Actiniaria: Actiniidae)

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Actiniaria: Actiniidae) Cnidae sizes in the two morphotypes of the giant Caribbean anemone Condylactis gigantea (Actiniaria: Actiniidae) Ricardo González-Muñoz1*, Carlos Hernández-Ortiz2, Agustín Garese1, Nuno Simões3 & Fabián Horacio Acuña1 1. Laboratorio de Biología de Cnidarios, Instituto de Investigaciones Marinas y Costeras, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata-CONICET, Rodríguez Peña 4046, 7600 Mar del Plata, Argentina; [email protected], [email protected], [email protected] 2. Departamento de Acuacultura y Biología Marina, Escuela de Ciencias Aplicadas del Mar, Universidad de Oriente, Núcleo Nueva Esparta, Boca de Río, Isla de Margarita, Venezuela; [email protected] 3. Unidad Multidisciplinaria de Docencia e Investigación en Sisal (UMDI-Sisal), Facultad de Ciencias, Universidad Nacional Autónoma de México (UNAM), Puerto de Abrigo, Sisal, 97356, Yucatán, México; International Chair of Coastal and Marine Studies in Mexico, Harte Research Institute, Texas A&M at Corpus Christi, TX, USA; Laboratorio Nacional de Resiliencia Costera (LANRESC); [email protected] Received 15-II-2018. Corrected 29-V-2018. Accepted 27-VI-2018. Abstract: The sea anemone Condylactis gigantea is an ecologically important member of the benthic com- munity in coral reefs of the tropical Atlantic, and displays two morphotypes with respect to the color in their tentacular tips: the green tip morphotype and the pink/purple tip morphotype. Although some molecular and ecological differences have been found between these morphotypes, no other morphological distinctions have been reported, and currently both are still considered a single taxonomic species. In the present study, we perform an exploration on the variability in the size of cnidae between these two morphotypes and performed statistical analyses to compare the 10 categories of cnidae from specimens hosted in the Cnidarian Collection of Gulf of Mexico and Mexican Caribbean, of the Universidad Nacional Autónoma de México, which were previ- ously collected in several coral reefs localities of the Yucatán Peninsula. Results reveal no significant variation in cnidae size between the two morphotypes, but significant variations were found within each morphotype. In addition, we update the composition of the cnidom of C. gigantea, and the utility of the size of cnidae to dis- tinguish between morphotypes or closely related species is discussed. Rev. Biol. Trop. 66(3): 1055-1064. Epub 2018 September 01. Key words: Cnidaria; Anthozoa; cnidom; coral reefs; morphotype; Condylactis gigantea. The giant Caribbean sea anemone Condy- shrimps (Silbiger & Childress, 2008; Colom- lactis gigantea (Weinland, 1860) (Actiniaria, bara, Quinn, & Chadwick, 2017), as well as Actiniidae) is one of the most common and symbiotic associations with some species of well-known actiniarian species that inhabits Caribbean fishes (Hanlon & Kaufman, 1976; in coastal and coral reefs environments of the Hanlon & Hixon, 1986). Moreover, this spe- Western Atlantic Ocean, and is distributed cies is also recognized as an important source from Bermuda to southeast Brazil, and along of biologically active compounds (e.g. Billen, the Gulf of Mexico and the Caribbean Sea Debaveye, Béress, Garateix, & Tytgat, 2010; (González-Muñoz, Simões, Sánchez-Rodrí- Romero et al., 2010; Santos et al., 2013) and, guez, Rodríguez, & Segura-Puertas, 2012). due to its brightly colors and attractive forms, it This species is an ecologically important is much appreciated in the aquarium trade (Chi- member of the benthic community providing appone, Swanson, & Miller, 2001; Sheridan, habitat for several species of caridean cleaner Fautin, & Garret, 2015). Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 66(3): 1055-1064, September 2018 1055 Fig. 1. Morphotypes of Condylactis gigantea: a. Green morphotype. b. Pink/Purple morphotype. Cnidom of C. gigantea per tissue type. Actinopharynx: c. small basitrich. d. large basitrich. Column: e. small basitrich. f. large basitrich. Filaments: g. small basitrich. h. large basitrich. i. microbasic b-mastigophore. j. microbasic p-mastigophore. Tentacles: k. basitrich. l. spirocysts. Condylactis gigantea displays two main Cook, Lang, Petelle, & Blumstein, 2012). morphotypes, particularly with respect to color However, no other morphological differences in the tentacular tips, which can be categorized between these two morphotypes have been as the green tip (Fig. 1A) and the pink/purple reported and both are currently considered a tip morphotypes (Fig. 1B), although some indi- single taxonomic species. viduals with whitish, yellowish, or bluish ten- Despite the size of cnidae alone is not tacular tips can be also rarely found. Previous necessarily a conclusive taxonomic character genetic comparisons with DNA sequence data to differentiate between species due to its vari- have found some differences between these ability within conspecific individuals (Fautin, two morphotypes from specimens of different 2009; Garese, Carrizo, & Acuña, 2016), some depths and reef areas, as well as differences in studies consider them as an additional specific UV absorbance capacities, suggesting reduced taxonomic characteristic to distinguish between gene flow and ecological differentiation among closely related species, but only when these the green and the pink/purple morphotypes at differences are accompanied by other mor- a small geographic scale (Stoletzki & Schi- phological or ecological distinctions (Fautin, erwater, 2005). Other differences between 1988). Some studies including quantitative these morphotypes have also been observed on analyses of the size of cnidae to distinguish personality and habitat segregation (Hensley, between closely related species or between 1056 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 66(3): 1055-1064, September 2018 morphotypes suggests results of two opposite actinopharynx, mesenteric filaments, and mid sorts. Some studies found significant statistical column. Terminology follows Östman (2000). differences in cnidae sizes comparisons (e.g. From each of the four squash preparations, the Allcock, Watts, & Thorpe, 1998; Watts & Thor- length and width of 30 undischarged capsules pe, 1998; Manchenko, Dautova, & Latypov, (replicates) of each type of cnidae were ran- 2000; Watts, Allcock, Lynch, & Thorpe, 2000), domly measured using DIC microscopy 1 000x while other attempts did not find statistical sup- oil immersion (following Williams, 1996). Cni- port to clearly separate species or morphotypes dae preparations were deposited in the same based on cnidae size differences (e.g. Solé- Cnidarian collection. Cava & Thorpe, 1987; Chintiroglou & Karalis, Cnidae samples were ordered in a bi- 2000; González-Muñoz et al., 2015; González- dimensional space using principal component Muñoz, Garese, Tello-Musi, & Acuña, 2017). analysis (PCA). Differences in ordination In the present study, we performed an given by morphotype and individual specimens exploration on the variability in the size of cni- within each morphotype were analyzed using a dae and statistically analyze them to compare permutational multivariate analysis of variance between the two morphotypes of C. gigantea, (PERMANOVA) procedure (Anderson, 2001). from preserved specimens of the Collection of Differences among cnidae size were analyzed Cnidarians of the Gulf of Mexico and Mexi- for each type of cnidae and tissue separately. can Caribbean, of the Universidad Nacional The PERMANOVA procedure was applied on Autónoma de México (UNAM). In addition, resemblance matrices based on the Euclidian we update the composition of the cnidom of C. distance between samples. Although length and gigantea, and discuss about the utility of the width of the capsules were in the same mea- size of cnidae to distinguish between closely related species or between morphotypes. surement unit, data were normalized prior to analyses. The statistical model used was given by: Yijkl= Mi+I(M)j(i)+Tk+MTik+I(M)Tj(i)k+eijkl, MATERIALS AND METHODS where Y is the response matrix with n samples * P = 2 variables (number of columns: length and Five specimens of each of the two mor- width); M is a fixed factor representing mor- photypes were selected from the Collection photype (with two levels); I is a random factor of Cnidarians of the Gulf of Mexico and representing individuals nested in M (with five Mexican Caribbean Sea (Registration code: YUC-CC-254-11) of the Unidad Multidisci- levels); T is the fixed factor representing type plinaria de Docencia e Investigación - Sisal of cnidae and is orthogonal to M and I; MT and (UMDI-Sisal) at the UNAM. Specimens were I(M)T are corresponding interactions terms; selected based on the photographs of the living and e is the residual matrix. Permutation pro- specimens which are included in each record cedures were applied to obtain appropriate dis- of the collection. All specimens were collected tributions for the pseudo-F statistic under the in coastal coral reefs localities along the Yuca- null hypothesis. All analyses were performed tán Peninsula (Appendix). Pedal disc diam- using 999 permutations of residuals under the eter and column height of the samples were reduced model. The experimental design was measured from living specimens (Appendix); balanced in every case, and the partitioning of however, comparisons between morphotypes variation was achieved so that the statistic test were made regardless the size of the specimens represents the proportion of the variation in the
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