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Species of dendrophylliid corals analysed for this study. A) Dendrophyllia cornigera; B) D. laboreli; C) Astroides calycularis; D) D. ramea. Photographs A-C by PJL-G, D by Diego Moreno Lampreave.
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Scleractinian corals are widely distributed in all oceans and at all bathymetric levels. Corals are among the most important bio-building organisms in marine ecosystems. The systematics of this hexacoral group is currently undergoing much change owing to studies that combine molecular analyses with morphological research on the calcareous skeletons...
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... of the dendrophylliid species studied here are Dendrophyllia ramea (Linné, 1758), D. cornigera (Lamarck, 1816), andD. laboreli Zibrowius & Brito, 1984, and the fourth is the only species recognised in the genus genus Astroides, viz., A. calycularis (Pallas, 1766) (Fig. 1). All the examined colonies of Dendrophyllia ramea were collected in Rota (Cádiz, Spain, Atlantic side of the Strait of Gibraltar, Jun. 2009) at a depth of 20 m. One of the examined colonies of Dendrophyllia cornigera was collected during the CORAL 8 expedition (Deep-coral, Stn 16, Sep. 2007), in the submarine cannon of Cap de Creus ...
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... interspecific relationships among the species studied here (also including information previously obtained from two European Balanophyllia species) showed six distinct groups, each including all sampled specimens from each examined dendrophylliid species (Fig. S1, available as Supplementary Material on the journal website). The similarity for all examined dendrophylliids was over 70%, increasing to 85 and 95% at intraspecific level. Other scleractinians (Mussiidae), corallimorpharians, and actiniarians included in the analysis are widely separated by less than 45% ...
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... In that contribution the examination of basitrichs by image analysis (as basitrichs and microbasic b-mastigophores) of nine different cnidarian species, demonstrated that these basitrichs were clearly distinguishable from one another. These authors emphasised that this kind of analysis is able to contribute to the systematics of cnidarians. Fig. 10. Example concerning all hypothetical evolutionary pathways producing the different holotrich composition characters found in a tissue (e.g pharynx, three holotrichs, H1 to ...
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... this study we consider that the diversification and appearance of the different holotrichs categories in a tissue is only related to themselves. In Fig. 10, we show a hypothetical example concerning all possible evolutionary pathways producing the different holotrich composition categories found in a tissue (e.g pharynx, as in this study it has three possible holotrich categories, H1, H2, and H3). In this figure '1' and '0' only indicate presence or absence, respectively, without any ...
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... Fig. 10, we show a hypothetical example concerning all possible evolutionary pathways producing the different holotrich composition categories found in a tissue (e.g pharynx, as in this study it has three possible holotrich categories, H1, H2, and H3). In this figure '1' and '0' only indicate presence or absence, respectively, without any indication of polarity. Thus, the configuration (100) indicates the presence of a single character (H1) while the configuration (011) indicates the presence of two characters (H2 and H3). ...
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Citations
... Studies on cnidae and their distribution and size have been widely developed, particularly in anthozoans (Weill 1934, Carlgren 1940, Cutress 1955, Schmidt 1969, 1972, Mariscal 1974, 1984, den Hartog 1977, Östman 1988, england 1991, Fautin and Mariscal 1991, Pires and Pitombo 1992, Pires 1997, Acuña et al. 2003, Terrón-Sigler and lópez-gonzález 2005, Fautin 2009, Picciani et al. 2011, garese et al. 2016). The analysis of morphometrical data of cnidae from statistical approaches (Thomason 1988, Zamponi and Acuña 1991, 1994, Allcock et al. 1998, Watts et al. 2000, Ardelean and Fautin 2004, Williams 1996, 1998, Acuña and Zamponi 1997, Chintiroglou 1996, Chintiroglou et al. 1997, Östman 2000, Francis 2004, Kramer and Francis 2004, Acuña et al. 2004, 2007, 2011, garese et al. 2016, as well as the study of intraspecific variations of cnidae in response to ecological conditions (Martínez-Baraldés et al. 2014, gonzález-Muñoz et al. 2015, 2017, are among the most studied topics related with the utility of cnidom to taxonomic purposes. ...
The cnidae are the exclusive diagnostic structures of phylum Cnidaria. The inventory of all
cnidae types of a particular species is called the cnidom. The study of cnidae has been
widely addressed in all classes of cnidarians. Particularly in the order Actiniaria (sea
anemones), the study of the composition, size and distribution of cnidae is essential to the
identification and description of species. In the present study, we examine the cnidom of
the sea anemone Aulactinia marplatensis in three different stages of development
throughout its life cycle. We found that the composition and abundance patterns are very
similar between the adult and juvenile stages, although significant differences in the size
capsules were found between both stages and in all cnidae types observed, being bigger
those from the adult forms. The planula larvae stage presents a less diverse cnidom in
comparison to the juvenile and adult stages; however, it present an exclusive cnidae type
(the mesobasic p-mastigophore) which is the biggest in size of all the cnidae types observed
in the species. These results highlight the importance of considering the stage of
development when cnidae is used as a diagnostic character, and the particular relevance of
the study of the cnidom in larval stages.
... Recent works demonstrated that soft-tissue traits are taxonomically valid and increased phylogenetic resolution. although some caveats should be made (Martinez-Barald es et al., 2014;Cordie and Budd, 2016). Embryogenetic morphological characters distinguish "complex" from "robust" corals, suggesting that patterns of early development and gastrulation are phylogenetically informative rather than being correlated with habitat or reproductive mode (Okubo et al., 2013;Okubo, 2016). ...
Extant biodiversity can easily be underestimated owing to the presence of cryptic taxa, even among commonly observed species. Scleractinian corals are challenging to identify because of their ecophenotypic variation and morphological plasticity. In addition, molecular analyses have revealed the occurrence of cryptic speciation. Here, we describe a new cryptic lobophylliid genus and species Paraechinophyllia variabilis gen. nov., sp. nov., which is morphologically similar to Echinophyllia aspera and E. orpheensis. The new taxon occurs in Mayotte Island, Madagascar, the Gulf of Aden and the Red Sea. Six molecular markers (COI, 12S, ATP6‐NAD4, NAD3‐NAD5, histone H3 and ITS) and 46 morphological characters at three different levels (macromorphology, micromorphology and microstructure) were examined. The resulting molecular phylogenetic reconstruction showed that Paraechinophyllia gen. nov. represents a distinct group within the Lobophylliidae that diverged from the lineage leading to Echinophyllia and Oxypora in the Early Miocene, approximately 21.5 Ma. The morphological phylogenetic reconstruction clustered Paraechinophyllia gen. nov., Echinophyllia and Oxypora together in a single clade. A sole morphological character, calice relief, discriminated Paraechinophyllia gen. nov. from the latter two genera, suggesting that limited morphological variation has occurred over a long period. These results highlight the importance of cryptic taxa in reef corals, with implications for population genetics, ecological studies and conservation.
... Although cnidae size alone is not generally considered a specific taxonomic diagnostic character due to its variability within conspecific individuals (Fautin 1988(Fautin , 2009Acuña et al. 2003Acuña et al. , 2004Acuña and Garese 2009;Garese et al. 2016), several studies suggest that quantitative analyses of cnidae size could help to distinguish between morphotypes or species when correlate with other morphological or ecological characters (Allcock et al. 1998;Watts and Thorpe 1998;Manchenko et al. 2000;Watts et al. 2000;Acuña et al. 2003;Martínez-Baraldés et al. 2014). ...
Lebrunia coralligens is among the most common sea anemone species found inhabiting in coral reefs environments along the Caribbean Sea and Gulf of Mexico. This species exhibit two distinct morphotypes characterized mainly by differences in the location of vesicles on its pseudotentacles. The morphology and cnidae size ranges between representatives of the two morphotypes, as well as with those from specimens of its congeners Lebrunia neglecta, were compared to identify taxonomic distinctions that would enable separation of the morphotypes into different species. Statistical variability in specimens and cnidae size ranges was found between the two morphotypes, but a greater variability was found when compared with cnidae size ranges of L. neglecta. These results suggest that pseudotentacular form and cnidae size differences are due to broad phenotypic plasticity within L. coralligens. In addition, results also suggest that quantitative comparisons of cnidae could reinforce the taxonomic distinction of closely related species as L. coralligens and L. neglecta.
... Although the intraspecific variation suggests decreasing taxonomic value of cnidocysts, their study is still useful in other kinds of research, such as comparisons between different populations of the same species (Acuña and Zamponi 1997), differentiation of morphotypes of a species (Allcock et al. 1998;Watts et al. 2000; Gonzále-Muñoz pers. com), or to establish more precise differences among closely related species (Watts et al. 2000;Martínez-Baraldés et al. 2014). Internal variation in a data set does not preclude a comparison with another data set because the variation between them could be different; however and hence these types of studies must be statistically well supported and suitable statistical tools should be used. ...
Many studies have dealt with data on the sizes of cnidae within different groups of anthozoans, such as Actiniaria, Corallimorpharia and Zoantharia. Statistical treatments of these data have been variable, according to the evaluation of the normality; hence the use of parametric or non-parametric tests. The normality of cnidocyst size data was assumed or proved by some authors; who used parametric tests to make comparisons. Other authors carried out non-parametric tests, or even proposed alternative analytical methods, such as the use of generalized linear models. Despite controversy about the statistical distribution of cnidae sizes, there has never been an attempt to study the normality of cnidocyst size data involving a significant volume of samples, using several specimens from various different species and using the same statistical approach. The objective of this paper is to evaluate statistical adjustment to a normal distribution of cnidocyst length from four sea anemone and one corallimorpharian species. The cnidoms of all species are detailed and the hypothesis of no intra-specific variation of cnidae sizes tested as a study case. Normality was accepted in 36.42% (sd = 17.91) of all data sets of all cnidocyst types analyzed from all studied species, while for the rest it was rejected. The evidence suggests that both normal and non-normal data sets are possible, although non-normality is slightly more frequent. Intra-specific variation of cnidocyst sizes is shown in 96.82% of the analyzed data sets. This paper provides a simple and detailed methodology to perform comparisons of cnidae size data.
Publication free access available at http://rdcu.be/npLO
... The first information about the cnidocysts of Astroides calycularis was studied by Carlgren (1940). Nevertheless, recently the whole cnidom of A. calycularis of populations from South of Spain has been studied (Martínez- Baraldés et al. 2014). These authors have showed that cnidom is comprised of two types of cnidocysts: spirocysts (Sp) and nematocysts. ...
... Tentacles comprised spirocysts, Bs3, Ht2 and Mpm3; while pharynx presented Bs2, Ht2 and Mpm1, and mesenterial filaments comprised only Ht1 and Mpm2. Significantly, scapus and columella obtained the same nematocyst composition: Bs1, Ht1, Ht4 and Mpm1 (see Martínez-Baraldés et al. 2014). ...
... In addition, other authors have highlighted the use of cnidocysts as "organic characters" as a new taxonomic informative source and the phylogenetic implications of using such characters (Martínez-Baraldés et al., 2014;Picciani et al., 2011;Pires, 1997;Terron-Sigler and Lopez-Gonzalez, 2005). ...
... Squash preparations of tissues from different parts of the polyp (scapus, tentacles, pharynx and mesenterial filament) were made for cnidocyst examinations. Observations, measurements and categorizations of undischarged capsules, using a Leica DMLB and/or Zeiss Axio Scope.A1 microscope with Nomarski interference contrast optics at 1000x magnification, were made following the methodology described in Godknecht and Tardent (1989) and Martínez-Baraldés et al. (2014). Two main types of cnidae, spirocysts (Sp) and nematocysts, were analysed. ...
... Similarity relationships among specimens collected at the different sampling stations were obtained by the Bray-Curtis index (Bray and Curtis, 1957), using the average length of each cnida in each tissue type. Original data from Terron-Sigler and Lopez-Gonzalez (2005) and Martínez- Baraldés et al. (2014) were also included to show the interspecific relationship. A hierarchical agglomerative clustering method was applied using the PRIMER v6 program (Clarke and Gorley, 2006). ...
Although zooxanthellate corals are well known for their ecophenotypic variations, there is increasing evidence that azooxanthellate species also harbour a high degree of plasticity. Desmophyllum dianthus, a widespread solitary coral, exhibits a high degree of morphological variation in corallum forms that has never been analysed quantitatively. To assess if the clear morphological variation of D. dianthus follows a specific pattern based on environmental (or others) variables, this study combines three different morphometric approaches: 1) classical linear external morphology, 2) use of threedimensional coordinates landmarks, and 3) linear measurements and counts made of cnidocyst features. Comparative morphological characterization of D. dianthus specimens shows a pattern of intraspecific variation over a wide ecological and biogeographic range. However, additional future studies on this and other proposed cosmopolitan species, including a similar sampling effort in localities and specimens, will be useful to explore the existence of common global patterns of morphological variability.
Hypotheses for intraspecific polymorphism are discussed to explain the incongruence between the obtained results and the seemingly high morphological variability observed within D. dianthus.
... In addition, other authors have highlighted the use of cnidocysts as "organic characters" as a new taxonomic informative source and the phylogenetic implications of using such characters (Martínez-Baraldés et al., 2014;Picciani et al., 2011;Pires, 1997;Terron-Sigler and Lopez-Gonzalez, 2005). ...
... Squash preparations of tissues from different parts of the polyp (scapus, tentacles, pharynx and mesenterial filament) were made for cnidocyst examinations. Observations, measurements and categorizations of undischarged capsules, using a Leica DMLB and/or Zeiss Axio Scope.A1 microscope with Nomarski interference contrast optics at 1000x magnification, were made following the methodology described in Godknecht and Tardent (1989) and Martínez-Baraldés et al. (2014). Two main types of cnidae, spirocysts (Sp) and nematocysts, were analysed. ...
... Similarity relationships among specimens collected at the different sampling stations were obtained by the Bray-Curtis index (Bray and Curtis, 1957), using the average length of each cnida in each tissue type. Original data from Terron-Sigler and Lopez-Gonzalez (2005) and Martínez- Baraldés et al. (2014) were also included to show the interspecific relationship. A hierarchical agglomerative clustering method was applied using the PRIMER v6 program (Clarke and Gorley, 2006). ...
Although zooxanthellate corals are well known for their ecophenotypic variations, there is increasingevidence that azooxanthellate species also harbour a high degree of plasticity. Desmophyllum dianthus,a widespread solitary coral, exhibits a high degree of morphological variation in corallum forms thathas never been analysed quantitatively. To assess if the clear morphological variation of D. dianthus fol-lows a specific pattern based on environmental (or others) variables, this study combines three differentmorphometric approaches: (1) classical linear external morphology, (2) use of three-dimensional coor-dinates landmarks, and (3) linear measurements and counts made of cnidocyst features. Comparativemorphological characterization of D. dianthus specimens shows a pattern of intraspecific variation over awide ecological and biogeographic range. However, additional future studies on this and other proposedcosmopolitan species, including a similar sampling effort in localities and specimens, will be useful toexplore the existence of common global patterns of morphological variability. Hypotheses for intraspe-cific polymorphism are discussed to explain the incongruence between the obtained results and theseemingly high morphological variability observed within D. dianthus.
... Dendrphylliidae/19165), a unique arrangement of the septa, is typical of the Dendrophylliidae, although only corals of some genera display it in adult stage (Wells 1933;Vaughan & Wells 1943;Alloiteau 1952;Cairns 2001). The monophyly of the family was further confirmed by its unique composition of cnidocysts (Picciani et al. 2011;Mart ınez-Barald es et al. 2014). ...
Recent molecular phylogenetic studies have shown that most traditional families of zooxanthellate shallow-water scleractinians are polyphyletic, whereas most families mainly composed of deep-sea and azooxanthellate species are monophyletic. In this context, the family Dendrophylliidae (Cnidaria, Scleractinia) has unique features. It shows a remarkable variation of morphological and ecological traits by including species that are either colonial or solitary, zooxanthellate or azooxanthellate, and inhabiting shallow or deep water. Despite this morphological heterogeneity, recent molecular works have confirmed that this family is monophyletic. Nevertheless, what so far is known about the evolutionary relationships within this family, is predominantly based on skeleton macromorphology, while most of its species have remained unstudied from a molecular point of view. Therefore, we analysed 11 dendrophylliid genera, four of which were investigated for the first time, and 30 species at molecular, micromorphological and microstructural levels. We present a robust molecular phylogeny reconstruction based on two mitochondrial markers (COI and the intergenic spacer between COI and 16S) and one nuclear (rDNA), which is used as basis to compare micromorphogical and microstructural character states within the family. The monophyly of the Dendrophylliidae is well supported by molecular data and also by the presence of rapid accretion deposits, which are ca. 5 μm in diameter and arranged in irregular clusters, and fibres that thicken the skeleton organized in small patches of a few micrometres in diameter. However, all genera represented by at least two species are not monophyletic, Tubastraea excluded. They were defined by traditional macromorphological characters that appear affected by convergence, homoplasy and intraspecific variation. Micromorphogical and microstructural analyses do not support the distinction of clades, with the exception of the organization of thickening deposits for the Tubastraea clade.
Scleractinian systematics have undergone rapid changes due to increased use of molecular phylogenetics and new perspectives on skeletal morphology from micromorphology and microstructure. Despite this increase in characters there are still unresolved clades in the phylogeny, indicating that more characters are needed. This study investigates a new source of morphological data within the soft tissue of Indo-Pacific scleractinian corals. Features of tissue layers, especially cnidocytes, are described in hematoxylin and eosin stained thin sections. Based on this new histological data source, a combined analysis with mitochondrial DNA and skeletal data is performed using parsimony and Bayesian analysis. Parsimony analysis yields three most-parsimonious trees similar to trees based on Bayesian analysis. Character maps are also produced that show origination of histomorphological traits at deep nodes within the phylogeny. In general, both analyses retain the previously designated families Lobophylliidae and Merulinidae, but some genera are found to be paraphyletic. Nonetheless, the combined analysis produces a highly resolved and well-supported phylogeny, which could lead to more effective use of biological conservation metrics based on evolutionary distinctiveness. These results show for the first time that inclusion of histomorphological characters improves the resolution of phylogenetic analyses of reef corals. J. Morphol., 2016. © 2016 Wiley Periodicals, Inc.
