Philip A. Hastings

A molecular phylogeny of the groupers of the subfamily Epinephelinae (Serranidae) with a revised classification of the Epinephelini

The phylogenetic relationships among the fishes in the perciform tribe Epinephelini (Serranidae) have long been poorly understood, in large part because of the numerous taxa that must be considered and the large, circumtropical distribution of the group. In this study, genetic data from two nuclear (Tmo-4C4 and histone H3) and two mitochondrial (16S and 12S) genes were gathered from 155 serranid and acanthomorph species as a means of developing a phylogenetic hypothesis using both maximum-likelihood and -parsimony criteria. The maximum-parsimony analysis recovered 675 most parsimonious trees of length 5703 steps (CI = 0.2523, HI = 0.7477, RI = 0.6582), and the maximum-likelihood analysis recovered 1 tree at −lnLikelihood = 28279.58341. These phylogenetic hypotheses are discussed in light of previous morphological evidence to evaluate the evolutionary history of the group and their implications for the currently recognized taxonomy. Our results question the monophyly of the Serranidae, as well as the genera Cephalopholis, Epinephelus, and Mycteroperca as currently defined. The Serranidae is monophyletic only with the exclusion of the genera Acanthistius and Niphon. We propose a revised classification of the tribe Epinephelini that reflects the hypothesized shared ancestry of the group and recognizes 11 genera: Alphestes, Cephalopholis, Dermatolepis, Epinephelus, Gonioplectrus, Hyporthodus (which is resurrected for 11 species of deep-bodied groupers), Mycteroperca (including 7 species heretofore allocated to Epinephelus), Plectropomus, Saloptia, Triso, and Variola.

Female choice and male reproductive success in the angel blenny, Coralliozetus angelica (Teleostei: Chaenopsidae)

Abstract Correlates of reproductive success of male Coralliozetus angelica were studied in field populations in the central Gulf of California by collecting males with their shelters (vacant barnacle tests) and counting the number of eggs defended. Male body size was positively correlated with the number of eggs defended. This apparently resulted from female preference for large males since females preferred to mate with the larger of two males in laboratory choice experiments. Condition of males, as assessed by their relative head width, was also positively correlated with the number of eggs defended. Fit of males in their shelter entrance, male colour intensity, and male size relative to immediately surrounding males were not correlated with reproductive success. The quality of shelters was an important determinant of male reproductive success since females apparently avoided mating with males in heavily fouled shelters. Large shelters tended to be more heavily fouled than small shelters and male size and shelter size were positively correlated. The resulting negative correlation of male size and shelter quality apparently constrained female preference for large males and may be an important factor limiting the degree of polygyny in this species.

Correlates of male reproductive success in the browncheek blenny, Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

SummaryAcanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.

A novel sexual pattern in serranid fishes: simultaneous hermaphrodites and secondary males in Serranus fasciatus

SynopsisGulf of California populations of Serranus fasciatus are composed of functional simultaneous hermaphrodites and males. This is the first serranid known to have this sexual pattern which is functionally intermediate between the typical serranid patterns of simultaneous and protogynous hermaphroditism. Males of S. fasciatus are derived from hermaphrodites by resorption of ovarian tissue and proliferation of the extant testicular band. Distinct sexual roles are evident in spawning events. Hermaphrodites gain female function by pair spawning with males (124 and 125 observed spawns) and rarely with other hermaphrodites (1 of 125). Hermaphrodites gain male function by sneak spawning (9 of 125) and rarely by pair spawning with other hermaphrodites. Males exclusively pair spawned with hermaphrodites. Despite its unusual sexual pattern, S. fasciatus appears allied with other Serranus species based on similarities in gonad morphology.

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.

The Digital Fish Library: Using MRI to Digitize, Database, and Document the Morphological Diversity of Fish

Museum fish collections possess a wealth of anatomical and morphological data that are essential for documenting and understanding biodiversity. Obtaining access to specimens for research, however, is not always practical and frequently conflicts with the need to maintain the physical integrity of specimens and the collection as a whole. Non-invasive three-dimensional (3D) digital imaging therefore serves a critical role in facilitating the digitization of these specimens for anatomical and morphological analysis as well as facilitating an efficient method for online storage and sharing of this imaging data. Here we describe the development of the Digital Fish Library (DFL, http://www.digitalfishlibrary.org), an online digital archive of high-resolution, high-contrast, magnetic resonance imaging (MRI) scans of the soft tissue anatomy of an array of fishes preserved in the Marine Vertebrate Collection of Scripps Institution of Oceanography. We have imaged and uploaded MRI data for over 300 marine and freshwater species, developed a data archival and retrieval system with a web-based image analysis and visualization tool, and integrated these into the public DFL website to disseminate data and associated metadata freely over the web. We show that MRI is a rapid and powerful method for accurately depicting the in-situ soft-tissue anatomy of preserved fishes in sufficient detail for large-scale comparative digital morphology. However these 3D volumetric data require a sophisticated computational and archival infrastructure in order to be broadly accessible to researchers and educators.

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.

Colour variation is incongruent with mitochondrial lineages: cryptic speciation and subsequent diversification in a Gulf of California reef fish (Teleostei: Blennioidei).

The Gulf of California endemic reef fish, Acanthemblemaria crockeri (Blennioidei, Chaenopsidae), reportedly has two colour morphs, one with melanic lateral spots (‘Gulf’ morph) and one with orange spots (‘Cape’ morph). In this study, we recorded colour morph in both males and females and collected mitochondrial DNA sequence data for cytochrome c oxidase I (COI) and tRNA‐Pro/D‐loop of specimens from throughout the Gulf to explore the genetic basis of the colour morphs. Two highly divergent (HKY + I distance = 11.9% for COI), reciprocally monophyletic lineages were identified, consistent with the presence of two parapatric species. A 30‐km gap between the distributions of mitochondrial lineages roughly corresponds to a hypothesized former seaway across the Baja California peninsula north of La Paz, although the estimated divergence time (1.84 million years ago) is more recent than the hypothetical seaway (3–4 million years ago). Surprisingly, the distribution of mitochondrial species is not congruent with the distribution of either male or female colour morphs. Our analysis also revealed significant population differentiation within both species and no shared haplotypes among populations. The northern Gulf species includes four populations (NB, CB, NM and CM) corresponding to northern and central Baja and northern and central mainland sites, while the Cape species includes two populations (SB and SM) corresponding to the Baja and mainland sides of the southern Gulf. The NB/CB division corresponds to a hypothesized Plio–Pleistocene mid‐peninsular seaway. The level of genetic divergence documented in this lineage is extraordinary for a marine fish with a pelagic larval stage within a semi‐enclosed basin.

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.