http://doc.rero.ch View metadata,citationandsimilarpapersatcore.ac.uk hc hudb ie orfrt thiswork. which shouldbecitedtorefer 2007" Published in"Facies53(4):615-623, previous views on growth forms, diversity patterns, and nutrient algalreefs levels.Overall, Guizhouin challenge reported hereinseemingly grew under conditions of low ofthe growthdecisive in Canadian Arctic Archipelago. Whereas nutrients appear and to that of time-equivalent examples reported from the ern environmentofa depositionalmod- margins, tothat similar cement arenoteworthy.Algal reefs developed atplatform sive marine cements (up to 50%) including botryoidal sea the actual dominated “meadows” algal phylloid that views previous to andalgal thalli)numerous large brachiopods.This contrasts with coralsalgae (someofwhich of of thesize aretwice sity of organisms other than phylloids: e.g., the intergrowth di surfacesoften and gravity defying. Algalreefs in Guizhou matrix dense, partly is laminated with marine and cement peloidal micrite.The peloidal e-mail: [email protected] Switzerland Fribourg, 1700 6, du Musée Chemin Department of Geosciences, University of Fribourg, ( Samankassou E. China 110004, Shenyang, University, Northeastern Geology, of Department · Baoliang S. Yongli · Z. · G. Changqing G. Enpu thalli form whose adensecarpet frameworkare pores reefalgalThe facies dominated is by boundstone. Algal cyathiform (cup-shaped) and leaf-like (undulate plates). play awide spectrum of growth forms, but arecommonly mlateralextension. Algae,th 50 mthicknessand 30– report onreefs ranging from2to 8 spread andwellexposed insouth Guizhou, China. Herewe Abstract Baoliang Sun · Yongli Zhang · · Changqing Guan Samankassou Gong· Elias Enpu China Guizhou, in south buildups Paleoecology ofPennsylvanianphylloidalgal V er from examples reported to date by the high biodiver- Halimeda Pennsylvanian phylloid algal reefs are wide- mounds in Java, Australia and o X & oor, excludingperva-biota. otherAlso, the ) Halimeda or clottedwithirregular e maincomponents, dis- reefs, algal reefs V Bahamas, W lled 1 inAcieao(euhm tal. dian Archipelago (Beauchamp et Permian sequences in southern China (GGMRB and Pennsylvanian with studies numerous dealt Although Introduction Guizhou · Pennsylvanian Keywords partly puzzling biogenic structures. depositional environments and add to the spectrum of these the interpretation and discus polished andof slabs thinsectionsfurnish databasefor the these emergingobservation, models. Field andpetrography have experienced new reports and models (e.g., Forsythe case thus represents a signi Samankassou Wahlman Wahlman largest provinceLate for Paleozoic carbonaterocks (see surprisinglyrarely reported from China, whichis among the 2003 1989 phylloid algae and these associated organisms. between relationships possible and algae, phylloid with structures, features, internal accessoryfossils associated theof allow study growth and preservation ditions 1969 Fan and Rigby ( and Fan Northern America (Pray and Wray Wray and (Pray America Northern in elsewhere—e.g., widespread so buildups algal sylvanian ied byFan andRigby ( algal stud- reefs those previously including south in China, The habit,growthmode, andecology ofphylloid algae The present study focuses onwell-exposedphylloid ), the Carnic Alps in Austria and Italy (Flügel (Flügel Italy and Austria in Alps Carnic ), the ; Toomey ; Samankassou andWest 2002 Phylloid algae · Buildups · Paleoecology · ·Paleoecology ·Buildups algae Phylloid 2003 ). The short report of phylloid algal reefs by reefs algal phylloid of report short The ). 1976 1994 ), and North Africa (Toomey , ) is the unique description to date. 1983 1994 ; among others), the Arctic Cana- sion inthe present paper. W ). Theexcellentexposure con- cant test for the extension of extension testfor the cant 2002 1963 , 2003 1989 ; Heckel and Cocke and Heckel ; provided byRERODOCDigitalLibrary ). The Chinese aise al. et ; Davies 1991 1987 brought toyouby )—were ), Penn- 1979 ; CORE http://doc.rero.ch the Guizhou province (Fig. km west to the town Houchang,Ziyun County in about 4 hou Province. Province. hou a 1 Fig. numerous carbonate buildups (GGMRB fossiliferous Weining and Maping Formations that include stratigraphy. TheCarboniferoustheofconsist sequences the region has been afocus of research on Carboniferous the abundanttoChina.fossils occurring rocks, Due inthese rocks characterizethe south Guizhou region, southern Extended outcrops of CarboniferousPermian andcarbonate stratigraphy and Setting and Permian Stratig limestones. appearing sequence of bedded and massive Pennsylvanian respectively Lower Permian, Upper Permian,and Lower Triassic rock series, 26 25 ° ° Reefs crop out extensively close to the village Bianping, the to village close cropoutextensively Reefs Kabutang Y u n n a n Panxian Zhongxinzhai Location of the studied area west of Houchang, South Guiz- Xingyi P 1 b C HINA 105 C Study area Xingren 2 ° , P G uizho Qinglong 1 Longlin , P 2 Anlong , and ,T and G uangxi Bianping Zhenfeng C Wengdao 1 2 1 Ceheng Anshun ). Reefs occur inathick, cyclic represent the Upper Pennsylvanian, Upper the represent Zhijin raphically, the algal reefs 106 0 106 ° Wangmo ° Ziyun 500m P 2 Houchang 1987 Huishui Lumazhai Luodian Leye 60 km ). GUIYANG T 1 107 107 26 25 27 ° ° ° ° ° 2 ( like (undulate plates) interpreted by Samankassou and West growth forms.However, cyathiform (cup-shaped)leaf- and Algae in the studied reefs display a wide spectrum of Growth mode of algae description of their internal obliterated bythe pervasive recrystallization, precluding blages. The The blages. Permian rocks inthe studied area, based on fusulinid and coral assem- 2 Fig. n 0cmlong inaverage (Fig. and 10 Fn tal. (Feng et Formation compared tothat shallower and smaller during thedeposition of the Maping i-eoin(i tal. Mid-Devonian (Qin et basin. Therock basin recordto canbetracedback ofthis shallow-water carbonate respectively, dominate (see also Baars and Torres trace accurately in the culture, the size and forms are, however, partly di form composite bodies. to Due vegetation cover and agri- (mound-shape) or,locally, superposed on eachanother to mlateral extension. Theyisolated, arelens-like and 30–50 m thickness Reefs vary in theirsize, ranging from 2to 8 Shape and size of reefs Reef description Baars Baars Formation (Fig. Formation elian, uppermost Pennsylvanian) that overlies the Wining MapingGzh- the Zone,belong Formation (theTricitites to 2003

U.PENNSYLVANIAN L. PERMIAN mmthick The often curled orundulating leaves are0.8 FORMATION FORMATION ) as constructional or accumulational growth forms, 1992 Stratigraphic scheme of Upper Pennsylvanianand Lower WEINING MAPING dark bar dark orse al. ; Torres et 1998 2 ). indicates the stratigraphic position of algal reefs algal of position stratigraphic the indicates ).Formation TheMaping ispartthe of Fusulina-Fusulinella Pseudoschwagerina Assemblage Zone uldeni Sub-zone Montiparus Triticites W Zone Zone eld. 1992 record that OAIIEACORALS FORAMINIFERA 1996 structure. The rarely better- The rarely structure. ; Torres T. schwageriniformis of theWeining Formation T. paramontiparus ). However, the basin is thebasin). However, mosquensis Sub-Zone Sub-Zone 3 W ). Thalli areoverall 1995 lled the Ziyun-Yadu Nephelophyllum simplex ). Antheria Assemblage Pseudotimania- Sub-Zone Zone Y cult to cult 1991 ; http://doc.rero.ch gastropods (Fig. Further components include bryozoan fragments and rare onophyllum thalli occur. Tuberitina fusulinids, and smaller foraminifers (calcitornellids, reefs. crinoids,Brachiopods, fragments of algal thalli, The substrate of reefs consists of packstone and grain- Reef substrate o slightly the collapse brecciationby indicated thalli thatareonly ( preserved specimens showing cortical pores (Fig. radiaxial of large, curled phylloid algal thalli ( 3 Fig. far from thelifehabitat because of theirsize (Fig. Some ofthethalli arebroken, but obviously nottransported gest that some might belongto phylloid algal thalli and multiple generations of cements (cements as stone, passing over into clotted wackestone toward the toward wackestone clotted into over passing stone, PM ), and voids Photographs of polished slabs of the reef facies. facies. reef the of slabs polished of Photographs W V brous cement ( cement brous set. sp., (Konishi andWray W lled with mosaic cement ( Bradyina 5 a, b). Coating and micritization of algal of micritization and Coating b). a, RFC sp.) and peloids are common. ), inhomogeneous peloidal micrite arrows 1961 Neoanchicodium ikade al. ; Kirkland et ), botryoidal cement ( MC ). ). b Boundstone of Boundstone a Boundstone and 3 b, c) and b, c) 5 c) sug- 1993 Eug- BC ). ), 3 marine cements (Fig. mm in diameter). long and 0.1 with marine cementand peloidal micrite (Fig. tion of marine space(Fig. cements in interstitial Tubiphytes/Shamovella peloidal matrixincludes foraminifers, bryozoan fragments, W peloids,someof which aresurrounded by thin marine irregular surfaces and often gravity defying. Individual with or clotted laminated partly dense, is matrix peloidal thalli form adensecarpetwhose framework pores are The algal reefis facies dominated by boundstone. Algal Reef composition tation ( Figs. latter are partly encrusted prior to precipitation of marine cements (see in occur. foraminifers ( brous cements, are in average 0.1 mm in mm diameter.The brous cements, areinaverage 0.1 a In distinct layers, algal thalli are draped by conspicuous Secondary encrusters including ). Note that the algal thalli were encrusted prior to marine cemen- 5 d, d, arrow 7 b) ). c Cementstone including fragments of algal thalli. The Tuberitina 3 c) or encrusted prior to the precipita- , and unde and endothyrids) commonly endothyrids) and W Tubiphytes e ue 0203mm ned tubes (0.2–0.3 and smaller 5 3 a, b). The b). W lled http://doc.rero.ch framework cavities of phylloid algal boundstone, are larger than the than larger are boundstone, algal of phylloid cavities framework Samankassou and West ( (Fig. thalli (Figs. manyorganisms exceeds the twice thatof phylloid algal assemblages arenot conassemblages Section through alarge solitary coral. Such corals, not con 4 Fig. seen reefs insimilar elsewhere(Toomey co-occurring auloporid corals locally form patches con- lina dium ated with theated algal with phylloid facies. The high-diversity entalis sisting of boundstone (Fig. Nephelophyllum Fusulinids ( sp., and sp., 4 Photographs of polished slabs of coral-dominated facies. facies. coral-dominated of slabs polished of Photographs ), and brachiopods ( , and Diversiphyllum Orthotichia marmerea 4 a and Triticites Fusulinella sp., 5 ). layers Some withintheor pockets Pseudotimania sp.), solitary and colonial corals 2002 W sp., sp., Choristes ned to as cavities commonly sp., and auloporid corals) 4 b). Furthermore, the size of ; Forsythe Pseudosta ) are commonly associ- sp., sp., sp., V Neospirifer ori- 2003 ella Streptophylli- 1976 sp., sp., ): e.g., the e.g., ): , W Fusu- 1980 ned to ned a ; 4 co-occurring phylloid algal thalli ( thalli algal phylloid co-occurring loporid coral colonies ( colonies coral loporid Framework pores are are pores Framework ( ( peloids large algal thalli ( thalli algal large envelopes. ( brachiopods 5 Fig. ments of phylloid algae ( algae phylloid of ments frag- include Bioclasts packstone. of bioclastic composed facies strate for all all for di rated from bioclasts. ( voids phylloid algal boundstone. boundstone. algal phylloid facies. The evidence of framework is demonstrated by cement- by demonstrated is framework of evidence The facies. noid ( noid loids and early marine cements are ubiquitous. pe- facies, algal-dominated phylloid for As facies. boundstone in the Au V erent-sized, early cemented peloids ( peloids cemented early erent-sized, ) coral boundstone. Dense corallite enclose framework voids as in as voids framework enclose corallite Dense boundstone. coral ) Cr W arrow Photomicrographs of reef substrate and core. core. and substrate reef of Photomicrographs gures Pe ) stem) alongphylloid with algal thalli ( c–d ), peloidalcrusts ( ) and the irregular surface ofpeloidal-clotted areas sepa- Br Phylloidalgal boundstone faciestypically composed o ). Peloids are common. Some grains show micritic PH ) enclosing framework voids variously variously voids framework ) enclosing h W Enlargement of peloidal area from lled by cements ( arrows PH f Bryozoans ( Bryozoans ), fusulinids ( fusulinids ), arrow )and phylloid algal thalli ( PH ), and cements ( cements and ), ). ). b Ce arrow Boundstonecomposed ofau- arrow ) arrow ) are major components major are ) ). g Scale bar Well-preserved cri- ), crinoids ( PH Ce ) in boundstone a–b ). e upper part upper Auloporid i . mm 0.5 is c Reef-sub- W , showing lled with Cr ) an W lled ). d f ᭤ http://doc.rero.ch 5 http://doc.rero.ch cements (Fig. fusulinids) (Fig. reefs are particularly rich in foraminifers (predominantly and nids and includes fragments of thealgae Botryoidal, acicular Beresella (Fig. 7 ) represent 20–30, locally over 50% of the 6 a). The top of the reef isrich in fusuli- 6 b, c). W brous, and radiaxial Epimastopora W brous 6 ᭣ vious studies (Pray and Wray Wray and (Pray studies vious equivalent phylloid algal mounds, as reported in many pre- characterizes time- assemblage A low-diversity fossil Biodiversity, ecological community the pinpointing of some valuable information. blagestextural coupledrocks analysisof withmay allow speculative. However, criteriadeduced from fossil assem- ecological parameters drawn from any analysis may appear Phylloid algaelack modern representatives. Thus, most of discussion and Paleoecology: interpretation furtherherein. developed be not will and authors the by preparation in currently tion theiroccurrencetance of arethefocus of another publica- cements. The widespread marine cements and the impor- rock volume. botryoidalOverall, cements predate radiaxial West E growth. a 6 Fig. support this assumption (Flügel (Flügel assumption this support throughout Pennsylvanian and Permian deposits strongly occurring mounds algal Low-diversity colonized. rapidly organisms away from the areas that they successfully and might haveproduced poisonous substances that kept other sella unique and novel case to date. ofterms biodiversity, examplethe Ziyun represents a in Thus, hypothesis.” the“poisoning contradicts studied numerous large brachiopods in the algal-dominated reefs (somecorals of which twice the sizeof algal and thalli) ous sessile organisms, e.g., theintergrowth ofalgae with tion,1998). high However,the overall diversity gregari- of and other environments (G. Gerdes, personal communica- Wilson Wilson umented from studies of recent reefs (Hay Cocke ostee al. Forsythe et kassou and West ostracodes) (Toomey (e.g., brachiopods,attached toalgalleaves fusulinids,and ows were con water organisms. Thus, biota within algal-dominated mead- dominated theliving space, excluding mostshallow- other that phylloid algae colonized the actualsea bottom and and Babcock pimastopora One can thatlikemodernspeculate, algae, phylloid algae ( arrow 2003 b 1969 Fusulinid-rich intervals occur at certain levels during mound during levels certain at occur intervals Fusulinid-rich 1975 Other fossilsin such intervals include fragments of the alga ). ). Indeed, chemical defense used by algaeis doc- (Ep) and and (Ep) Scale bar Scale ; Crowley Crowley ; , 1983 1977 2002 W ned to those toned in and/or those living cavities 2003 ; Pol ; Toomey ; Samankassou; is 0.5 mm for all for mm 0.5 is c rare fragments of the problematic alga alga problematic the of fragments rare ). 1976 1969 1985 ostee al. ; Forsythe et ; Toomey and Winland 1963 1976 ). Toomey ( Toomey ). ; Gray ; Gray 2003 W ; Flügel ; Flügel 1979 gures ; Samankassou and ; Toomey 1967 1997 1976 1979 2002 ; Heckel ; Heckel and ; Paul ; Paul ) assumed ; Toomey ; Saman- 1997 1973 1991 Bere- ) ; ; http://doc.rero.ch priorto marine cementation ( b marine cements ( A ments separated bythin micritic layers( mound core. Cements include multiple generations of botryoidal ce- 7 Fig. , and 0.1 mm for for mm 0.1 and , FC ), and radiaxial radiaxial and ), Early marine cementation is conspicuous throughout the throughout conspicuous is cementation marine Early c , c arrow W brous ( brous ). Locally, phylloid algal thalli are encrusted are thalli algal phylloid ). Locally, b b , , arrow RFC )cements. Peloids show rims o ). a Scale bar , arrows ), acicular is 0.5 mm for for mm 0.5 is W brous ( brous a and a f , 7 fossilsdescribed above presence (especially the corals) of do in low-latitudes during Pennsylvanian (Scotese (Scotese during Pennsylvanian low-latitudes in Furthermore,the paleogeographic position ofsouth China water depthstens ofseveral phylloid algae with phylloid Although non-conclusive,comparemadewere attempts to Nutrients data). unpublished (author’s chiopod and shells marinecement marine indicate value thermore, carbonoxygen and isotopic compositionof bra- with the overall depositional environment (see below). Fur- Province, and (Drew Australia Abel Davies Davies seDhrye al. (see Doherty et to close the lower limit of the wave activity (the shallowest) zonethe photic of lower limit algal reefs interpretedis to have takenplace betweenthe the photiczo ronment within envi- depositional ashallow-water to point may this nids: fusuli- and corals colonial with algae,are associated green Phylloidalgae in south China,assumedly photosynthetic depth Temperature andwater normal-marine conditions (Wilson pods, foraminifers, bryozoans, and corals points to open, The high-diversity biotic asso Salinity tant waterdepth. allow forof theassumption lig reefs in Guizhou. Subsequently, nutrients wereobviously not not point to eutrophic conditions during the growth of algal during Pennsylvanian (Heckel m despite sea-level eustatic changesthe on order of100 were not subaeriallyreefsphylloid exposed studied algal deduced.depthbetherealistically cannot Thefactthat Giles Giles identi paleogeographic reconstructions theprecise do not allow upwelling and, tohighle thus, compare tomodern aise al. Davies et ons(.. 010mfor mounds (e.g., 20–100 the widedepth ranges ofmodern green algaeand algal i Rbrse al. (Robertssia et of deep, nutrient-rich waters; e.g., inthe Java Sea, Indone- occurrences of tions concerning nutrients canbe made. Indeed, some ern Javaern Sea;Robertsand Phipps Settings of phylloid algal might be linked to zones of W 1999 cation ofsuch zones.However, high-diversity the 1988 ocisie al. ; Joachimskiet ). 2004 Halimeda ). Ifphylloid algalreefs, to some extent, 1987 2002 Halimeda Halimeda , ; Samankassou mounds were relatedto upwelling 1988 Halimeda of meters during reef growth. mounds, then some assump- ) and GreatBarrier the Reef ne. The growth of phylloid ht penetration into animpor- (the deepest) and the zone (thedeepest) and 1986 ciation including brachio- 2006 (e atne al. (see Martinet vels of nutrients. Current 1988 , )points to important 1975 1994 mounds in the east- 1988 2003 ), aprecise water ), inagreement ; Soreghan and ; Marshall ; Marshall and ). Considering ). 1997 ) may 1997 ; http://doc.rero.ch meda dimentary marinecementation seawater thatrequiresintense other examples (e.g., Samankassou (Davies in volume and importance (including botryoidal cement the CanadianArctic Archipelago, whichmarine in cements inferredthat agreesmounds forphylloid wellwith algal in (Heckel (Heckel vasive LatePaleozoic glacio-eustatic sea-level algal mounds occur in cyclic sequences that re neous deposits nearly worldwide. Along the same line, Pennsylvanian deposits in south China, like in contempora- reefssedimentaryalgal arepartsthe ofPhylloid record of Conclusion holm andKelley energy levels. layer foraminifers indistinct (Wilson elsewhere, shallowing commonly interrupts reef growth dian examples (Davies obtainedthe Cana- from the values compare with and well the Pennsylvanian(author’s seawater unpublished data) studied reefs point to “normal” marinevalues assumed for pic compositions ofmarine Paleozoic phylloid algalmounds. Carbon andoxygenisoto- modern algalbuildups occurring o ideal fossilideal analogs fortheupwelling-driven modern inGuizhou. phylloidalgae Also, reefsstudied algal arenot decisive fortheoccurrence extensive and growth ofphylloid position despiteof glacio-eustatic sea-level sea actual the kept that environment tional subsidenceand/or important for waterdepth thedeposi- et al. al. et modern seaward platform margins (e.g., o widespread occurrence ofbotryoidal in analogy cements, to the asdoes interpretation, ofthis support in argument solid circulation in open, normal-marine conditions represents a exposure was identi was exposure e aise al.( et led Davies loids. This contrasts to previous views that phylloid algal phylloid that views to previous contrasts This loids. highdate bythe biodiversityorganismsof other than phyl- for areastudiedthe (GGMRB A depositional environment at the shelf margin was assumed Depositional environment Algal Algal reefs grew on stabilized shoals, as observed in Furthermore, the depositional environment assumed Algal reefs in Guizhou di reefs. 1976 1977 1994 1972 ; JamesandGinsburg euhm tal. ; Beauchamp et ). However, no horizon exhibiting subaerial , 1975 2001 2004 W ), as evidenced by accumulation of ed, pointing tooverall high ratesof ). As for most oftheircounterparts 1977 ) to draw some analogiesbetween ) todrawsome in stabilizingthe reefstructure ). V s interpreted as result of high- interpreted result s as ) representcomponentmajor a cements originating from the er fromer examples reported to 1987 1979 1989 ). The ubiquitous synse- ubiquitous The ). ). V 2001 Australiaand Late ). Suchresemblance X ; see also Chis- also ; see V oor in a subtidal X Belize; James Belize; uctuations. X X ect the per- uctuations Hali- 8 mounds in Java, o tional environment simila worthy. cements (up to50%)including botryoidal cement arenote- which improved the initial draft of the manuscript. journal reviewers for comments and A.Freiwaldfor editorial advice anonymous two thank We Samankassou. Elias to 200021-107505) (Project Foundation Science National Swiss the and Enpu Gong to 40572014) No. (Grant China of Foundation Science National the Feng Z, Yang Y, Bao Z, Jin Z, Zhang H, Wu X, Qi D (1998)Lithofacies X, Qi D Y, BaoZ, Yang Jin ZhangH, Wu Feng Z, Z, in mounds algal phylloid Carboniferous Upper (1994) JK Rigby J, Fan reser- Outcrop-based (2002) JP Castagna GS, Soreghan PD, Doherty Acknowledgements 1987 Drew EA,AbelKM(1988) Studieson (1989) WW Nassichuk B, Beauchamp BC, Richards GR, Davies deep Holocene (2004) JM Webster M, Lund JC, Braga PJ, Davies submarine aragonite and calcite magnesian Former (1977) GR Davies Limestone Wyandotte in complex Algal-bank (1969) DJ Crowley Chisholm JRM, Kelley (2001) R Marine ecology:worms start thereef- Carbonifer- Upper (1989) WW Nassichuk GR, Davies B, Beauchamp a algae: phylloid Paleozoic Late (1991) AM Torres DL, Baars (1992) DL Baars References growth of need tobe put into awider perspective. the high-diversity co-o biota factors. Furthermore, theextensive marine cementation and spread in the investigated ar understand thespectrum ofalgal-dominatedwide- reefs environments of nutrientlow levels. algal reefs reported herein seemingly grew in depositional biota (Toomey aise al. Davies et “meadows”actual sea dominated the Algal reefs developedat platform margins, adeposi- Furtherneededare detailedstudies torecognize and English abstract). Geological Publishing House, Beijing, 119 pp Beijing, Publishing Geological House, English abstract). paleogeography of Carboniferous in South China Chinese(in with 40:17–24 Southern Guizhou,China. Brigham Young UnivGeol Stud 86:779–795 Bull AAPG Mexico). (New Basin Orogrande western mound, phylloid-algal acomposite characterization: voir Canad Soc Petrol Geol Mem 13:565–574 Mem Geol Petrol Soc Canad areas. adjacent and Canada in reefs Permian and Carboniferous 19:598–609 water algal buildupson the Eastern Australian Shelf.Palaios mary. Geology 5:11–15 cements in UpperPaleozoic reefs of the Canadian Arctic:sum- a 198:1–52 Bull Surv Geol Kansas Kansas. eastern in Pennsylvanian) (Late 409:152 Nature process. building 13:590–599 Mem Geol Petrol Soc Canad Archipelago. Arctic Canadian Permian Lower to ous 6:513–515 Palaios review. pragmatic 66:697–701 JPaleontol genus. algal , 1988 ; Freile et al. al. ; Freileet Halimeda 2004 Kansaphyllum 1976 ). nutrients weredecisiveinthe Whereas V We thankfullyWe acknowledge the Bhms n utai Rbrse al. (RobertsAustralia Bahamas, and et , reefs (Marshall and Davies 1980 Palaeoaplysina 1995 r to that ofmodern , a new Late Pennsylvanian phylloid ea and to sort out thetriggeringea andtosortout ). Also, thepervasive marine ccurring withphylloid algae ; Marshalland Davies Halimeda X -phylloid algal buildups, oor, excluding other . Coral Reefs 6:195–205 Reefs Coral . W nal support of Halimeda 1988 1988 ), ; http://doc.rero.ch Qin J, Wu Y, Yan Y, Zu Z (1996) Hercynian-Indosinian sedimentary- Hercynian-Indosinian Z(1996) Zu Y, Yan Y, Wu J, Qin Marshall JF, Davies PJ (1988) (1988) PJ Davies JF, Marshall Konishi K, Wray JL (1961) (1961) JL Wray K, Konishi ara- Well-preserved, (1993) JAD Dickson Jr, CH Moore BL, Kirkland on Constraints (2006) W Buggisch PH, Bitter von MM, Joachimski Martin J, Braga J, Riding R (1997) Late Miocene Miocene Late (1997) R Riding J, Braga J, Martin Pray LC, Wray JL (1963) Porous algal facies (Pennsylvanian) Honaker (Pennsylvanian) facies algal Porous (1963) JL Wray LC, Pray phyl- (Virgilian) Pennsylvanian anUpper of Sedimentation (1985) JC Pol Paul VJ (1997) Secondary metabolites and calcium carbonate as de- James NP, Ginsburg RN, Marszalek DS, Choquette PW (1976) Facies (1976) PW Choquette DS, Marszalek RN, Ginsburg NP, James James NP, Ginsburg(1979) RN The seaward margin ofBelize barrier Heckel PH, Cocke JM(1969)Phylloid algal-mound complexes in out- control eustatic glacial for evidence of Evaluation (1994) PH Heckel Heckel(1986) PH Sea-levelcurve for Pennsylvanian eustaticmarine Hay ME (1997) Calci (1997) ME Hay Gray RS (1967) Cache Cache (1967) RS Gray GGMRB (Guizhou Geology, Mineral Resource Bureau) (1987) The FreileD, Milliman JD,Hillis(1995)bank L Leewardmargin in spawning Mass (2002) JAD Dickson R, Wood GTW, Forsythe Permo-Carboniferous of synthesis new A (2003) GTW Forsythe Flügel (1979) E Paleoecology and microfacies ofPermian, Triassic rary research and applications. Springer, New York, pp 188–207 pp York, New Springer, applications. and research rary west country, Paso El Mountains, Hueco complex, mound loid algal 70:99–107 tectonic evolution of the Nanpanjiang Basin. Acta Geol Sin 204–234 pp USA, CO, Durango, Society, Geological Corners Permian algal genus. J Paleontol 35:659–666 JPaleontol genus. algal Permian Mexico. Palaios 8:111–120 Palaios Mexico. New Mountains, Sacramento Formation, Holder Pennsylvanian ( algae phylloid gonitic 34:277–280 Geology apatite. conodont of topes iso- oxygen using changes sea-level glacioeustatic Pennsylvanian 46:523–544 Petrol JSedimentary reefs. Honduras) (British Great Barrier Reef. Coral Reefs 6:139–148 carbonates, Paradox Basin (4th (4th Basin Paradox carbonates, Trail, San Juan Canyon, Utah. In: Bass RO, Sharps SL (eds) Shelf Paleoalgology: contempo- (eds) MH Nitecki DF, Toomey In: Texas. pp707–712 I, vol 1996, June 8thInternational Coral Reef Symposium, PanamaCity, Panama, the of Proceedings In: reefs. coral on algae calcareous of fenses 44:441–456 Sedimentology Spain. Basin, Sorbas Mediterranean marginal the in reefs segment microbial and fabric speci fabric and Sedimentologists, Surrey, UK, vol 3, 191 pp and atoll reefs.Special Publication, International Association of Bull 53:1058–1074 AAPG Mid-Continent. of rocks Pennsylvanian Upper cropping 4:65–87 Paleontol consideration of possible tectonic e over marine Pennsylvanian cyclothems North in America and outcropbelt, North America.Geology 14:330–334 transgressive-regressive depositional cycles along midcontinent 713–718 pp I, vol 1996, InternationalReefSymposium, Coral City, Panamy Panama, June 8th the of Proceedings habitats. as value and relationships, trophic western Colorado. AAPG 51:1959–1978 Bull AAPG Colorado. western Geology Publishing House, Beijing, 698 pp abstract). English with Chinese (in geology regional Guizhou 14:27–33 Reefs Coral slope. Bank Bahama Great western meadows and draperies and their sedimentary importance on the algae. Palaios 17:615–621 Palaios algae. ancientreef communities: fromLate evidence Paleozoic phylloid 78:171–188 Publ Spec SEPM ecology. reef algal phylloid 3:569–587 taine Elf-Aqui- Explor-Product Recherch Centres Bull Alps. the from and Jurassic algalcommunities of platformand reef carbonates W W city of early subsea cements in shallow Belize ed seaweeds on coral reefs: complex defenses, complex reefs: coral on seaweeds ed W eld: a Pennsylvanian algal reservoir in south- in reservoir algal aPennsylvanian eld: Eugonophyllum Eugonophyllum Halimeda W eld conference guidebook). Four guidebook). conference eld V bioherms ofthe northern ects. Concepts Sedimentol Concepts ects. , a new Pennsylvanian and Pennsylvanian anew , , Udoteaceae) from the from Udoteaceae) , Halimeda Halimeda alga- 9 Roberts HH, Phipps CV, E CV, Phipps HH, Roberts Samankassou E (2003) Upper Carboniferous-Lower Permian buildups Samankassou E (2001) Internal structure and depositional environ- Roberts HH, Phipps CV (1988) Proposed oceanographic controls on controls oceanographic Proposed (1988) CV Phipps HH, Roberts Roberts HH,Aharon P, Phipps CV (1988) Morphology and sedimen- Torres AM, West RR, Sawin RS (1992) (1992) RS Sawin RR, West AM, Torres Torres AM (1995) Wilson JL (1977) Regional distribution of phylloid algal mounds in late New Springer, history. geologic in facies Carbonate (1975) JL Wilson Virgilian in sedimentation reciprocal and Cyclic (1972) JL Wilson Wahlman GP (2002) Upper Carboniferous-Lower Permian (Bashkirian- Toomey DF, Winland HD (1973) Rock and biotic facies associated algal Paleozoic and Precambrian (1983) JAE Babcock DF, Toomey Toomey DF (1991) Late Permian reefs of southern Tunisia: facies pat- Toomey DF (1983) The paleoecology of a “Middle Limestone algal phylloid Carboniferous aLate of History (1980) DF Toomey dominated plant aPermian of Paleosynecology (1976) DF Toomey Soreghan GilesGS, KA(1999) Amplitudes of Late Pennsylvanian progress PALEOMAP atlas, Paleogeographic CR(1997) Scotese accumulational and Constructional (2003) RR West E, Samankassou Samankassou E, West RR (2002) Construction versus accumulation in soonal setting. In: Proceedings of the 6th international coral reef coral international 6th ofthe Proceedings In: setting. soonal Cantabrian Mountains, Spain. Sediment Geol 145:235–252 Geol Sediment Spain. Mountains, Cantabrian ment of late Carboniferous mounds from the Cármenes Syncline, Cármenes the from mounds Carboniferous late of ment 15:371–374 Geology Indonesia. Sea, Java eastern symposium, Townsville, Australia, August 1988, vol 3, pp 529–534 pp 3, vol 1988, August Australia, Townsville, symposium, modern Indonesian reefs:turn-oa 6:161–172 Reefs Coral sia). of tology of the Carnic Alps, Austria-Italy. In: Ahr WM, Harris PM, Mor- 69:381–387 Paleontol J cortices. dimorphic with alga membranous codiacean County. West Texas Geol Soc Publ New Mexico 77(68):1–7 Mexico New Publ Soc Geol Texas West County. In: Butler JH (ed) Geology of the Sacramento Mountains, Otero Mexico. New southern of strata Wolfcampian and Pennsylvanian pp 471 York, 72(60):82–99 Publ Soc Geol clic Sedimentation in the Permian Basin, 2nd ed. West Texas strata of southern NewMexico. In: Elam JG,Chuber S(eds) Cy- J (eds) Phanerozoic reef patterns. SEPM Spec Publ 72:271–338 Kungurian) mounds and reefs. In: Kiessling W, Flügel E, Golonka 66:678–681 JPaleontol alga. calcareous Paleozoic Late membranous a new Lucia Lucia with Middle Pennsylvanian (Desmoinesian) algal buildup,Nena MinesProf Contrib 11:1–345 carbonates, west Texas–southern New Mexico. Colorado School 25:119–146 Facies States. ternsand comparisonwith the Capitan reef, southwesternUnited cyclothem, midcontinent, USA. Facies 8:113–190 (Stephanian) Carboniferous of Upper an (Leavenworth) Member” 13:249–267 Lethaia Mexico. New northeastern in complex bank 152:1–18 Abhandlung Paläontol Geol Jahrb Neues community. marine 27:255–258 Geology glacioeustasy. Arlington, USA, 37 pp at Texas of University Geology, of Department 90–0497. report 78:219–237 Publ Spec SEPM/AAPG reefs. and platforms ous Harris PM, Morgan WA,Somerville ID(eds) Permo-Carbonifer- WM, Ahr In: USA. Midcontinent, Kansas, eastern in buildups modes offabrics selected in Pennsylvanian algal-dominated 185:379–389 Palaeoecol tol Palaeoclima- Palaeogeogr USA. Kansas, of Pennsylvanian the in mound constructed asmall of example an mounds: algal phylloid 78:201–217 Publ Spec SEPM/AAPG reefs. and gan WA,Somerville ID (eds) Permo-Carboniferous platforms W eld, Nolan County, Texas. Bull AAPG 57:1053–1074 Halimeda Ivanovia tebagaensis Ivanovia bioherms from the eastern Java Sea (Indone- Sea Java eastern the from bioherms V endi L (1987) L(1987) endi V /turn-on mechanism in a mon- Calcipatera cottonwoodensis Calcipatera wasa cyathiform Permian Halimeda bioherms of the ,