Kenyon C. Lindeman

Mangroves enhance the biomass of coral reef fish communities in the Caribbean

Mangrove forests are one of the worlds most threatened tropical ecosystems with global loss exceeding 35% (ref. 1). Juvenile coral reef fish often inhabit mangroves, but the importance of these nurseries to reef fish population dynamics has not been quantified. Indeed, mangroves might be expected to have negligible influence on reef fish communities: juvenile fish can inhabit alternative habitats and fish populations may be regulated by other limiting factors such as larval supply or fishing. Here we show that mangroves are unexpectedly important, serving as an intermediate nursery habitat that may increase the survivorship of young fish. Mangroves in the Caribbean strongly influence the community structure of fish on neighbouring coral reefs. In addition, the biomass of several commercially important species is more than doubled when adult habitat is connected to mangroves. The largest herbivorous fish in the Atlantic, Scarus guacamaia, has a functional dependency on mangroves and has suffered local extinction after mangrove removal. Current rates of mangrove deforestation are likely to have severe deleterious consequences for the ecosystem function, fisheries productivity and resilience of reefs. Conservation efforts should protect connected corridors of mangroves, seagrass beds and coral reefs.

A global baseline for spawning aggregations of reef fishes.

Species that periodically and predictably congregate on land or in the sea can be extremely vulnerable to overexploitation. Many coral reef fishes form spawning aggregations that are increasingly the target of fishing. Although serious declines are well known for a few species, the extent of this behavior among fishes and the impacts of aggregation fishing are not appreciated widely. To profile aggregating species globally, establish a baseline for future work, and strengthen the case for protection, we (as members of the Society for the Conservation of Reef Fish Aggregations) developed a global database on the occurrence, history, and management of spawning aggregations. We complemented the database with information from interviews with over 300 fishers in Asia and the western Pacific. Sixty-seven species, mainly commercial, in 9 families aggregate to spawn in the 29 countries or territories considered in the database. Ninety percent of aggregation records were from reef pass channels, promontories, and outer reef-slope drop-offs. Multispecies aggregation sites were common, and spawning seasons of most species typically lasted <3 months. The best-documented species in the database, the Nassau grouper (Epinephelus striatus), has undergone substantial declines in aggregations throughout its range and is now considered threatened. Our findings have important conservation and management implications for aggregating species given that exploitation pressures on them are increasing, there is little effective management, and 79% of those aggregations sufficiently well documented were reported to be in decline. Nonetheless, a few success stories demonstrate the benefits of aggregation management. A major shift in perspective on spawning aggregations of reef fish, from being seen as opportunities for exploitation to acknowledging them as important life-history phenomena in need of management, is urgently needed.

Historical biogeography and speciation in the reef fish genus Haemulon (Teleostei: Haemulidae).

The high biodiversity of tropical marine hotspots has long intrigued evolutionary biologists and biogeographers. The genus Haemulon (grunts) is one of the most important (numerically, ecologically, and economically) reef fish groups in the New World and an excellent candidate to test hypotheses of speciation and diversity generation in the Greater Caribbean, the richest Atlantic biodiversity hotspot, as well as the eastern Pacific. To elucidate the phylogenetic relationships among the species of Haemulon, we obtained a combined total of 2639 base pairs from two mitochondrial genes (cytochrome b and cytochrome oxidase I), and two nuclear genes (TMO-4C4 and RAG2) from all nominal species. Parsimony, Maximum likelihood, and Bayesian analyses resulted in a well-resolved phylogeny with almost identical topologies. Previous phylogenetic hypotheses based on adult morphology, such as the close relationship among H. aurolineatum, H. boschmae, and H. striatum were not supported, whereas others using developmental characters, such as the relationship between H. plumieri and H. sciurus, were confirmed. Our data also indicate that the populations of the nominal H. steindachneri from the two sides of the Isthmus of Panama are genetically divergent at the species level in each ocean, and that the boga, Inermia vittata (family Inermiidae), belongs in the genus Haemulon. This evidence implies that there are 21 valid species of Haemulon, two more than previously recognized. The Amazon barrier and the Isthmus of Panama seem to have played roles in allopatric speciation of Haemulon. However, the majority of sister species pairs have completely overlapping distributions, indicating that vicariance is not the only process driving speciation in this genus. We conclude that both vicariance between biogeographic provinces, and ecological mechanisms of speciation within provinces contribute to species richness in the genus Haemulon.

Red snapper management in the Gulf of Mexico: science- or faith-based?

The most controversial fishery in U.S. waters of the Gulf of Mexico (Gulf) is for northern red snapper Lutjanus campechanus, which collapsed in the late 1980s when stock biomass became too low to be fished commercially in the eastern Gulf. Red snapper management began in 1989; the stock is now showing signs of recovery. The Gulf of Mexico Fishery Management Council has been slow to sufficiently reduce catches of the directed fisheries to rebuild the stock in a timely fashion, although compliance with the Magnuson-Stevens Reauthorization Act of 2006 (MSRA) required substantial cuts in the harvest of red snapper beginning in 2007. In our opinion, this could have been avoided if conservative management practices had been adopted earlier. We believe that ‘faith-based fisheries’ arguments have been used to defer effective management of red snapper in the Gulf, which in turn has strained the relationship between science, management, and stakeholders there. We provide a simple empirical argument and alternate interpretations of a recently published perspective on the historical fishery of red snapper in the Gulf to conclude that the preponderance of evidence used in the agency stock assessment process, and the simple arguments made here, do not support the perspective that the red snapper stock has increased in size sufficiently to defer compliance with the MSRA.

Transforming management of tropical coastal seas to cope with challenges of the 21st century

Over 1.3 billion people live on tropical coasts, primarily in developing countries. Many depend on adjacent coastal seas for food, and livelihoods. We show how trends in demography and in several local and global anthropogenic stressors are progressively degrading capacity of coastal waters to sustain these people. Far more effective approaches to environmental management are needed if the loss in provision of ecosystem goods and services is to be stemmed. We propose expanded use of marine spatial planning as a framework for more effective, pragmatic management based on ocean zones to accommodate conflicting uses. This would force the holistic, regional-scale reconciliation of food security, livelihoods, and conservation that is needed. Transforming how countries manage coastal resources will require major change in policy and politics, implemented with sufficient flexibility to accommodate societal variations. Achieving this change is a major challenge - one that affects the lives of one fifth of humanity.

Management of Spawning Aggregations

This chapter examines the needs and tools for managing reef fish spawning aggregations. We present a global overview of the management of aggregations, and explore management options. We evaluate conventional fishery management and marine protected area options in relation to aggregation conservation, and examine examples of management successes and failures. Most management to date has been reactive, and there remains an overwhelming need for proactive management of aggregations. Long-term monitoring, appropriate fishery policy and extensive fisher and community consultation and outreach are key elements in instituting effective and adaptive management of spawning aggregations.

Juveniles of the Caribbean’s largest coral reef snapper do not use reefs

Coral Reefs (2005) 24: 359 DOI 10.1007/s00338-005-0015-3 Reef sites Fig. 1 In situ pigmentation of juvenile cubera snapper. Estimated total length: 12 cm. Photographed at a depth of 1 m at the base of red mangrove roots in a 4-m deep channel, Big Torch Key, FL The cubera snapper, Lutjanus cyanopterus (Lutjanidae), is widely distributed in the Caribbean and can reach 160 cm total length (TL), the largest snapper in the region (Allen 1985). However, the early life stages have never been recorded on reefs or photographed in situ in any habitat. Juvenile records are limited to museum specimens collected only from vegetated estuarine habitats or observations of larger subadults during field surveys (Christensen 1965; Starck 1970). From offshore reef areas, juveniles below 15 cm TL still remain uncollected and uncensused. Distinctive in situ pigmentation characterizes juveniles recently observed in the Florida Keys (Fig. 1). Diagnostic characters to at least 4 cm TL (smaller sizes based on museum materials) include approximately 9 pale vertical bands, a narrow body depth, and oblique dorsal scale rows (Lindeman et al. 2005). These characters differ from similar congeners such as gray snapper, L. griseus, which can co-occur in shallow, vegetated habitats. Additional diagnostics include an absence of yellow fin pigment which, coupled with a lower body depth, distinguish juvenile schoolmaster snapper, L. apodus, which also have vertical bands. Information on juvenile ecology is limited, although adults occur in several well-studied regions. All evidence suggests that cubera settle in vegetated or softbottom habitats in shallow-water and not on reef structures. Subsequent to settlement, they occupy mangrove habitats from at least 5 cm to 30 cm TL. Juvenile age and growth in cubera is unstudied. Estimates of juvenile growth in the similar gray snapper vary substantially among studies (Claro et al. 2001). It is within the ranges of these studies to estimate that a 15 cm TL cubera is 1–2 years of age. The exact age at which cubera snapper first migrate to offshore reefs is also not known. Starck (1970) observed individuals above 35 cm TL in deep inshore channels. Subadults may begin to migrate offshore to deeper areas with more hard structure by age 2–3. It is common for marine species using estuarine habitats to be termed estuarine dependent. The term dependence implies obligate use and testable hypotheses. For example, if a species often occurs in estuarine habitats but all life stages are also recorded from non-estuarine areas (facultative = opportunistic use), dependence is logically excluded. In the southeast United States, the juveniles of many snapper species can occur in bays and typically display some type of ontogenetic cross-shelf migration. However, conspecifics of the same age can often be found outside of coastal bays or in outer, euhaline portions of bays. Therefore, these snappers, including gray snapper, are estuarine opportunistic, not dependent (Lindeman et al. 2000). Juveniles of the cubera snapper cannot yet be excluded by such tests in the southeast United States. Notably, the significance of estuarine habitats to reef fish ecology is not diminished; continued reductions in area or water quality of estuaries will negatively impact the early life history of a wide array of reef fishes by reducing the quantity and quality of shelter and feeding resources during the highly vulnerable early life stages, whether inshore habitat use is facultative or obligate.

Depth-Variable Settlement Patterns and Predation Influence on Newly Settled Reef Fishes (Haemulon spp., Haemulidae)

During early demersal ontogeny, many marine fishes display complex habitat-use patterns. Grunts of the speciose genus Haemulon are among the most abundant fishes on western North Atlantic coral reefs, with most species settling to shallow habitats (≤12 m). To gain understanding into cross-shelf distributional patterns exhibited by newly settled stages of grunts (<2 cm total length), we examined: 1) depth-specific distributions of congeners at settlement among sites at 8 m, 12 m, and 21 m, and 2) depth-variable predation pressure on newly settled individuals (species pooled). Of the six species identified from collections of newly settled specimens (n = 2125), Haemulon aurolineatum (tomtate), H. flavolineatum (French grunt), and H. striatum (striped grunt) comprised 98% of the total abundance; with the first two species present at all sites. Prevalence of H. aurolineatum and H. flavolineatum decreased substantially from the 8-m site to the two deeper sites. In contrast, H. striatum was absent from the 8-m site and exhibited its highest frequency at the 21-m site. Comparison of newly settled grunt delta density for all species on caged (predator exclusion) and control artificial reefs at the shallowest site (8-m) revealed no difference, while the 12-m and 21-m sites exhibited significantly greater delta densities on the caged treatment. This result, along with significantly higher abundances of co-occurring piscivorous fishes at the deeper sites, indicated lower predation pressure at the 8-m site. This study suggests habitat-use patterns of newly settled stages of some coral reef fishes that undergo ontogenetic shifts are a function of depth-variable predation pressure while, for at least one deeper-water species, proximity to adult habitat appears to be an important factor affecting settlement distribution.

Timing and locations of reef fish spawning off the southeastern United States

Managed reef fish in the Atlantic Ocean of the southeastern United States (SEUS) support a multi-billion dollar industry. There is a broad interest in locating and protecting spawning fish from harvest, to enhance productivity and reduce the potential for overfishing. We assessed spatiotemporal cues for spawning for six species from four reef fish families, using data on individual spawning condition collected by over three decades of regional fishery-independent reef fish surveys, combined with a series of predictors derived from bathymetric features. We quantified the size of spawning areas used by reef fish across many years and identified several multispecies spawning locations. We quantitatively identified cues for peak spawning and generated predictive maps for Gray Triggerfish (Balistes capriscus), White Grunt (Haemulon plumierii), Red Snapper (Lutjanus campechanus), Vermilion Snapper (Rhomboplites aurorubens), Black Sea Bass (Centropristis striata), and Scamp (Mycteroperca phenax). For example, Red Snapper peak spawning was predicted in 24.7–29.0°C water prior to the new moon at locations with high curvature in the 24–30 m depth range off northeast Florida during June and July. External validation using scientific and fishery-dependent data collections strongly supported the predictive utility of our models. We identified locations where reconfiguration or expansion of existing marine protected areas would protect spawning reef fish. We recommend increased sampling off southern Florida (south of 27° N), during winter months, and in high-relief, high current habitats to improve our understanding of timing and location of reef fish spawning off the southeastern United States.

Science Needs for Sea-Level Adaptation Planning: Comparisons among Three U.S. Atlantic Coastal Regions

To identify priority information needs for sea-level rise planning, we conducted workshops in Florida, North Carolina, and Massachusetts in the summer of 2012. Attendees represented professionals from five stakeholder groups: federal and state governments, local governments, universities, businesses, and nongovernmental organizations. Over 100 people attended and 96 participated in breakout groups. Text analysis was used to organize and extract most frequently occurring content from 16 total breakout groups. The most frequent key words/phrases were identified among priority topics within five themes: analytic tools, communications, land use, ecosystem management, and economics. Diverse technical and communication tools were identified to help effectively plan for change. In many communities, planning has not formally begun. Attendees sought advanced prediction tools yet simple messaging for decision-makers facing politically challenging planning questions. High frequency key words/phrases involved fine spatial scales and temporal scales of less than 50 years. Many needs involved communications and the phrase “simple messaging” appeared with the highest frequency. There was some evidence of geographic variation among regions. North Carolina breakout groups had a higher frequency of key words/phrases involving land use. The results reflect challenges and tractable opportunities for planning beyond current, geophysically brief, time scales (e.g., election cycles and mortgage periods).