First visual occurrence data for deep-sea cnidarians in the South-western Colombian Caribbean.

BACKGROUND: Attention to the deep-sea environment has increased dramatically in the last decade due to the rising interest in natural resource exploitation. Although Colombia holds a large submerged territory, knowledge of the seabed and its biodiversity beyond 1,000 m depth is very limited. During 2015-2017, Anadarko Colombia Company (ACC) carried out hydrocarbon exploratory activities in the South-western Colombian Caribbean, at depths between 375 m and 2,565 m. NEW INFORMATION: Capitalising on available data resources from these activities, several cnidarian species were observed in ROV and towed camera surveys. We analysed over nine hours of video and 5,066 still images from these surveys, identifying organisms to the lowest possible taxonomic level. The images and associated data presented here correspond to 108 observations of deep-sea cnidarians, including seven new records for the Colombian Caribbean. Given the paucity of research and funding to explore the deep-sea in Colombia, the present dataset comprises the largest deep-sea Cnidaria imagery inventory to date for the Colombian Caribbean.

The invasive snowflake coral (Carijoa riisei) in the Tropical Eastern Pacific, Colombia / El coral copo de nieve invasivo (Carijoa riisei) en el Pacífico Oriental Tropical, Colombia

Carijoa riisei (Octocorallia: Cnidaria), a western Atlantic species, has been reported in the Pacific as an invasive species for nearly forty years. C. riisei has been recently observed overgrowing native octocorals at several rocky-coral littorals in the Colombian Tropical Eastern Pacific-(TEP). C. riisei has inhabited these reefs for at least 15 years but the aggressive overgrowth on other octocorals have been noted until recently. Here, we surveyed for the first time the distribution and inter-specific aggression by C. riisei in both coastal and oceanic areas colonized in the Colombian TEP (Malpelo, Gorgona and Cabo Corrientes), including preliminary multiyear surveys during 2007-2013. We observed community-wide octocoral mortalities (including local extinction of some Muricea spp.) and a steady occurrence of competing and overgrowing Pacifigorgia seafans and Leptogorgia seawhips. In Gorgona Island, at two different sites, over 87% (n=77 tagged colonies) of octocorals (Pacifigorgia spp. and Leptogorgia alba) died as a result of C. riisei interaction and/or overgrowth between 2011 and 2013. C. riisei overgrows octocorals with an estimate at linear growth rate of about 1cm m-1. The aggressive overgrowth of this species in TEP deserves more attention and regular monitoring programs. Rev. Biol. Trop. 62 (Suppl. 1): 199-207. Epub 2014 February 01.

Carijoa riisei (Octocorallia: Cnidaria), una especie nativa del Atlántico Occidental. Sin embargo, se ha reportado como especie exótica invasora en el Pacífico desde hace unos cuarenta años y en la costa occidental de Colombia hace 15-20 años. Recientemente C. riisei se ha observado sobrecreciendo octocorales nativos en varios litorales rocosos coralinos en el Pacífico Oriental Tropical-(POT) de Colombia. C. riisei ha habitado estos arrecifes durante al menos 15 años, pero este comportamiento agresivo frente a otros octocorales no se habían notado hasta hace unos pocos años. En este estudio, se evalúa por primera vez, en las zonas costeras y oceánicas, la distribución e incidencia C. riisei en otros octocorales del POT colombiano (Malpelo, Gorgona y Cabo Corrientes), incluyendo información observaciones entre 2007-2013. Hemos observado mortalidad en toda la comunidad de octocorales (incluyendo la extinción local de algunas especies de Muricea) y una constante competencia y cubrimiento de abanicos de mar Pacifigorgia y gorgonáceos Leptogorgia. C. riisei presento una tasa lineal de crecimiento sobre otros octocorales de aproximadamente 1 cm m-1. El comportamiento agresivo de C. riisei en el POT merece más atención y programas regulares de monitoreo.

A semi-automatic method to extract canal pathways in 3D micro-CT images of Octocorals.

The long-term goal of our study is to understand the internal organization of the octocoral stem canals, as well as their physiological and functional role in the growth of the colonies, and finally to assess the influence of climatic changes on this species. Here we focus on imaging tools, namely acquisition and processing of three-dimensional high-resolution images, with emphasis on automated extraction of canal pathways. Our aim was to evaluate the feasibility of the whole process, to point out and solve - if possible - technical problems related to the specimen conditioning, to determine the best acquisition parameters and to develop necessary image-processing algorithms. The pathways extracted are expected to facilitate the structural analysis of the colonies, namely to help observing the distribution, formation and number of canals along the colony. Five volumetric images of Muricea muricata specimens were successfully acquired by X-ray computed tomography with spatial resolution ranging from 4.5 to 25 micrometers. The success mainly depended on specimen immobilization. More than [Formula: see text] of the canals were successfully detected and tracked by the image-processing method developed. Thus obtained three-dimensional representation of the canal network was generated for the first time without the need of histological or other destructive methods. Several canal patterns were observed. Although most of them were simple, i.e. only followed the main branch or "turned" into a secondary branch, many others bifurcated or fused. A majority of bifurcations were observed at branching points. However, some canals appeared and/or ended anywhere along a branch. At the tip of a branch, all canals fused into a unique chamber. Three-dimensional high-resolution tomographic imaging gives a non-destructive insight to the coral ultrastructure and helps understanding the organization of the canal network. Advanced image-processing techniques greatly reduce human observer's effort and provide methods to both visualize and quantify the structures of interest.

Phylogenetic hypotheses of gorgoniid octocorals according to ITS2 and their predicted RNA secondary structures.

Gorgoniid octocorals taxonomy (Cnidaria; Octocorallia; Gorgoniidae) includes diagnostic characters not well defined at the generic level, and based on the family diagnosis some species could be classified in either Gorgoniidae or Plexauridae. In this study, we used sequences from the Internal Transcribed Spacer 2 (ITS2) and their predicted RNA secondary structure to both correct the alignment and reconstruct phylogenies using molecular morphometrics for 24 octocorals mostly from the Atlantic. ITS2 exhibited the six-helicoidal ring-model structure found in eukaryotes, and provided 38 parsimony-informative characters. The proposed phylogenies, though differing between sequence- and structure-base results, provided consistent support for several clades. Genera considered part of the polyphyletic genus Leptogorgia, such as Filigorgia, were distantly related to the former in all phylogenetic hypotheses. Main differences among the hypotheses consisted in the placement of Muriceopsis (previously considered from the Plexauridae family) and Filigorgia. Excluding Muriceopsis and an undescribed octocoral from Tobago, Plexaurella and Pterogorgia grouped together as a sister branch of Pinnigorgia spp. but long-branch attraction was evident for the grouping of Plexaurella nutans (another plexaurid) and Pterogorgia citrina. Unexpected results were the divergence between Caribbean genera, Gorgonia and Pseudopterogorgia, which were placed basal respect to Pacifigorgia and Leptogorgia (=Lophogorgia). ITS2 provided support to corroborate observations based on sclerite morphology: species with "capstan sclerites" (e.g., Pacifigorgia and Leptogorgia) were characterized by a long helix IV with one internal loop and a helix V with four internal loops; "scaphoid sclerites" had a predominantly long helix V if compared to helix IV; "asymmetric spiny sclerites" (Muriceopsis, Pinnigorgia and the undescribed octocoral) exhibited one or two lateral bulges in the V helix. Remarkably, Muriceopsis and Pinnigorgia were supported by a complete Compensatory Base Change (CBC) (A-U to G-C) in helix V. Filigorgia with simple "spindles" had a short helix IV and a large central ring. DNA sequences from the nuclear ITS2 region, including information from predicted RNA secondary structure, despite their reduced length, provided numerous characters and phylogenetic information among Gorgoniidae genera and species.

Do multi-branched colonial organisms exceed normal growth after partial mortality?

One of the advantages of modular colonial growth is the capability to recover after partial mortality. Tolerance to partial mortality is a known property of some resistant species of plants that respond to mortality with vigorous regrowth or overcompensation. It is not clear whether modular marine invertebrates such as octocorals overcompensate. This study provides evidence that following injury to colonies (by breaking apical dominance), new growth exceeds normal rates of branching, as observed in some plants, in a degree correlated to the original multi-branched network setting (e.g. the number of original branches connected to main stem), in colonies of the Caribbean gorgonian octocoral Pseudopterogorgia bipinnata. This can be explained by the network of communicating vessels and canals inside octocoral colonies, which provide the structure for effective allocation of resources to regenerating parts.

Branching and self-organization in marine modular colonial organisms: a model.

Despite the universality of branching patterns in marine modular colonial organisms, there is neither a clear explanation about the growth of their branching forms nor an understanding of how these organisms conserve their shape during development. This study develops a model of branching and colony growth using parameters and variables related to actual modular structures (e.g., branches) in Caribbean gorgonian corals (Cnidaria). Gorgonians exhibiting treelike networks branch subapically, creating hierarchical mother-daughter relationships among branches. We modeled both the intrinsic subapical branching along with an ecological-physiological limit to growth or maximum number of mother branches (k). Shape is preserved by maintaining a constant ratio (c) between the total number of branches and the mother branches. The size frequency distribution of mother branches follows a scaling power law suggesting self-organized criticality. Differences in branching among species with the same k values are determined by r (branching rate) and c. Species with r<r/2 or c>r>0). Ecological/physiological constraints limit growth without altering colony form or the interaction between r and c. The model described the branching dynamics giving the form to colonies and how colony growth declines over time without altering the branching pattern. This model provides a theoretical basis to study branching as a simple function of the number of branches independently of ordering- and bifurcation-based schemes.

Determinate growth and modularity in a gorgonian octocoral.

Growth rates of branches of colonies of the gorgonian Pseudopterogorgia elisabethae were monitored for 2 years on a reef at San Salvador Island, Bahamas. Images of 261 colonies were made at 6-month intervals and colony and branch growth analyzed. Branch growth rates differed between colonies and between the time intervals in which the measurements were made. Colonies developed a plumelike morphology through a pattern of branch origination and determinate growth in which branch growth rates were greatest at the time the branch originated and branches seldom grew beyond a length of 8 cm. A small number of branches had greater growth rates, did not stop growing, and were sites for the origination of subsequent "generations" of branches. The rate of branch origination decreased with each generation of branching, and branch growth rates were lower on larger colonies, leading to determinate colony growth. Although colonial invertebrates like P. elisabethae grow through the addition of polyps, branches behave as modules with determinate growth. Colony form and size is generated by the iterative addition of branches.

Patterns of morphological integration in marine modular organisms: supra-module organization in branching octocoral colonies.

Despite the relative simplicity of their modular growth, marine invertebrates such as arborescent gorgonian octocorals (Octocorallia: Cnidaria) generate complex colonial forms. Colony form in these taxa is a consequence of modular (polyp) replication, and if there is a tight integration among modular and supramodular traits (e.g. polyp aperture, inter-polyp spacing, branch thickness, internode and branch length), then changes at the module level may lead to changes in colony architecture. Alternatively, different groups of traits may evolve semi-independently (or conditionally independent). To examine the patterns of integration among morphological traits in Caribbean octocorals, we compared five morphological traits across 21 species, correcting for the effects of phylogenetic relationships among the taxa. Graphical modelling and phylogenetic independence contrasts among the five morphological characters indicate two groups of integrated traits based on whether they were polyp- or colony-level traits. Although all characters exhibited bivariate associations, multivariate analyses (partial correlation coefficients) showed the strongest integration among the colony-level characters (internode distance and branch length). It is a quantitative demonstration that branching characters within the octocorals studied are independent of characters of the polyps. Despite the universally recognized modularity of octocorals at the level of polyps, branching during colony development may represent an emergent level of integration and modularity.