Christopher Kelley

The feeding ecology of three species of Caribbean angelfishes (family Pomacanthidae)

SynopsisThe foraging behavior and associated morphology of the feeding apparatus of three sympatric species of angelfishes, Holacanthus tricolor, Pomacanthus arcuatus and Pomacanthus paru were studied at St. Croix, U.S. Virgin Islands. All three had overlapping diets, consisting of algae and numerous species of sponges. The two Pomacanthus species also fed on gorgonians. The morphology of the dentition, jaws and gill rakers was similar in all three species. Male Holacanthus tricolor defended territories overlapping the foraging areas of two to four females. Within the males territory, females defended smaller territories against other females of the same size, but tolerated smaller females. In contrast, both Pomacanthus spp. formed pairs which defended intraspecific feeding territories.

Genetic Analyses and Simulations of Larval Dispersal Reveal Distinct Populations and Directional Connectivity across the Range of the Hawaiian Grouper (Epinephelus quernus)

Integration of ecological and genetic data to study patterns of biological connectivity can aid in ecosystem-based management. Here we investigated connectivity of the Hawaiian grouper Epinephelus quernus, a species of management concern within the Main Hawaiian Islands (MHI), by comparing genetic analyses with simulated larval dispersal patterns across the species range in the Hawaiian Archipelago and Johnston Atoll. Larval simulations revealed higher dispersal from the MHI to the Northwestern Hawaiian Islands (NWHI) than in the opposite direction and evidence for a dispersal corridor between Johnston and the middle of the Hawaiian Archipelago. Genetic analyses using mitochondrial DNA (mtDNA) control region sequences and microsatellites revealed relatively high connectivity across the Hawaiian Archipelago, with the exception of genetically distinct populations and higher mtDNA diversity in the mid-Archipelago. These analyses support the preservation of the mid-archipelago as a source of genetic diversity and a region of connectivity with locations outside the Hawaiian Archipelago. Additionally, our evidence for directional dispersal away from the MHI lends caution to any management decisions that would rely on the NWHI replenishing depleted MHI stocks.

Histology of the gonads and observations on the social behavior of the Caribbean angelfish Holacanthus tricolor

Male and female specimens of Holacanthus tricolor were collected in May 1981 and September 1982 off the Island of St. Croix. U.S. Virgin Islands, and their gonads were examined histologically. All testes examined showed vasa deferentia which were characteristic of secondary males, suggesting protogynous sex change. A transitional individual supported this interpretation. Ovarian histology and size-specific egg counts revealed seasonal differences in spawning frequency and clutch size. Females appeared to spawn every day in May. The timing of final maturation and hydration of the oocytes indicated that spawning occurred in the late afternoon or early evening. Not all females spawned every day in September, and clutch sizes were smaller than those of May. Behavioral observations of H. tricolor in the field revealed that males defended harems of two to four smaller females. Within each harem, females of different sizes had overlapping home ranges, while the home ranges of females of similar size did not overlap. Besides size dimorphism between the sexes, H. tricolor exhibited a previously unreported sexual dichromatism.

Phylogeographic analyses of submesophotic snappers Etelis coruscans and Etelis "marshi" (family Lutjanidae) reveal concordant genetic structure across the Hawaiian Archipelago.

The Hawaiian Archipelago has become a natural laboratory for understanding genetic connectivity in marine organisms as a result of the large number of population genetics studies that have been conducted across this island chain for a wide taxonomic range of organisms. However, population genetic studies have been conducted for only two species occurring in the mesophotic or submesophotic zones (30+m) in this archipelago. To gain a greater understanding of genetic connectivity in these deepwater habitats, we investigated the genetic structure of two submesophotic fish species (occurring ∼200–360 m) in this archipelago. We surveyed 16 locations across the archipelago for submesophotic snappers Etelis coruscans (N = 787) and E. “marshi” (formerly E. carbunculus; N = 770) with 436–490 bp of mtDNA cytochrome b and 10–11 microsatellite loci. Phylogeographic analyses reveal no geographic structuring of mtDNA lineages and recent coalescence times that are typical of shallow reef fauna. Population genetic analyses reveal no overall structure across most of the archipelago, a pattern also typical of dispersive shallow fishes. However some sites in the mid-archipelago (Raita Bank to French Frigate Shoals) had significant population differentiation. This pattern of no structure between ends of the Hawaiian range, and significant structure in the middle, was previously observed in a submesophotic snapper (Pristipomoides filamentosus) and a submesophotic grouper (Hyporthodus quernus). Three of these four species also have elevated genetic diversity in the mid-archipelago. Biophysical larval dispersal models from previous studies indicate that this elevated diversity may result from larval supplement from Johnston Atoll, ∼800 km southwest of Hawaii. In this case the boundaries of stocks for fishery management cannot be defined simply in terms of geography, and fishery management in Hawaii may need to incorporate external larval supply into management plans.

Establishing species–habitat associations for 4 eteline snappers with the use of a baited stereo-video camera system

With the use of a baited stereo-video camera system, this study semiquantitatively defined the habitat associations of 4 species of Lutjanidae: Opakapaka (Pristipomoides filamentosus), Kalekale (P. sieboldii), Onaga (Etelis coruscans), and Ehu (E. carbunculus). Fish abundance and length data from 6 locations in the main Hawaiian Islands were evaluated for species-specific and size-specific differences between regions and habitat types. Multibeam bathymetry and backscatter were used to classify habitats into 4 types on the basis of substrate (hard or soft) and slope (high or low). Depth was a major influence on bottomfish distributions. Opakapaka occurred at depths shallower than the depths at which other species were observed, and this species showed an ontogenetic shift to deeper water with increasing size. Opakapaka and Ehu had an overall preference for hard substrate with low slope (hard-low), and Onaga was found over both hard-low and hard-high habitats. No significant habitat preferences were recorded for Kalekale. Opakapaka, Kalekale, and Onaga exhibited size-related shifts with habitat type. A move into hard-high environments with increasing size was evident for Opakapaka and Kalekale. Onaga was seen predominantly in hard-low habitats at smaller sizes and in either hard-low or hard-high at larger sizes. These ontogenetic habitat shifts could be driven by reproductive triggers because they roughly coincided with the length at sexual maturity of each species. However, further studies are required to determine causality. No ontogenetic shifts were seen for Ehu, but only a limited number of juveniles were observed. Regional variations in abundance and length were also found and could be related to fishing pressure or large-scale habitat features.

Acoustic backscattering by Hawaiian lutjanid snappers. I. Target strength and swimbladder characteristics

The target strengths and swimbladder morphology of six snapper species were investigated using broadband sonar, x rays, and swimbladder casts. Backscatter data were obtained using a frequency-modulated sweep (60-200 kHz) and a broadband, dolphinlike click (peak frequency 120 kHz) from live fish, mounted and rotated around each of their three axes. X rays revealed species-specific differences in the shape, size, and orientation of the swimbladders. The angle between the fishs dorsal aspect and the major axis of its swimbladder ranged from 3 degrees to 12 degrees and was consistent between individuals within a species. This angle had a one-to-one relationship with the angle at which the maximum dorsal aspect target strength was measured (r2 = 0.93), regardless of species. Maximum dorsal aspect target strength was correlated with length within species. However, the swimbladder modeled as an air-filled prolate spheroid with axes measured from the x rays of the swimbladder predicted maximum target strength significantly better than models based on fish length or swimbladder volume. For both the dorsal and lateral aspects, the prolate spheroid models predictions were not significantly different from the measured target strengths (observed power >0.75) and were within 3 dB of the measured values. This model predicts the target strengths of all species equally well, unlike those based on length.

Acoustic backscattering by deepwater fish measured in situ from a manned submersible

An outstanding problem in fisheries acoustics is the depth dependence of scattering characteristics of swimbladderbearing fish, andthe effects of pressure on the target strength of physoclistous fish remain unresolved . In situ echoes from deepwater snappers were obtained with a sonar transducer mounted on a manned submersible next to a low-light video camera, permitting simultaneous echo recording and identification of species, fish size and orientation. The sonar system, consisting of a transducer, single board computer, hard disk, and analog-to-digital converter, used a 80ms, broadband signal (bandwidth 35 kHz, center frequency 120 kHz). The observed relationship between fish length and in situ target strength shows no difference from the relationship measured at the surface. No differences in the speciesspecific temporal echo characteristics were observedbetween surface andin situ measures. This ind icates that the size and shape of the snappers’ swimbladders are maintained both at the surface and at depths of up to 250 m. Information obtainedthrough controlledbackscatter measurements of tethered , anesthetizedfish at the surface can be appliedto free-swimming fish at depth. This is the first published account of the use of a manned submersible to measure in situ scattering from identified, individual animals with known orientations. The distinct advantage of this technique comparedwith other in situ techniques is the ability to observe the target fish, obtaining accurate species, size, and orientation information. r 2003 Elsevier Science Ltd. All rights reserved.

An integrative systematic framework helps to reconstruct skeletal evolution of glass sponges (Porifera, Hexactinellida)

BackgroundGlass sponges (Class Hexactinellida) are important components of deep-sea ecosystems and are of interest from geological and materials science perspectives. The reconstruction of their phylogeny with molecular data has only recently begun and shows a better agreement with morphology-based systematics than is typical for other sponge groups, likely because of a greater number of informative morphological characters. However, inconsistencies remain that have far-reaching implications for hypotheses about the evolution of their major skeletal construction types (body plans). Furthermore, less than half of all described extant genera have been sampled for molecular systematics, and several taxa important for understanding skeletal evolution are still missing. Increased taxon sampling for molecular phylogenetics of this group is therefore urgently needed. However, due to their remote habitat and often poorly preserved museum material, sequencing all 126 currently recognized extant genera will be difficult to achieve. Utilizing morphological data to incorporate unsequenced taxa into an integrative systematics framework therefore holds great promise, but it is unclear which methodological approach best suits this task.ResultsHere, we increase the taxon sampling of four previously established molecular markers (18S, 28S, and 16S ribosomal DNA, as well as cytochrome oxidase subunit I) by 12 genera, for the first time including representatives of the order Aulocalycoida and the type genus of Dactylocalycidae, taxa that are key to understanding hexactinellid body plan evolution. Phylogenetic analyses suggest that Aulocalycoida is diphyletic and provide further support for the paraphyly of order Hexactinosida; hence these orders are abolished from the Linnean classification. We further assembled morphological character matrices to integrate so far unsequenced genera into phylogenetic analyses in maximum parsimony (MP), maximum likelihood (ML), Bayesian, and morphology-based binning frameworks. We find that of these four approaches, total-evidence analysis using MP gave the most plausible results concerning congruence with existing phylogenetic and taxonomic hypotheses, whereas the other methods, especially ML and binning, performed more poorly. We use our total-evidence phylogeny of all extant glass sponge genera for ancestral state reconstruction of morphological characters in MP and ML frameworks, gaining new insights into the evolution of major hexactinellid body plans and other characters such as different spicule types.ConclusionsOur study demonstrates how a comprehensive, albeit in some parts provisional, phylogeny of a larger taxon can be achieved with an integrative approach utilizing molecular and morphological data, and how this can be used as a basis for understanding phenotypic evolution. The datasets and associated trees presented here are intended as a resource and starting point for future work on glass sponge evolution.

The implementation of rare events logistic regression to predict the distribution of mesophotic hard corals across the main Hawaiian Islands

Predictive habitat suitability models are powerful tools for cost-effective, statistically robust assessment of the environmental drivers of species distributions. The aim of this study was to develop predictive habitat suitability models for two genera of scleractinian corals (Leptoserisand Montipora) found within the mesophotic zone across the main Hawaiian Islands. The mesophotic zone (30–180 m) is challenging to reach, and therefore historically understudied, because it falls between the maximum limit of SCUBA divers and the minimum typical working depth of submersible vehicles. Here, we implement a logistic regression with rare events corrections to account for the scarcity of presence observations within the dataset. These corrections reduced the coefficient error and improved overall prediction success (73.6% and 74.3%) for both original regression models. The final models included depth, rugosity, slope, mean current velocity, and wave height as the best environmental covariates for predicting the occurrence of the two genera in the mesophotic zone. Using an objectively selected theta (“presence”) threshold, the predicted presence probability values (average of 0.051 for Leptoseris and 0.040 for Montipora) were translated to spatially-explicit habitat suitability maps of the main Hawaiian Islands at 25 m grid cell resolution. Our maps are the first of their kind to use extant presence and absence data to examine the habitat preferences of these two dominant mesophotic coral genera across Hawai‘i.

The largest sponge in the world

Sponges (phylum Porifera) are essential components of benthic marine communities that rival reef-building corals in terms of abundance and diversity (Van Soest et al. 2012; Kerksen et al. 2014). Several marine sponges are known to attain massive sizes, thereby providing key ecosystem services such as filtering large amounts of seawater, as well as providing important habitat to a myriad of invertebrate and microbial species (McMurray et al. 2008; Kerksen et al. 2014). The largest dimensions reported for any sponge were those of a colony of Aphrocallistes vastus Schulze, 1887, found in shallow waters (<25 m) off western Canada. The colony measured 3.4 m in length, 1.1 m in height and 0.5 m in width (Austin et al. 2007). Large sponges are known to exist in deep water as well—for example, Monorhaphis chuni, which forms giant basal spicules that can reach lengths of up to 3 m (Wang et al. 2009). During a recent expedition to the Northwestern Hawaiian Islands aboard the R/V Okeanos Explorer, we encountered a massive sponge (Fig. 1), whose length, height and width exceeded the dimensions of the largest specimens reported in the literature. The sponge was captured on high-definition video from the two-body remotely operated vehicle (ROV) system (Deep Discoverer and Seirios) during a dive on a ridge extending north from Bank 9 (27.13082°N, 175.57085°W 2117 m, 12 August 2015), within the Papahānaumokuākea Marine National Monument. The height and width of the sponge were measured from horizontal images taken from the Deep Discoverer ROV while two parallel laser beams were projected onto the sponge, whereas its length was measured from a Seirios ROV overhead image using the dimensions of the ROV (3.15 m×1.85 m) for calibration. ImageJ image analysis software (Wayne Rasband, National Institutes of Health, Bethesda, MD) was used to obtain the measurements. The sponge was determined to be over 3.5 m in length, 2.0 m in width and 1.5 m in width, thus exceeding the dimensions of the largest sponge previously known (Austin et al. 2007). While the massive colony was not sampled, we collected a specimen of what we believe to be the same species dur ing a p rev ious d ive a t a nea rby loca t ion (27.14316°N, 176.22617°W, 2104 m). Based on a microscopic examination of spicules, that specimen was identified as belonging to the hexactinellid family Rossellidae and subfamily Lanuginellinae. Aside from these two specimens, we encountered no individuals of this non reef-building species as part of the expedition, during which we conducted a total of 18 ROV dives to depths ranging from 1096 to 4829 m. The finding of such an enormous and presumably old sponge inside the Papahānaumokuākea Marine National Monument underscores the need to protect this area using the highest conservation measures available. While not much is known about the lifespan of sponges, some massive Communicated by P. Martinez Arbizu