Regular mode of increase, and constrained but variable growth, in the Silurian rugose coral Stauria favosa

Stauria favosa exhibits a typical pattern of axial division (tripartite or quadripartite). The four protosepta first appear with a definite polarity in offset corallites, and metasepta are inserted serially, following Kunth's rule. There are no variations in essential modes of asexual reproduction. However, at the corallum periphery, up to four offset corallites appear to have been derived by discrete offsetting within the parent calices. Detailed observations of vertical morphological changes, however, indicate that this apparently unique mode of increase is derived from axial division at the sites of the four protosepta, as usual in this species. Just after the initiation of division, the contraction of polyps occurred in response to somewhat accidental, deteriorating habitat conditions. The combination of asexually divided corallites and rejuvenescence resulted in phaceloid daughter corallites, which subsequently underwent partial mortality at the corallum periphery. Stauria favosa always exhibits not only regular modes of asexual reproduction and septal insertion under the strict control of phylogenetic and developmental constraints, but also variable growth forms of corallites and coralla. This variability, though within limited options, is palaeoecologically significant for successfully coping with unpredictable habitats.     

Life-history strategies of a species of Catenipora (Tabulata; Upper Ordovician; southern Manitoba, Canada)

Detailed study of coralla by transverse serial sections permits the determination and evaluation of life-history strategies (survival and growth characteristics) in response to different physical environments, for Catenipora foerstei Nelson, 1963 from the Selkirk Member, Red River Formation, in Manitoba. We recognize various modes of corallite increase: one type of axial increase, four types of lateral increase, and agglutinated patches of corallites in association with normal, undamaged corallites; and one type of axial increase, one type of lateral increase, and temporary agglutinated patches from the recovery processes of corallites damaged by sediment or bioclast influx. In addition, the formation of new ranks by lateral increase is the most effective method for rapid growth of a corallum or for reconstructing part of a corallum damaged by physical disturbances. Fluctuations in the tabularial area of corallites occur in cycles over vertical intervals ranging from 3.20-7.90 mm. We consider each cycle to represent annual growth. Average annual growth of the three coralla ranges from 4.20-6.27 mm. According to correlations between annual growth cycles and other growth characteristics, a high frequency of offsetting is associated with rapid vertical growth. Specifically, annual growth is relatively high in association with episodes of sediment or bioclast influx, probably generated by storms. In some coralla, however, annual growth is highest in the cycle characterized by few new corallites or by extraordinarily high rates of offsetting by normal, undamaged corallites as well as damaged corallites. This suggests that vertical growth could also be affected by factors other than storm-related disturbance.     

桂西和黔南中三叠世珊瑚

文内描述中三叠世安尼阶(Anisian)珊瑚化石分别采自桂西田阳板纳组和黔南贵阳青岩青岩组。板纳组珊瑚群中建立Pentasmilia guangxiensisgen.etsp.nov.,该新属种有5条第一周期原生隔片甚至有5条第二周期后生隔片,以此不同于有6条第一周期原生隔片和有6条第二周期后生隔片的石珊瑚类(scleractinians)的珊瑚。产于板纳组的另一新属种Radiophyllia astylatusgen.etsp.nov.,其隔片的排列格式与前人归纳的隔片周期插入示意图甚似,但它却发生在中生代最早期(Anisian)珊瑚的骸体内。文中还对青岩组的Pamiroseris silesiaca作了重新描述。     

Life‐history strategies of Manipora amicarum Sinclair, 1955 (Tabulata; Upper Ordovician; southern Manitoba,Canada)

Based on detailed study of transverse serial sections, we recognize various modes of corallite increase in a multichain cateniform coral, Manipora amicarum from the Selkirk Member, Red River Formation, in Manitoba. One type of axial increase and four types of lateral increase involve normal, undamaged corallites, and one type of axial increase and one type of lateral increase occur during recovery processes of corallites damaged by sediment or bioclast influx. All but one of these types of increase are comparable to those in a single‐chain coral, Catenipora foerstei, which we previously documented from the same stratigraphic unit and locality. In M. amicarum, the formation of double ranks and agglutinated patches of corallites by normal corallites, and by recovery processes following corallite damage, is common and presumably genetically controlled. Agglutinated patches originate differently in C. foerstei, occurring sporadically or temporarily in only some coralla. Average annual vertical corallum growth in M. amicarum, as indicated by cyclic fluctuations of tabularial area, is higher than in C. foerstei, which has comparatively smaller corallites. In general, annual growth in M. amicarum is positively correlated with average tabularial area, negatively correlated with frequency of damaged corallites, and is not related to the frequency of corallite increase. In C. foerstei, however, there is a positive association between annual growth rate and the frequency of increase by damaged corallites, related to episodes of sediment or bioclast influx probably generated by storms. In comparison with C. foerstei, M. amicarum has a low frequency of corallite termination and extensive partial mortality is rare. It seems that the relatively rapid overall vertical corallum growth in M. amicarum was effective for protecting the coral from unfavourable situations, possibly by maintaining the growth surface higher above the substrate than in C. foerstei. Although these two species show many similarities in the types of corallite increase, their reactions and strategies in relation to physical disturbance were quite different.     

Modes of regeneration and adaptation to soft‐bottom substrates of the free‐living solitary scleractinian Deltocyathoides orientalis

Scleractinian corals adapt to various substrate conditions with a variety of growth morphologies and modes of life. The azooxanthellate solitary scleractinian Deltocyathoides orientalis exhibits slightly flattened, bowl‐shaped corallites. This study describes in detail the modes of skeletal regeneration after fragmentation in association with exquisitely adaptive strategies of the corals for life on soft substrates. Larger fragments of individuals retaining almost two‐thirds to five‐sixths of the original skeletal area inherit the densely dilated, lower central skeleton, so as to keep a stable life position on soft substrates and regenerate the lost parts promptly. Even highly fragmented individuals preserving less than 10% of the original skeleton still regenerate and repair. Fragmented individuals with almost one‐sixth to one‐third original skeleton actively maintain a posture with the oral disc upward using movements of remaining tentacles. Damaged and missing soft tissues are then efficiently regenerated to form a mouth and gastrovascular cavity near the new centre of the corallum. Every regenerated individual reuses skeleton and soft tissues, and is capable of burrowing before the completion of growth morphology. The mode of regeneration characteristic of D. orientalis is thus effective and adaptive for maintenance of a stable life position on soft substrates for this solitary scleractinian. As fragmentation in deeper‐water, soft‐bottom settings is likely due to predation rather than turbulence, the rapid corallum regeneration and burrowing strategy may both represent adaptive strategies for life on soft substrates and exploitation of new niches, such as an infaunal mode of life, in a predator‐rich environment.     

Regular budding modes in a zooxanthellate dendrophylliid Turbinaria peltata (Scleractinia) revealed by X‐ray CT imaging and three‐dimensional reconstruction

The zooxanthellate dendrophylliid coral, Turbinaria peltata (Scleractinia), exhibit various growth forms that increase the photoreception area through the development of coenosteum skeletons. Because it is difficult to make detailed observations of the internal structures, we visualized inner skeletal structures using nondestructive microfocus X‐ray computed tomography (CT) imaging. After removal of the coenosteum skeletons from the X‐ray CT images, three‐dimensional 3D‐models were reconstructed for individual corallites. Regular budding was observed from the 3D‐model and cross‐sectional images as follows: 1) lateral corallites occurred only near the two primary septa on one side, apart from a directive primary septum with distinct polarity; 2) the budding occurred upward at acute angles; and 3) these regular structures and polarity were repeated throughout growth with every generation. Even in zooxanthellate dendrophylliids, the same budding modes as observed in azooxanthellate equivalents control the colonial growth. These characteristics provide clues for understanding the mechanisms that regulate the shapes of modular marine organisms. J. Morphol. 276:1100–1108, 2015. © 2015 Wiley Periodicals, Inc.     

Regularity and polarity in budding of the colonial scleractinian Dendrophyllia ehrenbergiana: consequences of radio‐bilateral symmetry of the scleractinian body plan

Sentoku, A. & Ezaki, Y. 2012: Regularity and polarity in budding of the colonial scleractinian Dendrophyllia ehrenbergiana: consequences of radio‐bilateral symmetry of the scleractinian body plan. Lethaia, Vol. 45, pp. 586–593. Regularities and polarity in budding of the azooxanthellate scleractinian Dendrophyllia ehrenbergiana were examined with the aim of understanding the developmental constraints on the formation of colonies. Its mode of budding, in light of the orientations of directive septa of offsets and the inclination angle of budding, is consistent with that of other dendrophyllids; however, the offsets of D. ehrenbergiana only occur near the two primary septa on the convex side of individual corallites, showing a plane of bilateral symmetry with a distinct polarity. These regularities and polarity are seen in the axial and its derived corallites throughout growth. Of note, the polarity at individual corallites is clearly reflected in subsequent colony growth by the branching pattern and corallite number. These characteristics imply the presence of radio‐bilateral symmetrical constraints on the asexual reproduction of the Scleractinia and give us invaluable clues to the understanding of shape‐making mechanisms of marine modular organisms. □Asexual reproduction, azooxanthellate coral, budding, colony, Dendrophyllia ehrenbergiana, polarity.     

Growth characteristics of Protoheliolites norvegicus (Tabulata; Upper Ordovician; Estonia)

Protoheliolites is an early heliolitine coral characterized by closely spaced corallites separated in places by sparse coenenchyme. Growth characteristics in the type species, P. norvegicus, are revealed by detailed analysis based on serial peels and thin sections of coralla from the uppermost Katian of north‐western Estonia. Colonies of this species had a strong ability to recover from damage and partial mortality, resulting in various forms of rejuvenation, regeneration, fusion and reorganization of corallites; in some cases, this involved relatively large areas of undifferentiated soft parts. The shells of commensal cornulitids became enclosed in host coralla during colony growth. Coralla of P. norvegicus exhibit distinctive growth cycles due to responses to seasonal changes. The production of new corallites by coenenchymal increase usually occurred in low‐density bands, in which corallites generally display round to subrounded transverse outlines. In high‐density bands, the corallites became crenulated, their wall thickness increased, septal development was more pronounced, and the amount of coenenchyme increased. In addition to these cyclomorphic changes, there were significant astogenetic changes during growth. Compared with the early stage of colony development, distinctive characteristics in the late astogenetic stage include a decrease in the growth rate of the colony, better coordination among corallites, maximum development of corallite crenulations and septa in high‐density bands, more numerous coenenchymal tubules and a greater proportion of corallum area occupied by coenenchyme. In general, the role of polyps in determining morphological characteristics of individual corallites, such as tabularium area, corallite crenulations and wall thickness, was subordinate to the astogeny of the colony. Growth characteristics including colony‐wide coordination of polyp behaviour and subjugation of individuals to restore the colony following damage suggest a strong astogenetic control and high level of colony integration. Protoheliolites probably arose from a heliolitine genus rather than from a nonheliolitine group as some authors have proposed.     

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.     

Regularity in budding mode and resultant growth morphology of the azooxanthellate colonial scleractinian Tubastraea coccinea

Scleractinia exhibit a variety of growth forms, whether zooxanthellate or azooxanthellate, according to factors that control asexual reproduction and ensuing coral growth. The azooxanthellate branching scleractinian Dendrophyllia arbuscula shows regular modes of budding in terms of the locations of budding sites, the orientations of directive septa, and the inclination angle of budding throughout colonial growth. This study reports that such regularities are also found in the apparently different growth form of the massive dendrophylliid Tubastraea coccinea, which shows the following growth features: (1) the offsets (lateral corallites) always occur near four primary septa, except the two directive primary septa, meaning that the lateral corallites do not appear in the sectors of the two directive septa; (2) the two directive septa in lateral corallites tend to be oriented subperpendicular to the growth direction of the parental corallites; (3) the lateral corallites grow approximately diagonally upwards; and (4) these regularities are seen in the axial and derived lateral corallites among all generations during colony growth. Large differences in growth form are found between the branching D. arbuscula and massive T. coccinea, irrespective of the presence of specific regularities. It is likely that subtle modifications of certain parameters (e.g., budding interval, branch length, corallite size, and inclination angle of lateral corallites) have a strong effect on the overall growth morphology. A precise understanding of such regularities, which occur regardless of generation or taxonomic position, would contribute to understanding the “shape-controlling mechanism” of corals, which are an archetypal modular organism.     

Large dwellers of the Silurian Halysites biostrome: rhizosessile life strategies of cystiphyllid rugose corals from the Llandovery of Gotland

An exceptionally well‐preserved, unusual biostrome composed of the framebuilding cateniform tabulate coral Halysites catenularius (Linnaeus, 1767) bears an assemblage of the relatively large solitary cystiphyllid rugosan Cystiphyllum visbyense Wedekind, 1927. The corallites of solitary cystiphyllids are embedded within the ranks of the halysitid colonies, which developed on a soft, muddy substrate and in relatively turbid water. The cystiphyllid larvae successively settled mostly on the ranks of halysitid colonies and on colonies of the tiny phaceloid rugose coral Nanophyllum ramosum Johannessen, 1995, whereas calice‐in‐calice recruitment was not successful for these cystiphyllid corals. Further growth of C. visbyense was supported by rhizoid structures, which were most frequently developed on the cardinal (convex) side of the corallite. The process of formation of the rhizoid structures is here discussed and explained in detail, showing that they were formed by the extension of the basal ectodermal tissue of the polyp. The cystiphyllids, which settled on the walls of living corallites of halysitid colonies, used sweeper tentacles to kill the smaller polyps of the colony to maintain the space around them and expand. Hence, they ultimately used the halysitid colonies only as a hard substrate to stabilize their position on the soft muddy sediment.     

Constraints on the formation of colonies of the extant azooxanthellate scleractinian coral Dendrophyllia arbuscula

Sentoku, A. & Ezaki, Y. 2011: Constraints on the formation of colonies of the extant azooxanthellate scleractinian coral Dendrophyllia arbuscula. Lethaia, Vol. 45, pp. 62–70. Scleractinia display a variety of growth forms, whether zooxanthellate or azooxanthellate, as the consequence of the combined effects of both intrinsic and extrinsic factors. New modules arise in colonial corals through asexual reproduction, including budding and division. The azooxanthellate, branching dendrophylliid Dendrophyllia arbuscula van der Horst 1922 , is a good species to investigate intrinsic regularities in budding, because: (1) the lateral corallites always occur in the vicinity of four primary septa, excluding the two directive primary septa; (2) the two directive septa in lateral corallites tend to be oriented almost perpendicular to the growth orientation of parental corallites; (3) the lateral corallites grow more‐or‐less diagonally upwards; and (4) these regularities are retained from the axial to the derived lateral corallites during colony growth. Accordingly, a colony of apparently complex dendroid corals is formed according to certain universal rules that apply to successive generations of corallites. The presence of two opposite sectors in which budding do not occur seems to be common to other azooxanthellate scleractinian families. Regularities, other than the orientation of the directive septa, are also commonly found at least in other azooxanthellate dendrophylliid genera. These regularities suggest the presence of strict developmental constraints on the asexual reproduction of the Scleractinia, both extant and extinct. These regularities by azooxanthellate scleractinians, as one of the representative colonial metazoan groups, provide us with fundamental data with which we can understand how colonies are constructed. □Azooxanthellate coral, budding, colony, Dendrophyllia arbuscula, regularity.     

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.     

Microstructural characteristics of the stony coral genus Acropora useful to coral reef paleoecology and modern conservation

Identification of fossil corals is often limited due to poor preservation of external skeleton morphology, especially in the genus Acropora which is widespread across the Indo‐Pacific. Based on skeleton characteristics from thin section, we here develop a link between the internal skeleton structure and external morphology. Ten characteristics were summarized to distinguish Acropora and five related genera, including the type and differentiation of corallites, the skeleton nature of corallites (septa, columellae, dissepiments, wall), and calcification centers within septa. Acropora is distinctive for its dimorphic corallites: axial and radial. Isopora is similar to Acropora but possess more than a single axial corallites. Montipora and Astreopora (family Acroporidae) have monomorphic corallites and a synapticular ring wall, with clustered calcification center in the former and medial lines in the latter. Pocillopora and Porties are classified by distinctive dissepiments, columellae and septa. These microstructural skeleton characteristics were effective in the genus identification of fossil corals from drilled cores in the South China Sea. Eighteen detailed characteristics (ten of axial corallites, four of radial corallites, and four of coenosteum) were used in the Acropora species classification. The axial corallites size and structure (including corallite diameter, synapticular rings, and septa), the septa of radial corallites, and the arrangement of coenosteum were critical indicators for species identification. This identification guide can help paleoenvironmental and paleoecological analyses and modern coral reef conservation and restoration.     

Intra-colony variation in the growth of Acropora formosa: extension rates and skeletal structure of white (zooxanthellae-free) and brown-tipped branches

The presence or absence of zooxanthellae near the tip of Acropora formosa branches is correlated with apical skeletal structure and extension rates. White (zooxanthellae-free) tips are lightly calcified, possess thin, widely spaced skeletal elements and bear only a few, poorly developed radial corallites. Brown tips are heavily calcified, possess smaller axial corallites and larger, more numerous radial corallites. White tips exhibit a range of normally distributed extension rates. Brown tips do not extend, but radial growth and internal calcification continue. These processes progressively alter the appearance and density of brown tipped branches. In addition, the axial corallite of brown tips becomes progressively smaller and is eventually indistinguishable from adjacent radial corallites. Although brown and white tips can change from one form to the other, with a corresponding change in extension rate, it is hypothesized that in brown tips with degenerated axial corallites, a new axial corallite must develop before extension can resume. Brown tips predominate in the interior of arborescent colonies, where space for continued extension is limited. They may therefore represent a means of coordination of growth within a colony. Field and experimental evidence suggest that brown tips may develop in response to micro-environmental conditions. White, zooxanthellae-free zones are also characteristic of other branched and plate-forming species, which exhibit rapid extension in a localized region of the colony.     

Growth characteristics in co‐occurring Upper Ordovician species of the tabulate Catenipora from southern Manitoba,Canada

Four species of the tabulate coral Catenipora are present in the Selkirk Member of the Red River Formation at Garson, Manitoba. They provide an opportunity to compare the growth characteristics of multiple, co‐occurring species that produced cateniform coralla. Corallite increase, cyclomorphism and other growth features show high variability within and/or among the species. A total of five types of lateral increase and two types of axial increase are recognized. Lateral increase accounts for over 80% of all occurrences of corallite increase in each species, with the four species differing significantly in the relative frequency of the various types of lateral increase. The type of axial increase, megacorallites and agglutinated patches of corallites that developed from normal, undamaged corallites in C. foerstei are species specific. In all species, cyclic fluctuations in the tabularial area of corallites are considered to be annual, and the variable growth rates within colonies and species are attributed to differences in astogenetic stages or environmental conditions. Average annual vertical growth was positively correlated with average tabularial area in C. foerstei, C. cf. robusta and C. rubra. Catenipora cf. agglomeratiformis, however, which had the lowest average tabularial area and greatest sensitivity to sediment influx, had a high average growth rate comparable to that of C. rubra, which had the largest average tabularial area. The formation of ranks or lacunae by certain types of lateral increase seems to have been the most effective strategy for maintaining and/or expanding the colony growth surface in all four species, and was most common in C. cf. agglomeratiformis. A reptant growth pattern, characterized by creeping ranks, permitted effective recovery of damaged parts as well as quick formation of new ranks or lacunae. The growth surface of these species was situated near the sediment–water interface. □Growth characteristics, intraspecific variation, interspecific variation, palaeobiology, tabulate corals.     

Echinophyllia tarae sp. n. (Cnidaria,Anthozoa, Scleractinia), a new reef coral species from the?Gambier Islands,French Polynesia

A new shallow water scleractinian coral species, Echinophyllia tarae sp. n., is described from the Gambier Islands, French Polynesia. It is characterized by an encrusting corallum, a few large and highly variable corallites with protruding walls, and distinctive costosepta. This coral was observed in muddy environments where several colonies showed partial mortality and re-growth. The new species has morphological affinities with both Echinophyllia echinata and with Echinomorpha nishihirai, from which it can be distinguished on the basis of the diameter and the protrusion of the largest corallite, the thickness of the septa, and the development of the size of the crown of paliform lobes.     

河南济源晚石炭世太原组的一种复体四射珊瑚

Ａｃｒｏｃｙａｔｈｕｓ是一类块状体或丛状体的四射珊瑚，在北美分布于下石炭统，我国见于上石炭统，但丛状体的在我国系首次发现。在连续切片上，可见两个生长阶段，其繁殖方式有侧芽繁殖和边缘泡沫板内繁殖，系统分类上另置独立的科级分类。根据形态分析，该珊瑚栖息在动水，能量略高及食料丰富的浅水环境。当前标本为一新种：Ａ．ｊｉｙｕａｎｅｎｓｉｓｓｐ．ｎｏｖ．。     

Corallite size and spacing as an aspect of niche-partitioning in tabulate corals of Silurian reefs, Racine Formation, North America

Tabulate corals are common in reefs of the Silurian (Wenlockian) Racine Formation in Wisconsin and Illinois, North America. Variation in size and spacing of corallites in this fauna represents an aspect of niche-partitioning that is probably related to feeding. Corallite morphospace, represented by a plot of corallite diameter versus number of corallites per square cm, is characteristically partitioned among favositines, alveolitines, halysitines, syringoporids, and heliolitines, usually with minimal overlap between these major taxonomic groups. Within all groups except alveolitines, morphospace occupied by each major taxon is partitioned further between forms with small corallites and forms with larger corallites. This is probably related to differences in feeding, with ­larger corallite forms specializing in tentacular capture of larger prey, and smaller ­corallite forms specializing in smaller prey involving capture by cilia-directed sheets of mucus as well as by tentacles. Feeding-based differences among tabulates augmented niche-partitioning effected by colony form and relation to substrate. Cerioid, cateniform, coenenchymal, and fasciculate colony types in the Racine fauna were primarily adapted to a soft substrate. Ragged edges of colonies indicate growth during episodic sedimentation, and colonies were partially buried during life. Most tabulates are ­scattered through wackestone and packstone and were not major contributors to reef growth.     

Vent and mound rugose coral associations from the Middle Devonian of Hamar Laghdad (Anti-Atlas, Morocco)

Peculiar associations of small, solitary, deep-water rugose corals are described from the Middle Devonian buildups situated in the easternmost part of Hamar Laghdad area of southern Morocco. The most of them are monospecific and consist of simplified taxon “Amplexus” florescens but one is polyspecific and composed of specimens belonging to four different species representing three families. These rugosan associations form isolated nest-like aggregations where numerous densely packed specimens are arranged mostly in life position. The polyspecific and two monospecific associations are interpreted as growing in close proximity to venting fields. They reveal a unique “calice-in-calice” recolonization pattern expressed by successive settlement of juvenile specimens in the calice of dead individuals. This pattern was presumably a consequence of selective survival of coral larvae settling in extreme vent habitats. Although, the “calice-in-calice” pattern is common in both, mono- and polyspecific associations, there are differences expressed in the character of larval attachments and various types of the calice fillings. The trophic interaction between corals and ostracods is discussed. Additionally, associations of “Amplexus” florescens, not displaying “calice-in-calice” pattern of growth have been found within the mound where polyspecific association occurs. These are interpreted as growing away from venting fields. Comparisons of Amplexus-type coral faunas with the other North African and European corals allow the classification of these ampleximorph rugosan taxa as characteristic biotic components of the Middle Devonian mound environments influenced by venting activity. Two new genera and species, Weyeraia prima and Vesiculolasma erfoudi, are introduced.