An enigmatic new octocoral species (Anthozoa, Octocorallia, Malacalcyonacea) from Isla del Coco National Park.

Alienaparvagen. et sp. nov. is described from Cocos Island, Costa Rica. The species was found at various islets and rocky outcrops north and northwest of the island, 20-30 m in depth. The genus is characterised by polyps, retracting into calyces, that form thin encrusting mats extending on dead or live substrates. Sclerites are mostly asymmetrical spindles. Anthocodial rods are arranged in points, not forming a collaret. Colonies and coenenchymal sclerites are red, and polyps are transparent. Using an integrative taxonomic approach, we found the new genus to morphologically and genetically differ from all other described taxa. The molecular phylogenetic analyses provide strong support for the placement of this new genus in the family Pterogorgiidae. Morphologically it is unlike any of the other members of this family, necessitating an amendment to the diagnosis of Pterogorgiidae. Like several other known taxa of octocorals with encrusting growth forms, Alienagen. nov. appears to have evolved from a gorgonian ancestor by loss of an internal skeletal axis. It is the first member of Pterogorgiidae to be reported from the eastern Pacific, contributing further to the knowledge of marine biodiversity in the eastern tropical Pacific and to the octocoral biodiversity of Cocos Island in particular.

Nearshore marine biodiversity of Osa Peninsula, Costa Rica: Where the ocean meets the rainforest.

Osa Peninsula in remote southwest Costa Rica harbors 2.5% of global terrestrial biodiversity in only 1,200 km2 and has the largest remaining tract of Pacific lowland wet forest in Mesoamerica. However, little is known about the marine ecosystems of this diverse region. Much of the coastline consists of soft sediment exposed to strong wave action. Three major hard bottom habitat types define this region, including: 1) coral reefs around Isla del Caño Biological Reserve, a no-take marine protected area (MPA) of 52 km2, 2) coastal rocky reefs and islets along the peninsula, including Corcovado National Park, and 3) submerged pinnacles just outside the Isla del Caño MPA. Average coral cover at Isla del Caño was 21%, composed primarily of Porites lobata and Pocillopora elegans. In contrast, coastal rocky reefs were dominated by turf algae (39.8%) and macroalgae (20.7%) with low coral cover (1.1%). Submerged pinnacles were dominated by crustose coralline algae (33.3%) and erect coralline algae (25.7%). Fish assemblage characteristics (species richness, abundance, biomass) were significantly higher at the pinnacles compared to the other habitats and was dominated by schooling species such as Haemulon steindachneri, and the herbivores Kyphosus ocyurus, and Acanthurus xanthopterus. Top predators, primarily Triaenodon obesus, Caranx sexfasciatus, and Lutjanus argentimaculatus, were also most abundant at these pinnacles and accounted for the largest differences in fish trophic structure among habitats. Despite Isla del Caño being fully protected from fishing, biomass was similar to fished areas along the coast and lower than the adjacent submerged pinnacles outside the reserve. Similarly, Corcovado National Park includes 20.3 km2 of no-take MPAs; however, there is limited enforcement, and we noted several instances of fishing within the park. The unique configuration of healthy offshore coral reefs and pinnacles connected to coastal habitats provides corridors for many species including large predators such as sharks and other marine megafauna, which warrants additional protection.

Rhodolitica on rhodoliths: a new stoloniferan genus (Anthozoa, Octocorallia, Alcyonacea).

Rhodolitica occulta gen. nov. et sp. nov. (Clavulariidae) is described from Cocos Island National Park, Pacific Ocean, Costa Rica. The species was found at various islets and rocky outcrops around the island, 20-55 m in depth. The genus is characterised by tubular, single, erect anthosteles interconnected by thin basal ribbon-like stolons on the surfaces of living rhodoliths. The anthosteles are devoid of fused sclerites, which are only present in the stolons. Coenenchymal sclerites are mostly spindles of various shapes, with a characteristic cylindrical warty type in the outer layer, crosses and radiates. Anthocodiae are armed with points, lacking collarets. Colonies and sclerites are red. Using an integrative taxonomic approach, we separate the new genus from similar genera through both morphological comparison and a molecular phylogenetic analysis. This research is a contribution to the knowledge of the octocoral biodiversity in Cocos Island and marine biodiversity in the eastern tropical Pacific.

A revision of the genus Psammogorgia Verrill, 1868 (Cnidaria, Anthozoa, Octocorallia) in the tropical eastern Pacific Ocean.

The species of the genus Psammogorgia Verrill, 1868 from the shallow waters of the tropical eastern Pacific were mainly described from 1846 to 1870. Very few contributions were published subsequently. Recently, the genus was revisited with the addition of two new species. However, a comprehensive generic study is still missing for the eastern Pacific. Psammogorgia is characterised by having axes cores without mineralisation, mainly coarse irregular spindles and thorny, leafy or tuberculate clubs coenenchymal sclerites and the anthocodial armature with distinct collaret and points arrangements. Herein a taxonomic revision of the genus is presented based on type material which was morphologically analysed and illustrated using optical and scanning electron microscopy. Comparative character tables are provided for comparison among species in the genus, along with a taxonomic key. Moreover, the taxonomic status of each species was analysed. The genus Psammogorgia comprises six valid species and two varieties, and three lectotypes and a new combination are proposed to establish the taxonomic status of these species.

New records of Swiftia (Cnidaria, Anthozoa, Octocorallia) from off the Pacific Costa Rican margin, including a new species from methane seeps.

Exploration of the deep sea off the Pacific margin of Costa Rica has resulted in the discovery of a number of new species and reports for the region. Here, we report on the occurrence of the octocoral genus Swiftia, and describe a new species collected by the Alvin submersible off the Pacific coast of Costa Rica. The new species has been observed at around 1000 m depth, growing on authigenic carbonates near methane seeps. Swiftia sahlingi sp. nov. is characterised by having bright red colonies that are with limited branching, with slightly raised polyp-mounds, thin coenenchyme mainly composed of long warty spindles, and conspicuous plates. A molecular phylogenetic analysis supports the differences between this new taxon and the closest Swiftia species. The new species represents the first record of the genus from Costa Rica and in fact for the Eastern Tropical Pacific.

Revision of the genus Adelogorgia Bayer, 1958 (Cnidaria: Anthozoa: Octocorallia) with the description of three new species.

The genus Adelogorgia is distinguished from other holaxonians in having conspicuously ornamented double-disc sclerites and leaf clubs in the coenenchyme, and non-mineralised axis cores. The two eastern Pacific species currently recognised as Adelogorgia are diagnosed and illustrated. Three new species for the genus are described from new localities and depth ranges. Analysis of external and internal characters, especially sclerite colours and sizes, and colony colour, shape and branching, allows separating the species. An identification key to the five species is provided, as well as a character table for comparisons. This study was based on newly collected specimens from 50 to 200 m deep, and re-examination of all historical material. We conclude that the genus comprises five valid species with a wider distribution than previously reported. This research is a contribution to the octocoral systematics and biodiversity from mesophotic and deep waters.

A new Muricea species (Cnidaria, Anthozoa, Octocorallia) from the eastern tropical Pacific.

The genus Muricea is considered abundant and widely distributed along the eastern Pacific. Its occurrence in shallow waters has been recognised; however species from deeper than 30 m have been rarely recorded. During the 2005 R/V Urracá expedition along the north and central Pacific coast of Costa Rica several octocoral specimens were collected by bottom trawling from 30 to 150 m yielding new species and new records. Herein we describe a new species of Muricea from deeper than 30 m. The morphological characters of the species were analysed and illustrated by optic and scanning electron microscopy. Muricea subtilissp. n. can be distinguished from the other species in the genus by its thin spiny branches, non-imbricate calyces, white colony and sclerites, and the size and composition of sclerites. Comparative character tables are provided for the closest Muricea species-group. This new species increases the number in the genus to 26, and contributes to the knowledge on the diversity and distribution of mesophotic soft corals in the eastern Pacific.

A revision of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part II.

The species of the genus Muricea were mainly described from 1846 to 1870. After that very few contributions were published. Although the highest richness of Muricea species is in the eastern Pacific shallow waters, a comprehensive systematic study of the genus does not exist. Recently we started a taxonomic review of the genus in order to validate the status of four species previously included in the genus Eumuricea. Herein we present the second part of the Muricea revision dealing with the species-group characterised by shelf-like calyces instead of tubular-like calyces (the Muricea squarrosa-group). Original type material was morphologically analysed and illustrated using optical and scanning electron microscopy. Comparative character tables are provided for the genus. The taxonomic status of the species was analysed and established by designating lectotypes, alternatively by recognising a holotype by monotypy. We conclude that the genus Muricea comprises 20 valid species, including the previous four in the Muricea squarrosa-group. We propose 10 lectotypes, a new combination and three more species groups for the genus based on morphology: the Muricea fruticosa-group, Muricea plantaginea-group and Muricea austera-group.

Homoplasious colony morphology and mito-nuclear phylogenetic discordance among Eastern Pacific octocorals.

Octocorals are a diverse and ecologically important group of cnidarians. However, the phylogenetic relationships of many octocoral groups are not well understood and are based mostly on mitochondrial sequence data. In addition, the discovery and description of new gorgonian species displaying unusual or intermediate morphologies and uncertain phylogenetic affinities further complicates the study of octocoral systematics and raises questions about the role played by processes such as plasticity, crypsis, and convergence in the evolution of this group of organisms. Here, we use nuclear (i.e. 28S rDNA) and mitochondrial (mtMutS) markers and a sample of Eastern Pacific gorgonians thought to be remarkable from a morphological point of view to shed light on the morphological diversification among these organisms. Our study reveals the loss of the anastomosed colony morphology in two unrelated lineages of the seafan genus Pacifigorgia and offers strong evidence for the independent evolution of a whip-like morphology in two lineages of Eastern Pacific Leptogorgia. Additionally, our data revealed one instance of mito-nuclear discordance in the genera Leptogorgia and Eugorgia, which may be the results of incomplete lineage sorting or ancient hybridization-introgression events. Our study stresses the importance of comprehensive taxonomic sampling and the use of independent sources of evidence to address the phylogenetic relationships and clarifying the evolution of octocorals.

Corrigenda: Breedy O, Guzman HM (2015) A revision of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part I: Eumuricea Verrill, 1869 revisited. ZooKeys 537: 1-32. doi: 10.3897/zookeys.537.6025.

[This corrects the article DOI: 10.3897/zookeys.537.6025.].

A new alcyonacean octocoral (Cnidaria, Anthozoa, Octocorallia) from Chilean fjords.

A new species, Swiftia comauensis, is described from Chile. It occurs in shallow waters from 18 to 59 m in the Patagonian fjord region and seems to be endemic to the northern part of the region. The species is characterized by having straggly colonies with sparse branching and long drooping branches, prominent polyp mounds, and long, thin spindles; the colonies are bright orange with pale yellow polyp mounds. A sharp decline in colony abundance was observed between 2003 and 2013, and in January 2014 a proposal was submitted to the IUCN for the addition of this taxon to the Red List of Threatened Species.

A revision of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part I: Eumuricea Verrill, 1869 revisited.

Muricea is an amphi-American genus. Verrill proposed dividing the species from the Pacific Ocean into three genera and established the genus Eumuricea for five eastern Pacific species with tubular calyces. Eumuricea is basically characterized by colonies with elongate, cylindrical calyces with truncate margins and star-like opercula, and the occurrence of unilateral spinous spindles. According to these characteristics, Eumuricea does not show enough difference from Muricea to be treated as a separate genus. Original type material of Eumuricea was morphologically analysed and illustrated using optical and scanning electron microscopy. We conclude that the eastern Pacific species should be placed in the genus Muricea and form a group characterised by tubular calyces that comprises four species at present, Muricea acervata, Muricea hispida, Muricea squarrosa, and Muricea tubigera and a dubious species Muricea horrida. Lectotypes were designated for Muricea squarrosa and Muricea hispida to establish their taxonomic status. The genus Eumuricea has also been misunderstood by former authors who erroneously assigned species to it. For these species we propose new combinations: Swiftia pusilla, Astrogorgia splendens and Astrogorgia ramosa.

Two new species of gorgonian octocorals from the Tropical Eastern Pacific Biogeographic Region (Cnidaria, Anthozoa, Gorgoniidae).

The gorgoniid Eugorgia is exclusively an eastern Pacific genus. It has a wide geographic and bathymetric range of distribution, found from California to Perú and extends down to 65 m deep. Two new species are herein described. The morphological characters were analyzed and illustrated by light and scanning electron microscopy. Eugorgia beebei sp. n. can be distinguished by its white, ascending, sparse colony growth. Eugorgia mutabilis sp. n. can be distinguished by its white colony that changes color after collection, and the conspicuous sharp-crested disc sclerites. From a morphological point of view the new species are related to the daniana-group, the rubens-group and the siedenburgae-group of Eugorgia; their affiliations, and the proposal of a new group are discussed. These new species increases the number of species in the genus to 15, and contribute to the knowledge of the eastern Pacific octocoral biodiversity.

Species composition and bathymetric distribution of gorgonians (Anthozoa: Octocorallia) on the Southern Mexican Pacific coast.

Gorgonians are important components of coastal ecosystems, as they provide niches, natural compounds with medical applications and are used as bioindicators. Species composition and assemblage structure of gorgonians (Anthozoa: Octocorallia) were studied along a bathymetric profile in the Southern Mexican Pacific coast. Species composition was based on specimens collected within a depth range of 0-70 m in 15 sites. The relative abundance of species was determined in six sites at four depths (5, 10, 20 and 25 m) using three 10 m2 transects at each depth level. Twenty-seven species of gorgonians belonging to six genera and three families were registered. The species composition varied with depth: 11 species were distributed between 0-25m depth, while 17 species were found between 40-70 m depth interval. The shallow zone is characterized by a relatively large abundance of gorgonians, dominated by colonies of Leptogorgia cuspidata and L. ena. In contrast, the deepest zone was characterized by relatively low abundance of gorgonians, dominated by L. alba, the only species observed in both depth intervals. The similarity analysis showed differences in the composition and abundance of species by depth and site, suggesting that the main factor in determining the assemblage structure is depth. Results of this study suggest that the highest richness of gorgonian species in the study area may be located at depths of 40-70 m, whereas the highest abundances are found between 5 and 10 m depth. This study represents a contribution to the poorly known eastern Pacific gorgonian biota.

Species composition and bathymetric distribution of gorgonians (Anthozoa: Octocorallia) on the Southern Mexican Pacific coast / Composición de especies y distribución batimétrica de gorgonios (Anthozoa: Octocorallia) en la costa sur del Pacífico mexicano

Gorgonians are important components of coastal ecosystems, as they provide niches, natural compounds with medical applications and are used as bioindicators. Species composition and assemblage structure of gorgonians (Anthozoa: Octocorallia) were studied along a bathymetric profile in the Southern Mexican Pacific coast. Species composition was based on specimens collected within a depth range of 0-70m in 15 sites. The relative abundance of species was determined in six sites at four depths (5, 10, 20 and 25m) using three 10m2 transects at each depth level. Twenty-seven species of gorgonians belonging to six genera and three families were registered. The species composition varied with depth: 11 species were distributed between 0-25m depth, while 17 species were found between 40-70m depth interval. The shallow zone is characterized by a relatively large abundance of gorgonians, dominated by colonies of Leptogorgia cuspidata and L. ena. In contrast, the deepest zone was characterized by relatively low abundance of gorgonians, dominated by L. alba, the only species observed in both depth intervals. The similarity analysis showed differences in the composition and abundance of species by depth and site, suggesting that the main factor in determining the assemblage structure is depth. Results of this study suggest that the highest richness of gorgonian species in the study area may be located at depths of 40-70m, whereas the highest abundances are found between 5 and 10m depth. This study represents a contribution to the poorly known eastern Pacific gorgonian biota.

La composición de especies y estructura de la comunidad de gorgonáceos se determinó a lo largo de un perfil batimétrico en la costa suroeste del Pacífico Mexicano. La composición de especies se realizó a través de una revisión y recolecta de especímenes entre 0 y 70m de profundidad en 15 sitios de muestreo. La caracterización de la estructura de la comunidad se realizó en seis sitios, en los cuales se colocaron tres transectos de 10m² cada uno a 5, 10, 20 y 25m de profundidad. Se encontraron 27 especies de gorgonáceos pertenecientes a seis géneros y tres familias; aunque seis especies requieren ser confirmadas. La mayor riqueza de especies se registró entre 40 y 70m. La estructura de la comunidad entre 5 y 25m de profundidad mostró la existencia de dos zonas con la misma composición de especies pero con diferente abundancia relativa: una somera entre 5 y 10m de profundidad, caracterizada por una mayor abundancia de gorgonáceos y dominado por colonias de Leptogorgia cuspidata y Leptogorgia ena y una zona profunda entre 20 y 25m de profundidad, caracterizada por una baja abundancia de gorgonáceos y dominada por colonias de Leptogorgia alba.

Characterization of deepwater invertebrates at Isla del Coco National Park and Las Gemelas Seamount, Costa Rica / Caracterización de invertebrados de aguas profundas en el Parque Nacional Isla del Coco y Monte Submarino Las Gemelas, Costa Rica

The deepwater faunas of oceanic islands and seamounts of the Eastern Tropical Pacific are poorly known. From 11-22 September 2009, we conducted an exploration of the deepwater areas around Isla del Coco National Park and Las Gemelas Seamount, located about 50km southwest of Isla del Coco, Costa Rica using a manned submersible to survey the seafloor habitats. The goal of the exploration was to characterize the habitats and biota, and conduct quantitative surveys of the deepwater portions of Isla del Coco National Park and Las Gemelas. We completed a total of 22 successful submersible dives, spanning more than 80hr underwater, and collected a total of 36hr of video. With respect to invertebrates, our objectives were to gather quantitative information on species composition, density, distribution and habitat associations as well as to compare the invertebrate communities between the two sites. A total of 7 172 invertebrates were counted from analysis of the video collected on this project. Larger organisms were counted and placed into 27 taxonomic groups to characterize the deepwater invertebrate fauna of Las Gemelas Seamount and Isla del Coco National Park. The Shannon-Weiner Index for biodiversity (H’) was calculated to be 0.14 ± 0.02 for Isla del Coco and 0.07 ± 0.03 for Las Gemelas surveys. Although richness was fairly equal between the two sites, evenness was greater at Isla del Coco (J = 0.04 ± 0.006) when compared to Las Gemelas (J = 0.02 ± 0.01). This lower level of evenness in the community at Las Gemelas was a result of high densities of a few dominant species groups, specifically sea urchins and black corals. We also evaluated invertebrate percent cover at both Isla del Coco and Las Gemelas Seamount with respect to habitat type, slope and rugosity. Results indicated that highly rugose habitats contained the highest frequencies of all invertebrates at both sites, with the exception of glass sponges and polychaetes at Isla del Coco, which were found in greater quantities at intermediate levels of rugosity. Information obtained from these submersible surveys indicate that seamounts in the tropical eastern Pacific Ocean may be an important source of biodiversity and that more quantitative surveys are needed to characterize the fauna of the region.

La fauna de aguas profundas de islas oceánicas y de montes submarinos del Pacífico Tropical Oriental son muy poco conocidas. Para caracterizar las faunas de aguas profundas del Parque Nacional Isla del Coco y el Monte Submarino Las Gemelas, Costa Rica, llevamos a cabo un estudio cuantitativo de los hábitats y su fauna. Se tomaron videos de transectos desde un sumergible entre 50 y 402m de profundidad del 11 al 22 de setiembre 2009. Se recolectó información cuantitativa de la composición de especies de invertebrados, densidad, distribución y hábitats asociados en ambas localidades y se comparó. Se contaron 7,172 invertebrados en los videos analizados, y se ubicaron en 27 categorías taxonómicas. El Índice de Shannon-Weiner (H’) fue de 0.14 ± 0.02 para la Isla del Coco y 0.07 ± 0.03 para Las Gemelas. La riqueza fue parecida en ambos sitios pero la equitabilidad fue mayor en la Isla del Coco (J = 0.04 ± 0.006) comparado con Las Gemelas (J = 0.02 ± 0.01). Este menor nivel de equitabilidad en Las gemelas de debió a la alta densidad de unos pocos grupos dominantes, específicamente erizos de mar y coral negro. También evaluamos el porcentaje de cobertura de los invertebrados con respecto a tipo de hábitat, pendiente y rugosidad del sustrato. Los resultados indican que hábitats con alta rugosidad contiene más invertebrados en ambos sitios, excepto la esponjas silíceas y los poliquetos en la Isla del Coco, que se encontraron en mayor cantidad en niveles intermedios de rugosidad. La información obtenida con este estudio indican que los montes submarinos en el Pacífico Tropical Oriental pueden ser una fuente importante de biodiversidad y que se necesitan más estudios cuantitativos para caracterizar la fauna de la región.

How many kinds of sclerite? Towards a morphometric classification of gorgoniid microskeletal components.

Gorgoniid octocorals constitute a diverse group of organisms that inhabit a wide range of marine environments. The group is currently defined by the presence of calcareous sclerites that are less than 0.3 mm in length with regularly arranged warts. Generic and specific classification schemes are based on the presence/absence of different sclerite classes in the sampled specimen as well as the frequency in which each class occurs in the sample. Sclerite classification typically has been difficult because a continuum of sclerite forms is found within and between species. Thus, the use of sclerites for phylogenetic inference and classification is problematic. Herein, we present a methodology to obtain quantitative measurements of large numbers of sclerites and used finite mixture modeling to assess the number of statistically different sclerite classes present in the eastern Pacific octocoral genus Pacifigorgia. We also test the ability of simple neural classifiers (perceptrons) to sort sclerites into the classes traditionally used in octocoral taxonomy. This methodology can be used for other gorgoniids and can be further extended to include shape quantifiers for groups other than those studied here.