Population viability, ecological processes and biodiversity: Valuing sites for reserve selection

No-take reserves constitute one tool to improve conservation of marine ecosystems, yet criteria for their placement, size, and arrangement remain uncertain. Representation of biodiversity is necessary in reserve planning, but will ultimately fail for conservation unless factors affecting species’ persistence are also incorporated. This study presents an empirical example of the divergent relationships among multiple metrics used to quantify a site’s conservation value, including those that address representation (habitat type, species richness, species diversity), and others that address ecological processes and viability (density and reproductive capacity of a keystone species, in this case, the black chiton, Katharina tunicata). We characterized 10 rocky intertidal sites across two habitats in Barkley Sound, British Columbia, Canada, according to these site metrics. High-richness and high-production sites for K. tunicata were present in both habitat types, but high richness and high-production sites did not overlap. Across sites, species richness ranged from 29 to 46, and adult K. tunicata varied from 6 to 22 individuals m⁻². Adult density was negatively correlated with species richness, a pattern that likely occurs due to post-recruitment growth and survival because no correlation was evident with non-reproductive juveniles. Sites with high adult density also contributed disproportionately greater potential reproductive output (PRO), defined by total gonad mass. PRO varied by a factor of five across sites and was also negatively correlated with species richness. Compromise or relative weighting would be necessary to select valuable sites for conservation because of inherent contradictions among some reserve selection criteria. We suspect that this inconsistency among site metrics will occur more generally in other ecosystems and emphasize the importance of population viability of strongly interacting species.     

Rapid diversity and abundance decline in a Caribbean coral reef sponge community

Sponges are abundant and diverse on coral reefs, and play key functional roles; but virtually nothing is known of their dynamics. This is the first report of coral reef sponge community dynamics documented by a series of censuses in which volume and species of every individual were recorded. At the start of the 14 year study, there were 1395 sponge individuals, representing 39 species in nine orders, and a total sponge volume of 33,721 cm³ in the censused area of a shallow Caribbean reef in San Blas, Panama. The most striking results of the 5 censuses were steady disappearance of species (51.3%) throughout the study period, and a steep drop in total sponge volume (42.6%). Species in keratose orders and with massive growth forms were lost disproportionately. Sponge losses could not be attributed to predators, physical disturbance (including a hurricane), extreme episodes of other abiotic factors, or disproportionately great loss of rare species due to random fluctuations. Disease may have played a role.High loss rates documented at this and nearby sites could be a local phenomenon, but scattered reports of disease and mass mortality of sponges from other sites suggest these data may reflect region-wide losses. Monitoring programs designed for corals and mobile unitary organisms can miss changes in sponges because very high sponge species diversity, facile fragmentation and fusion, and quick and complete disappearance of dead sponges, impedes perception of changes if there are no prior censuses. Whether or not sponge declines are extensive will remain unknown until repeat censuses have been accomplished at additional sites. Sponges can increase water clarity, bind live corals to the reef frame, and facilitate reef regeneration, suggesting that loss of sponges could accelerate declines of coral reefs.     

Effects of fishing and protection on Brazilian reef fishes

The vast Brazilian coast harbors unique and diverse reef fish communities. Unfortunately, relatively little is known about the impact of fishing on these fish species, and few management or conservation efforts are being made to protect them. Here, we examine the effect of different levels of protection on the composition, abundance, and size structure of reef fish species along a 2500 km portion of the Brazilian coastline, noting in particular the relative abundance of endemics and the effect of protection on these species. Pairwise comparisons of sites with different protection status (more versus less protected) were used to determine the potential responses of reef fishes to the establishment of marine protected areas. Highly targeted species (top predators and large herbivores) were significantly more abundant and larger in size within sites with a higher degree of protection, indicating that they benefit from protection, while lightly fished and unfished species were not. These results are consistent with past work documenting the responses of species to protection. Here, we use our results in particular to suggest strategies and provide expectations for managing and protecting Brazilian reef fisheries. Because this biogeographic province lies entirely within the jurisdiction of a single nation, there may be unique and significant opportunities to effectively manage and conserve these fish species.     

Priority areas for conservation of Western Australian coastal fishes: A comparison of hotspot, biogeographical and complementarity approaches

Western Australia has an extensive coastline ranging from the tropical north to the temperate south, with a high diversity of neritic marine fishes. Distribution data of 1855 neritic fish species were used to compare a range of methods for identifying priority areas for their conservation. Species richness and endemism richness hotspots, biogeographic zoning and complementarity analysis were tested for their efficiency at representing the total suite of species. The hotspot approaches demonstrated low efficiency, as the sections of coastline selected were grouped together in isolated geographic locations and relatively few of the total suite of neritic fish species were represented. Biogeographic zoning divided the coast of Western Australia into six regions, and as such, priority areas were selected around the entire coastline, which resulted in fairly high levels of efficiency. However, the complementarity analysis proved to be the most efficient method, as >95% of all neritic fish species could be represented in six, appropriately located, 100 km long sections of coastline. Complementarity analysis indicated a total of 26 priority areas for neritic fish conservation, which were spread around the entire coast of Western Australia. However, as current marine conservation measures in Western Australia are focused on the west and northwest coasts, this study highlighted the need for marine conservation efforts to be extended to cover the north and south coasts.     

The potential value of indicator groups in the selection of marine reserves

Efforts to systematically select networks of marine reserves to conserve biodiversity may be constrained by limited information on the distribution of biodiversity. A potential solution is the use of surrogates, such as indicator groups, that effectively represent other elements of biodiversity. The potential of macroalgae and molluscs to act as indicator groups for all species in the selection of intertidal marine reserves was tested for a sample of rocky shore locations in south-east Australia. Fifteen locations were surveyed between November 1999 and January 2000, and reserve selection for each of the potential indicator groups and for all species was undertaken using richness-based and rarity-based algorithms. The sets of locations chosen by both algorithms to conserve each of the indicator groups included around 95% of all species. There was 54% spatial correspondence of the sets of locations chosen for each of the indicator groups. Molluscs appeared to be a reliable indicator for areas of reservation ranging from 6.7 to 73.3% of the locations surveyed, including a similar percentage of all species as the locations selected for all species. Unlike macroalgae, locations chosen for molluscs included significantly more species than a set of locations chosen at random, and more species than the set of locations chosen for randomly selected subsets of species of the same richness as molluscs. There is significant scope for further research into the value of indicator groups in reserve selection in marine systems.     

Storm-generated coral fragments - A viable source of transplants for reef rehabilitation

Coral reefs throughout the world have been damaged by storms, diseases, coral predators, temperature anomalies, and human activities. During the past three decades, recovery has been limited and patchy. Although a damaged coral reef cannot be restored to its original condition, interest in reef restoration is increasing. In a pilot project in the Caribbean (US Virgin Islands), storm-produced fragments of Acroporapalmata, A. cervicornis, and Porites porites were collected from donor reefs and transplanted to nearby degraded reefs. Sixty coral fragments were attached to dead-coral substrate (usually A. palmata skeletons), at similar depths from which they had been collected (1-3.5 m), using nylon cable ties. Seventy-five intact colonies were designated as controls. Study colonies were assessed at 6-month intervals for 2 years (1999-2001) and annually thereafter (through 2004). One-fourth of the 135 colonies and fragments monitored were alive at the conclusion of the 5-year study. Survival of control and transplanted A.cervicornis and P. porites was very low (median survival 2.4 and 1.8 years, respectively), with no significant differences between transplant and control colonies. Site and depth did not contribute significantly to A. palmata colony survival, but colony size and transplant/control status did. Probability of survival increased with colony size. Median survival for A. palmata was 1.3 years for transplant and 4.3 years for natural colonies when not controlled for size. A. palmata was the only viable candidate for reef rehabilitation. Storm swells were the primary cause of mortality.     

Incorporating ontogenetic dispersal, ecological processes and conservation zoning into reserve design

Computational methods for marine reserve design are frequently used as decision-support tools for the identification of conservation areas. Most reserve-selection algorithms minimise the cost of the reserve system whilst aiming to meet specified biodiversity objectives. Here, we extend a widely-used selection algorithm, Marxan, to incorporate several important considerations related to biodiversity processes and management. First we relax the scorched earth assumption to allow conservation features in non-reserve zones to contribute explicitly to conservation objectives. To achieve this, we generate conservation targets at landscape scales rather than focusing purely on the representation of features within reserves. Second, we develop the notion of spatial dependencies further to incorporate spatial heterogeneity in the value of individual conservation features such as habitat types. We use the example of ontogenetic migrations of fish from mangroves to coral reefs because it nicely demonstrates how spatial ecological processes generate predictable heterogeneity in habitat value that should be considered in the reserve design process. Lastly, we show how habitat value can be disaggregated into ecosystem processes and services. Using a case study for the Belize Barrier Reef we compare reserve networks generated using our new approach with the results of traditional analyses. Consideration of the contribution of different protection zones, connectivity among habitats and more complex management goals resulted in up to a 52% increase in the mean biomass of commercially and ecologically-important fish species represented in the landscape. Our approach strengthens the ecological basis of reserve-design algorithms and might facilitate the uptake of ecosystem-based management into reserve design.     

The cost of postponing biodiversity conservation in Mexico

Though Neotropical countries are the most species rich in the world, their biodiversity is threatened by the loss of native vegetation. Land conversion in Mexico during the last 30 years has been extensive and is representative of that of other developing countries. However, the effects of land use change on the required size and configuration of an adequate biological conservation area network are largely unknown. It is shown here that endemic mammals in Mexico could have been protected considerably more economically if a conservation plan had been implemented in 1970 than is possible today due to extensive conversion of primary habitats. Analysis of the distributions of 86 endemic mammal species in 1970, 1976, 1993, and 2000 indicates that the distributions of 90% of the species shrank during this 30-year period. At each time step, optimal conservation area networks were selected to represent all species. 90% more land must be protected after 2000 to protect adequate mammal habitat than would have been required in 1970. In addition, under a realistic conservation budget, 79% fewer species can be represented adequately in a conservation area network after 2000 compared to 1970. This provides an incentive for rapid conservation action in Mexico and other biodiversity hotspots with comparable deforestation rates, including Burma, Ecuador, Indonesia, the Philippines, and Sri Lanka. Due to ongoing habitat degradation, the efficiency of a conservation plan decreases with delays in its implementation.     

Connectivity of reef fish between mangroves and coral reefs: Algorithms for the design of marine reserves at seascape scales

Many species of coral reef fish undertake ontogenetic migrations between seagrass beds, mangroves, and coral reefs. A recent study from the Caribbean found that the availability of mangrove nursery habitat had a striking impact on the community structure and biomass of reef fish in their adult, coral reef habitat. The biomass of several species more than doubled when the reefs were connected to rich mangrove resources (defined as having at least 70 km of fringing Rhizophora mangle within a region of 200 km²). Here, the results of this large-scale empirical study are translated into a series of algorithms for use in natural resource management planning. Four algorithms are described that identify (i) the relative importance of mangrove nursery sites, (ii) the connectivity of individual reefs to mangrove nurseries, (iii) areas of nursery habitat that have an unusually large importance to specific reefs, and (iv) priority sites for mangrove reforestation projects. The algorithms generate a connectivity matrix among mangroves and coral reefs that facilitates the identification of connected corridors of habitats within a dynamic planning environment (e.g., reserve selection algorithms).     

Community-based conservation results in the recovery of reef fish spawning aggregations in the Coral Triangle

In the Coral Triangle community-based marine protected areas (MPAs) are being established at a prolific rate. Their establishment can benefit both fisheries and biodiversity, and they provide both a socially and economically acceptable means of managing coral reefs in developing nations. However, because such MPAs are typically small (usually <0.5 km²), they will rarely provide protection to large mobile fishes. An exception to this limitation may exist when community-based MPAs are established to protect small sites where vital processes occur, such as fish spawning aggregations (FSAs). To test the effectiveness of small (0.1–0.2 km²) MPAs for protecting FSAs, we monitored three FSA sites where brown-marbled grouper (Epinephelus fuscoguttatus), camouflage grouper (Epinephelus polyphekadion) and squaretail coralgrouper (Plectropomus areolatus) aggregate to spawn. Sites were monitored during peak reproductive periods (several days prior to each new moon) between January 2005 and November 2009. All three sites are located in New Ireland Province, Papua New Guinea, and had been exploited for decades, but in 2004 two sites were protected by the establishment of community-based MPAs. The third site continued to be exploited. Over the monitoring period densities of E. fuscoguttatus and E. polyphekadion increased at both MPAs, but not at the site that remained open to fishing. At one MPA the densities of E. polyphekadion increased tenfold. Our findings demonstrate that community-based MPAs that are appropriately designed and adequately enforced can lead to the recovery of populations of vulnerable species that aggregate to spawn.     

Evidence for a depth refuge effect in artisanal coral reef fisheries

Natural refuges can play a fundamental role in protecting species from overexploitation but have not been adequately quantified in the marine environment. We quantified the effect of a depth refuge on all fish species in an artisanal coral reef fishery in Zanzibar, Tanzania by comparing changes in fish species richness and relative abundance with depth at five fished and three unfished reefs across the region. Commercial species richness was depleted by 15.1% at shallow depths in fished reefs, but there was no difference between the reefs deeper than seven metres. Non-commercial species were not affected by fishing or depth. Evidence for similar patterns in fish communities in other countries and depth-limitations to artisanal fishing methods imply this effect is widespread. The depth refuge effect could be sustaining coral reef fisheries and should be taken in to account before implementing policies or subsidies that encourage or allow fisheries to exploit deeper waters.     

Effectiveness of five small Philippines’ coral reef reserves for fish populations depends on site-specific factors, particularly enforcement history

Community-managed, no-take marine reserves are increasingly promoted as a simple, precautionary measure to conserve biodiversity and sustain coral reef fisheries. However, we need to demonstrate the effects of such reserves to those affected by the loss of potential fishing grounds and the wider scientific community. We surveyed changes in fish communities in five small marine reserves in the central Philippines and three distant Control sites over seven years. We conducted underwater visual censuses of 53 fish families within the reserve (Inside), with a kilometre of the boundary (Outside) and at Control sites. We found significant differences between fish communities Inside and Outside the reserve only at the two sites with strictest compliance with fishing prohibition, while there were significant differences to distant Control sites in all cases. The strongest responses to reserve protection were found in predatory fishes (groupers and breams) and in butterflyfish. Other abundant fish families showed weak effects of protection. For all taxa analysed, we found significant effects of reserve Site and Site × Treatment interactions. The detection of fish responses to reserves is complicated by potential spillover effects, site-specific factors, particularly compliance, and the difficulty of identifying appropriate control areas.     

Effects of habitat characteristics and sedimentation on performance of marine reserves in St. Lucia

This study examines factors affecting the rate and extent of biomass build-up among commercially important groupers, snappers, grunts, parrotfish and surgeonfish in a network of four marine reserves in southwest St. Lucia, Caribbean. Reserves constituted 35% of the total reef area originally available for fishing. Protection was instigated in 1995 after a baseline survey with annual or biennial censuses performed until 2002. Each survey consisted of 114 fifteen minute fish counts in reserves and 83 in fishing grounds, at depths of 5 m and 15 m in a 10 m diameter counting area. Estimates of number and size (cm) of target species were used to calculate fish family biomass. Data were analysed using three-way ANOVA in a before-after-control-impact pairs (BACIP) design. All families increased significantly in biomass over time at nearly all sites. Increases were greater in reserves than fishing grounds, except for grunts, and responses were strongest in parrotfish and surgeonfish. The combined biomass of families more than quadrupled in reserves and tripled in fishing grounds between 1995 and 2002. During this period coral cover declined by 46% in reserves and 35% in fishing grounds. Multiple regression showed that neither habitat characteristics nor habitat deterioration significantly affected rates of biomass build-up. The key factor was protection from fishing, which explained 44% of the variance in biomass growth. A further 28% of the variance was explained by sedimentation, a process known to stress reef invertebrates, significantly reducing the rate of biomass build-up. St. Lucia’s reserves succeeded in producing significant gains to fish stocks despite coral cover and structural complexity falling steeply over the period of the study.     

Methods for reserve selection: Interior point search

Spatial reserve design concerns the planning of biological reserves for conservation. Typical reserve selection formulations operate on a large set of landscape elements, which could be grid cells or irregular sites, and selection algorithms aim to select the set of sites that achieves biodiversity target levels with minimum cost. This study presents a completely different optimization approach to reserve design. The reserve selection problem can be considerably simplified given the reasonable assumptions that: (i) maximum reserve cost is known; (ii) the approximate number of new reserves to be established is known; (iii) individual reserves need to be spatially contiguous. Further assuming the ability to construct a set of reserves in an efficient and close to optimal manner around designated reserve locations, the reserve selection problem can be turned into a search for a single interior point and area for each reserve. The utility of the proposed method is demonstrated for a data set of seven indicator species living in an conservation priority area in Southern Australia consisting of ca 73,000 selection units, with up to 10,000 cells chosen for inclusion in a reserve network. Requirements (ii) and (iii) above make interior point search computationally very efficient, allowing use with landscapes in the order of millions of elements. The method could also be used with non-linear species distribution models.     

The importance of in situ site loss in nature reserve selection: Balancing notions of complementarity and robustness

Because the threat of habitat destruction can never be entirely eliminated, there is a legitimate concern that some reserve networks, especially highly complementary ones with minimal species overlap, may be predisposed to severe losses in species representation if one or more core reserve sites are destroyed. In order to address this problem in a systematic way, we propose the use of two different optimization models for designing complementary reserve networks that are also highly robust to possible site losses. Given limited budgets, the first maximizes expected species representation over all possible site loss patterns while the second maximizes a combination of representation given all sites and remaining representation following the worst-case loss of a restricted subset of reserve sites. By incorporating reserve loss in fundamentally different ways, these two models provide a range of options in terms of information requirements, assumptions about risk aversion, and structural complexity. We compare both of these methods to a more standard approach, which completely ignores the inherent risk posed by reserve site loss. Results confirm that significantly more robust solutions can be obtained for a marginal decrease in initial species representation within the reserve system.     

Alleviating impacts of anthropogenic activities by traditional conservation measures: can a small reef reserve be sustainedly managed?

Zoning schemes that are becoming an important management tool in large marine reserves, are difficult to implement in small reef areas. At the 3.4 km long reef of Eilat (Red Sea), a small (ca. 350 m of coastline) enclosure strategy has been enforced since 1992, while the remaining reef was left open to intense human activities. Here we have investigated for 2.5 y three populations of the branching coral Stylophora pistillata (3605 colonies) in a locality within the enclosed area (site NR) and in two areas open to the public, by tossing random quadrats at the shallow lagoonar zone (0.5-1.5 m depth). In the two open sites we found significantly higher levels of colony breakage (14-34% vs. 4-9% in the enclosed site), lower partial mortality levels of colonies (in the first 1.5 y; 7-9% vs. 23-30% at NR), higher recruitment (up to 3.0 vs. up to 0.9 colonies/m 2), 50% reduction in coral life span (10 vs. 20 y) and an estimated extinction period of 9-10 y for new cohorts as compared to >20 y in the enclosed site. Average colony size and maximal colony size were about half in the open sites. Live coverage fluctuated widely in all sites but was 3 times higher in the enclosed area (1.0-3.0% vs. 0.3-1.1%). Log-transformed size frequency distributions revealed, at the open sites, a shift from small towards medium-size classes and at the enclosed site, a shift from larger to medium size classes. We conclude that the enclosure of a limited core zone, although improved some ecological parameters, was not sufficient to compensate for stress imposed by anthropogenic activities. It is suggested to employ active restoration approaches, such as the “gardening concept”, as supplementary management tools.     

The quest for cryptic creatures: Impacts of species-focused recreational diving on corals

Coral reefs are popular with ecotourists, but the impact of divers on reefs is cause for concern. In this study, we assessed the damage to corals caused by divers seeking cryptic but charismatic fish such as seahorses (family Syngnathidae) and frogfishes (Antennariidae), which are found on reefs around the world. These fish are closely associated with the reef substratum, thus bringing divers into close proximity to coral. We found that when in the vicinity of frogfish and seahorses, divers made unintentional contact with corals significantly more often and for longer periods than when these species were absent. This change in diver behaviour resulted in a greater frequency of coral breakage and scarring at seahorse/frogfish sites than at ecologically equivalent control sites. However, the spatial extent of damage appeared limited. Beyond 1-3 m from the seahorse or frogfish, coral breakage and scarring rate decreased to levels similar to those of control sites. None of the coral species, which suffered the most damage, was particularly rare, suggesting that the habitat conservation concern of these marked shifts in diver behaviour is limited. Nevertheless, the use of pre-dive briefings and smaller dive group sizes could minimise the damage caused by divers approaching cryptic species of interest near the reef.     

Entanglement of monofilament fishing lines and coral death

This study examined the extent of damage by monofilament fishing lines on the cauliflower coral (Pocillopora meandrina) colonies at a popular cast fishing site in Oahu, Hawaii in March 1998. Sixty-five percent of coral colonies had fishing lines on their surface, and 80% of colonies were either entirely or partially dead. The percentage of entirely and partially dead colonies was positively correlated with the percentage of colonies entangled with fishing lines. Percent dead coral surface area was larger for colonies with fishing lines than colonies without fishing lines. In colonies with fishing lines, percent dead coral surface area was positively correlated with percent area with fishing lines. The high incidence of coral colonies with fishing lines and the three sets of significant results indicated destructive effects of monofilament fishing lines to P. meandrina corals. There is thus a clear case that reefs should be protected not only from the effects of excessive removal of fishes, but also from the effects of unsuccessful fishing that results in fishing-line entanglement on corals.     

Complementarity analysis: Mapping the performance of surrogates for biodiversity

Efficient planning for biodiversity conservation requires a consideration of complementarity when assessing the value of adding new areas for management. Unfortunately, complementarity in biodiversity across all groups cannot usually be measured directly, so methods are needed to choose good surrogates (or ‘indicators’) for predicting this overall complementarity. Previous attempts at assessment of biological surrogates have measured dissimilarity among biotas, or congruence between sets of selected areas, or species representation within a set of selected areas, all of which can seriously misrepresent the strength of a surrogate relationship across all areas. Therefore, we propose a new approach to complementarity analysis. We show that the pattern of complementarity among all biotas can be assessed in terms of the frequency of false high and false low predictions by the surrogates. We also show how the spatial pattern in these false predictions can be mapped and discuss their usefulness. On the one hand, areas on these maps associated with many false high predictions are overvalued and would be an inefficient investment for scarce conservation resources. On the other hand, areas associated with many false low predictions are undervalued and unlikely to attract conservation action, so we need to know whether they are particularly likely to be highly threatened. These geographical patterns can be used to identify habitat-associated biases in the performance of surrogate groups.