Soft body reconstructions of Palaeozoic corals: implications for the system of Anthozoa (Coelenterata)

The soft bodies and individual developmental sequences of the polyps of Palaeozoic corals are reconstructed on the basis of the functional, engineering and constructional morphology of Recent Anthozoa. Four types of body construction can be distinguished by the arrangement of mesenteries, the number of mesentery insertion zones and characteristic septal insertion patterns: (1) serial insertion of single mesenteries in four sectors, which has resulted in the serial insertion of septa, (2) symmetrical insertion of single mesenteries in four sectors, which has resulted in a symmetrical septal pattern or pseudo radial septal patterns, (3) exponential insertion of paired mesenteries in four sectors, which has resulted in septa bifurcation, and (4) exponential insertion of paired mesenteries in six sectors, which has resulted in a radial septal pattern. Suggestions for a constructional taxonomy are given on the basis of the arrangement of mesenteries and on individual development of the reconstructed polyps. Accordingly, new superorders are introduced: 'Seriales' for the first body construction and 'Symmetricales' for the second. The third body construction is that of the Heterocorallia and the fourth that of the Hexacorallia. Following a constructional taxonomy, new and stricter definitions are given for the new and already known taxonomic categories of Rugosa, Heterocorallia and Hexacorallia. A byproduct of this re-definition is that several Palaeozoic corals represent the body construction of the Hexacorallia and should therefore be revised as Hexacorallia; consequently, Hexacorallia must already have been present in the Palaeozoic.     

Abstammung und systematische Stellung der Rugosa und Auloporida

The study of the morphology of the rugose corals and the acception of a solitary sessil sexual polyp, as the stem form of the Anthozoa (B. WERNER) lead to the conclusion that the stem form of the rugose corals was a solitary sessil polyp without a base disc and with four or more pairs of mesenteries from which two were directive pairs. Therefore it seems correct to supply the name Rugosa by Tetracorallia Haeckel 1866. The evolution of the Auloporida from such a stem form follows another trend, which is characterised by the development of colonies. It seems possible, that this trend was attented with the reduction and the lost of the four mesenteries of the hypothetic stem form. Therefore it is not sure, that the Auloporida belong to the Anthozoa.     

Die Weichkörperkonstruktion vonCalceola sandalina (Rugosa, Anthozoa) — Konstruktionszusammenhänge, Ontogenese, Evolution und Funktionsweisen

The slipper-like corallum ofCalceola sandalina comprises the calyx and the lid. It was generated by an anthozoan-like coral. Anthozoan polyps have a barrel-like shape that is produced and maintained by the activity of internal tethering elements and muscular structures in the body wall. All these structures counteract the hydrodynamics of the gastrovascular cavity. In this reconstruction of the soft parts ofCalceola the functional aspects pointed out byRichter (1929) will be reviewed and revised and partially corrected. Calceola was reconstructed on the basis of the model of rugose polyps. Consequently it consisted of single mesenteries which were added in four sectors in a serial manner. The lid was formed on the counter side and is explained as a bulge of the basal parts of the soft body. The lid was connected coherently by the mesenteries to the whole construction. It could be closed by contraction of the mesenteries and opened by regeneration of the hydraulic volume in the gastric cavity. As a mechanical consequence of moving the lid a straight hinge was developed between the calyx and the lid. The soft body ofCalceola developed by quite simple structural modifications of a rugose coral. The most important modification was the bulging of the basal parts of the soft body on the counter side which formed the lid. This bulge enlarged the surface for adhesion to the substrate and might have been an advantage in survival as well. The soft body generated its own substrate giving it the specific slipper shape. This shape preventedCalceola from sinking into soft or muddy substrate by the principle of a snowshoe which also enabledCalceola to conquer new territories. The lid meant protection where enemies and pollution were concerned. The slipper shape and the lid ofCalceola indicated a complex evolutionary pathway, but they evolved by simple modifications of an ancestor with a round diameter. The slipper shape is just a mechanical consequence of the possession of a flappable lid.     

扬子区中、晚奥陶世和兰多维利世四射珊瑚起源、扩散及生物古地理关系

扬子区中奥陶世—志留纪兰多维利世四射珊瑚产出丰富 ,尤以兰多维利世最为繁盛。目前已报道的1 2 3属 (包括中奥陶世 4属 ,晚奥陶世 2 5属 ,兰多维利世 94属 )中 ,有 30属最早出现在扬子区 ,尔后扩散到欧洲、北美和其它邻区。例如 :Calostylis最早出现在川南的中奥陶世 (Llandeilo)地层中 ;Aphyllum和Cantrillia最早出现在浙西的晚奥陶世中Ashgill地层 ;扭心珊瑚类Briantelasma ,Pycnactis和Tunguselasma等 ,最早出现在黔东北晚Ruddanian地层 ;柱珊瑚类Ceriaster、Stauria、Amplexoides、Synamplexoides等 ,泡沫珊湖类Maikottia ,Rhizophyl lum等均最早出现在黔东北的中兰多维利统 (MiddleLlandovery)。根据这些资料 ,我们认为扬子区应该是奥陶纪和志留纪四射珊瑚起源中心之一。文中论述扬子区中奥陶世—兰多维利世四射珊瑚动物群特征及其生物古地理关系。该区中奥陶世珊瑚以穿孔珊瑚类的Calostylis和Yohophyllum为特征。下扬子区浙西晚奥陶世三衢山组 (中Ashgill)四射珊瑚群有某些澳大利亚分子Hillophyllum和Bowanophyllum ;而上扬子区晚Ashgill观音桥层的四射珊瑚动物群与北欧同期珊瑚群有高度的相似性。这表明扬子区当时与欧洲具有较密切的古生物地理关系。扬子区兰多维利世四射珊瑚群与西伯利亚。     

Diversity of Saint Helena Island and zoogeography of zoantharians in the Atlantic Ocean: Jigsaw falling into place

Diversity surveys in isolated sites, such as oceanic islands, provide biogeographic data that can improve our analyses and knowledge of evolutionary processes in the oceans. Zoantharians (Cnidaria: Anthozoa) are common and widespread components of shallow-water reefs, but distributional analyses are scarce for this group. In this study, we collected Zoantharia specimens from around Saint Helena Island (STH) in the mid-Atlantic and identified species using external morphology and molecular data. Moreover, we compiled and analysed the most comprehensive distributional data for zoantharian species in the subtropical and tropical Atlantic Ocean to date. Our results show eight zoantharian species in STH, which includes seven new records for STH waters. Furthermore, all families and genera of the suborder Brachycnemina recorded are widespread in the Atlantic Ocean, including at least four amphi-Atlantic species. The Caribbean is the richness centre in the Atlantic Ocean for zoantharian species, a pattern similar to that observed for many other subtropical/tropical marine taxa. However, Zoantharia may have a lower endemism rate in some areas than other common reef animals, for example zooxanthellate scleractinian corals and reef fishes. Moreover, zoantharian species have a more extensive distribution than close-related taxa such as zooxanthellate scleractinian corals and hydrocorals in the Atlantic Ocean.     

从底栖生物化石再释贵州独山泥盆系龙洞水段的时代

贵州独山的猴儿山组龙洞水段是一套浅海相碳酸盐沉积,盛产底栖牛物化石.通过对该段地层中所含四射珊瑚、床板珊瑚和腕足类等化石属种的详尽分析,再次证实龙洞水段的地质时代仍应属中泥盆世早期(Eifelian).而不是像某些根据痕迹化石或层序地层学的研究那样将它置于下泥盆统.     

中国造礁石珊瑚分类厘定

造礁石珊瑚是珊瑚礁框架建造者, 具有维持珊瑚礁生态系统功能和稳定性的重要作用, 其分类对于造礁石珊瑚和珊瑚礁的研究与保护至关重要。目前, 随着分子系统学的不断发展, 造礁石珊瑚的分类体系发生改变, 伴随着出现大量同物异名。近年来也出现许多无中文学名的中国造礁石珊瑚新记录种, 这些都给物种认定和命名带来困难, 阻碍了中国造礁石珊瑚的研究与保护工作。为此, 本文收集了中国造礁石珊瑚物种记录文献资料, 采用最新的造礁石珊瑚分类体系, 确认同物异名, 形成中国造礁石珊瑚物种名录, 并对中国造礁石珊瑚物种的中文名进行统一的规范和命名。结果表明, 中国共有造礁石珊瑚2个类群16科77属445种。与《中国动物志·腔肠动物门·珊瑚虫纲·石珊瑚目·造礁石珊瑚》相比, 科级分类阶元新增7个科, 变更5个科; 属级分类阶元新增26属, 变更1属, 合并3属; 种级分类阶元新增291种, 变更13种, 合并20种, 新命名305个物种的中文名。并且筛选出187个同物异名。此外, 由于造礁石珊瑚分类体系现仍有部分争议, 文章也进行了讨论说明。     

Heavy metals distribution in the coral reef ecosystems of the Northern Red Sea

Concentrations of seven heavy metals (Cu, Zn, Pb, Cd, Ni, Co and Fe) were measured in the seawater, sediments, common scleractinian reef-building corals and soft corals (Octocorallia : Alcyonacea) at seven reef sites in the Northern Red Sea: I (Hurghada), II (Ras Za’farana), III (El-Ain Al-Sukhna), IV (El-Tur), V (Sha’b Rashdan), VI (Sharm El-Sheikh) and VII (Dahab). Levels of heavy metals were considerably elevated in seawater, sediments and corals collected from reef sites exposed to increased environmental contamination, as a result of diversified natural and anthropogenic inputs. Soft corals of genera Lithophyton, Sarcophyton and Sinularia showed higher concentrations of Zn, Pb, Cd and Ni than hard coral genera Acropora and Stylophora. Soft coral Sarcophyton trocheliophorum collected from El Ain Al-Suhkna (Gulf of Suez) had greater concentration of Cu, followed by hard corals Acropora pharaonis and Acropora hemprichi. The elevated levels of Zn, Cd and Ni were reported in the dry tissue of soft coral Sinularia spp. On the other hand, the soft coral Lithophyton arboreum displayed the highest concentration of Pb at Sha’b Rashdan (Gulf of Suez) and elevated concentration of Zn at Sharm El-Sheikh. Sediments showed significantly higher concentration of Fe than corals. The higher levels of Fe in hard corals than soft corals reflected the incorporation of Fe into the aragonite and the chelation with the organic matrix of the skeleton. The greater abundance of soft corals in metal-contaminated reef sites and the elevated levels of metals in their tissue suggesting that the soft corals could develop a tolerance mechanism to relatively high concentrations of metals. Although the effects of heavy metals on reef corals were not isolated from the possible effects of other stresses, the percentage cover of dead corals were significantly higher as the concentrations of heavy metals increased.     

Unusual growth pattern in the Frasnian alveolitids (Tabulata) from the Holy Cross Mountains (Poland)

Abstract: Growth periodicity is a phenomenon occurring in fossil and modern corals. The most apparent feature is growth banding, and environmental changes are broadly accepted as controls on this phenomenon. If environment controls the growth, then all corallites within a colony should repeat the same growth pattern, as individuals are clones and must have shared the same environment. A study on several species of Alveolitidae (Anthozoa, Tabulata) from the Late Devonian (Early Frasnian) of the Holy Cross Mountains (Poland) shows that the growth pattern varies between neighbouring individuals within the same corallum. This contradicts observations of closely related Favositida as demonstrated on Pachyfavosites sp. from the Givetian of Avesnois, France, where neighbouring individuals repeat the same pattern. Therefore, environmental control on growth rhythm in Alveolitidae can be excluded; the causes of differences between individuals remain unknown.     

Upper Permian (Murghabian) rugose corals from Oman (Ba’id area,Saih Hatat): Community structure and contributions to reefbuilding processes

Summary Coral-dominated communties are rare in Upper Permian reefs. The study of Murghabian rugose and tabulate corals from allochtonous carbonates (‘Oman exotics’) of the Hawasina Complex and autochthonous carbonates of the Saih Hatat area/Arabian Platform (Oman) provides evidence for a significant contribution of rugose corals to the formation of Late Permian reefs. The corals are described with respect to taxonomy, microfacies and community structure. 8 genera and 7 species were recognized.Monothecalis minor n.sp.,Praewentzelella regulare n.sp. andWentzelella katoi magna n.ssp. are new. The corals represent three communities: (1)Praewentzelella community (Hawasina Complex), (2) cerioid coral community (Hawasina Complex), and (3)Waagenophyllum community (Hawasina Complex and Saih hatat). The corals from the Hawasina Complex and the Saih Hatat flourished in significantly different environments: Rugosa from the Hawasina Complex are representatives of reefs, whereas their counterparts from the Saih Hatat lived in level-bottom communities. Coral-bearing reefal boundstones are characterized by a diverse assemblage of sphinctozoans, inozoans, chaetetids, bryozoans, crinoids,Tubiphytes, Archaeolithoporella and algae. These communities produced bafflestones or framestones and were part of a sponge reef complex. The level-bottom community of the Saih Hatat is low-diverse only comprising rugose and tabulate corals. These of isolated colonies locally acted as bafflers.     

The Rugosa–Scleractinia gap re-examined through microstructural and biochemical evidence: A tribute to H.C. Wang

More than sixty years ago, H.C. Wang carried out an extensive study of skeletal microstructures of the Paleozoic corals and concluded that a “direct descent” may have existed between the two coral suborders: the Paleozoic Rugosa and the younger Scleractinia that had been established as distinct phyla by Haeckel (1896). Skeletal microstructures and three-dimensional reconstructions of walls and septa have revealed remarkable similarities between some Permian and Triassic corals, but it is only during the recent years that significant relationships were established between the structural properties of coral skeletons and their control by the biological process. Supported by recent genetic studies of calcareous biomineralization among various invertebrate phyla, the Wang's opinion now appears a reasonable working hypothesis.     

贵州省惠水县王佑中泥盆世晚期的四射珊瑚(二)

贵州省惠水县王佑镇至翁赖村一带出露中泥盆统上部浅海底栖相翁赖组(新名),该组以陆源碎屑沉积居多、沉积厚度较大、群体珊瑚很少而单体珊瑚却比较发育为主要特征。其中,翁赖组上部层位与贵州独山剖面独山组鸡窝寨段相当,时代为中泥盆世吉维特期的晚期(late Givetian)。翁赖组上部之底非常特征的中华海绵珊瑚化石(Sinospongophyllum)与中泥盆统上部吉维特阶标准的腕足类化石鸮头贝(Stringocephalus)共栖。该属与华南地区中泥盆世晚期标准的内板珊瑚(Endophyllum)的内部骨骼构造非常相似,仅外部形态不同。在华南地区,内板珊瑚和中华海绵珊瑚的层位非常稳定,大致相当于吉维特阶的下varcus带。文中描述翁赖组上部四射珊瑚8属13种,包括Calceola intermediata inflata Yoh,,Cystiphylloides kwangsiense Yoh,C.secundus (Goldfuss),C.sp.,Paramixogonaria wangyouensis sp.nov.,Grypophyllum? sp.,Mictophyllum shaw...     

Generation of bilateral symmetry in Anthozoa: a model

Polyps of Anthozoa usually display bilateral symmetry with respect to their mouth opening, to their pharynx, and in particular to the arrangement of their mesenteries. Mesenteries, which are endodermal folds running from the apical to the basal end of the body, subdivide the gastric cavity into pouches. They form in a bilateral symmetric sequence. In this article I propose that early in polyp development the endoderm subdivides successively into three different types of compartments. A mesentery forms at the border between compartments. Two of the compartments are homologous to those of Scyphozoa. They form by mutual activation of cell states that locally exclude each other. The third compartment leads to siphonoglyph formation and is an evolutionary innovation of the Anthozoa. The mechanism that controls the number and spatial arrangement of the third type of compartment changes the radial symmetry into a bilateral one and occasionally into a different one. The dynamics of its formation indicate an activator-inhibitor mechanism. Computer models are provided that reproduce decision steps in the generation of the mesenteries.     

Patterns in the distribution of soft corals across the central Great Barrier Reef

Distribution patterns of soft coral genera were examined at 11 reefs situated in a broad transect from inshore to the Coral Sea in the central region of the Great Barrier Reef. Twenty-five genera representing the Orders Alcyonacea and Stolonifera were recorded, and the survey also included one genus of the Order Gorgonacea. Total living soft coral cover is greatest on outershelf reef slopes, and is often less than and inversely related to the cover by stony corals. Soft coral diversity is generally low on reef flats, where soft coral cover is low or nil except in protected, inshore areas. The most diverse assemblages occur on reef slopes in midshelf and outershelf areas, where Efflatounaria and nephtheid genera predominate, and widely distributed alcyoniid genera are common. These richer assemblages are less well represented in the Coral Sea, while innershelf reefs support a less diverse fauna of somewhat different generic composition. Distribution patterns of soft corals across the transect broadly match similar variations in the distributions of stony corals and fishes, inshore reefs being generally depauperate. Such variations across the continental shelf are closely associated with changes in prevailing environmental conditions, but further research will be required to elucidate the effects of environmental parameters on benthic community structure.     

Diversity and evolution of coral fluorescent proteins

GFP-like fluorescent proteins (FPs) are the key color determinants in reef-building corals (class Anthozoa, order Scleractinia) and are of considerable interest as potential genetically encoded fluorescent labels. Here we report 40 additional members of the GFP family from corals. There are three major paralogous lineages of coral FPs. One of them is retained in all sampled coral families and is responsible for the non-fluorescent purple-blue color, while each of the other two evolved a full complement of typical coral fluorescent colors (cyan, green, and red) and underwent sorting between coral groups. Among the newly cloned proteins are a "chromo-red" color type from Echinopora forskaliana (family Faviidae) and pink chromoprotein from Stylophora pistillata (Pocilloporidae), both evolving independently from the rest of coral chromoproteins. There are several cyan FPs that possess a novel kind of excitation spectrum indicating a neutral chromophore ground state, for which the residue E167 is responsible (numeration according to GFP from A. victoria). The chromoprotein from Acropora millepora is an unusual blue instead of purple, which is due to two mutations: S64C and S183T. We applied a novel probabilistic sampling approach to recreate the common ancestor of all coral FPs as well as the more derived common ancestor of three main fluorescent colors of the Faviina suborder. Both proteins were green such as found elsewhere outside class Anthozoa. Interestingly, a substantial fraction of the all-coral ancestral protein had a chromohore apparently locked in a non-fluorescent neutral state, which may reflect the transitional stage that enabled rapid color diversification early in the history of coral FPs. Our results highlight the extent of convergent or parallel evolution of the color diversity in corals, provide the foundation for experimental studies of evolutionary processes that led to color diversification, and enable a comparative analysis of structural determinants of different colors.     

The biology and physiology of alga-invertebrates symbioses. II. The density of symbiotic algal cells in a number of hermatypic hard corals and alcyonarians from various depths

Ten genera of hermatypic hard corals and two genera of soft corals were found to contain similar numbers of symbiotic algal cells per unit area irrespective of genus, depth of growth, growth form, or locality. An average of 1.45 cells/cm² (range 0.90–2.48) was found with some reduction in very shallow and very deep waters. The hard coral Plerogyra sp. contained substantially more algal cells per unit area of corallite, a feature correlated with its structure and habit, but its effective algal cell density during daylight approachs that of the other corals examined.     

Some toxicological characteristics of three venomous soft corals from the Red Sea

Three common Red Sea soft corals (Cnidaria: Anthozoa), Nephthea sp, Dendronephthya sp and Heteroxenia fuscescens sting humans. Nematocyst venoms of each animal are lethal to mice and hemolytic to human erythrocytes. However, these hemolysins are partially inhibited by known anti-hemolytic agents. Venoms and their gel chromatography-separated fractions have different dermonecrosis and vasopermeability potency in mouse skin. The venom of Heteroxenia fuscescens (Hf) was more lethal (LD50: 0.7 mg/kg), with one prominent 97-kDa protein fraction (LD50: 0.55 mg/kg). Hf venom was more hemolytic, more dermonecrotic, and had more vasopermeable factors than that of the two other species. SDS polyacrylamide gel electrophoresis of soft coral whole venoms and fractions showed different protein molecular masses ranging from 200 to less than 6 kDa. High IgG titers were assayed from venom-sensitized mice blood sera. Enzyme-linked immunosorbent assays (ELISA) marked significant immunological cross-reaction between the studied soft coral venoms and their bioactive fractions.     

Ordovician corals of the Siberian and Mongolian basins: Taxonomic diversity,morphogenesis, and occurrence

Taxonomic diversity, structural peculiarities, and occurrence of the Ordovician corals Tabulatoidea, Heliolitoidea, Rugosa, and Cyrtophyllida from the Siberian and Mongolian basins are analyzed. The corals settled in the Siberian basin almost for 1.5 million of years earlier than in the Mongolian basin. First corals in the Siberian basin were found in the lowermost Muktei Horizon (Llanvirn, i.e. Darriwilian Stage) while first corals in the Mongolian basin are known from the lower part of the Bairim Horizon (Late Caradoc, i.e. lower part of the Katian Stage). First corals in both basins belonged to the subclass Tabulatoidea.     

MODES OF REPRODUCTION IN RUGOSE CORALS

A consideration of reproduction among fossil compound and 'solitary' rugose corals leads to the conclusion that(1) compound corals belonging to the order Rugosa must have been dimorphic and alternated between an asexual generation and a sexual generation; and (2) 'solitary' forms of rugose corals, although dominantly sexually-reproducing, included some individuals in which evidence of a repressed asexual generation is present.
The presence of a sexually-reproducing generation among compound corals cannot be demonstrated by clear morphological evidence, but is deduced from the observation that this is the most likely explanation for the origin of the initial corallite ('protocorallite') of a compound corallum.
It is proposed to restrict the term 'solitary' to non-compound individual corallites in which asexual budding is not observed and which therefore are presumed to have reproduced sexually. Simple corallites in which budding is observable are referred to as 'simple budding' forms. Thus, several described species of non-colonial rugose corals include both solitary individuals and simple budding individuals, including 'Lonsdaleoides' nishikawai Hayasaka & Minato, Timania rainbowensis Rowett, and 'Clisaxophyllum' awa atetsuense Minato & Nakazawa.