Brad Erisman

Large recovery of fish biomass in a no-take marine reserve

No-take marine reserves are effective management tools used to restore fish biomass and community structure in areas depleted by overfishing. Cabo Pulmo National Park (CPNP) was created in 1995 and is the only well enforced no-take area in the Gulf of California, Mexico, mostly because of widespread support from the local community. In 1999, four years after the establishment of the reserve, there were no significant differences in fish biomass between CPNP (0.75 t ha−1 on average) and other marine protected areas or open access areas in the Gulf of California. By 2009, total fish biomass at CPNP had increased to 4.24 t ha−1 (absolute biomass increase of 3.49 t ha−1, or 463%), and the biomass of top predators and carnivores increased by 11 and 4 times, respectively. However, fish biomass did not change significantly in other marine protected areas or open access areas over the same time period. The absolute increase in fish biomass at CPNP within a decade is the largest measured in a marine reserve worldwide, and it is likely due to a combination of social (strong community leadership, social cohesion, effective enforcement) and ecological factors. The recovery of fish biomass inside CPNP has resulted in significant economic benefits, indicating that community-managed marine reserves are a viable solution to unsustainable coastal development and fisheries collapse in the Gulf of California and elsewhere.

Operationalizing the social-ecological systems framework to assess sustainability

Significance Meeting human needs while sustaining ecosystems and the benefits they provide is a global challenge. Coastal marine systems present a particularly important case, given that >50% of the world’s population lives within 100 km of the coast and fisheries are the primary source of protein for >1 billion people worldwide. Our integrative analysis here yields an understanding of the sustainability of coupled social-ecological systems that is quite distinct from that provided by either the biophysical or the social sciences alone and that illustrates the feasibility and value of operationalizing the social-ecological systems framework for comparative analyses of coupled systems, particularly in data-poor and developing nation settings. Environmental governance is more effective when the scales of ecological processes are well matched with the human institutions charged with managing human–environment interactions. The social-ecological systems (SESs) framework provides guidance on how to assess the social and ecological dimensions that contribute to sustainable resource use and management, but rarely if ever has been operationalized for multiple localities in a spatially explicit, quantitative manner. Here, we use the case of small-scale fisheries in Baja California Sur, Mexico, to identify distinct SES regions and test key aspects of coupled SESs theory. Regions that exhibit greater potential for social-ecological sustainability in one dimension do not necessarily exhibit it in others, highlighting the importance of integrative, coupled system analyses when implementing spatial planning and other ecosystem-based strategies.

Spatio-temporal dynamics of a fish spawning aggregation and its fishery in the Gulf of California

We engaged in cooperative research with fishers and stakeholders to characterize the fine-scale, spatio-temporal characteristics of spawning behavior in an aggregating marine fish (Cynoscion othonopterus: Sciaenidae) and coincident activities of its commercial fishery in the Upper Gulf of California. Approximately 1.5–1.8 million fish are harvested annually from spawning aggregations of C. othonopterus during 21–25 days of fishing and within an area of 1,149 km2 of a biosphere reserve. Spawning and fishing are synchronized on a semi-lunar cycle, with peaks in both occurring 5 to 2 days before the new and full moon, and fishing intensity and catch are highest at the spawning grounds within a no-take reserve. Results of this study demonstrate the benefits of combining GPS data loggers, fisheries data, biological surveys, and cooperative research with fishers to produce spatio-temporally explicit information relevant to the science and management of fish spawning aggregations and the spatial planning of marine reserves.

A Phylogenetic Test of the Size‐Advantage Model: Evolutionary Changes in Mating Behavior Influence the Loss of Sex Change in a Fish Lineage

The size‐advantage model asserts that mating behavior influences the incidence and direction of sex change in animals. Selection for protogyny (female to male sex change) occurs in mating systems in which large males monopolize and pair spawn with females; however, gonochorism (no sex change) is favored when adults spawn in groups and sperm competition is present. Despite widespread empirical and theoretical support for the model, these predictions have not been tested within a phylogenetic context. Here we show that the loss of sex change within a lineage of reef fishes is influenced by evolutionary changes in two traits related to their mating behavior: mating group structure and sperm competition intensity. Phylogenetic reconstructions of the reproductive evolution of groupers (Epinephelidae) indicate that protogyny and paired spawning are the ancestral conditions for the lineage; both gonochorism and group spawning evolved independently at least four times in three different genera. Evolutionary transformations from protogyny to gonochorism (loss of sex change) are associated with equivalent transformations in mating group structure from paired to group spawning, and sperm competition is considerably higher in gonochoric species than in protogynous species. These results provide explicit phylogenetic support for predictions of the size‐advantage model, demonstrating that selection for protogynous sex change decreases as mating group size and sperm competition intensity increase.

Evidence of gonochorism in a grouper, Mycteroperca rosacea, from the Gulf of California, Mexico

The sexual pattern and sexual development of the leopard grouper, Mycteroperca rosacea, were investigated from 483 specimens collected from the Gulf of California, Mexico. Histological and population data indicated a gonochoric sexual pattern. Some juveniles passed through an immature bisexual phase of gonadal development, but no evidence of post-maturational sex change was found. The immature bisexual phase is believed to be associated only with male development. The size distribution and size at sexual maturity were similar for both males and females. In accordance with predictions of the size-advantage model, the gonochoric sexual pattern of M. rosacea is likely influenced by its group-spawning mating system.

Courtship and Spawning Behavior in the California Sheephead, Semicossyphus Pulcher (Pisces: Labridae)

We recorded the courtship and spawning behavior of a protogynous fish, the California sheephead, Semicossyphus pulcher, throughout their spawning season at Bird Rock, Santa Catalina Island, California. We made additional observations at the Monterey Bay Aquarium and confirmed the details of behavior seen in the field. Large males held spawning territories in which females congregated approximately 1 h before sunset. Courtship commenced shortly before sunset and involved the male approaching each female, making lateral contact and leading her in a circular pattern. Smaller males attempted to court females within the territories, prompting large males to abort spawns and engage in chasing behavior with small males. Females visited several territorial males throughout the day, indicating that they are not part of a strict harem. Both field and aquarium observations confirm that the mating system can be successfully predicted from the size advantage model. Current regulations on the sheephead fishery, which allow for the removal of large, rare males, could have significant effects on the social structure, reproductive output, and mating processes of local populations. Moreover, understanding the mating system of the California sheephead illustrates the need for creating management strategies that better complement the unique life histories of marine fishes with alternative reproductive strategies.

Phylogenetic Perspectives on the Evolution of Functional Hermaphroditism in Teleost Fishes

Hermaphroditism is taxonomically widespread among teleost fishes and takes on many forms including simultaneous, protogynous, and protandrous hermaphroditism, bidirectional sex change, and androdioecy. The proximate mechanisms that influence the timing, incidence, and forms of hermaphroditism in fishes are supported by numerous theoretical and empirical studies on their mating systems and sexual patterns, but few have examined aspects of sex-allocation theory or the evolution of hermaphroditism for this group within a strict phylogenetic context. Fortunately, species-level phylogenetic reconstructions of the evolutionary history of many lineages of fishes have emerged, providing opportunities for understanding fine-scale evolutionary pathways and transformations of sex allocation. Examinations of several families of fishes with adequate data on phylogeny, patterns of sex allocation, mating systems, and with some form of hermaphroditism reveal that the evolution and expression of protogyny and other forms of sex allocation show little evidence of phylogenetic inertia within specific lineages but rather are associated with particular mating systems in accordance with prevalent theories about sex allocation. Transformations from protogyny to gonochorism in groupers (Epinephelidae), seabasses (Serranidae), and wrasses and parrotfishes (Labridae) are associated with equivalent transformations in the structure of mating groups from spawning of pairs to group spawning and related increases in sperm competition. Similarly, patterns of protandry, androdioecy, simultaneous hermaphroditism, and bidirectional sex change in other lineages (Aulopiformes, Gobiidae, and Pomacentridae) match well with particular mating systems in accordance with sex-allocation theory. Unlike other animals and plants, we did not find evidence that transitions between hermaphroditism and gonochorism required functional intermediates. Two instances in which our general conclusions might not hold include the expression of protandry in the Sparidae and the distribution of simultaneous hermaphroditism. In the Sparidae, the association of hypothesized mating systems and patterns of sex allocation were not always consistent with the size-advantage model (SAM), in that certain protandric sparids show evidence of intense sperm competition that should favor the expression of gonochorism. In the other case, simultaneous hermaphroditism does not occur in some groups of monogamous fishes, which are similar in ecology to the hermaphroditic serranines, suggesting that this form of sex allocation may be more limited by phylogenetic inertia. Overall, this work strongly supports sexual lability within teleost fishes and confirms evolutionary theories of sex allocation in this group of vertebrates.

Asymmetric connectivity of spawning aggregations of a commercially important marine fish using a multidisciplinary approach

Understanding patterns of larval dispersal is key in determining whether no-take marine reserves are self-sustaining, what will be protected inside reserves and where the benefits of reserves will be observed. We followed a multidisciplinary approach that merged detailed descriptions of fishing zones and spawning time at 17 sites distributed in the Midriff Island region of the Gulf of California with a biophysical oceanographic model that simulated larval transport at Pelagic Larval Duration (PLD) 14, 21 and 28 days for the most common and targeted predatory reef fish, (leopard grouper Mycteroperca rosacea). We tested the hypothesis that source–sink larval metapopulation dynamics describing the direction and frequency of larval dispersal according to an oceanographic model can help to explain empirical genetic data. We described modeled metapopulation dynamics using graph theory and employed empirical sequence data from a subset of 11 sites at two mitochondrial genes to verify the model predictions based on patterns of genetic diversity within sites and genetic structure between sites. We employed a population graph describing a network of genetic relationships among sites and contrasted it against modeled networks. While our results failed to explain genetic diversity within sites, they confirmed that ocean models summarized via graph and adjacency distances over modeled networks can explain seemingly chaotic patterns of genetic structure between sites. Empirical and modeled networks showed significant similarities in the clustering coefficients of each site and adjacency matrices between sites. Most of the connectivity patterns observed towards downstream sites (Sonora coast) were strictly asymmetric, while those between upstream sites (Baja and the Midriffs) were symmetric. The best-supported gene flow model and analyses of modularity of the modeled networks confirmed a pulse of larvae from the Baja Peninsula, across the Midriff Island region and towards the Sonoran coastline that acts like a larval sink, in agreement with the cyclonic gyre (anti-clockwise) present at the peak of spawning (May–June). Our approach provided a mechanistic explanation of the location of fishing zones: most of the largest areas where fishing takes place seem to be sustained simultaneously by high levels of local retention, contribution of larvae from upstream sites and oceanographic patterns that concentrate larval density from all over the region. The general asymmetry in marine connectivity observed highlights that benefits from reserves are biased towards particular directions, that no-take areas need to be located upstream of targeted fishing zones, and that some fishing localities might not directly benefit from avoiding fishing within reserves located adjacent to their communities. We discuss the implications of marine connectivity for the current network of marine protected areas and no-take zones, and identify ways of improving it.

Reproductive biology of the Panama graysby Cephalopholis panamensis (Teleostei: Epinephelidae).

The reproductive biology of the Panama graysby Cephalopholis panamensis was studied from collections and behavioural observations made in the Gulf of California from 2001 to 2006. Histological examinations, particularly the identification of gonads undergoing sexual transition, confirmed a protogynous hermaphroditic sexual pattern. The population structure and mating behaviour provided further support for protogyny. Size and age distributions by sex were bimodal, with males larger and older than females and sex ratios biased towards females. Mating groups consisted of a large male and several smaller females, and courtship occurred in pairs during the evening. Results on spawning periodicity and seasonality were incomplete, but histological data, monthly gonado-somatic indices (I(G)) and behavioural observations suggest that adults spawned around the full moon from May to September. Certain aspects of their reproductive biology (e.g. protogyny and low egg production) indicate that C. panamensis is particularly vulnerable to fishing and would benefit from a management policy in Mexico.

Evolutionary Transitions in the Sexual Patterns of Fishes: Insights from a Phylogenetic Analysis of the Seabasses (Teleostei: Serranidae)

Abstract The evolution of hermaphroditism in fishes has intrigued scientists for over a century, but few have studied fine-scale evolutionary transitions between sexual patterns within the context of detailed hypotheses regarding phylogenetic relationships. Our phylogenetic reconstruction of sexual patterns in the seabasses (Teleostei: Serranidae) using a composite tree of 47 species is consistent with the hypothesis that protogyny is the ancestral condition in the family from which other sexual patterns evolved. Under this scenario, members of the Anthiinae retained protogynous hermaphroditism, as did the serranine genera Centropristis and Cheilidoperca. Gonochorism evolved once in the genus Paralabrax, and simultaneous hermaphroditism evolved once in the lineage that includes species of Hypoplectrus, Serranus, Serraniculus, and Diplectrum. Androdioecy evolved once within the genus Serranus and was derived from simultaneous hermaphroditism. Results from this study differ from previous hypotheses on the evolution of sexual patterns in seabasses and suggest that sexual patterns in fishes can evolve in several directions within single lineages and do not require functional intermediates.