Peter J. Etnoyer

Habitat-forming deep-sea corals in the Northeast Pacific Ocean

We define habitat-forming deep-sea corals as those families of octocorals, hexacorals, and stylasterids with species that live deeper than 200 m, with a majority of species exhibiting complex branching morphology and a sufficient size to provide substrata or refugia to associated species. We present 2,649 records (name, geoposition, depth, and data quality) from eleven institutions on eight habitatforming deep-sea coral families, including octocorals in the families Coralliidae, Isididae, Paragorgiidae and Primnoidae, hexacorals in the families Antipathidae, Oculinidae and Caryophylliidae, and stylasterids in the family Stylasteridae. The data are ranked according to record quality. We compare family range and distribution as predicted by historical records to the family extent as informed by recent collections aboard the National Oceanic of Atmospheric Administration (NOAA) Office of Ocean Exploration 2002 Gulf of Alaska Seamount Expedition (GOASEX). We present a map of one of these families, the Primnoidae.

Habitat Availability and Heterogeneity and the Indo-Pacific Warm Pool as Predictors of Marine Species Richness in the Tropical Indo-Pacific

Range overlap patterns were observed in a dataset of 10,446 expert-derived marine species distribution maps, including 8,295 coastal fishes, 1,212 invertebrates (crustaceans and molluscs), 820 reef-building corals, 50 seagrasses, and 69 mangroves. Distributions of tropical Indo-Pacific shore fishes revealed a concentration of species richness in the northern apex and central region of the Coral Triangle epicenter of marine biodiversity. This pattern was supported by distributions of invertebrates and habitat-forming primary producers. Habitat availability, heterogeneity, and sea surface temperatures were highly correlated with species richness across spatial grains ranging from 23,000 to 5,100,000 km2 with and without correction for autocorrelation. The consistent retention of habitat variables in our predictive models supports the area of refuge hypothesis which posits reduced extinction rates in the Coral Triangle. This does not preclude support for a center of origin hypothesis that suggests increased speciation in the region may contribute to species richness. In addition, consistent retention of sea surface temperatures in models suggests that available kinetic energy may also be an important factor in shaping patterns of marine species richness. Kinetic energy may hasten rates of both extinction and speciation. The position of the Indo-Pacific Warm Pool to the east of the Coral Triangle in central Oceania and a pattern of increasing species richness from this region into the central and northern parts of the Coral Triangle suggests peripheral speciation with enhanced survival in the cooler parts of the Coral Triangle that also have highly concentrated available habitat. These results indicate that conservation of habitat availability and heterogeneity is important to reduce extinction of marine species and that changes in sea surface temperatures may influence the evolutionary potential of the region.

Biodiversity Assessment of the Fishes of Saba Bank Atoll, Netherlands Antilles

Biodiversity surveys were conducted on Saba Bank, Netherlands Antilles, to assess ichthyofaunal richness and to compare with published surveys of other Caribbean localities. The primary objective was to estimate the total species richness of the Saba Bank ichthyofauna. A variety of sampling techniques was utilized to survey the fish species of both the visually accessible megafauna and the camouflaged and small-sized species comprising the cryptic ichthyofauna. Based on results presented herein, the number of species known on Saba Bank is increased from 42 previously known species to 270 species. Expected species-accumulation curves demonstrate that the current estimate of species richness of fishes for Saba Bank under represents the actual richness, and our knowledge of the ichthyofauna has not plateaued. The total expected fish-species richness may be somewhere between 320 and 411 species. The Saba Bank ichthyofaunal assemblage is compared to fish assemblages found elsewhere in the Caribbean. Despite the absence of shallow or emergent shore habitats like mangroves, Saba Bank ranks as having the eighth highest ichthyofaunal richness of surveyed localities in the Greater Caribbean. Some degree of habitat heterogeneity was evident. Fore-reef, patch-reef, and lagoonal habitats were sampled. Fish assemblages were significantly different between habitats. Species richness was highest on the fore reef, but 11 species were found only at lagoonal sites. A comprehensive, annotated list of the fishes currently known to occur on Saba Bank, Netherland Antilles, is provided and color photographs of freshly collected specimens are presented for 165 of the listed species of Saba Bank fishes to facilitate identification and taxonomic comparison with similar taxa at other localities. Coloration of some species is shown for the first time. Preliminary analysis indicates that at least six undescribed new species were collected during the survey and these are indicated in the annotated list.

Conservation and management implications of deep-sea coral and fishing effort distributions in the Northeast Pacific Ocean

The conservation of deep-sea corals is of growing interest in the United States. A range of issues including biodiversity protection, conservation of seafloor habitats, and the role of deep-sea corals as essential fish habitat places greater significance on understanding the distributions of these corals and fishing activities. At the same time overfishing of some groundfish populations highlights the need for ecosystem-based management. Here we present records of habitat-forming deep-sea corals from the United States Pacific Fishery Management Council region that we analyze in relation to differential ecological impacts of demersal fishing gears. We use an ecological footprint approach combining groundfish catch by gear type with a previously published ecological severity ranking of fishing gears.

A new species of Isidella bamboo coral (Octocorallia: Alcyonacea: Isididae) from northeast Pacific Seamounts

Abstract A new species of deep-sea gorgonian, Isidella tentaculum, from 720–1050 m depths on Northeast Pacific seamount peaks, continental slopes, and shelf canyons is described and illustrated. The octocoral colonies were observed alive and in situ using a manned submersible and remotely operated vehicles. The new species is a large (up to 132 cm high), abundant, and conspicuous habitat former. It differs from its closest sister species (I. trichotoma, I. longiflora, I. lofotensis, and I. elongata) in size and stature, polyp size, and polyp arrangement. Distinctive characteristics of I. tentaculum include thornstar-shaped sclerites and large, closely-spaced, dimorphic polyps, never before reported in the family Isididae. Living colonies exhibit long basal zooids trailing from the trunk, similar in appearance to scleractinian ‘sweeper tentacles’. Large colonies have long tentacles (∼40 cm). The flabellum has whorls of 4–5 large (6–9 mm high, 2–3 mm diameter), closely packed (2–4 mm apart) autozooid polyps. Long needle-shaped sclerites project from the septa between the pinnate tentacles. Small rod and platelet sclerites are evident in pinnules, and thornstars are evident in the pharynx of the polyps. Thornstars have been reported in Acanella, but not in Isidella.

Rapid Assessment of Octocoral Diversity and Habitat on Saba Bank, Netherlands Antilles

Saba Bank is a large submerged platform (∼2200 km2), average depth 30 m, located 4 km southwest of Saba Island in Netherlands Antilles, Caribbean Sea. Ships traveling to and from oil terminals on nearby St. Eustatius routinely anchor on the Bank, damaging benthic megafauna. Gorgonian octocorals are vulnerable to anchor damage, and they are common and conspicuous in shallow water (15–50 m) around the banks. This prompted a rapid assessment of octocoral habitat and diversity. The primary objectives were to estimate total species richness and to characterize habitats vis a vis gorgonians. Landsat imagery and multibeam bathymetry were employed to identify random sites for quantitative transects. A Seabotix LBV200L remotely operated vehicle (ROV) and SCUBA were used to collect and survey to 130 m. A total of 14 scuba dives and 3 ROV dives were completed in 10 days. During that time, 48 octocoral species were collected, including two likely undescribed species in the genera Pterogorgia and Lytreia. Gorgonian richness was exceptional, but not all species were collected, because the species accumulation curve remained steeply inclined after all surveys. Two shallow-water gorgonian habitat types were identified using multidimensional scaling and hierarchical cluster analyses: 1) a high diversity, high density fore-reef environment characterized by Eunicea spp., Gorgonia spp., and Pseudopterogorgia spp. and 2) a low diversity, low density plateau environment characterized by Pseudopterogorgia acerosa, Pterogorgia guadalupensis, and Gorgonia mariae. The analyses support hypotheses of broad (∼15 km) habitat homogeneity (ANOSIM, P>0.05), but a significant difference between fore-reef and plateau environments (ANOSIM, P<0.05). However, there was some indication of habitat heterogeneity along the 15 km study section of the 50 km platform edge along the southeast rim. Our results highlight the complexity and biodiversity of the Saba Bank, and emphasize the need for more scientific exploration.

Rapid assessment of stony coral richness and condition on Saba Bank, Netherlands Antilles.

The benthic habitats of Saba Bank (17°25′N, 63°30′W) are at risk from maritime traffic, especially oil tankers (e.g., anchoring). To mitigate this risk, information is needed on the biodiversity and location of habitats to develop a zone use plan. A rapid survey to document the biodiversity of macro-algae, sponges, corals and fishes was conducted. Here we report on the richness and condition of stony coral species at 18 select sites, and we test for the effects of bottom type, depth, and distance from platform edge. Species richness was visually assessed by roving scuba diver with voucher specimens of each species collected. Coral tissue was examined for bleaching and diseases. Thirty-three coral species were documented. There were no significant differences in coral composition among bottom types or depth classes (ANOSIM, P>0.05). There was a significant difference between sites (ANOSIM, P<0.05) near and far from the platform edge. The number of coral species observed ranged from zero and one in algal dominated habitats to 23 at a reef habitat on the southern edge of the Bank. Five reef sites had stands of Acropora cervicornis, a critically endangered species on the IUCN redlist. Bleaching was evident at 82% of the sites assessed with 43 colonies bleached. Only three coral colonies were observed to have disease. Combining our findings with that of other studies, a total of 43 species have been documented from Saba Bank. The coral assemblage on the bank is representative and typical of those found elsewhere in the Caribbean. Although our findings will help develop effective protection, more information is needed on Saba Bank to create a comprehensive zone use plan. Nevertheless, immediate action is warranted to protect the diverse coral reef habitats documented here, especially those containing A. cervicornis.

Toxicity of oil and dispersant on the deep water gorgonian octocoral Swiftia exserta, with implications for the effects of the Deepwater Horizon oil spill

Benthic surveys of mesophotic reefs in the Gulf of Mexico post Deepwater Horizon (DWH) showed that Swiftia exserta octocorals exhibited significantly more injury than in years before the spill. To determine the vulnerability of S. exserta to oil and dispersants, 96h toxicity assays of surrogate DWH oil water-accommodated fractions (WAF), Corexit® 9500 dispersant, and the combination of both (CEWAF) were conducted in the laboratory. Fragment mortality occurred within 48h for some fragments in the dispersant-alone and oil-dispersant treatments, while the WAF group remained relatively unaffected. The 96h LC50 values were 70.27mg/L for Corexit-alone and 41.04mg/L for Corexit in CEWAF. This study provides new information on octocoral sensitivity to toxins, and indicates that combinations of oil and dispersants are more toxic to octocorals than exposure to oil alone. These results have important implications for the assessment of effects of the DWH spill on deep-water organisms.

Growth and feeding of deep-sea coral Lophelia pertusa from the California margin under simulated ocean acidification conditions

The global decrease in seawater pH known as ocean acidification has important ecological consequences and is an imminent threat for numerous marine organisms. Even though the deep sea is generally considered to be a stable environment, it can be dynamic and vulnerable to anthropogenic disturbances including increasing temperature, deoxygenation, ocean acidification and pollution. Lophelia pertusa is among the better-studied cold-water corals but was only recently documented along the US West Coast, growing in acidified conditions. In the present study, coral fragments were collected at ∼300 m depth along the southern California margin and kept in recirculating tanks simulating conditions normally found in the natural environment for this species. At the collection site, waters exhibited persistently low pH and aragonite saturation states (Ωarag) with average values for pH of 7.66 ± 0.01 and Ωarag of 0.81 ± 0.07. In the laboratory, fragments were grown for three weeks in “favorable” pH/Ωarag of 7.9/1.47 (aragonite saturated) and “unfavorable” pH/Ωarag of 7.6/0.84 (aragonite undersaturated) conditions. There was a highly significant treatment effect (P < 0.001) with an average% net calcification for favorable conditions of 0.023 ± 0.009% d−1 and net dissolution of −0.010 ± 0.014% d-1 for unfavorable conditions. We did not find any treatment effect on feeding rates, which suggests that corals did not depress feeding in low pH/ Ωarag in an attempt to conserve energy. However, these results suggest that the suboptimal conditions for L. pertusa from the California margin could potentially threaten the persistence of this cold-water coral with negative consequences for the future stability of this already fragile ecosystem.

Distribution of deep-water corals, sponges, and demersal fisheries landings in Southern California, USA: implications for conservation priorities

Deep-sea corals in Southern California are diverse and abundant but subject to multiple stressors, including bottom-contact fisheries using mobile and fixed gear. There is a need for more information on the distribution of these taxa in relation to the distribution of demersal fishing effort, and the distribution of marine protected areas, in order to improve spatial planning. There are many marine managed areas in Southern California, including essential fish habitat (EFH) areas, conservation areas, and a national marine sanctuary, but specific areas of overlap between bottom fishing and benthic epifauna are poorly known. Groundfish surveys were conducted by the National Marine Fisheries Service using a remotely operated vehicle throughout Southern California between 2003 and 2011 to document abundance and distribution of deep-water rockfish and flatfish to a depth of 500 m. Corals and sponges were also common in these images, providing an opportunity to examine these communities. Analyses of 34,792 still images revealed abundance and diversity of coral and sponge taxa, as well as frequency of fishing debris. The occurrence data were overlaid in a geographic information system with landings data for deep-water (>50 m) demersal fisheries to identify areas of spatial overlap. Corals or sponges were observed in 23% of images. A total of 15 coral genera and six sponge morphotypes were identified. A total of 70 species codes were targeted by deep-water demersal fisheries operating below 50 m for years 2007–2011. A novel priority-setting algorithm was developed to identify areas of high richness, abundance, and fishing intensity (RAFi). Several highly-ranked areas were already protected as EFH (Footprint, Piggy Bank). Other highly-ranked sites (West Catalina Island, San Clemente Island, 9-Mile Bank, Santa Rosa Flats) were encompassed by transient gear restrictions, such as Rockfish conservation areas, but are now recommended for permanent protection by the Pacific Fishery Management Council.