Life strategies of solitary undissepimented rugose corals from the upper member of the Picos de Europa Formation (Moscovian, Carboniferous, Cantabrian Mountains, NW Spain)

Rugose corals belonging to the orders Metriophyllina, Stereolasmatina and Plerophyllina have been identified in the upper member of the Picos de Europa Formation (Moscovian). Corals occur in crinoidal limestones containing common bryozoans. The coral assemblage shows a high diversity. Mode of preservation and spatial distribution of corals demonstrate close relationships with biogenic components such as crinoids, bryozoans, brachiopods and bivalves. The morphology of corals provides valuable data for identifying their life strategies, four of which have been identified: liberosessile with straight growth, liberosessile with curved growth, fixosessile with straight growth and fixosessile with curved growth. Most corals from the Picos de Europa Formation appear to have been fixosessile with straight growth; they attached themselves to bioclasts and subsequently developed radiciform processes (mainly talons) to remain straight.     

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.     

贵州省惠水县王佑中泥盆世晚期的四射珊瑚(一)

贵州省惠水县王佑至翁赖一带出露的中泥盆世晚期浅海底栖相地层翁赖组(新名)陆源碎屑发育,沉积厚度巨大,单体珊瑚居多。由于受到当地穹窿地质构造的影响,只出露了中泥盆统的最上部,即相当于独山剖面的独山组鸡窝寨段和宋家桥段这二段地层,而相当于鸡泡段及其以下的地层都被掩埋在地下。根据岩性和珊瑚群组合面貌,翁赖组可以分成上、下两个部分:下部砂岩比较多,也有少许灰岩和泥质砂岩,大致可与独山剖面的独山组宋家桥段相当;上部虽然也有砂岩和泥质砂岩,但灰岩和泥质灰岩明显增加,最富有生物地层学意义的是在上部之底发现了中泥盆世晚期的标准珊瑚化石Sinospongophyllum,表明它可与独山剖面的独山组鸡窝寨段进行对比。无独有偶,独山县的泥盆系剖面是以群体的内板珊瑚(Endophyllum)层作为鸡窝寨段的底界的,Endophyllum层之上是鸡窝寨段(灰岩);Endophyllum层之下则是宋家桥段(砂岩)。Sinospongophyllum(单体珊瑚)与Endophyllum(群体珊瑚),虽然它们外部形态不同,但它们的骨骼内部构造基本上是相同的或者是非常相似的,而它们外形的差异主要是受不同海洋生态环境(如海浪的强弱和陆源碎屑物的多少)影响的结果。文中描述了翁赖组下部的四射珊瑚,共有6属7种,含2个新种,分别是:Grypophyllum cf.primum Wedekind,Grypophyllum wangyouense(Li in Kong and Huang),Cyathophyllum wenglaiense sp. nov.,Sinodisphyllum litvinovitshae(Soshkina),Temnophyllum majus Walther,Argutastrea quadrigemina(Goldfuss)和Ceratophyllum simplex sp. nov.。它们都是华南以及亚洲、欧洲、澳大利亚和北美西部等地中泥盆世晚期常见的一些珊瑚属种。     

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 shawoziense He,M.sp.,Sinospongophyllum planotabulatum Yoh,Temnophyllum waltheri Yoh,T.poshiense Wang,T.complicatum Wang和Thamnophyllum crassum (Kong)等。它们与华南其他地区以及亚洲、欧洲、北非、澳大利亚和北美西部的同期分子都有十分密切的关系,同属于老世界生物地理大区(Old World Realm)。     

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.     

新疆北部密西西比亚纪早期无鳞板的单体珊瑚

新疆北部准噶尔盆地东缘巴里坤、西缘托里与西北缘和布克赛尔等地的密西西比亚纪早期地层为一套较深水相的黑色页岩与生物屑泥晶灰岩, 产浮游生活的菊石、牙形类和营底栖固着生活的腕足类、腹足类、双壳类、苔藓虫、无鳞板的单体珊瑚以及孢子等化石。根据鉴定, 该地层时代应属于密西西比亚纪(或称早石炭世)早期, 相当于国际地层表中的杜内阶(Tournaisian)。产出珊瑚分别为: Amplexus gennarenensis sp. nov., Cyathaxonia stereoseptata, Hapsiphyllum crissum, Meniscophyllum irregulare, Rotiphyllum xinjiangense sp. nov., Zaphrentites parallelus (Carruthers), Neozaphrentis sp., fam. gen. et sp. indet.。     

THE NEW ORDOVICIAN GENUS HILLOPHYLLUM AND THE EARLY HISTORY OF RUGOSE CORALS WITH ACANTHINE SEPTA

Two species of the new rugose coral genus Hillophyllum are described from Ordovician limestones of central-western New South Wales. The larger species, H. priscum sp. nov., is the earliest rugosan to be found in New South Wales successions, first appearing in horizons of probable Gisbornian age (about middle Lower Caradocian), and seems to be the earliest record of a solitary to fasciculate species exhibiting monacanthine septa. A smaller species, H. sp., occurs with H. priscum and the first species of Palaeophyllum in higher horizons, of probable Lower Eastonian age (approximately upper Lower Caradocian). The history of the Ordovician rugose corals exhibiting acanthine septa is reviewed. Hillophyllum shows relationships with the massive, cerioid rugose species of Foerstephyllum type, and with the solitary, non-tabulated Lambeophyllum, Coelostylis , and Primitophyllum. Together they appear to be the earliest representatives of the Suborder Cystiphyllina.     

滇东北昭通箐门的泥盆纪珊瑚(三)———晚泥盆世早期的四射珊瑚*

云南昭通市东北郊箐门出露了一条比较完整的泥盆纪浅海相地层剖面,下、中、上三个统均有出露,浅海底栖生物化石丰富,研究基础也相对较好,是研究滇东北地区泥盆纪地层的一条理想剖面。由于地层剖面比较长,沉积厚度比较大,此前笔者已经描述了早泥盆世和中泥盆世的四射珊瑚,本文继续描述该剖面的晚泥盆世的地层及其四射珊瑚,讨论它们的地层划分、珊瑚群对比与古生物地理。昭通箐门剖面的上泥盆统主要由白云质灰岩组成,与滇东其它地方的宰格组的岩性基本一致。过去由于宰格组的化石稀少,其地层时代难于确定,因此,昭通箐门宰格组底部珊瑚化石的发现具有重要意义。文中首次描述了箐门放羊冲剖面宰格组底部的四射珊瑚5属7种,包括Disphyllum catenatum,Hexagonaria davidsoni,H.magna,Sinodisphyllum simplex,S.variabile,Temnophyllum poshiense和Truncicarinulum temeniophylloides等,总体显示了晚泥盆世弗拉早期(early Frasnian)的特点。它除了与华南各省的珊瑚群有密切关系外,也与加拿大西部的分子有一定的联系。     

Late Moscovian (mid-Pennsylvanian) rugose corals from Wadi Araba (Egypt,Eastern Desert): Taxonomy,palaeoecology and palaeobiogeography

An important late Moscovian rugose coral association is described from the Rod El Hamal Formation of the Wadi Araba area (northern Eastern Desert, Egypt). The upper part of the formation yielded a moderately abundant but poorly diversified coral fauna composed of large dissepimented solitary rugose corals. In total, six species belong to the Bothrophyllidae and the Geyerophyllidae, including three new species: Bothrophyllum suezensis, Bothrophyllum cylindricum, and Amygdalophylloides omarai. Three undeterminable Bothrophyllidae taxa are left in open nomenclature. The corals were attached to the soft substrate by talons and rootlets, either rooted and growing upward as mud-stickers or as secondary recliners. They show frequent rejuvenations and constrictions. Encrustation and/or bioerosion are scarce. Carbonate microfacies indicate a habitat in littoral and inner neritic zones adjacent to a low terrigenous hinterland. Colonial taxa are missing and tabulate corals are rare. The corals from the Rod El Hamal Formation are the only example of a late Moscovian coral fauna on the southern margin of the Palaeotethys, that is to say along the northern margin of Gondwana. The association shows similarities with northwestern Spain (Cantabrian Mountains), the Donets and Moscow basins, and thus a general attribution to the Palaeotethys realm.     

Palaeoenvironmental analysis based on rugose corals and microfacies: a case study at La Cornuda section (early Serpukhovian, Guadiato Area, SW Spain)

Combined analysis of microfacies and rugose coral features provides a useful tool for palaeoenvironmental studies in areas where outcrops are not appropriate for field observations. A detailed study of Serpukhovian rugose corals from La Cornuda section (Guadiato Area, SW Spain) by means of thin sections allowed the identification of environments where they lived. All corals were collected in unit 1 of the section where three different but intimately connected environments have been identified. Corals developed mainly in small mounds built jointly by microbial communities, algae and corals. Some corals also lived in calcareous shoals mainly composed of echinoderm plates. Finally, some corals occur in Oncoidal limestone that represents a shallow ramp, but they were mainly transported from shoals and mounds.     

华南中泥盆世艾菲尔期浅海相四射珊瑚的组合特征和古生物地理区系

虽然在华南中泥盆世艾菲尔期(Eifelian)较深水或斜坡相地层中产有许多国际标准的牙形类带化石,但在广阔的浅海相地层中却很难寻觅到其踪影。华南浅海相沉积地区是否缺失艾菲尔阶?在野外众多剖面的实地考察中表明,从下泥盆统埃姆斯阶到中泥盆统吉维特阶都是连续沉积,中间并没有发现地层的缺失或间断现象。浅海相的艾菲尔阶与其上覆的吉维特阶和下伏的埃姆斯阶都是连续沉积,说明可能是由于海水太浅,不太适合那些国际标准分子的生存而已。艾菲尔期中期末发生一次生物灭绝事件(Mid-Eifelian event),favositids,heliolitids和许多古老类型的珊瑚在地球上灭绝。华南艾菲尔期四射珊瑚可以划分成下、上两个完全不同的组合:1)Utara-tuiasinen sis-Sociop hyllum minor组合(牙形类partitus带—costatus带);2)Columnaria spinosa-Dendrostella praerhenana组合(牙形类australis带—kocklianus带)。     

新疆准噶尔盆地周边晚石炭世的珊瑚*

新疆北部准噶尔盆地周边石炭系广泛出露, 其中上石炭统发育海相火山—沉积序列, 包括西准噶尔地区的哈拉阿拉特组、阿腊德依克赛组, 东准噶尔地区的石钱滩组以及盆地南部的祁家沟组、奥尔吐组等。本文研究在上述各地层的灰岩层或夹层、碎屑岩中获得的四射珊瑚9属17种和床板珊瑚2属3种, 对石钱滩组中的3个新种进行了详细的系统古生物学描述。上述珊瑚化石的地质时代主要为巴什基尔期至莫斯科期。在珊瑚动物群的组成上, 哈拉阿拉特组中新发现Caninophyllum属分子, 结合盆地东、西部共同产出的床板珊瑚Cystodendropora 属分子, 说明整个准噶尔盆地周边的动物群面貌大致类似, 而西准噶尔地区的化石丰度、分异度显著低于盆地东部和东南部。综合前人资料, 目前在准噶尔盆地周边发现的珊瑚化石有约一半为地方性分子。本文认为研究区珊瑚化石的分布和组成特征与晚石炭世准噶尔洋走向封闭或半封闭的演化趋势密切相关。     

新疆准噶尔盆地周边晚石炭世的珊瑚*

新疆北部准噶尔盆地周边石炭系广泛出露, 其中上石炭统发育海相火山—沉积序列, 包括西准噶尔地区的哈拉阿拉特组、阿腊德依克赛组, 东准噶尔地区的石钱滩组以及盆地南部的祁家沟组、奥尔吐组等。本文研究在上述各地层的灰岩层或夹层、碎屑岩中获得的四射珊瑚9属17种和床板珊瑚2属3种, 对石钱滩组中的3个新种进行了详细的系统古生物学描述。上述珊瑚化石的地质时代主要为巴什基尔期至莫斯科期。在珊瑚动物群的组成上, 哈拉阿拉特组中新发现Caninophyllum属分子, 结合盆地东、西部共同产出的床板珊瑚Cystodendropora 属分子, 说明整个准噶尔盆地周边的动物群面貌大致类似, 而西准噶尔地区的化石丰度、分异度显著低于盆地东部和东南部。综合前人资料, 目前在准噶尔盆地周边发现的珊瑚化石有约一半为地方性分子。本文认为研究区珊瑚化石的分布和组成特征与晚石炭世准噶尔洋走向封闭或半封闭的演化趋势密切相关。     

Lucky rugose corals on crinoid stems: unusual examples of subepidermal epizoans from the Devonian of Morocco

Berkowski, B & Klug, C. 2011: Lucky rugose corals on crinoid stems: unusual examples of subepidermal epizoans from the Devonian of Morocco. Lethaia, Vol. 45, pp. 24–33. In the fossil record, evidence for true epizoans, i.e. living animals inhabiting other living host‐animals, is rather rare. A host reaction is usually needed to proof the syn vivo‐settling of the epizoan. Herein, we provide a first report of such an epizoan biocoenosis from various strata of the Early Devonian of Hamar Laghdad, the world‐renowned Moroccan mud‐mound locality. In this case, solitary rugose corals settled as larvae on crinoid stems, perhaps at a spot where the epidermis was missing for some reason (injury, disease). Both the crinoid and the coral began to grow around each other. By doing so, the affected crinoid columnals formed a swelling, where ultimately only an opening slightly larger than the coral orifice remained. We discuss both macroecological and small‐scale synecological aspects of this biocoenosis. The coral profited from its elevated home because it reached into more rapid currents providing the polyp with more food than at the densely populated seafloor, which was probably covered by a coral‐meadow around the mounds and hydrothermal vents. □Corals, crinoids, Early Devonian, epizoans, Morocco, Rugosa.     

黔北晚奥陶世和早志留世四射珊瑚新资料

报道黔北早志留志中期（Middle Llandovery)地层中新发现的四射珊瑚7属--Cantrillia,Neocantrillia,Prototryplasma,Pycnostylus,Dalmanophyllum?.Rhegmaphyllum,Schlotheimophyllum,并描述产自黔东北石阡奥陶纪末期（Late Ashgill)观音桥层四射珊瑚2种--Grewingkia cf.bilateralis Neuman,Brachyelasma cf.medioseptatum(Neuman)及晚奥陶世（Early Ashgill期）涧草沟组中首次发现的四射珊瑚Crassilasma sp.。全文共描述四射珊瑚13属18种,其中6新种,它们是Crassilasma fenggangense sp.nov.,C.crebrumseptatum sp.nov.,Dinophyllum insolitum sp.nov.,Neocantrillia silurica sp.nov.,Prototryplasma guizhouense sp.nov.,Schlotheimophyllum regeneranum sp.nov.。     

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.     

Diversity,speciation, endemism and extinction in Devonian reef and level-bottom communities,eastern North America

Lower Devonian late Emsian (Bois Blanc and Clear Creek Limestones; Schoharie Formation) level-bottom communities in New York, Michigan and Illinois were moderately cosmopolitan and diverse and dominated by brachiopods and solitary rugose corals. Subsequently (Early Eifelian?), there was an important episode of cratonal patch reef building in New York (Edgecliff Member, Onondaga Limestone), southwestern Ontario (Formosa Reef Limestone, lower Detroit River Group), and the Hudson Bay Lowland (Kwataboahegan Formation) by highly diverse endemic communities. The Edgecliff reefs were built by corals whereas the Formosa and Kwataboahegan reefs were built primarily by stromatoporoids. The strong correlation between high diversity and high endemism during the reef-building episode suggests that these communities contained numberous, small species populations belonging to several major taxa — an example of rapid speciation by geographic isolation and genetic drift.     

Rugose corals and associated carbonate microfossils from the Brigantian (Mississippian) of Castelsec (Montagne Noire, southern France)

The disused quarry east of Castelsec offers a view of shallow-marine carbonates of the poorly known Uppermost Mississippian of the Montagne Noire. At Castelsec, sections are studied in two characteristic facies types (bioclastic wackestone and microbial dominated boundstone) of the Upper Mississippian. The succession is rich in rugose corals and carbonate microfossils. Six genera with seven species belonging to a rugose coral fauna consisting of at least eight genera with several species are described herein; Dibunophyllum castelsecensis sp. nov. is described as new. Twenty-seven carbonate microfossils of different groups have been identified. The Castelsec succession is Brigantian in age, based on the stratigraphic occurrence of rugose corals, foraminifers, and calcareous algae observed in both sections. The rugose coral fauna shows relationships with the well-known fauna of northwestern Europe and the Ouralian-Asian Province. Typical elements of northwestern Europe are missing at Castelsec and vice versa. This differentiation between north and south is interpreted as responses to different palaeolatitudes and tectonic settings.     

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。     

黔南紫云、晴隆晚石炭世及早二叠世四射珊瑚

通过对黔南地区晚石炭世及早二叠世早期四射珊瑚的研究,建立4个四射珊瑚组合,自下而上：1）Antheria-Pseudotimania组合;2）Nephelophyllum-Eokepingophyllum组合;3）Parawentzellophyllum-Kepingophyllum组合;4）Anfractophyllum组合。组合1、2以Cyathopsidae科分子和Kepingophyllidae科的原始分子为主;组合3以Kepingophyllidae科的典型分子为代表;组合4以Kepingophyllidae的进化型分子为特征。文中描述四射珊瑚9新种及1未定种。