

The distribution of mobile species in dynamic systems can vary greatly over time and space. Estimating their population size and geographic range can be problematic and affect the accuracy of conservation assessments. Scarce data on mobile species and the resources they need can also limit the type of analytical approaches available to derive such estimates. We quantified change in availability and use of key ecological resources required for breeding for a critically endangered nomadic habitat specialist, the Swift Parrot (Lathamus discolor). We compared estimates of occupied habitat derived from dynamic presence‐background (i.e., presence‐only data) climatic models with estimates derived from dynamic occupancy models that included a direct measure of food availability. We then compared estimates that incorporate fine‐resolution spatial data on the availability of key ecological resources (i.e., functional habitats) with more common approaches that focus on broader climatic suitability or vegetation cover (due to the absence of fine‐resolution data). The occupancy models produced significantly (P < 0.001) smaller (up to an order of magnitude) and more spatially discrete estimates of the total occupied area than climate‐based models. The spatial location and extent of the total area occupied with the occupancy models was highly variable between years (131 and 1498 km²). Estimates accounting for the area of functional habitats were significantly smaller (2–58% [SD 16]) than estimates based only on the total area occupied. An increase or decrease in the area of one functional habitat (foraging or nesting) did not necessarily correspond to an increase or decrease in the other. Thus, an increase in the extent of occupied area may not equate to improved habitat quality or function. We argue these patterns are typical for mobile resource specialists but often go unnoticed because of limited data over relevant spatial and temporal scales and lack of spatial data on the availability of key resources. Understanding changes in the relative availability of functional habitats is crucial to informing conservation planning and accurately assessing extinction risk for mobile resource specialists.     

Comparative study of the structure of some tropical and temperate marine soft-bottom macrobenthic communities

Macrobenthic sampling of a transect of six stations on the North coast of Central Java in October 1985 revealed three distinct communities which can be regarded as paraliels of those from north temperate waters: (i) An Amphioplus (Lymanella) laevis/Lovenia sp. community parallel with the Amphiura chiajei/Brissopsis lyrifera community. (ii) A Tellinides timorensis community parallel with Tellina spp. communities. (iii) A Laciolina philippinarum community parallel with Macoma spp. communities. A strict comparison of the structure of the first two of these communities with their counterparts in England shows that within-habitat diversity is similar in both regions. Diversity at two shallower subtidal stations is higher than at two deeper stations: this is attributed to the effects of natural, low-level, physical disturbance which maintains the communities in a sub-climax stage. Total abundance and biomass of the macrobenthic fauna was lower at the Javanese stations than in their temperate counterparts.     

Movement within and between different habitats by the portunid crab Scylla serrata

Tagging experiments were carried out on Scylla serrata in three different types of habitat in Moreton Bay, Queensland, Australia, during 1976–1981. A total of 6 233 crabs were tagged and 1 180 recaptured. Two categories of movement were found, a free ranging type and an offshore migration by females. Crabs in a narrow creek with mangrove-covered banks displayed little movement. In areas with large intertidal flats bare of mangroves, crabs underwent more movement and adults (carapace width 150 mm or greater) and subadults (carapace width 100 to 149 mm) moved similar distances (mean 3.9 km). In an area with direct access to the sea, males and females moved equal distances, but in a long channel behind an island, mean female movement (6.6 km) was significantly more than that of males (mean 3.7 km). The distance between tag and recapture site was not greatly affected by the time at liberty over a period of 1 to 36 wk. Tag recaptures showed an exchange between the populations of a mangrove creek and those in the neighbouring bay. There was very limited exchange between the population in an estuary and the adjacent bay and no exchange was found between neighbouring areas separated by a region of habitat unsuitable for S. serrata. None of the more than 3 000 females captured in the study area was carrying eggs, but two tagged ovigerous females were caught at sea after having moved out of the study area. Eight other females were recaptured in other inshore or estuarine areas 20 to 65 km from their release site.     

Spatial trends in abundance of long-finned pilot whales, white-beaked dolphins and harbour porpoises in West Greenland

Data from an aerial line transect survey conducted off West Greenland during August–September 2007 were used to estimate the abundance of long-finned pilot whales (Globicephala melas), white-beaked dolphins (Lagenorhynchus albirostris) and harbour porpoises (Phocoena phocoena). The abundance of each species was estimated using mark-recapture distance sampling techniques to correct for perception bias, and correction factors for time spent at the surface were applied. The fully corrected abundance estimates were 8,133 long-finned pilot whales, 11,984 white-beaked dolphins and 33,271 harbour porpoises. Based on density surface modelling methods, a count model with a generalised additive model formulation was used to relate abundance to spatial variables. Response curves indicated that the preferred habitats were deep offshore areas in Midwest Greenland for pilot whales, deep water over steep seabed slopes in South Greenland for white-beaked dolphins and relatively shallow inshore waters in Midwest–South Greenland for harbour porpoises. The abundance estimates and spatial trends for the three species are the first obtained from Greenland.     

Combined Effects of Levels of Protection and Environmental Variables at Different Spatial Resolutions on Fish Assemblages in a Marine Protected Area

Abstract: The links between species–environment relations and species’ responses to protection are unclear, but the objectives of marine protected areas (MPAs) are most likely to be achieved when those relations are known and inform MPA design. The components of a species’ habitat vary with the spatial resolution of the area considered. We characterized areas at two resolutions: 250 m² (transect) and approximately 30,000 m² (seascape). We considered three categories of environmental variables: substrate type, bottom complexity, and depth. We sought to determine at which resolution habitat characteristics were a better predictor of abundance and species composition of fishes and whether the relations with environmental variables at either resolution affected species’ responses to protection. Habitat features accounted for a larger proportion of spatial variation in species composition and abundances than differences in protection status. This spatial variation was explained best by habitat characteristics at the seascape level than at the transect level. Species’ responses to protected areas were specific to particular seascape characteristics, primarily depth, and bottom complexity. Our method may be useful for prioritizing marine areas for protection, designing MPAs, and monitoring their effectiveness. It identified areas that provided natural shelter, areas acting as buffer zones, and areas where fish species were most responsive to protection. The identification of such areas is necessary for cost‐effective establishment and monitoring of MPAs.     

Influence of habitat on abundance and size structure of a large temperate-reef fish, Achoerodus viridis (Pisces: Labridae)

The relationship between densities of Achoerodus viridis (Pisces: Labridae) and reef habitats at various localities within New South Wales (NSW), Australia was examined. Types of habitats were quantified at inner, mid and outer estuarine locations in each of two estuaries (Botany Bay and Port Jackson) to determine whether purported patterns of movement from estuaries could be related to differences in habitat. Although the same types of habitat were generally found at all locations, differences in the proportion of habitat types were found between shallow and deep reefs and among inner, mid and outer estuarine locations for both estuaries. Shallow habitats were usually dominated by Ecklonia radiata, turf and/or fringe habitat in Botany Bay, whereas deep sites were generally dominated by urchin-grazed barrens habitat and, sometimes, sponge- and ascidian-dominated deep reef. Shallow sites in Port Jackson were dominated by a mixture of habitats, as were deep reefs at mid-estuarine locations. Other deep reefs in Port Jackson were dominated by E. radiata (inner estuarine) or barrens (outer estuarine) habitat. Thus, patterns of habitat cover were not consistent between estuaries and numbers of fish could not be related to proportional representation of habitat on reefs along estuarine gradients. Univariate and multivariate analyses showed that there was little evidence that any size class of fish was correlated with the proportional representation of a particular habitat or group of habitats. Counts of fish that focused on barrens and E. radiata forest habitats over a period of 10 yr showed that similar numbers and all sizes of fish were found in the two types of habitat. Greater numbers of small fish were, however, found in the E. radiata forest habitat than in the barrens habitat. Estimates of abundance along the coast of NSW (100s to 1000 km) in a range of habitats (e.g. ascidian-dominated reef, kelp forest, urchin-grazed barrens) showed that there was no indication that a particular habitat consistently had greater numbers of A. viridis than other habitats. Therefore, A. viridis of a range of sizes appears to be flexible in its use of habitats on reefs. Received: 24 December 1997 / Accepted: 23 June 1998     

Effects of landscape design of forest reserves on Saproxylic beetle diversity

Increasing the density of natural reserves in the forest landscape may provide conservation benefits for biodiversity within and beyond reserve borders. We used 2 French data sets on saproxylic beetles and landscape cover of forest reserves (LCFR) to test this hypothesis: national standardized data derived from 252 assessment plots in managed and reserve stands in 9 lowland and 5 highland forests and data from the lowland Rambouillet forest, a forested landscape where a pioneer conservation policy led to creation of a dense network of reserves. Abundance of rare and common saproxylic species and total saproxylic species richness were higher in forest reserves than in adjacent managed stands only in highland forests. In the lowland regional case study, as LCFR increased total species richness and common species abundance in reserves increased. In this case study, when there were two or more reserve patches, rare species abundance inside reserves was higher and common species richness in managed stands was higher than when there was a single large reserve. Spillover and habitat amount affected ecological processes underlying these landscape reserve effects. When LCFR positively affected species richness and abundance in reserves or managed stands, >12‐20% reserve cover led to the highest species diversity and abundance. This result is consistent with the target of 17% forested land area in reserves set at the Nagoya biodiversity summit in 2010. Therefore, to preserve biodiversity we recommend at least doubling the current proportion of forest reserves in European forested landscapes.     

Distribution and abundance of finless porpoise in the Inland Sea of Japan

Aerial sighting surveys were conducted in 2000 to evaluate the distribution and abundance of finless porpoise Neophocaena phocaenoides in the Inland Sea of Japan. We flew 60 north–south transects (2,218 km in total) at intervals of 6.43 km over the study area between 131°3′ and 134°59′ E. In total, 148 groups were detected by two observers. The average group size was 1.56 individuals. The effective strip half-width of each observer was estimated to be 107 m (coefficient of variation [CV] = 8.26%). Porpoise abundance was estimated at 7,572 individuals and the density was as low as 0.506 individuals/km² (CV = 17.3%). The sea is among the Japanese waters with the lowest density in spite of its favorable topographical conditions. In the western stratum of the study area, where the estimated density was 1.31 individuals/km², we observed a relatively regular distribution. In the central-eastern stratum, which had an estimated density of 0.208 individuals/km², we confirmed a clumped distribution that was restricted to inshore waters or near islands. No individuals were observed in waters between 132°51′ and 133°11′E, and between 133°43′ and 133°59′E, where sand dredging and other human activities have been active, suggesting that habitat fragmentation has occurred.