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1.
Micrabaciids are solitary, exclusively azooxanthellate deep‐sea corals belonging to one of the deepest‐living (up to 5,000 m) scleractinian representatives. All modern micrabaciid taxa (genera: Letepsammia, Rhombopsammia, Stephanophyllia, Leptopenus) have a porous and often very fragile skeleton consisting of two main microstructural components known also from other scleractinians: rapid accretion deposits and thickening deposits. However, at the microstructural level, the skeletal organization of the micrabaciids is distinctly different from that of other scleractinians. Rapid accretion deposits consist of alternations of superimposed “microcrystalline” (micrometer‐sized aggregates of nodular nanodomains) and fibrous zones. In contrast to all shallow‐water and sympatric deep‐water corals so far described, the thickening deposits of micrabaciids are composed of irregular meshwork of short (1–2 μm) and extremely thin (ca. 100–300 nm) fibers organized into small, chip‐like bundles (ca. 1–2 μm thick). Longer axes of fiber bundles are usually subparallel to the skeletal surfaces and oriented variably in this plane. The unique microstructural organization of the micrabaciid skeleton is consistent with their monophyletic status based on macromorphological and molecular data, and points to a diversity of organic matrix‐mediated biomineralization strategies in Scleractinia. J. Morphol.,2011. © 2010 Wiley‐Liss, Inc.  相似文献   

2.
The skeletal ontogeny of the Micrabaciidae, one of two modern basal scleractinian lineages, is herein reconstructed based on serial micro‐computed tomography sections and scanning electron micrographs. Similar to other scleractinians, skeletal growth of micrabaciids starts from the simultaneous formation of six primary septa. New septa of consecutive cycles arise between septa of the preceding cycles from unique wedge‐shaped invaginations of the wall. The invagination of wall and formation of septa are accompanied by development of costae alternating in position with septa. During corallite growth, deepening invagination of the wall results in elevation of septa above the level of a horizontal base. The corallite wall is regularly perforated thus invaginated regions consist of pillars inclined downwardly and outwardly from the lower septal margins. Shortly after formation of septa (S2 and higher cycles) their upper margins bend and fuse with the neighboring members of a previous cycle, resulting in a unique septal pattern, formerly misinterpreted as “septal bifurcation.” Septa as in other Scleractinia are hexamerally arranged in cycles. However, starting from the quaternaries, septa within single cycles do not appear simultaneously but are inserted in pairs and successively flank the members of a preceding cycle, invariably starting from those in the outermost parts of the septal system. In each pair, the septum adjacent to older septa arises first (e.g., the quinaries between S2 and S4 before quinaries between S3 and S4). Unique features of micrabaciid skeletal ontogeny are congruent with their basal position in scleractinian phylogeny, which was previously supported by microstructural and molecular data. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.  相似文献   

3.
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.  相似文献   

4.
Recent molecular analyses have challenged the traditional classification of scleractinian corals at all taxonomic levels suggesting that new morphological characters are needed. Here we tackle this problem for the family Mussidae, which is polyphyletic. Most of its members belong to two molecular clades composed of: (1) Atlantic Mussidae and Faviidae (except Montastraea) and (2) Pacific Mussidae (Cynarina, Lobophyllia, Scolymia, Symphyllia) and Pectiniidae. Other Pacific mussids (e.g. Acanthastrea) belong to additional clades. To discover new characters that would better serve as phylogenetic markers, we compare the skeletal morphology of mussid genera in different molecular‐based clades. Three sets of characters are considered: (1) macromorphology (budding; colony form; size and shape of corallites; numbers of septal cycles), (2) micromorphology (shapes and distributions of septal teeth and granules), and (3) microstructure (arrangement of calcification centres and thickening deposits within costosepta). Although most traditional macromorphological characters exhibit homoplasy, several new micromorphological characters are effective at distinguishing clades, including the shapes and distribution of septal teeth and granules, the area between teeth, and the development of thickening deposits. Arrangements of calcification centres and fibres differ among clades, but the fine‐scale structure of thickening deposits does not.  相似文献   

5.
6.
The crystallographic orientation of structural elements in skeletons of representatives of Carboniferous Syringoporicae (Auloporida) has been analysed by scanning electron microscopy (SEM), petrographic microscopy and electron backscatter diffraction (EBSD) on specimens from the Iberian Peninsula. The skeletons of the tabulate corals of the Syringoporicae consist of biogenic calcite crystals, and their microstructure is composed of lamellae, fibres and granules, or of a combination of these. Independent of the microstructure, the c‐axis is oriented towards the lumen, quasi‐perpendicular to the growth direction of the skeleton (perpendicular to the morphological axis lamellae, parallel to fibres). Most phaceloid taxa have a turbostratic distribution, as a biogenic response to prevent the cleavage of crystals. Cerioid and some phaceloid corals, whose microstructure is conditioned by wall elements, do not exhibit turbostratic distribution. Wall elements are determined by the biology of each taxon. Holacanth septal spines are composed of fibres arranged in a cone‐shape structure, sometimes clamped to the external part of the corallite and show a complex crystallography. Monacanth septal spines are spindle shaped and composed of bundles of fibres. Tabulae are composed of lamellae. Their development and crystallographic orientation depends on the position of the epithelium in each case. Shared walls are formed by a combination of the walls of two independent corallites with a median lamina, composed of granules; these have a crystallographic orientation between that of the two corallites. The growth of the microstructure is derived by a coordinated stepping mode of growth, similar to other groups of organisms such as molluscs and scleractinians. The nucleation and formation of packages of co‐oriented microcrystals suggest a growth mode similar to mineral bridges with a competitive growth mode between each crystal. The growth pattern of corallites suggests that the growth direction is divided into two main components: a horizontal growth direction towards the lumen and a vertical direction towards the top.  相似文献   

7.
Phymaturus comprises 44 species mainly distributed along the south‐west of South America on both sides of the Andes. In this study we present a phylogenetic analysis of Phymaturus of the palluma group, one of its two large clades, including almost all described species. This analysis duplicates the number of in‐group taxa compared with previous contributions. We performed a total‐evidence analysis, combining molecular and morphological characters: sequencing fragments of cytochome b (cytb), 12S, and ND4, for all terminals; describing 45 new morphological characters; and incorporating all DNA sequences available from GenBank. Separate analyses of morphology and DNA partitions are presented and discussed in detail. Seven subclades are recognized here. We named three new subclades and redefined another, found to be paraphyletic. In order to recognize lineages within the traditional Phymaturus palluma group we proposed to treat it as a natural group, containing within it the ranks of clade, subclade, and lineages, respectively. The palluma group is composed by the vociferator and the bibronii clades. The vociferator clade, composed of Chilean and Argentinean species, would be the most basal in the group. Within the bibronii clade, the roigorum subclade includes the Phymaturus verdugo lineage, whereas the mallimaccii subclade would consist of 13 terminal taxa, for which three Chilean species have been added. In this study, morphological apomorphies are identified for all clades and the evolution of ‘male head melanism’ is discussed. © 2015 The Linnean Society of London  相似文献   

8.
Spatially intimate symbioses, such as those between scleractinian corals and unicellular algae belonging to the genus Symbiodinium, can potentially adapt to changes in the environment by altering the taxonomic composition of their endosymbiont communities. We quantified the spatial relationship between the cumulative frequency of thermal stress anomalies (TSAs) and the taxonomic composition of Symbiodinium in the corals Montipora capitata, Porites lobata, and Porites compressa across the Hawaiian archipelago. Specifically, we investigated whether thermally tolerant clade D Symbiodinium was in greater abundance in corals from sites with high frequencies of TSAs. We recovered 2305 Symbiodinium ITS2 sequences from 242 coral colonies in lagoonal reef habitats at Pearl and Hermes Atoll, French Frigate Shoals, and Kaneohe Bay, Oahu in 2007. Sequences were grouped into 26 operational taxonomic units (OTUs) with 12 OTUs associated with Montipora and 21 with Porites. Both coral genera associated with Symbiodinium in clade C, and these co‐occurred with clade D in M. capitata and clade G in P. lobata. The latter represents the first report of clade G Symbiodinium in P. lobata. In M. capitata (but not Porites spp.), there was a significant correlation between the presence of Symbiodinium in clade D and a thermal history characterized by high cumulative frequency of TSAs. The endogenous community composition of Symbiodinium and an association with clade D symbionts after long‐term thermal disturbance appear strongly dependent on the taxa of the coral host.  相似文献   

9.
Phymaturus is a clade of lizards that occurs at moderate to high elevations in western Argentina and the adjacent central region of Chile, as well as in various volcanic plateaus of the Patagonian region of Argentina. This genus had previously been divided into two groups: the patagonicus and the palluma groups. In this study, we analyzed relationships within the patagonicus group. The data set was built for 23 species plus nine other terminal taxa of undetermined taxonomic status. In total, 10,631 bp (ND4, Cytb, 12S, COI, five protein coding nuclear genes and seven anonymous nuclear loci) and 254 morphological characters were analyzed in a combined data set for 35 ingroup taxa and nine outgroups. We also ran separate DNA sequence and morphological data sets. We identified four main clades, and revealed congruencies and incongruences with previous studies. The indistinctus clade is recovered as the most basal within the patagonicus group in the strict parsimony analysis, while the somuncurensis clade is the most basal under Bayesian inference. The previously recovered calcogaster clade resulted paraphyletic in both analyses and part of their species are included in a redefined somuncurensis clade. We found low support at basal nodes provoked in part by contradictory evidence shown by rogue taxa. We show the phylogenetic information given by each partition/marker and how they contribute to relationships found in the total evidence analysis. We discuss the phylogenetic position of Phymaturus manuelae, Phymaturus tenebrosus, and Phymaturus patagonicus.  相似文献   

10.
Hydroids in the genus Zanclea are a recently discovered component of the fauna associated with reef‐building corals. The phylogenetic relationships among these species are not well known. The present work is based on field surveys in the Republic of Maldives, and for the first time, morphological and molecular analyses are integrated to distinguish a new hydroid species and provide new information on the ecology of this symbiosis. This new hydroid, Zanclea gallii sp. n., was associated with the scleractinian Acropora muricata; it was living sympatrically with its congener Zanclea sango, which was observed for the first time at this locality on the new scleractinian host Pavona varians. The relationships between these two hydroids and other available scleractinian‐associated Zanclea were investigated using two molecular markers, nuclear 28S rDNA and mitochondrial 16S rRNA. Zanclea gallii sp. n. and Z. sango were recovered as distinct lineages within a monophyletic group of scleractinian‐associated Zanclea based on both molecular and morphological data. All Zanclea species that were observed living in association with scleractinians belong to the ‘polymorpha group’ and share the morphological characteristic ‘polymorphic colony’. The genus Leptoseris is the 16th host coral identified for Zanclea. Compared with the frequency of the Z. gallii sp. n. association with A. muricata and Z. sango with the scleractinian P. varians, the latter is twice as common; however, the former exhibited higher Zanclea polyps concentrations over the colony surface. Overall, the Zanclea survey indicates that these diminutive hydroids are more commonly associated with coral than previously known.  相似文献   

11.
Buchwitz, M., Witzmann, F., Voigt, S. and Golubev, V. 2012. Osteoderm microstructure indicates the presence of a crocodylian‐like trunk bracing system in a group of armoured basal tetrapods. —Acta Zoologica (Stockholm) 93 : 260–280. The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel‐fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel‐fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio‐ or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian‐like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.  相似文献   

12.
Kitahara, M.V., Cairns, S.D., Stolarski, J. & Miller, D.J. (2012). Deltocyathiidae, an early‐diverging family of Robust corals (Anthozoa, Scleractinia). —Zoologica Scripta, 00, 000–000. Over the last decade, molecular phylogenetics has called into question some fundamental aspects of coral systematics. Within the Scleractinia, most families composed exclusively by zooxanthellate species are polyphyletic on the basis of molecular data, and the second most speciose coral family, the Caryophylliidae (most members of which are azooxanthellate), is an unnatural grouping. As part of the process of resolving taxonomic affinities of ‘caryophylliids’, here a new ‘Robust’ scleractinian family (Deltocyathiidae fam. n.) is proposed on the basis of combined molecular (CO1 and 28S rDNA) and morphological data, accommodating the early‐diverging clade of traditional caryophylliids (represented today by the genus Deltocyathus). Whereas this family captures the full morphological diversity of the genus Deltocyathus, one species, Deltocyathus magnificus, is an outlier in terms of molecular data, and groups with the ‘Complex” coral family Turbinoliidae. Ultrastructural data, however, place D. magnificus within Deltocyathiidae fam. nov. Unfortunately, limited ultrastructural data are as yet available for turbinoliids, but D. magnificus may represent the first documented case of morphological convergence at the microstructural level among scleractinian corals. Marcelo V. Kitahara, Centro de Biologia Marinha, Universidade de São Paulo, São Sebastião, S.P. 11600‐000, Brazil. E‐mail: kitahara@usp.br  相似文献   

13.
Marine sessile benthic organisms living on hard substrates have evolved a variety of attachment strategies. Rhizotrochus (Scleractinia, Flabellidae) is a representative azooxanthellate solitary scleractinian coral with a wide geographical distribution and unique attachment structures; it firmly attaches to hard substrates using numerous tube‐like rootlets, which are extended from a corallum wall, whereas most sessile corals are attached by stereome‐reinforced structures at their corallite bases. Detailed morphological and constructional traits of the rootlets themselves, along with their evolutionary significance, have not yet been fully resolved. Growth and developmental processes of spines in Truncatoflabellum and rootlets in Rhizotrochus suggest that these structures are homologous, as they both develop from the growth edges of walls and are formed by transformation of wall structures and their skeletal microstructures possess similar characteristics, such as patterns of rapid accretion and thickening deposits. Taking molecular phylogeny and fossil records of flabellids into consideration, Rhizotrochus evolved from a common free‐living ancestor and invaded hard‐substrate habitats by exploiting rootlets of spines origin, which were adaptive for soft‐substrate environments.  相似文献   

14.
Calcium carbonate biomineralization of scleractinian coral recruits is fundamental to the construction of reefs and their survival under stress from global and local environmental change. Establishing a baseline for how normal, healthy coral recruits initiate skeletal formation is, therefore, warranted. Here, we present a thorough, multiscale, microscopic and spectroscopic investigation of skeletal elements deposited by Pocillopora damicornis recruits, from 12 h to 22 days after settlement in aquarium on a flat substrate. Six growth stages are defined, primarily based on appearance and morphology of successively deposited skeletal structures, with the following average formation time‐scales: A (<24 h), B (24–36 h), C (36–48 h), D (48–72 h), E (72–96 h), and F (>10 days). Raman and energy dispersive X‐ray spectroscopy indicate the presence of calcite among the earliest components of the basal plate, which consist of micrometer‐sized, rod‐shaped crystals with rhomboidal habit. All later CaCO3 skeletal structures are composed exclusively of aragonite. High‐resolution scanning electron microscopy reveals that, externally, all CaCO3 deposits consist of <100 nm granular units. Fusiform, dumbbell‐like, and semispherulitic structures, 25–35 µm in longest dimension, occur only during the earliest stages (Stages A–C), with morphologies similar to structures formed abiotically or induced by organics in in vitro carbonate crystallization experiments. All other skeletal structures of the basal plate are composed of vertically extending lamellar bundles of granules. From Stage D, straight fibrils, 40–45 nm in width and presumably of organic composition, form bridges between these aragonitic bundles emerging from the growing front of fusing skeletal structures. Our results show a clear evolution in the coral polyp biomineralization process as the carbonate structures develop toward those characterizing the adult skeleton. J. Morphol. 275:1349–1365, 2014. © 2014 Wiley Periodicals, Inc.  相似文献   

15.
Hydrozoans of the genus Zanclea have been acknowledged only recently as a fundamental component of the highly diverse fauna associated with reef‐building scleractinian corals. Although widely distributed in coral reefs and demonstrated to be important in protecting corals from predation and diseases, the biodiversity of these hydrozoans remains enigmatic due to the paucity of available morphological characters, incomplete morphological characterisations and the possible existence of cryptic species. Recently, molecular techniques have revealed the existence of multiple hidden genetic lineages not yet supported by diagnostic morphological characters. In this work, we further explore the morpho‐diversity of three genetic lineages, namely Zanclea associated with the coral genera Goniastrea (clade I), Porites (clade II) and Pavona (clade VI). Aside from providing a complete classical characterisation of the polyp and medusa stage of each clade, we searched for new potential taxonomic indicators either on symbiotic hydroids or on host corals. On the hydroids, statistical analyses on almost 7,000 nematocyst capsules revealed a significant difference in terms of nematocyst size among the three Zanclea clades investigated. On each host coral genus, we identified peculiar skeletal modifications related to the presence of Zanclea symbionts. Lastly, we discussed the potential diagnostic value of these footprints in the characterisation of Zanclea–scleractinian associations.  相似文献   

16.
The phylogeny of subtribe Gorteriinae (Asteraceae‐Arctotideae) is investigated by means of cladistic analysis of morphological characters. Two sister groups are formed, namely a Gorteria clade also containing Hirpicium and Gazania, and a Berkheya clade, which also contains Cullumia, Cuspidia, Didelta and Heterorhachis. The Gorteria clade has strong jackknife support and is diagnosed by four morphological characters (leaves with longitudinally striate hairs, fringed anther apical appendages, pollen of the “Gazania‐type”, and subulate‐ensiform, ascending style sweeping hairs) that are unique within the Asteraceae. The Berkheya clade is moderately supported and diagnosed by two characters without contradiction (spiny leaves, and mamillate, large style sweeping hairs). Hirpicium and Berkheya are paraphyletic, with the other, morphologically more homogeneous genera (Gorteria, and Gazania, Cullumia, Cuspidia, Didelta and Heterorhachis, respectively) nested within them. There is some evidence for a radiation of species of the summer rainfall area of South Africa and tropical Africa and the corresponding species are nested within a grade confined to the Cape Floristic Region. © The Willi Hennig Society 2006.  相似文献   

17.
18.
A new marine species of naked lobose amoebae Pseudoparamoeba garorimi n. sp. (Amoebozoa, Dactylopodida) isolated from intertidal marine sediments of Garorim Bay, Korea was studied with light and transmission electron microscopy. This species has a typical set of morphological characters for a genus including the shape of the locomotive form, type of subpseudopodia and the tendency to form the single long waving pseudopodium in locomotion. Furthermore, it has the same cell surface structures as were described for the type species, Pseudoparamoeba pagei: blister‐like glycostyles with hexagonal base and dome‐shaped apex; besides, cell surface bears hair‐like outgrowths. The new species described here lacks clear morphological distinctions from the two other Pseudoparamoeba species, but has considerable differences in the 18S rDNA and COX1 gene sequences. Phylogenetic analysis based on 18S rDNA placed P. garorimi n. sp. at the base of the Pseudoparamoeba clade with high PP/BS support. The level of COX1 sequence divergence was 22% between P. garorimi n. sp. and P. pagei and 25% between P. garorimi n. sp. and P. microlepis. Pseudoparamoeba species are hardly distinguishable by morphology alone, but display clear differences in 18S rDNA and COX1 gene sequences.  相似文献   

19.
20.
Recent molecular phylogenies conflict with traditional scleractinian classification at ranks ranging from suborder to genus, challenging morphologists to discover new characters that better agree with molecular data. Such characters are essential for including fossils in analyses and tracing evolutionary patterns through geologic time. We examine the skeletal morphology of 36 species belonging to the traditional families Faviidae, Merulinidae, Pectiniidae, and Trachyphylliidae (3 Atlantic, 14 Indo‐Pacific, 2 cosmopolitan genera) at the macromorphological, micromorphological, and microstructural levels. Molecular analyses indicate that the families are not monophyletic groups, but consist of six family‐level clades, four of which are examined [clade XV = Diploastrea heliopora; clade XVI = Montastraea cavernosa; clade XVII (“Pacific faviids”) = Pacific faviids (part) + merulinids (part) + pectiniids (part) + M. annularis complex; clade XXI (“Atlantic faviids”) = Atlantic faviids (part) + Atlantic mussids]. Comparisons among molecular clades indicate that micromorphological and microstructural characters (singly and in combination) are clade diagnostic, but with two exceptions, macromorphologic characters are not. The septal teeth of “Atlantic faviids” are paddle‐shaped (strong secondary calcification axes) or blocky, whereas the septal teeth of “Pacific faviids” are spine‐shaped or multidirectional. Corallite walls in “Atlantic faviids” are usually septothecal, with occasional trabeculothecal elements; whereas corallite walls in “Pacific faviids” are usually trabeculothecal or parathecal or they contain abortive septa. Exceptions include subclades of “Pacific faviids” consisting of a) Caulastraea and Oulophyllia (strong secondary axes) and b) Cyphastrea (septothecal walls). Diploastrea has a diagnostic synapticulothecal wall and thick triangular teeth; Montastraea cavernosa is also distinct, possessing both “Pacific faviid” (abortive septa) and “Atlantic faviid” (paddle‐shaped teeth) attributes. The development of secondary axes is similar in traditional Atlantic faviids and mussids, supporting molecular results placing them in the same clade. Subclades of “Pacific faviids” reveal differences in wall structure and the arrangement and distinctiveness of centers of rapid accretion. J. Morphol. 272:66–88, 2011. © 2010 Wiley‐Liss, Inc.  相似文献   

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