document04

Reef development at the Frasnian/Famennianmass extinction boundary

Paul Copper �

Department of Earth Sciences, Laurentian University, Sudbury, Canada P3E 2C6

Accepted 6 December 2001

Abstract

A newly compiled global reef database indicates that the 5^6 Myr long Frasnian (Late Devonian) metazoan reefepisode had relatively low diversity compared to Middle Devonian highs (with over 200 genera of calcitic rugose andtabulate corals). Following an initial early rise after Late Givetian coral and stromatoporoid extinctions, reefsexpanded for the last time during mid-Frasnian sealevel highstands, but declined markedly in the Late Frasnian(rhenana-linguiformis conodont zones), below the Frasnian/Famennian (F/F) boundary. Globally, metazoan reefs werewiped out by the end Frasnian: some Famennian reefs, while partly retaining the structure of the underlyingcarbonate platform, were built by cyanobacterial consortia such as Renalcis, Rothpletzella, Girvanella and Epiphyton.During the Famennian, foraminiferans with calcite walls became abundant for the first time in the Phanerozoic,adding a new dimension to carbonate platforms. Colonial rugose corals (phaceloid, cerioid and thamnasterioidmodules) were absent in the early post-extinction phases up into the mid-Famennian, and very rare and non-reef-building later, but solitary deep-water Lazarus corals survived locally. Coral^sponge reefs are unknown from the 21Myr long Famennian, also a time of very low platform carbonate production. Rare, small, isolated stromatoporoidsponge, and lithistid sponge patch reefs returned episodically during the Famennian in North America, westernEurope, Australia and China: the aragonitic stromatoporoids became extinct at the end of the Famennian. During aLate Devonian tectonically very active, collisional Caledonian mountain-building phase, oceanic and atmosphericcooling, accompanied by sealevel lowstand systems, exposed most carbonate platforms, accelerating coastal erosionand karsting. This increased the amount of clastics in the shelf-slope setting, in the last 1^3 Myr prior to the F/Fboundary, often burying reefs. Immediately following, there were protracted losses in nearly all major tropical shelf,benthic marine invertebrates, exceeded only by the end Permian extinctions in severity. There is no apparent linkbetween black, organic-rich horizons and reef demise at or close to the F/F boundary. The F/F boundary not alsomarks the largest change from widespread flooded Early and Mid-Paleozoic continental cratons to narrow, distalshelves, but also spikes the largest known global Phanerozoic shift in atmospheric O2 enrichment, and CO2 drawdown.This threshold matched the rise of the first tropical rainforests, and expansion of terrestrial biomes on the tropicalcoastal lowlands formerly occupied by carbonate platforms. 6 2002 Elsevier Science B.V. All rights reserved.

Keywords: Frasnian/Famennian boundary; coral^sponge reefs; extinctions; ocean^atmosphere system; climate; anoxia

1. Introduction

The Mid- to Late Devonian marked the Pha-

0031-0182 / 02 / $ ^ see front matter 6 2002 Elsevier Science B.V. All rights reserved.PII: S 0 0 3 1 - 0 1 8 2 ( 0 1 ) 0 0 4 7 2 - 2

* Fax: +1-705-673-6508.E-mail address: [email protected] (P. Copper).

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nerozoic acme of metazoan reef and carbonateplatform productivity (Copper, 1994; Kiesslinget al., 1999), with a second acme in the Mid-Si-lurian (Wenlock). During this ca. 80 Myr interval,sandwiched between two major ice ages and glob-al mass extinctions, coral and sponge reefsreached their maximum known geographic extent,with barrier reef complexes far exceeding in sizethose of today’s interglacial Holocene. Reefs wereperiodically projected well into latitudes 45‡S andpossibly 60‡N, as seasurface temperatures were10^15‡ higher than today’s interglacial 16‡C glob-al average. Exceptional global warming wascoupled to atmospheric CO2 levels 14^24U great-er than today, and more than double the CO2levels of the warm Mesozoic (Berner, 1994,1998, 1999b). High rates of weathering of terres-trial Ca/Mg silicates in the presence of CO2-richatmospheres ultimately meant a long term trans-fer of atmospheric CO2 to oceanic-stored CaCO3during the Early and Mid-Palaeozoic. Sealevelhighstands £ooded vast continental interiors,and epicontinental seas greatly exceeded in sizetheir modern tropical analogues, the relativelysmall South China, Java and Arafura seas. This£ooding created ample accommodation space forshallow-water, ‘skeleton framework reef facto-ries’, and coral^stromatoporoid sponge carpetsand meadows, suitable for accelerated calciteand aragonite production under warm tropicalclimates (e.g. as seen for the mid-Frasnian, Fig.1). Prime examples of such carbonate factoriesexisted on the areas fringing and covering Lau-rentia during the mid-Devonian: more than halfof the present 10 million km2 land area of Canadawas covered in tropical seas at that time, as NorthAmerica straddled the equator, and similar trop-ical oceans covered Russia and Siberia. As a re-sult, giant barrier, fringing, and patch reefs £our-ished into ‘super’ sizes. Reefs were dominated bytabulate and rugose corals, and cyanobacteriancalcimicrobes possessing calcite skeletons, andby the aragonitic stromatoporoid sponges. Oceanswere in a calcite mode, in contrast to the modernCenozoic and Late Palaeozoic ‘aragonite’ oceans(Sandberg, 1983). From the 5^6 Myr long Fras-nian into the 21 Myr long Famennian, the ocean^atmosphere climate system switched to an arago-

nite saturation system, triggered by rising O2 andcooler temperatures. The Famennian^Tournaisianwas also marked by a major radiation of the sili-ceous planktic radiolarians, identi¢ed by a riseand peak in the Tournaisian ‘hypersiliceous peri-od’ (Nazarov and Ormiston, 1986; Racki, 1999;Racki and Cordey, 2000).From the Silurian through Middle Devonian,

land vegetation had initially consisted of simple,low-spreading pteridophytes, or mosses and liver-worts, until the evolution of trunked and cano-pied trees (the large horsetails and clubmosses)led to the ¢rst appearance of Phanerozoic rain-forests in the Late Devonian. Examples of suchtree trunks are found preserved in mid-Frasnianreefs and inter-reef strata of Banks Island (Embryand Klovan, 1971), the ¢rst time this is seen in thegeologic record. Dramatic changes in spore sizesof the global land £ora at the Frasnian/Famen-nian (F/F) boundary are related to new strategiesfor plant reproduction, e.g. seeds and pre-pollen(Streel and Loboziak, 1995). These ¢rst rainfor-ests a¡ected soil development and the storage ofcarbon by soil microbiota, chemical rock weath-ering, and coastal sediment transport (Wilder,1989; Algeo et al., 1995; Berner, 1997). Fairlystable levels of 15^20% O2 prevailed in the Cam-brian through Mid-Ordovician, probably with O2generated primarily by marine phytoplankton.Levels of O2 then rose during the Late Ordovi-cian, as CO2 declined (Berner, 1998), with the ¢rstarrival of very simple small plant cover of bryo-phytes (Retallack, 1985). Oxygen levels then shotup dramatically to elevated levels, possibly near35% in the Late Carboniferous (Berner and Can-¢eld, 1989; Berner, 1999a), with a shift to land-based carbon sequestration, O2 ceilings being con-strained by upper wild¢re limits. Thus both LateOrdovician and Late Devonian oceanic coolingepisodes were marked by a reduction of CO2and an increase of O2 coinciding with plant evolu-tionary events (Retallack, 1997; FrancOois et al.,1997). In terrestrial lake, river and deltaic envi-ronments, and in the coastal mixed fresh- andsalt-water estuarine systems, there was a disas-trous loss of ¢sh during the multiple F/F crises,e.g. the thelodont, heterostracans, galeaspids,ptiuraspids, osteostracans, onychodonts, and po-

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rolepids all became extinct during the Frasnian,with placoderms and lobe-¢nned ¢sh dying out inthe Famennian (Long, 1995). This is the largestextinction and diversity loss of ¢sh known in thePhanerozoic. The only survivors appear to havebeen groups that migrated into marine environ-ments, that adapted to land as amphibian tetra-pods, or like the estivating lung¢sh, survived

harsh terrestrial river and lake climates by slimecocooning and burrowing. This strongly suggeststhat coastal lowland, terrestrial habitats wereequally as much a¡ected as shallow marine trop-ical successions in the Late Devonian, ruling outmarine anoxia as a prime cause for extinctions. Inturn, changing land conditions (e.g. the rise of the¢rst forests, rise of oxygen via photosynthesis,

Fig. 1. Time scale for global F/F reef development, showing major hiatuses at the F/F boundary coinciding with sealevel low-stands. Reefs expanded during the mid-Frasnian, but had largely ceased growth by the close of the rhenana CZ: reefs werepoorly developed during the ¢nal Frasnian linguiformis CZ. Famennian reefs were dominantly calcimicrobial consortia, with rarestromatoporoid, or lithistid patch reefs, e.g. in the Early Famennian of the Bonaparte-Canning basins, western Canada carbonateplatform. Following repeated regressive cycles and cooling events, mid- to late Famennian carbonate platforms declined, followedby transgressive intervals of isolated mudmounds to sponge patch reefs (e.g. Belgium, Poland small patch reefs in the marginiferaand postera-praesulcata CZ). In other parts of the world, giant siliciclastic deltas were built up, £ooding the shelf areas (NE Can-ada, Appalachians, Caledonides), or areas were subject to erosion and non-deposition (asterisks denote Frasnian coral^stromatop-oroid reefs, see Fig. 4).

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reduced atmospheric CO2, weathering and erosionpro¢les) had a run-o¡ impact on neighboringtropical marine systems.The Siluro-Devonian coral^sponge reef ecosys-

tem had a coral biodiversity of more than 200genera (Scrutton, 1998), 30% greater than thatof the scleractinian corals for the Cenozoic (Ve-ron, 1995; Budd, 2000). Complementing this weremore than 60 genera of aragonitic stromatoporoidsponges adapted to open, illuminated, fore- andback-reef settings (Stearn et al., 1999), comparedto only a handful of primarily cryptic corallinesponges today. Metazoans were accompanied bydiverse coralline algae and green algae. Frasnianreefs were dominated by alveolitid and thamno-porid tabulates, and giant phaceloid to aphroidrugosan corals, but with a diversity much lowerthan seen in the mid-Devonian (showing that£ourishing reef growth was not necessarily corre-lated with diversity). Many were capped by verylarge, meter-sized, platy to domal stromatopor-oids in the fore-reef and reef £at areas (Fager-strom, 1994). Calcimicrobes played a signi¢cantcementing, encrusting and cavity-¢lling role, asfrom the Miocene to today (Riding et al., 1991;Reid and Macintyre, 1992; Esteban, 1996). Fromthe close of the Frasnian, the metazoan, shallowtropical reef ecosystem never fully recovered insize or diversity in the Famennian, nor Late Pa-laeozoic. For tropical marine biota, the Late De-vonian was the second largest mass extinction inthe Phanerozoic, exceeding severity of Ordovicianevents for the metazoan reef ecosystem (wipingout more than 70% of the primary reef dwellersand builders and shallow carbonate platform bio-tas, as well as losing the Mid-Palaeozoic metazo-an, coral^sponge reefs). Famennian microbialreefs covered 6 10^20% of the shelf space seenin the Frasnian.F/F boundary events still remain controversial.

Controversy relates to (a) the timing of the eventswithin conodont zones (CZs), whether sudden(Claeys and Casier, 1994), or staggered, gradualor stepdown (Copper, 1984; Joachimski and Bug-gisch, 1996), (b) whether there was a single event(Goodfellow et al., 1988; Feist, 1991; Wang et al.,1996), or whether there were multiple events (Joa-chimski and Buggisch, 1993), including the timing

of these, and (c) whether only marine environ-ments were a¡ected (House, 1985; Becker,1993a,b), or whether both terrestrial and marinesystems were disturbed (Wilder, 1989; Algeo etal., 1995; Copper, 1998). Debate also swirlsaround causes, primarily whether these were ter-restrially internal in origin (climate change, oce-anic overturning via current systems, anoxia, vol-canism, tectonics, etc.), or astronomically forced(single or multiple impacts, orbital changes andcycles, etc.). When examining causal e¡ects, thequestion turns to whether there was a primary,triggering factor (e.g. did climate change kill reefs,or was it global ocean anoxia?), a secondary fac-tor triggered by another event (did warming orcooling trigger anoxia?, nutrient £ux?, sealevelchange), or whether there were multi-causal orcascading threshold factors (were sealevel draw-down, cooling and anoxia inter-linked, and inwhat order?). Can the same extinction factorhave multiple, opposing explanations? (e.g. wasanoxia due to stagnant, sluggish ‘hot’ oceans, orthe product of rapid overturn by sea surface tem-perature (SST) cooling, bringing up deep CO2-rich waters and nutrients?). For those favoringanoxia as a primary cause, were black shales orblack limestones the result of high oceanic surfaceproductivity under well-oxygenated bottoms, orthe result of bottom anoxia and burial factors?(i.e. were black shales the cause, or the e¡ect, ororganic burial events unrelated to mass extinc-tions?). Were pelagic events coupled to shelfevents, or not? Were atmospheric events linkedto oceanic events? Were some events simplysymptoms of change, or the real agents of massextinction, e.g. were sealevel lowstands just re-£ecting glacial cooling events, or did eustatic sea-level drain shelf areas, removing habitable spaceand biotas (e.g. Johnson, 1974)? Was the F/F ex-tinction due to anomalous nutrient (biogeochem-ical) cycling and eutrophication (e.g. Murphy etal., 2000), or were these end-products (symptoms)of extinction? How did reefs ¢t into this picture?Since the events are dated at or near the F/Fboundary, at ca. 376 Ma (Tucker et al., 1998;Okulitch, 1999), the oceanic crust that mighthave given us distinctive platinum^iridium signa-tures, has largely been lost by subduction. Despite

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exhaustive search, no de¢nitive impact crater, norboundary clay horizon, nor iridium anomalieshave been detected at the F/F boundary, thoughanomalies have been noted well above and belowthe boundary in both shallow and deeper watersections.It is generally agreed that the equatorial coastal

marine ecosystems, particularly shallow-waterreefs, from interior £ooded cratons to shelf mar-gins, were most strongly a¡ected by the F/Fevents. During the Middle Devonian (Eifelian^Givetian), metazoan reefs reached latitudes ofup to 60‡N (if the Mongolia plate is correctlyplotted), and 45^50‡S. The most severe extinc-tions marked the end of the Givetian, when tab-ulate and rugose corals (Scrutton, 1997, 1998),stromatoporoids (Stearn et al., 1999), and reefalto peri-reefal atrypid and pentamerid brachiopodsdeclined, with the loss of many families and evena whole suborder (Copper, 1998; Godefroid andHelsen, 1998). The terminal Frasnian extinctionsin these groups were less than the end-Givetianbiodiversity losses at the genus and subgenus lev-el, a fact which is generally buried in the litera-ture. During the Emsian and succeeding Eifelian^Givetian (Middle Devonian), reefs reached theirgreatest extent of the Phanerozoic, using four fac-tors: (1) overall size of reef tracts (cumulativethickness and areal extent), (2) pervasive occupa-tion of widely £ooded, tropical and subtropicalcontinental interiors and platform margins tohigh latitudes, (3) high coral and coralline spongediversity, with more than 200 genera of tabulateand rugose corals (Scrutton, 1997, 1998), and 60Devonian stromatoporoid genera (Stearn et al.,1999) and (4) peripheral invasion by deep-waterslope, and high-latitude, cool shallow-water mud-mounds. A number of mid-Devonian reef provin-ces were at least three to four times longer, andseveral times wider, than the modern Great Bar-rier Reef (e.g. the Emsian^Givetian carbonate reefplatform extending from Nevada into EllesmereIsland, some 5000 km along the northwesternmargin of Laurentia). In the Frasnian this reefbelt disappeared almost totally from the Cana-da^Greenland arctic, shrinking by nearly 2000km to only a 150 km belt along Banks island atthe western margin.

The Late Devonian encompassed two phases ofreef evolution, a restricted Frasnian episode ofcoral and stromatoporoid reefs, and a Famennianepisode of catastrophic metazoan reef collapse,and replacement by microbial reefs. During theFrasnian, the taxa that survived the severe coralbiodiversity losses of the Late Givetian (norrisiCZ), probably triggered by a phase of cooling,global sealevel drawdown, and exposure of theshallow tropical reef setting to subaerial karst ero-sion, had regrouped. Surviving tabulate coralsand colonial rugosans formed reef tracts whichwere regionally extensive, such as in the westernCanada sedimentary basin and the Urals. In thehigh 40^50‡S latitudes, for example NW Africa(Morocco and Algeria), and even in lower latitudeareas such as the Montagne Noire and Sardinia,reefs completely vanished during the Frasnian.Early Frasnian reefs were usually smaller, withperi-reefal settings of lower diversities: reefs in-cluded more mudmound reef types than seen ear-lier. By mid-Frasnian time, reefs had temporarilyrecovered to a considerable extent in size andareal extent, with cosmopolitan coral taxa,though with reduced diversity compared to theGivetian. Late Frasnian reefs were diminished inabundance and geographic distribution, and theseappear to have become even more restricted to-wards the F/F boundary, with few localitiesknown where reefs extended to the ¢nal extinctionevents at the close of the linguiformis CZ (lastzone below the F/F boundary). Commonly, latestFrasnian reefs were replaced by bryozoan bio-stromes or stromatolitic units in the rhenanathrough linguiformis zones. The Famennian wasmarked by major, worldwide, sealevel drawdownpulses, interrupted by rapid transgressions: meta-zoan reefs were replaced by calcimicrobial, Renal-cis-dominated reefs. Famennian coral reefs areunknown, but a few patch reefs with stromatop-oroid and lithistid sponges occurred regionally(Fig. 1). Though some stromatoporoids survivedas rare Lazarus taxa, especially the domed or lam-inar labechiids and some stick-like amphiporids,these were greatly reduced in diversity.This paper explores some of the possible sce-

narios worldwide in the decline and ultimate lossof the Mid-Palaeozoic reef ecosystem from the

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Frasnian through Famennian. To date no paperhas yet been devoted exclusively to looking atwhat happened to reefs worldwide at the F/Fboundary. Problems remain with the lack of pre-cise dating of reef terminations in a number ofareas. Since conodonts are scarce in reef settings,the precise age of reef cores may not be fullydeterminable, though the reef base and top maybe better dated. Data from the early 1990s sug-gested that the Frasnian was a relatively lengthyepisode lasting ca. 10 Myr, with the Famennianabout 4^5 Myr long. More recently, Okulitch(1999) has compiled radiometric data suggestingthe converse, a 5^6 Myr long Frasnian followedby a much longer Famennian lasting 21 Myr. Thissuggests that the end-Givetian through Frasnianextinction and diversity losses in reefs lasted 6 6Myr, but that the post-extinction phase wasnearly four times longer than previously sus-pected. It also changes the general picture ofwhat happened to reefs, with a considerablyshorter period of Frasnian reef expansion, and amuch more protracted phase of reef absence andrecovery. In addition, the accumulation rates ofpre- and post-F/F carbonate platforms must bere-interpreted (see below). The Devonian termi-nated about 355 Ma (Ciaou-Long et al., 1992).During the succeeding Carboniferous, mud-mounds became the preferred method of reef con-struction (Bridges et al., 1995).

2. Frasnian metazoan reefs: low diversity,cosmopolitanism and collapse

Though reefs were regionally abundant in themid-Frasnian, they never reached such high lati-tudes as those of the Eifelian^Givetian, being con-¢ned to ca. 45‡N and 30‡S of the equator, slightlybetter than those of the Holocene. Because manyFrasnian reefs are petroleum reservoirs (e.g. inwestern Canada and Timan^Pechora), these areoften better studied than Middle Devonian exam-ples. Broadly summarized, there were three globalFrasnian reef trends, probably related to majorsealevel cycles plotted by Johnson and Klapper(1992) and to climate change. Regressive LateGivetian trends were succeeded by transgressive

episodes with reef development controlled partlyby local tectonics, or limited by siliciclastic deltadevelopment (Canadian arctic, Catskill delta ofthe Appalachians), but with overall reef develop-ment remarkably coordinated worldwide. Reefsare summarized here on an area by area basis.A detailed region-by-region description and anal-ysis of global Frasnian reef provinces is not re-peated here (Copper, 2002; abbreviated in Appen-dix of Frasnian reef localities).The Early Frasnian (Montagne Noire CZ

(MNCZ) 1^4: Klapper, 1997, approximately tran-sitans-punctata zones of others) marked a some-what reduced episode of reef-building, and themid-Frasnian (MNCZ 5^11: approximately has-si-lower rhenana CZ of others) saw maximal reefdevelopment. The Late Frasnian (MNCZ 12^13:approximately upper rhenana-linguiformis CZ ofothers) saw the broad decline of reefs, with small-er sizes and generally more restricted faunas, ter-minating in regressions, or regional black shaleevents. A typical Middle Frasnian highstand car-bonate systems tract saw a broad passive marginwith barrier and platform reefs and isolated largereef mounds up to several kilometers in diameter(Fig. 2). The Frasnian interval represents the ‘dy-ing’ episodes and close of the Middle Palaeozoicmetazoan reef ecosystem, with reefs losing morethan 50% of their tabulate and rugose coral ge-neric diversity by the end Givetian. Only 39% ofthe Devonian coral suborders and superfamiliessurvived into the Carboniferous (Scrutton,1997). The stromatoporoid sponges, the other ma-jor reef builders, saw four orders surviving intoFamennian (Labechiida, Clathrodictyida, Stroma-toporida and Amphiporida), and with one possi-ble syringostromatid (Stearn, 1987). However, 10genera extended into the Famennian (with sevenothers more doubtfully ranging into the Famen-nian: Stearn et al., 1999). Total diversity of thestromatoporoids in the Middle Devonian was ca.64 genera, with 38 recorded from the Frasnian(including four new genera), so that extinctionof stromatoporoid genera by the end Givetianwas ca. 47%, and by the end Frasnian 74% ofthe remaining genera were lost. There are nopost-Famennian stromatoporoids, with Mesozoicspiculate forms probably independently derived.

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The Frasnian reefs of western Europe grew oncarbonate platform terrain that covered less thana million km2, but reef outcrops form only afraction of this southern border of the Old Redcontinent, i.e. western Baltica (Krebs, 1974;Burchette, 1981). Beginning in the Frasnian,transgression caused back-stepping of the carbon-ate platform northwards, and reef growth ex-panded, with reefs peaking in the mid-Frasnian.In southwest England, the eastern terminus of thisbelt, reef activity declined in the Early Frasnian,probably strongly in£uenced by nearby siliciclas-tics and volcanics (Scrutton, 1977), but there ap-pear to be no Middle to Late Frasnian reefs be-cause of local regression. In Belgium, for theclassic sections around Frasne and Famenne (Le-compte, 1936), limited Early Frasnian reefgrowth, was followed by maximal expansion inthe mid-Frasnian, with many microbial mud-mound-type reefs from 30 to 150 m thick. Re-stricted small reefs feature the Late Frasnian,with the latest Frasnian Matagne Shales (rhe-

nana-linguiformis CZ) shutting down reef growthin Belgium and the Aachen area of Germany.Boulvain and Herbosch (1996) demonstratedthat Frasnian mudmounds of Belgium were in£u-enced in their growth by bathymetry, as the mid-Frasnian rimmed shelf changed to a ramp by theLate Frasnian, signifying a slowdown in carbon-ate production. Reefs disappeared in Belgium atthe top of the F2j level, roughly at the base of theMatagne organic-rich shales, well below the F/Fboundary (De Jonghe and Mamet, 1988). Gode-froid (1970) and Godefroid and Helsen (1998)noted that atrypids and reefs in Belgium disap-peared together near the top of the NeuvilleFm., below the Matagne shales. Casier and De-vleeschouwer (1995) attributed decline of the Bel-gian reefs to anoxia and variations in sealevel,factors that he also used to explain the 75% lossof benthic ostracodes at the F/F boundary. InGermany, reefs were best developed east of theRhine, in part as shelf or ramp carbonates, andin part as isolated atoll-like, commonly ‘mud-

Fig. 2. Schematic picture of idealized mid-Frasnian reef development in a carbonate platform sealevel highstand setting (e.g.based on western Canadian and arctic examples). Reef ecosystems during sealevel highstands featured prograding sediments thatexpanded carbonate platform margins seawards, and back-stepped reefs inland. Note the coral^stromatoporoid complexes thatdominated the reefs. O¡-reef settings featured organic-rich carbonates and shales. Coastal lowlands had the ¢rst pteridophyterainforests.

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mound’ complexes in a tectonically active, vol-canic belt of the Rhenish Schiefergebirge (Jux,1960; Krebs, 1967, 1971, 1974). Rhenish reefswere best developed in the Early and MiddleFrasnian, commonly over Givetian highs, butmetazoan reef and microbial mudmound growthceased in the Late Frasnian (Weller, 1989; Gisch-ler, 1995). This has been related to regressivecycles of siliciclastics in shelf areas, capping layersmarking restricted facies (shallow water?), or de-position of hypoxic, lower Kellwasser black lime-stones in the deeper water facies of the Rhenishtrough (Krebs, 1974; Fuchs, 1990).In Poland, a 600 km long Frasnian carbonate

shelf, stretched along the eastern margin of theOld Red Continent, is typi¢ed by extensive coraland stromatoporoid patch reefs of Early to Mid-dle Frasnian age. Kadzielnia-type microbial Re-nalcis mounds were prominent in the Early Fras-nian (Racki, 1988). Calcimicrobial moundsappear to have locally become increasingly impor-tant upwards in the Frasnian succession ofPoland (Racki, 1992). Reef growth halted in thelower rhenana CZ, when condensation anddepositional hiatuses, with basin-wide facies re-structuring, and local karsting took over. Synse-dimentary tectonics complicated carbonate plat-form foundering, and sealevel drawdown, withshallowing sealevel signatures even in basinalsections in the highest linguiformis CZ (Szulczew-ski et al., 1996). A series of micro-plates or ter-ranes that today make up parts of southern andcentral Europe, i.e. southern France, Spain (in-cluding the French Pyrenees), the Balearic islands,Sardinia, northern Italy, Austria, the Czech Re-public, Slovakia, northern Italy, Austria, and theformer Yugoslavia were situated between Balticaand Gondwana in south subequatorial latitudes.Frasnian reef development was very limitedhere or locally wiped out, or possibly con¢nedto low diversity sponge and microbial mud-mounds, with other areas showing small coral^stromatoporoid patch reefs. In Moravia somereef growth continued to the F/F boundary, whilein the Montagne Noire and Prague Basin Fras-nian reefs were absent. Reef growth had ceased inwestern North Africa (Morocco, Algeria) by theend Givetian (Wendt, 1988), though possible ex-

ceptions are noted in the Atlas Mountains (Gen-drot, 1973).Frasnian reef development was relatively vigo-

rous on the eastern side of the Russian Platformor eastern Baltica (western slopes Urals from Vai-gach Island in the north through the south Volgaregion). Reefs are known both from outcrop andsubsurface, and many are petroleum-bearing(Menner et al., 1996; Antoshkina, 1997). Insouthern Timan, Frasnian stromatoporoid^coralreefs (Sirakhoi, Vezhavozh, Sedyu, Bolshoii Ke-ran reefal limestones) extended into the lower rhe-nana CZ, but highest Frasnian linguiformis strataare barren (Yudina and Moskalenko, 1997). Cor-al^stromatoporoid reefs ceased at the F/F bound-ary, and best development was in the mid-Fras-nian. In Siberia, two lower to Middle Frasnianbarrier and patch reef complexes are located inthe Kuznetsk Basin (Belskaya, 1960; Ivanova,1983). Stromatoporoid^coral reefs of the Glubo-kaya complex, associated with solenoporids andoncolites, range into the rhenana CZ, but are lack-ing in higher strata (Rzhonsnitskaya et al., 1992).The Kazakhstan Dzungar and Tyan-Shan hadsmall, lower Frasnian coral patch reefs, linkedto volcanic suites (Kim and Erina, 1984; Zado-roshnaya et al., 1990a). Neighbouring Afghanistanhad a coral^stromatoporoid tract about 100 kmlong (Mistiaen, 1985), and in Iran biostromes andsmall coral^stromatoporoid patch reefs occurredin the upper Givetian through mid-Frasnian tothe top of the jamieae CZ (Wendt et al., 1997;Mistiaen et al., 2000). These identify the shelfmargin of a separate plate positioned adjacentto northern Gondwana but in the subtropics.The North American craton was extensively

covered by giant reef complexes during the Mid-dle Devonian and to a reduced extent in the Fras-nian. The largest known Frasnian reefs are in Al-berta (spilling over into NE British Columbia andthe Yukon); many of these formed as rimmedplatform and isolated reef complexes. Canadianreef studies have placed emphasis on di¡erent as-pects, e.g. oil and gas resources (Davies, 1975;Reinson et al., 1993), tectonics, basement andgeography (Moore, 1988), platform-basin se-quence stratigraphy (Whalen et al., 2000), andcoral distribution correlated with conodonts with-

PALAEO 2824 31-5-02


in and around reefs (McLean and Klapper, 1998).A number of reef episodes, tied in part to trans-gressive/regressive (T/R) cycles, have been recon-structed. These were during the Early Frasnian,following the Late Givetian regressive episode(below the norrisi CZ), and in general three reefepisodes are de¢ned for the early, middle and lat-est Frasnian. Reefs particularly £ourished in theMiddle Frasnian, succeeded by major declines inthe Late Frasnian (CZ 12^13: Klapper, 1997).During the Frasnian, reefs vanished completelyfrom the central and eastern arctic (Victoriathrough Ellesmere islands, over 1500 km of shelf),as a result of giant delta construction. In the west-ern Canada sedimentary basin, Frasnian reefswere developed as barriers, banks, platforms, lin-ear trends, or large patch reef complexes manykilometers in diameter, or as arcuate fringing reefsaround land areas. Deep seismic surveys indicatethat Upper Devonian reefs of Alberta partly in-herited their regional distribution from underlyingPrecambrian and Cambrian basement highs, butsome reef trends show no correlation with preced-ing platform topography (Edwards and Brown,1999). Some rare Frasnian deeper water, slopereefs with renalcids or receptaculitid green algaesat in troughs or embayments between carbonateplatforms: these were probably still within thephotic zone 6 100 m deep (MacKenzie, 1967;Mountjoy and Riding, 1981; Pratt and Weissen-berger, 1989). Carbonate reef mega-breccias, reefolisthostromes and reef margin debris £ows alsoreveal evidence of nearby shallower water reefs,where adjacent reefs themselves are not evidentin drill-core data or outcrop (Cook et al., 1972).On the northwestern side of Laurentia, at, or

north of, the equator, the 1500+ km long, and ca.300^500 km wide central Frasnian reef belt ex-tended in the subsurface and outcrop from Alber-ta through British Columbia into the Yukon andNorthwest Territories (NWT). If distal outcropsof carbonate platform with reefs are added, thisbelt reached another 2000 km into southern Ari-zona, and 800 km north up to Banks Island. Fras-nian reefs were time-clustered in three cycles :(1) an Early Beaverhill Lake-Waterways cycle 4(which may include some Givetian strata, fromthe disparilis CZ through Frasnian CZ 4, a marine

transgressive phase, (2) the Woodbend (cycle 5)and (3) a generally regressive phase, the Winter-burn, cycle 6 (Reinson et al., 1993; Weissenber-ger, 1994). Woodbend cycle 5 (Middle Frasnian,CZ 5^11) saw an extensive reef-rimmed shelf com-plex developed on the southern Alberta Shelf, ex-panding wider with the 250 m thick Middle Fras-nian Leduc Reef complexes on the Cooking Lakeand Beaverhill carbonate platforms to centralnorth Alberta (Andrichuk, 1958), and fringingreefs around the Peace River Arch to the north-west. Many of these reefs were terminated at theend of cycle 5 (within CZ 11, the rhenana zone).Small reefs continued growth around the WestPembina area (Nisku reefs) during more regres-sive phases of succeeding Winterburn cycle 6 inthe Late Frasnian. However, few reefs, if any,seem to have extended up to the F/F boundary.Organic-rich, black shales and limestones accu-mulated in the intervening basins during most ofthe Frasnian (Geldsetzer and Morrow, 1992), pro-viding the source for petroleum trapped in thereefs. These Frasnian black limestones, compara-ble to the Givetian^Frasnian Domanik facies ofthe Russian Platform, and Lower Devonian Nan-dan facies of South China, were developed as in-ter-reef or back-reef facies throughout the Fras-nian. The mid-Frasnian of western arctic BanksIsland shows the northernmost Frasnian reefs inCanada, developed as a ca. 150 m thick carbonatebarrier bank on the distal margins of a giant deltacomplex s that stretched west from Greenland(Thorsteinsson and Tozer, 1962). Reefs becamescarce in the uppermost Frasnian (CZ 13, lingui-formis zone), largely disappearing with regressivefacies, or absent due to crustal £exing and deep-ening into open marine facies in the west andnorth (NWT and Yukon). Uppermost Frasnianreefs of the District of Mackenzie and NE BritishColumbia, belonging to the Kakisa Formation,were open marine, small coral patch reefs withan abundant and diverse rugose coral fauna, re-markably the most diverse Frasnian coral faunain western Canada with some 32 species (McLeanand Klapper, 1998). Uppermost Frasnian rugosecoral patch reefs up to 30 m thick also outcrop inthe Rocky Mountains, and succeeding Famennianstrata shallow into intertidal facies.

PALAEO 2824 31-5-02


The Carnarvon Basin, Lennard Shelf and Can-ning Basin of Australia developed patch reefs,barriers, banks and atolls through the Frasnian.Maximal reef development in the Canning BarrierReef came during the mid-Frasnian. Such reefsfeatured a £ourishing coral^stromatoporoid andcalcimicrobial community (Wood, 1998, 1999).Wood (2000a) suggested that the back-reef areaof the Canning Basin was dominated by a micro-bial community that also included branching andvery large domed stromatoporoids, the latter upto 5 m in diameter. Precise expansion and con-traction of the various Australian barrier, atoll,platform, patch and pinnacle reefs is not yetknown for the Frasnian, but the coral^stromatop-oroid reefs of the Pillara Formation were termi-nated at or close to the F/F boundary, and under-went karsti¢cation (Holmes and Christie-Blick,1993). South China’s (Guangxi, Guizhou, Hunan)Frasnian reefs parallel those of the Canning Basinin many respects, with the growth of coral^stro-matoporoid-rimmed banks, barriers and patchreefs curtailed by Famennian intertidal or re-stricted marine facies.

3. The F/F boundary and succeeding Famenniancalcimicrobial reefs

The long Famennian time interval eliminatedcoral^sponge reef habitats on a worldwide basis,with a protracted ‘winter’ of condensed shelf car-bonate productivity. The Famennian must havebeen a 21 myr long period of continued stress inthe tropical carbonate setting. In terms of CaCO3production (e.g. Timan^Pechora carbonate plat-form), the total thickness for the Frasnian was350 m, but only 160 m for the Famennian (Men-ner et al., 1996), which indicates that carbonatesediment production dropped from ca. 70 m perMyr to 8 m per Myr after the F/F extinction, adrop of nearly 90%. This di¡erence is comparableto average tropical carbonate production on theGreat Barrier Reef versus the temperate cool-water Great Australian Bight, and greater thanthe decline of platform reefs in the Late Miocenedue to temperature drops (Isern et al., 1996).Although other areas have not been compared,

this suggests that at the F/F boundary, carbonateproduction fell at rates as great as reef losses, andthat the systems were coupled (but contrast Kiess-ling et al., 2000). Area available for carbonateplatforms also shrank as sealevel lowstands werefavored (Fig. 3). Dramatic global biodiversity de-clines and extinctions, and absence of innovationof new species of reef faunas for the Late Fras-nian (McGhee, 1982) were the second factor atwork. Losses were not on a ‘decimation’ scale(i.e. a 10% loss), but a major ecosystem catastro-phe in the range of 60^85% of the skeleton- andreef-building invertebrate phyla, and of equatorialtaxa in general, primarily the corals and stroma-toporoid sponges. Famennian rugose corals weremostly solitary, with only rare colonial forms(Sorauf, 1989, 1992; Poty, 1999). Very few tabu-late corals are prominent (Smirnova in Simakovet al., 1983; Scrutton, 1998), and neither coralgroups were reef builders. Stromatoporoids werestragglers that limped through the Early Famen-nian, had a minor resurgence in the Late Famen-nian Strunian substage, and disappeared by theclose of the Famennian (Smirnova in Simakovet al., 1983; Stearn, 1987, 1988, 1997; Stock,1990, 1997; Mistiaen et al., 1998; Stearn et al.,1999).Few Famennian metazoan reefs are recorded,

and these were inhabited by low abundance, lowdiversity, Lazarus metazoans, such as labechiidstromatoporoids and lithistid sponges. Mostwere mudmound reefs, or occurred as microbialreef caps in open marine, shelf settings. Thoughlocal alpha diversity for lithistid sponges and cal-cimicrobes in some reefs was moderate in theCanning Basin (Wood, 2000b), this paled by com-parison to similar reefs in the Frasnian, and coralswere absent in these reefs. The ‘crisis-progenitormodel’ of Kau¡man and Harries (1996), e.g. thathere the calcimicrobes, lithistid and rare stroma-toporoid sponges were the post-extinction win-ners, generally holds true for the Famennian.The Famennian carbonate platform was typi¢edby calcimicrobes (Fig. 3), along with solenoporidred algae, dasyclad greens, and the ¢rst calcite-shelled foraminiferans. Thus, a new Late Palaeo-zoic, i.e. Carboniferous^Permian consortium, be-gan to operate (note that many Famennian out-

PALAEO 2824 31-5-02


crops were formerly dated as Carboniferous). Cal-cimicrobes and algae radiated rapidly into manynew genera in a not surprising burst of evolution,countering the pattern of extinction in nearly allother biota, as they became abundant to rock-forming (Bogush et al., 1990; Bolshakova et al.,1994). Reefs in the Famennian were predomi-nantly constructed by the Renalcis, Rothpletzella,Girvanella, Chabakovia, Renalcis, Parachaetetes,and Shuguria complex, that formed hard, resis-tant, frame-building, stromatolite-like crusts,mounds, pillars and digits. This post-F/F catas-trophe response was the result of the mass extinc-tion of the main invertebrate frame-builders, andthe loss of accommodation space due to globalregressive phases, and changing cooler climatesshifting into Late Palaeozoic icehouse modes.The top of the Frasnian carbonate shelf succes-

sion, virtually worldwide, is marked by a discon-formity and an erosional gap (sometimes extend-ing well up into the lower Famennian, and downinto the uppermost Frasnian). This is clearly evi-

dent, for example, in correlation charts and sec-tional reconstructions for large parts of the cen-tral and western Canada carbonate platform(Reinson et al., 1993). In an area covering over800 000 km2, extending from the southern NWT,through northern Alberta, over the Peace Riverarch, into central Alberta and the Williston Basin,the Frasnian succession is capped by such a dis-conformity. For the NWT, Geldsetzer et al.(1993) indicated that regional sealevel lowstandsbegan as early as the rhenana zone and persistedthrough the F/F boundary in shallow shelf set-tings: reefs were karsti¢ed, and developed ¢ssureswhich were in¢lled. In the central and easternarctic, from Victoria Island through Greenland,Famennian strata consist of siliciclastic delta com-plexes that prograded westwards and southwards,burying pre-existing mid-Frasnian reefs of BanksIsland to the far west (Goodbody, 1988; Embry,1991). No Late Frasnian, nor Famennian reefswere developed along a coastal lowland systemover 1500 km long, where reefs had once been

Fig. 3. Schematic setting for global, Early Famennian, lowstand setting, with calcimicrobial reef development, following the F/Fextinction boundary events: calcimicrobes inherited the platform architecture, or there was periodic build-up of mudmounds dur-ing intermittent highstands. The cyanobacterial consortium of Renalcis, Rothpletzella and Epiphyton dominated in carbonate plat-form margin and slope settings (retaining the structure of the barrier reef), with isolated relict stromatoporoid patch reefs devel-oped primarily in o¡-reef, muddy, quieter water settings.

PALAEO 2824 31-5-02


extensive during the Emsian through Givetian(Thorsteinsson and Mayer, 1987). Delta develop-ment was assisted by dominant recurring Famen-nian sealevel lowstands that enabled erosion offormer marine platforms, and renewed river inci-sion inland. The F/F boundary here can be de-¢ned only within terrestrial sandstone sequencescontaining miospores and plant remains (Embryand Klovan, 1976).

3.1. Western Europe (W. Baltica plate)

During the Famennian, the north Europeanplatform was located in the 15^20‡S latitudes,£anking the Old Red continent Laurussia (Fig.4). In general, Europe featured a major regressivemegasequence in the Famennian, interrupted by

shorter, periodic transgressive events, triggeredby regional epeirogenic movements as seen in dis-conformities, erosional gaps, evaporites, blackshales, extensive sandstones, and paleosols (Dree-sen et al., 1985a, 1988). Coral^sponge reefs of theBelgian Ardennes shelf progressively died outwithin the rhenana CZ, below the linguiformisCZ, and the F/F boundary (Dreesen et al.,1985b). Black shales mark the terminal Famen-nian Hangenberg event (praesulcata CZ), butironstones, ooids, and hardgrounds record ero-sional gaps, and turbulent, cyclic oceanic eventsthroughout the Famennian (Dreesen et al., 1988).Mid-Famennian mudmounds up to 100 m thick,formed by crinoids, dasyclad green algae, andsponges cemented by calcimicrobes, in an openmarine, marginal ramp setting, and possibly emer-

Fig. 4. Global reef base map for the Famennian, showing loss of reef coverage in terms of size, geographic distribution and low-er latitudes (modi¢ed from Copper, 2002). Two adjacent super-continents dominated, the equator straddling Old Red continent,Laurussia, separated by a narrow sea from Siberia to the north, and closed o¡ to the south by Gondwana, centered on thesouthern polar regions. Ocean currents were thus equatorially blocked, except by narrow straits, producing divergent warm-watermasses: nearly all reefs were in this ancient equatorial paleo-Tethys. Metazoan reefs were extremely scarce and patchy, with stro-matoporoid patch reefs known from Belgium, Poland, and equatorial W. Canada, and lithistid sponge reefs from Australia (£ow-ers): most reefs were calcimicrobial (solid circles). Post-crepida CZ reefs appear to be scarce, possibly due to mid-Famenniancooling episodes, with data mainly derived from localities in Belgium and Poland, e.g. isolated build-ups from the marginiferaand postera-praesulcata zones. Vacated and karsted reef platforms were partially recovered by calcimicrobial consortia presentduring Famennian transgressive cycles. Note the relatively southernly latitude Canning and Bonaparte basins of Australia, north-ernly location of Russia, absence of reefs in northeastern and north central Gondwana.

PALAEO 2824 31-5-02


gent in later stages, are described from the mar-ginifera CZ in Belgium and Germany (Kasig andWilder, 1983; Dreesen and Flajs, 1984; Dreesenet al., 1985a, 1991; Dreesen, 1989). The ¢nal stro-matoporoid patch reefs of Belgium occur withinthe Etroeungt (Strunian) praesulcata CZ, directlybelow the Carboniferous boundary (Dreesen,1989). Stromatoporoids occurred more as thicketsor biostromes, such as in the Epinette andEtroeungt formations of Belgium, rather than asprominent reefs (Paproth et al., 1986). Rare smallcoral patch reefs, 6 1^2 m thick, also occur in theStrunian of Belgium (Poty, 1999).During the latest Frasnian and Famennian,

some of the Frasnian reefs, e.g. the Iberg atollcomplex of Germany, built on a volcanic pile,were eroded, karsted, pierced by neptunian dykes,and depressions were in¢lled or covered withblack shales and phosphorite nodules (Franke,1973; Fuchs, 1990; Schindler, 1990; Gischler,1992. Fuchs (1990), who examined the Elberinge-rode reef complex, concluded that the atoll ulti-mately foundered and was covered by deep-waterlimestones. The subsiding atoll thus became a sea-mount, covered by deep-water coral banks duringthe marginifera CZ, but emergent for most of theremaining Famennian (Gischler, 1996). He alsoremarked that these ‘drowned’ atolls were pre-vented from reef revival by cold waters. This re-sponse is analogous to that of ‘drowned’ Creta-ceous atolls and carbonate platforms in thewestern Paci¢c ocean today, part of the sinkinghotspot system that a¡ects seamounts as they ei-ther become inactive volcanic piles, or move outof the equatorial belt (Wheeler and Aharon, 1991;Wilson et al., 1998). However, ancient platform‘drowning’ (i.e. loss or reduction of carbonateproduction) may be a direct response to coolingclimates, and have nothing to do with relativesealevel rise or tectonically sinking platforms(Isern et al., 1996; Schlager, 1999).In Poland, there is generally a major Early Fa-

mennian erosional gap above the uppermost Fras-nian shallow carbonate platform, with karst sur-faces developed as low as the Late Frasnianrhenana CZ, except in deeper water facies (Szulc-zewski, 1986; Narkiewicz, 1988). Casier et al.(2000) suggested that normal marine carbonate

facies, under a sealevel drop of 10 m, at leastlocally switched the platform to semi-restrictedback shoals and erosional surfaces at the F/Fboundary. Where present, the ¢rst recovering Pol-ish Famennian reefs of the crepida CZ were mud-mounds with crinoids, bryozoans, and algae (Ma-tyja, 1988; Racki, 1990). Other calcimicrobialmounds, or small, shallow-water stromatoporoidpatch reefs were present in higher rhomboidea topostera CZ, £anking peri-tidal carbonates (Nar-kiewicz, 1988; Szulczewski et al., 1996; Racki,1997). The Famennian of Poland ended in car-bonate shoaling and a further regressive discon-formity in the praesulcata CZ (Racki, 1997). Inthe Prague basin, reefs had disappeared by thebeginning of the Frasnian (Zukalova and Skocek,1979; Zukalova and Chlupac, 1982). For Mora-via, carbonate production declined from the EarlyFrasnian to Famennian (Hladil, 1986, 1988).Dvorak (1986) listed Moravian ‘Famennian’ reefs(now redated as Frasnian!) with rare stromatop-oroids, and condensed carbonates with calcimic-robes (algal nodules) and foraminiferans. Fras-nian reefs were terminated diachronously earlierin the north, forming emergent karst highs in theFamennian, or replaced by shoaling foraminiferalshelf facies, algal laminites and evaporites. Fa-mennian basinal facies in Moravia were markedby cherts with planktic radiolarians and benthicsponges (Chlupac, 1988).The stratotype F/F boundary is set in the Mon-

tagne Noire area of southern France, in deeperwater, ammonoid and conodont-bearing, pelagicfacies, lacking reefs or reefal mudmounds at theboundary (Klapper et al., 1993). Reefs ceased herein the Late Emsian. In the Montagne Noire, theFamennian ostracode survivors were suggested tohave come from oxygen oases in shallow waters,without reefs or mudmounds (Lethiers and Ca-sier, 1994). Spain and the French Pyrenees weredevoid of reefs by the Famennian, as platformswere converted to deeper water, black shale^lime-stone facies. In the Carnic Alps, F/F boundarystrata are phosphatic (up to 90%), and no Famen-nian mudmounds or reefs are known (Vai, 1963,1967; Bandel, 1972). In Morocco, the Famennianlacked metazoan reefs and mudmounds, with se-quences dominated by siliciclastics, turbidites, or

PALAEO 2824 31-5-02


erosional gaps (Gendrot, 1973; Wendt, 1988;Benbouziane et al., 1993).

3.2. Central and eastern Baltica (including RussianPlatform, Urals)

In the Timan^Pechora region, stromatoporoid^coral reefs ceased after the Frasnian and werereplaced by ‘algal bioherms and mud hills’ (Men-ner et al., 1996). In the northern Pechora Urals adiverse assemblage of calcimicrobes such as Re-nalcis, Epiphyton, Izhella and Shuguria formedframework in Famennian patch reefs, and wereaccompanied by inter-reef or lagoonal facies con-taining calcispheres (Belyaeva, 1986; Antoshkina,1997). Some of these Famennian Pechora reefsoccur as petroleum-bearing prospects in the sub-surface (Belyaeva, 1986). On the NE tip of No-vaya Zemlya Island, Bondarev et al. (1967, p. 110)identi¢ed Famennian calcimicrobial reefs in a re-gressive setting, con¢rmed by observations of An-dreeva et al. (1979). Famennian calcimicrobial‘Girvanella’, and red and green algal reefs, fromdrill-core data, were said to continue on tops ofhighs created by underlying Frasnian reefs in thesouthwestern Urals, and to be present in stratapossibly as young as the Tournaisian (Ulmishek,1988). A disconformity at the base of these Fa-mennian microbial subsurface reefs is not re-ported, though the system switched from reef toshelf limestones at the F/F boundary (Ulmishek,1988). In the Urals, some reefs were mudmoundswith an impoverished bryozoan, sponge or un-known microbial contribution, and minor skeletalremains, and in others, Famennian calcimicrobialpatch reefs with distinctive fabrics (Shuiskii,1986). Such reefs were also said to continue intothe Early Tournaisian (Mirchink, 1974). Famen-nian reefs disappeared from the shallow platformto the west, and were con¢ned to calcimicrobialbuild-ups along the new shelf edge margin, S andE of Pechora, in the black Domanik facies (Men-ner et al., 1996).For the central and southern Russian Platform,

Moskvich and Kruchek (1984) and Makhnach etal. (1986) ¢gured Early Famennian microbial (‘al-gal’) reefs, from the Pripyat High or ‘Beloruss’shelf, in a 250 km wide tract, covered by Late

Famennian dolostones and evaporites. No Fa-mennian black shales are reported as £oodingthis platform, but microbial (‘algal’) reefs wererepeated in the Carboniferous in the same, albeitreduced, area of Beloruss and Ukraine (Makh-nach et al., 1986). In the Precaspian region ofthe SE Russian Platform, Famennian (and Tour-naisian) microbial reefs, 6 10 m thick, are presentnear Saratov in a sequence with dolostones, oo-lites and conglomerates (Rusetskaya and Yaro-shenko, 1990). In the southern part of the Precas-pian Synclinorium, data from oil¢elds on theN margin of the Caspian Sea, 400^500 km E of theVolga delta (NW Kazakhstan, Karaton), identifysubsurface reefal limestones of Famennian andEarly Carboniferous age. These are of unde-scribed composition, in a succession some 80^90m thick (Rusetskaya and Yaroshenko, 1990). Inthe Caucasus to the south, a collisional continen-tal margin facies suppressed Famennian carbon-ates, which are frequently interrupted by erosionaldisconformities, slumped units, and sandstones.Much of the sequence is regressional, reef activityhaving ceased in the Givetian (Chegodaev et al.,1984).

3.3. Kazakhstan plate and Near East plate(s)

Famennian through Tournaisian reefs in Ka-zakhstan broadly conformed to patterns elsewhere(Zadoroshnaya et al., 1990a). In central Kazakh-stan, ca. 300 km NW of Lake Balkhash, a terrig-enous-volcanogenic pile of sediments incorpo-rated ‘algal’ reef massifs of Famennian age(Buzmakov and Shibrik, 1976). Though the fring-ing reefs are only a few meters thick (condensedFamennian carbonate section = 50^100 m), theyextend discontinuously for about 150 km; Tour-naisian build-ups are thicker. In the northernTyan-Shan Fold Belt (Karatau range, S. Kazakh-stan), Famennian reef belts stretch about 500 kmSE towards the Akshirak^Moldotau range, westof Lake Issuk Kul. These appear to be dolomi-tized, Famennian rimmed, ‘algal’ reef complexes,covering an area about 10 km2 but ‘algal’ texturesare diagenetic relicts (the Famenne section here isabout 500 m thick). The carbonates are near thetop of a 1^5 km thick, rapidly subsiding sequen-

PALAEO 2824 31-5-02


ces of molasse and volcanics, probably derivedfrom an actively uplifting area during the Devo-nian and Carboniferous (Zadoroshnaya et al.,1990b). From Bolshoi Karatau, the same generalarea, Cook et al. (1994) ¢gured Famennian‘Waulsortian’-type reef mounds, dominated bycrinoids, bryozoans and algae, up to 100 m thickand 2 km long. Tournaisian reefs continue thisnorth Tyan-Shan section. In the Caspian Basinof southern Kazakhstan, seismic modeling hasdiscovered a very large, horseshoe-shaped reefatoll (the Tengiz structure, 400 km2), initiated inthe Frasnian, interrupted by probable sealevellowstands but continuing into the Famennianand up to the Visean (Pavlov et al., 1988). Be-cause of its great depth at 4^5 km, the morethan 1 km thick Tengiz reefal units, with sixstratigraphic breaks, are not precisely dated, andthe nature of the Frasnian and Famennian com-ponents of the reef are not known. In the south-ern Tyan-Shan ranges of Zeravshan, Famennianstrata of the Yatavluk Fm., some 500^550 mthick, contain dolomitized stromatolitic moundsassociated with lagoonal or peri-tidal micrites(Kim et al., 1984). Such Famennian microbialreefs may also be present in the Chinese Tyan-Shan ranges, but remain to be discovered. NoFamennian reefs are known from Iran, Pakistanor Afghanistan, in areas that previously sup-ported limited Frasnian or Givetian reef develop-ment (Gaetani, 1968; Huckriede et al., 1972; Das-tanpour, 1996). Frasnian reefs were replaced inthe rhenana CZ by bryozoan and stromatolitecommunities (Mistiaen et al., 2000).

3.4. Siberia plate

Lower Famennian calcimicrobial (‘algal’) reefs,several meters thick, and locally associated withbryozoans, superceded Frasnian coral reefs alongthe Tom River north of Kemerovo (in the Kuz-netsk Basin), over a distance of about 60 km.These microbial reefs disappeared in higher strata(reefs illustrated by Belskaya, 1960, pp. 164^165,pl. 1, ¢g. 2). Ivanova (1983, ¢g. 3) illustrated twoFamennian ‘algal reef’, and rimmed bank com-plexes in the Kuznetsk Basin, west of Kemerovo,one ca. 45 km in diameter, and the other ca. 30

km, with a bryozoan patch reef facies in the on-shore Salair paleocontinent setting. Belskaya(1960, ¢gs. 51^52) described and ¢gured promi-nent Famennian ‘algal’ reefs, replacing earlier,more widely distributed Frasnian coral reefs inthe Kuznetsk Basin, close to Kemerovo, stretch-ing over a distance of nearly 100 km. Famennianand Early Carboniferous ‘algal’ reef massifs alsooccur in the Alai Ranges near Archaltur (Dronovand Natalin, 1990). Yolkin et al. (1994) pointedout that ‘powerful volcanism’ within the Altaimountains, the fold belt £anking the KuznetskBasin in the Late Devonian, may have disruptedsome of the reef development in the region.

3.5. Mongolia (Tuva), North China and NERussia plates

Famennian-age reefs are thus far unknown inthe far-eastern parts of Russia (Kolyma^Chukot)and in Mongolia (Tuva). Though a major reefbelt nearly 2000 km long on the Mongolian plateextended discontinuously from the Late Silurian(Ludlow) through Eifelian (Middle Devonian),this appears to have marked the end for condi-tions suitable for the growth of either microbialreefs, reefal mudmounds or stromatoporoid reefs(Sharkova, 1980, 1986a,b). Sharkova (1986a,b) at-tributed the termination of reef growth in the Ei-felian to submergence and turbidity as a result ofactive tectonism, but it is also possible that thewhole continent had simply moved northwardsout of the tropical reef belt, which was alreadyat relatively high north latitudes in the Devonian(Fig. 4: assuming the Tuva location is correct).There was no reef development in the Late De-vonian and Early Carboniferous, thus sheddingno direct clues on F/F reef demise in the area.In the Omolon Massif of far northeastern Russia,apparently only Famennian coral and stromato-poroid biostromes occur, but without metazoanreef, or microbial mounds (Simakov et al., 1983).Similarly, for the North China plate no MiddleDevonian nor Frasnian^Famennian reefs havebeen described to date. These areas without reefgrowth suggest possible drift of these plates out ofthe tropics, or unsuitable conditions for reefgrowth in the Famennian.

PALAEO 2824 31-5-02


3.6. Laurentia

The western Canada sedimentary basin, whichhad the world’s largest Frasnian reef tracts, lostits metazoan coral^sponge reefs by the Famen-nian. Calcimicrobial reefs are also not known,possibly due to shallow to emergent conditionsover much of the carbonate shelf. In Alberta,the weathered, karsted surface of the latest Fras-nian Southesk Formation, e.g. at Jasper Park inthe central Rockies, is typically overlain by theprimarily intertidal dolomitic siltstones of the Fa-mennian Sassenach Fm., with reefs ending in themid-Frasnian Mount Hawk Fm. (MacKenzie,1969). Reinson et al. (1993) identify the F/F breakat the top of their cycle C6, the WinterburnGroup, which is succeeded by cycle C7, the Wa-bamun Group. Stromatolites in the Wabamunsediments suggest highly restrictive and stressedconditions on the western side of Laurentia. TheF/F disconformity is followed in other areas bythe ‘Graminia Silt’, a regressive unit recording athickening siliciclastic wedge, the base of whichmarks the F/F boundary locally (Reinson et al.,1993). McLean and Klapper (1998), however,dated the regressive lower Graminia Silt as latestFrasnian: this suggests that uppermost marinestrata are discontinuous, and that regression be-gan in the latest Frasnian. The only known Fa-mennian reefs on the west £anks of North Amer-ica, over a distance of more than 5000 km, aresmall stromatoporoid patch reefs of mid- to lateFamennian age recorded from Alberta boreholedata (Stearn et al., 1987; Nishida, 1987; Stearn,1988; Halim-Dihardja and Mountjoy, 1988), andsome small outcropping mounds along Humming-bird Creek, probably in the crepida CZ (Stearn,1987). In basinal sequences, a regressive event isalso recorded in western Canada (Van Buchem etal., 1996). To the contrary, Geldsetzer et al. (1987)suggested that a basinal section near Jasper, Alber-ta, was marked by £ooding of anoxic waters,spiked by sharply positive N

34S values at the F/Fboundary, a feature not con¢rmed elsewhere.However, this level also has icriodid conodonts,indicating shallowing. In a 1200 km long beltfrom Nevada to Montana, the terminal Famennian(praesulcata CZ) is featured by vast beds of Girva-

nella^Wetheredella oncoids up to 3 m thick, over-lying black, cherty, conchostracan-rich mid-Fa-mennian shales (Rodriguez and Gutschick, 2000).In Iowa, in central Laurentia, Frasnian reefs

are missing, and there was a post-Middle Fras-nian platform regression marked by deep LateFrasnian shale ¢ssure in¢lls containing brachio-pods (Day, 1998). Frasnian reefs were absent ineastern North America (Fagerstrom, 1983). TheFamennian in southern and south central Lauren-tia (e.g. Michigan Basin, Ontario, New York,Pennsylvania) was marked by shallow, hypoxicto anoxic waters, pyrite and organic-rich, silici-clastic mud deposition, or local deepening (Gut-schick and Sandberg, 1991), partly as a responseof crustal £exing and the expansion of the Catskilldelta. Even bryozoan, or microbial mounds areunknown. The Famennian black shales (the An-trim, Kettle Point formations, etc.) were rich inwoody trunks, leaves, and fresh-water molluscsand insects, deposited in delta-distant, coastallowlands under fresh water, or estuarine to partlymarine conditions. Widespread local marine blackshale horizons, precisely dated by conodonts andbentonites, occur below, at and above the F/Fboundary in eastern North America, and thusKellwasser-type ‘anoxia’ here appears to mark lo-cal sedimentary anoxic or hypertrophic events ofdi¡erent ages (Over, 2000).

3.7. South China

Wang (1985) made no mention of the ChineseFamennian as an unusual episode in reef history,marked by regressions and loss of reef biotas.South China is assumed to have been locatedproximal to the Australian plate in the low lati-tudes 6 30‡S of the equator (Fig. 4). In southChina, Famennian (Xiwangshanian) strata weremost commonly restricted intertidal to supratidalfacies, though local deeper sediments mark a com-plete record of the F/F events, with collapse brec-cias recorded in the rhenana CZ (Hou et al.,1988). On the platform, inter-reef, shelf marginand slope, calcimicrobial reefs were dominatedvirtually exclusively by massive Renalcis and Epi-phyton (labeled as ‘algal’ mounds) on the platformmargins, such as those in Guangxi, eastern

PALAEO 2824 31-5-02


Yunnan and southern Guizhou (Hang, 1987;Tsien et al., 1988; Yu et al., 1991; Gao, 1991;Yu and Shen, 1998). Prominent, up to 35 m thickand ca. 50^180 m diameter calcimicrobial ‘Renal-cis^Epiphyton’ mounds of the upper YongshienFm. near Miaomen and Zhaijiang, west of Guilin(Guangxi province) have their surfaces intersectedby sediment-in¢lled ¢ssures showing contempora-neous erosion (Yu and Shen, 1998). Inter-reefsediments with gastropods, ostracodes and foramsindicate a restricted subtidal to peri-tidal settingfor these reefs. The famous Baisa reef in Guangxiis a Famennian calcimicrobial mound (Tsien etal., 1988; Yu and Shen, 1998).The South China calcimicrobial reefs replaced

stromatoporoid reefs of the Late Frasnian. Fa-mennian stromatoporoid reefs are scarce, thoughendemic stromatoporoids, including labechiidswere common to abundant, albeit not generallyreef-building. Rare, small and 6 5 m thick, upperFamennian stromatoporoid patch reefs are re-ported from the Dongcun Fm., at the village ofEtaoucun, south of Guilin (Milhau et al., 1997).In Xinjiang province (NE China) the colonial ru-gose coral fauna of the Late Frasnian collapsed,and the Famennian rugosans were simple, solitaryforms that appear to have been con¢ned to deeperwaters (Guo, 1990). In platform non-reefal facies,the uppermost Frasnian is a normal marine facies,and basal Famennian is marked by a peloidal,‘‘algal-laminated limestone representing a tidal£at environment’’ (Hou et al., 1988). In deeperwater, o¡-reef sections of Maanshan in Guangxiprovince, the top of the Frasnian is marked by abrachiopod bed of ‘‘shell-beach facies under highdynamic conditions [sic]’’, de¢ning a short shal-lowing event directly below the boundary. Thereis no record of any within-Famennian faunal ex-tinction in forams or microbes, nor any sedimen-tologic carbonate platform cessation in SouthChina, e.g. in the crepida CZ, as suggested byWang et al. (1994). Wang (1992) discovered glassymicrospherules, below a siderophile anomaly,within the crepida CZ from the northern marginof the South China carbonate platform, and sug-gested Taihu Lake near Shanghai as the impactsite : neither reefs, nor diversity losses, are re-ported in these strata. Muchez et al. (1996) com-

pared the South China and Belgian sections of theF/F boundary and suggested that the same eu-static sealevel fall occurred in both, coincidingwith the base of the triangularis CZ. In Vietnam,small stromatoporoid patch reefs are reportedfrom the latest Famennian (Strunian) by Nguyenand Mistiaen (1998).

3.8. Australia

There is an erosional gap at the top of theFrasnian, and possibly the earliest Famennian,in much of the Canning Basin, which made theshallow carbonate platform emergent (Playford,1984; Cockbain and Playford, 1988; Playford etal., 1989). Famennian reefs of the Nullara andWindjana limestones were dominated by back-stepping cycles with the cyanobacteria Renalcis,Shuguria, Girvanella and Rothpletzella in shallowwaters less than 45 m deep on both the platformand shallow slope (Playford and Cockbain, 1969;Begg, 1987). Some calcimicrobial reefs were prob-ably at the limit of the photic zone at 100 m, andlithistid sponge reefs existed in still deeper waters(Playford and Cockbain, 1969). Reef architecturewas largely inherited from the pre-existing car-bonate platform, and modi¢ed by regressivepulses. Neptunian fracturing and platform col-lapse may be partly related to rapid sealevel £uc-tuations at this time (Southgate et al., 1993). Itwas suggested by Wood (2000b) that a local novelreef ecology with a relatively diverse lithistid,sphinctozoan and calcimicrobial community dem-onstrated that Famennian reef recovery was in-stantaneous in the Canning Basin, and that thereef ecosystem had been fully re-established.Nevertheless, corals played no role in the reefs,as they did in the Frasnian, and stromatoporoidswere minor or absent.The iron-concentrating cyanobacterium Frutex-

ites marks a hypoxic condensation surface, and isfound on some reef and reef slope surfaces withinthe Famennian crepida CZ, at a level post-datingthe F/F extinction by possibly 0.5^1.5 Myr ormore. This con¢rms that the iridium anomaly isnot associated with any metazoan reef or faunalextinction here, contradicting Wang et al. (1994).No black shales are known from the carbonate

PALAEO 2824 31-5-02


platform. Carbonate platform and calcimicrobialreef growth seems to have persisted through theLate Famennian, at which time it stopped (Play-ford, 1980; Becker and House, 1997). Becker(1995) stated that ‘stromatolitic’ (calcimicrobial,e.g. Rothpletzella, etc.) biostromes and ‘mini-bio-herms’ were best developed in Famennian trans-gressive pulses on platform slopes and £ats.George et al. (1995) interpreted a mid-Famennianplatform margin collapse during a sealevel high-stand of the Canning Basin, marked by dislodgedreef blocks, and this was suggested to be due toover-steepening and/or tectonic activity. On thetectonically active eastern margins of Australiaand New Zealand, no post-Emsian reefs areknown.

4. Temperate to cool Gondwana (South America,southern Africa)

This region was located within the south polarcold to cool temperate climate regime, as seenfrom the dominance of siliciclastics, total absenceof carbonates, and declining abundance of marineshelf benthic faunas through the Devonian (Cop-per, 1977). From the Eifelian onwards, brachio-pod and mollusc faunas declined (bivalves, gastro-pods), and even the rare Early Devonian favositidcorals had vanished. Reefs and microbial mud-mounds were completely absent in this realmthroughout the Devonian, the last reefs being re-ported from the Middle Ordovician (Copper,1997). Glaciation commenced sometime in the Fa-mennian or even earlier, as seen from striatedpebbles, diamictites, dropstones, and other glacio-marine sediments (Caputo and Crowell, 1985;Barrett and Isaacson, 1988; Isaacson, 1997;Isaacson and Grader, 1997). The only thin car-bonates and low diversity coral faunas were de-veloped on the northern margin of South Amer-ica, but reefs were absent (Scrutton, 1977).

5. The deep-water, pelagic setting during the F/Fboundary

Deep-water settings during the F/F boundary

interval generally record continuous sedimenta-tion, and little biotic disturbance of their benthicfaunas: the zonal stratigraphic scheme is exclu-sively based on this environment. Sorauf and Ped-der (1986) demonstrated that deep-water solitaryrugose coral faunas persisted in the Famennian,and were almost undisturbed. There were virtuallyno Famennian colonial rugosans, though somereturned in the Strunian (Poty, 1999). For theostracodes, although shallow-water taxa werestrongly a¡ected by F/F extinctions, deep-waterostracodes were ‘almost untouched’ (Lethiersand Casier, 1996, 1999a,b). Famennian deep-water mudmounds appear to have been veryrare to absent, though the glass sponges and ra-diolarians saw a radiation in North American andEuropean slope to basin sections (McGhee, 1982;Racki, 1999; Racki and Cordey, 2000). The Mon-tagne Noire section at Coumiac, with condenseddark gray to black, carbonate^shale facies wasdeposited on a deep submarine rise (several 100^1000 m+ deep), continuously across the bound-ary. Both sections record planktic and nektic fau-nas consisting of ammonoids, conodonts, smallepiplanktic bivalves, palynomorphs, acritarchs,chitinozoans, microforams, and pelagic to benthicostracodes and trilobites (a number of the trilo-bites are blind: Feist, 1991). Through the upper-most Frasnian linguiformis CZ, about 2 m thick,and possibly lasting as long as 500 000 yr (Sand-berg et al., 1988), conodont abundance droppeddramatically from about 10 000 elements per kg ofrock to less than 100 per kg, about 1 m below theextinction boundary, thus over ca. 250 000 yr(Girard in Paris et al., 1998). At the same time,the tasmanitid £ora jumped in abundance (in con-junction with microforams), suggesting high sur-face productivity and nutrient abundance in theLate Frasnian pelagic system, as also seen in east-ern North America and Brazil at this time. Incontrast, chitinozoan plankton were relativelylow in abundance in the Late Frasnian, but roseto 19 000 specimens per g of rock in the EarlyFamennian, the highest known abundance of chi-tinozoans in the fossil record to date (Paris et al.,1998). Since the a⁄nity of chitinozoans is un-known, it is tempting to suggest that these wereremnants of detritivores (egg cases, resting

PALAEO 2824 31-5-02


stages?), feeding on the enriched planktic detritusderived from the ¢nal collapse at the F/F bound-ary (possibly a disaster biota). Paris et al. (1998)suggest basal Famennian chitinozoan abundancewas due to a ‘‘temporary lowering of ocean tem-peratureT and very low activity of the usual pred-ators’’. Girard and Albarede (1996), who lookedat conodont phosphorus, suggested that the ex-tinction in the Montagne Noire was due to trans-gression and anoxia.

6. Why did the Mid-Palaeozoic reef ecosystemcollapse?

The global nature of the ‘catastrophic’ F/F ex-tinction was ¢rst suggested by Copper (1966), andexpanded by McLaren (1970), who proposed anextra-terrestrial impact, associated with giant tsu-namis. Most of the theories regarding the F/Fextinction have been succinctly surveyed byMcGhee (1996), but no papers have focused onwhat happened globally to carbonate platformsand reefs at the time. To determine the causesof catastrophic reef losses during the Late Fras-nian, the following must be taken into account:(1) Reefs disappeared prior or close to the

boundary, within or prior to the last one or twoCZs, almost on a worldwide basis.(2) Reef losses were not simultaneous world-

wide, but show progressive stepdown declines,with smallest and fewest reefs towards the F/Fboundary, matching diversity declines (Copper,1984, 1986).(3) The terminal Frasnian extinction events

were marked by regressions in virtually all plat-form carbonate successions that can be dated, andthe overlying Famennian is initially also usuallyregressive, with the break usually marked by dis-conformities, erosional boundaries and sharpchanges in lithology.(4) The Frasnian^Famennian was marked by

rapidly oscillating sealevel lowstands and high-stands.(5) Deep-water rugose coral faunas, mostly soli-

tary forms, were relatively una¡ected (Sorauf andPedder, 1986; Oliver and Pedder, 1994).(6) Deeper and shallower water stromatopor-

oids declined markedly in the latest Frasnian, inthe rhenana CZ, below the F/F boundary (Stearn,1975), and survived the F/F extinction, only toperish at the end of the Famennian, with the cos-mopolitan Frasnian genera surviving preferen-tially (Stearn, 1987; Cockbain, 1989).(7) Exclusively tropical orders of brachiopods,

such as pentamerids and atrypids, were com-pletely eliminated, but the high-latitude and deep-er water brachiopods survived (Copper, 1977).(8) Bryozoans had few losses, though reef

dwellers su¡ered more than others (Bigey, 1988).(9) Tropical peri-reefal ammonoids showed dra-

matic declines or total extinction, and the onlysurviving family was that present in the cool-water malvinoka¡ric province (House, 1985;Becker, 1993a,b; Becker and House, 1994).(10) Tropical planktic and benthic foraminifer-

ans showed dramatic declines and change-overs,but a resurgence in the Famennian (Kalvoda,1986).(11) Reef calcimicrobes showed no decline to-

wards the F/F boundary, and thrived in the post-F/F events, dominating reefs (Copper, 1997).(12) Terrestrial fresh-water and estuarine ¢sh

faunas showed dramatic losses towards the closeof the Frasnian (Long, 1995).(13) Terrestrial £oras show major overturns at

the F/F boundary (Streel and Loboziak, 1995).(14) Dramatic decline of platform CaCO3 pro-

duction from the Frasnian into the Famennian.A number of causes for the mass extinction of

reef biota and loss of metazoan reefs towards theF/F boundary have been proposed (Copper,1998; Racki, 1998a for brachiopod losses). Somesuggested global warming for reef decline(Thompson and Newton, 1988; Brand, 1989).Nutrient (phosphorus) poisoning was a theme inother explanations (Schlager, 1981; Hallock andSchlager, 1986; Wood, 1993; Eliuk, 1998). Theo-ries for direct causes presently seem to revolvemainly around two schools of thought: (1) oneor two cycles of worldwide oceanic anoxia asso-ciated with the lower and upper Kellwasser lime-stones (Walliser, 1984, 1996) and supported byothers (e.g. Joachimski and Buggisch, 1993;May, 1995; Murphy et al., 2000), or (2) globalclimatic cooling and sealevel drawdown, a¡ecting

PALAEO 2824 31-5-02


both sea and land, ¢rst proposed by Copper(1975, 1977). A corollary of these two e¡ectsmight be rapidly alternating T/R cycles and cli-mate change, and a coupling e¡ect involving nu-trient cycling, between the two end-points of T/Rcycles (Buggisch, 1991). The database for F/Freefs is far from complete, and new discoveriesare bound to change the picture. Reefs fortu-nately do not su¡er from the ‘Signor-Lipps e¡ect’at extinction boundaries, i.e. the absence of speci-mens or species, especially planktic and nektictaxa in key sections or drill-core, leading to apostulated gradual or stepdown interpretation,when, in fact, all organisms may have died atthe same time. Unlike single samples, whose dis-tribution and abundance varies from site to site,reefs represent the rock record of an entire eco-system which is di⁄cult to miss in outcrop orboreholes. They also represent an entire commun-ity raised above the sea£oor, usually preserved asmassive, or thickly bedded stratigraphic units,highly unlikely to erode completely in outcrop,even if diagenetically altered. Thus the reef recordis as close to complete for any ecosystem, andmuch more complete than the geologic recordfor terrestrial rainforests.

6.1. Anoxia

How would anoxia, explained by water columnstrati¢cation during transgressive pulses, forcereef demise and the loss of marine biotas on thetropical, shallow shelves? Brongersma-Sanders(1966) suggested that anoxia could be created byupwelling, of nutrient-rich and O2-depleted watersvia strong winds, but considered this a local, notglobal model. The main dilemma is how giant‘burps’ of deep water from below the carbonatecompensation depth (CCD) could kill o¡ reefs ona worldwide basis, and what would trigger suchburps? And do black shale events have the sametiming as reef demise? Wignall and Hallam (1992)and Wignall and Twitchett (1996) favored theidea of anoxia as the prime killer for the endPermian extinctions, though Hallam and Wignall(1997) also added regression as a factor, perhapseven over-riding anoxia as the main cause. Theend Permian mass extinctions coincide with well-

dated £ood basalt events in NW Siberia, the larg-est basalt traps of the Phanerozoic, and possiblesources for atmospheric CO2 or H2S enrichment:no such large scale end Devonian £ood basaltsare known at present. Vogt (1989) suggested ina general model that volcanism drove anoxiaand reef death, but there is limited evidence forthis on the huge passive margin of Laurentia,though bentonites are common on the easternside (Tucker et al., 1998). Kellwasser eventswere related to transgressions by Walliser (1984)and Joachimski et al. (2001). Black shales andlimestones have traditionally been viewed asmarkers for high rates of carbon burial (hencethe development of suitable petroleum sourcerocks), and strongly thermally layered, relativelystagnant oceans with stable water column strati¢-cation, e.g. a deep CCD. Black shales thus havebeen regarded as proxies for transgressive sealevelhighstands and global warming. Nevertheless, thisis contradicted by new models for organic-richblack shales, which include those of the BlackSea, the western coast of Peru and Ecuador inSouth America, the Gulf of California, the Nor-wegian coast, and the west coast of Africa. Mod-ern counterparts for black muds occur mostly intemperate areas, and on the cool western shelvesof continents, marked by upwelling and mixing,and high nutrient spill-over on the shallower shelfor embayments. This led Pedersen and Calvert(1990), Calvert and Pedersen (1992) and Calvertet al. (1992) to suggest that organic-rich shales inshallow and deep areas could best be explained byhigh planktic productivity, stimulated by shelfand slope upwelling, and cooling. Murphy et al.(2000) suggested, to the contrary, that marine C,N and P biogeochemical cycles were decoupledfrom mass extinction: they preferred an anoxiceutrophication model, tied to global cooling, forthe mass extinction.How would reefs be a¡ected by global anoxia,

and is worldwide, simultaneous anoxia of thetropical shallow shelf feasible? A major problemwith the anoxia hypothesis is that it is di⁄cult toimagine how ‘giant megaburps’ of CO2 (andSO2)-enriched waters, brought up from belowthe CCD, could simultaneously spill over all theworld’s tropical shelf areas, where reefs were

PALAEO 2824 31-5-02


growing. No mechanism is known for this viawarming or transgression. At best, rapid oceanicoverturn seems possible only during major diver-sions in ocean currents and deep sea watermasses, accompanied by cooling and loss of shelfareas by regressions, as it did in the modern ice-house during the Pleistocene. Most reefs todayappear to be reasonably well adapted to livingon the margins of ecosystems which face theproblem of being ‘shot in the back’ by a stagnant,eutrophic lagoon or back-reef zone (Neumannand Macintyre, 1985). Throughout the Middleand Late Devonian, black shales and limestoneswere widespread in inter-reef basins, and in back-reef zones. Many patch reefs and their associatedcorals such as thamnoporids, and the delicate,stick-like amphiporid and stachyodid sponges,and speci¢c brachiopods (leiorhynchids, atrypids,productids, etc.) were adapted to oxygen-starvedsettings. Indeed, they form a distinctive Eifelianthrough Frasnian black, organic-rich marker fa-cies easily recognizable in drill-cores and reef out-crops throughout the world. The reef £at and out-er reefs fringed the margins of such shallow,hypoxic Devonian systems for over 20 Myr. An-oxic and hypoxic substrates were commonly rec-ognized in successions in North America, espe-cially in the Frasnian and Famennian. In otherareas, there is no evidence for anoxia and blackshales or limestones at the F/F boundary, e.g. inthe Canning reef complex, yet the metazoan reefsstill died out (Becker and House, 1997). In SouthChina, black shales are also missing at the F/Fboundary, yet metazoan reefs gave way to calci-microbial reefs (Yu and Shen, 1998). In Nevada,black shale events pre-date the F/F boundary, andare not tied to extinctions, nor carbonate plat-form changes (Bratton et al., 1999). In arctic Can-ada, Frasnian reefs died out due to sealevel low-stands and burial by deltaic siliciclastics (Embryand Klovan, 1971; Embry, 1991). There is noevident, direct relationship between black shalehorizons and reef disappearances in any sections,as the timing of these do not coincide. There maybe no direct link between black shales and bottomanoxia, except as indicators of high surface pro-ductivity in an icehouse climate setting. Blackshale horizons are widespread in the Late Fras-

nian, Famennian and Early Carboniferous, andbest tie in to widespread climatic cooling, andvigorous oceanic overturn (Caplan and Bustin,1999).Reefs are generally not highly nutrient tolerant,

having long term adaptations to tropical, oligo-trophic waters, since phosphorus is a major inhib-itor of carbonate skeletal precipitation, and sincehigh nutrient input may stimulate the growth ofsoft algal carpets, smothering reef metazoans.Hallock and Schlager (1986) proposed a nutrientkill model for reef demise, including the LateDevonian. Quinn and Johnson (1996), and others,however, have noted relatively strong tolerancefor some modern reefs in marginal settings to sea-sonal high nutrient loading. What was the sidee¡ect of upwelling nutrients and cold waters,which normally stimulate P, S and N production,and consequent SiO2 precipitation by plankton?For example, in cool upwelling areas of the trop-ical western Indian Ocean today, reefs have lowdiversity and low growth (Quinn and Johnson,1996; Coles, 1997). Is there evidence for phos-phorites or silica-producing biotas at the F/Fboundary, and how do these relate to the reefs?McGhee (1996) noted the coincidence of the ex-pansion of siliceous demosponges during the Fa-mennian in North America, Australia and Po-land. Racki (1999) and Racki and Cordey (2000)summarized the nature of the rise of siliceoussponges and radiolarians during extinction events,postulating a ‘hyper-siliceous’ connection, but re-lated this primarily to volcanism in NW Europe.However, widespread volcanism is not a featureelsewhere in the Devonian, particularly on passivemargin, stable carbonate platforms, nor in thegiant epicontinental seas of the Late Devonian.

6.2. Impacts

The impact origin for the F/F mass extinctionswas initially promulgated by McLaren (1970),who favored instant mass killing and destructionof ecosystems at the boundary by giant tsunamis.No evidence has yet been found for the tsunamiconcept. The impact theory has been stressed by anumber of other workers, but the geochemicalanomalies (Geldsetzer et al., 1987; Goodfellow

PALAEO 2824 31-5-02


et al., 1988; McLaren and Goodfellow, 1990;Nicoll and Playford, 1993; Wang et al., 1991,1994, 1996), and spherules (Wang, 1992) occurin the Famennian, well above the F/F boundary,and mostly within the crepida CZ, or below theF/F boundary in the rhenana CZ. Geochemicalanomalies within the two Frasnian Kellwasser ho-rizons also suggest an internal oceanic source foranomalous values of N13C (Joachimski, 1997). Amid-Famennian spherule horizon comes fromblack shales of eastern Germany (Marini et al.,1997). Belgian examples of ‘spherules’ at or belowthe F/F boundary were contaminated by roadmarker glass beads (Claeys et al., 1992, 1996).Re-examination of the boundary stratotype inthe Montagne Noire for Pt/Ir has failed to con-¢rm earlier anomalies suggested by Geldsetzer(Girard et al., 1997). The only postulated LateDevonian impact structure, the ‘Alamo Breccia’(Warme and Sandberg, 1996), has been dated asEarly Frasnian, probably 3^4 Myr prior to the F/F, and this had no known e¡ect on the carbonateplatform on which it landed in Nevada, nor onglobal or regional diversity (J.E. Sorauf, personalcommunication, regards these as slope-toe plat-form breccias, as they occur along strike formore than 150 km). Claeys and Casier (1994)and Claeys et al. (1996) have suggested the 55-km diameter Siljans crater in Sweden as a LateDevonian impact site, but the crater has not beenreliably dated (McGhee, 1996), and no proximal^distal e¡ects are identi¢ed. The Flynn crater ofeastern Laurentia is too small (6 5 km) to havebeen e¡ective in extinction or reef demise. Wang(1992) suggested Lake Taihu in China, nearShanghai, as a possible Late Devonian impactsite, but the crater age is not con¢rmed. Thusseveral possible impacts are known from the 26Myr long Late Devonian, none of which had ane¡ect on reefs, not even regional extinctions.

6.3. The ‘reef hypothesis’ and atmospheric CO2

budgets

Modern reefs use ‘keep-up’ and ‘catch-up’modes to match sealevel rise, as coral and reefcarbonate accretion is high in healthy reefs (Neu-mann and Macintyre, 1985), and there is every

indication that Devonian reef accretion rateswere high. What is the relation between high car-bonate production, via reefs, the demise of reefs,and the atmospheric CO2 budget? Berger (1982),Broecker and Peng (1987) and Opdyke andWalker (1992) suggested that reefs were net ex-porters of CO2 to the atmosphere, because ofthe CaCO3 equation, which dictates that CO2 isreleased, as CaCO3 is precipitated: this shouldtheoretically enhance atmospheric CO2 and globalwarming. This would not take into account a po-tential net surplus of O2 generated via reef-build-ing symbionts such as dino£agellates, which had afossil record already in the Neoproterozoic. Wareet al. (1992) concluded that modern reefs were netsources of CO2 to the atmosphere, not sinks, evenwith symbionts. But this is certainly not clear forthe Mid-Palaeozoic calcite greenhouse system(Copper, 1997). The demise of metazoan reefs atthe F/F boundary coincides with a rapid rise ofO2 in the atmosphere, but it seems unlikely thatLate Devonian reefs ‘shot themselves in the back’(with net CO2 output), and caused their own col-lapse, because the rise of O2 matches the appear-ance of the ¢rst rain forests. In parallel, Kleypaset al. (1999) proposed that rapid increase in mod-ern CO2 was a major threat to reefs because ofincreased solubility of aragonite in more acidoceans, leading to a drop in CaCO3 production.However, they noted also that such CO2 increaseswould favor calcite-secretors, or the growth ofnon-skeletal organisms. The largest reefs of thePhanerozoic occurred at a time of maximal Pha-nerozoic CO2 levels and calcite-dominated oceans,suggesting that at this time reefs, dominated bycoral calcite precipitators (rugosans and tabu-lates), were capable of tolerating high CO2, andthat globally warm shelf seas over-rode the factorof oceanic CO2 concentrations. The three mostimportant factors favoring the growth of moderncoral reefs, in order of importance, are temper-ature, light and carbonate saturation (Buddemeierand Fautin, 1996). The fact the Siluro-Devonianmetazoan reefs showed high-temperature equato-rial and high-latitude distribution, high growthrates similar to those seen in modern reef biotas(Gao and Copper, 1997), and shallow-water pref-erence, indicates that the same three factors were

PALAEO 2824 31-5-02


important in the Palaeozoic. There is little evi-dence that Mid-Palaeozoic reefs su¡ered duringgreenhouse conditions with much higher SSTsthan today, as seen in low diversity Panamianreefs on the west coast of the Americas (Glynn,1988; Glynn and D’Croz, 1990). Carbonate plat-form development and reefs in the Coral Sea, thenorthern extent of the Great Barrier reef, shrankby 50% when Late Miocene SSTs dropped toaverages of 18^20‡C (Isern et al., 1996), showingthat for modern reefs temperature is also theprime control. A parallel decline is seen in theLate Miocene tropical Porites^Tarbellastraea reefsof the semi-enclosed Mediterranean (Esteban,1996).

7. Conclusions

The Mid-Palaeozoic calcite-dominated coralreef ecosystem was adapted to tropical, carbonateplatform SSTs well above the Holocene intergla-cial icehouse norm. Calcite secretors are bu¡eredagainst excessively hot tropical shallow waters, incontrast to those which secrete aragonite, and canthus survive the ‘paradox’ of large reefs in a glob-al greenhouse (Kleypas et al., 1999). The alterna-tive to excessively high, or low tropical SSTs, is toretreat to small tropical oases (refugia), devolve askeleton, or to become extinct, leaving a gap inthe fossil record, as seen in the Famennian. Highequatorial temperatures leave the solution of ex-panding to higher latitudes (the Emsian^Give-tian); excessively low temperatures lead to vanish-ing habitats or extinction (the Famennian). Themain driving mechanism today for the productionof coral reefs and thick carbonate platforms at theequator, and up to 30‡N and S, is warm, tropicaltemperatures (Buddemeier and Fautin, 1996). Thedistribution pattern of tropical SSTs and coralreefs, as plotted by satellite data, and concomitantcarbonate saturation, is identical. The second fac-tor is light, therefore the requirement of shallowwaters for reef-dwelling symbionts that assisted inhigh rates of CaCO3 precipitation: Mid-Palaeozo-ic reefs favored shallow carbonate platforms. Thedistribution of Palaeozoic reefs matched and ex-ceeded these two main requirements, as tropical

temperatures were present at much higher lati-tudes than in the Cenozoic. The Mid-Palaeozoiccoral^sponge reef ecosystem evolved from theLate Ordovician into the Late Devonian, forover 100 Myr, and became strongly adapted tothe global greenhouse. Reefs survived short-livedglaciation events (6 1 Myr) in the ¢nal stage ofthe Ordovician, and repeated sealevel changes inthe Late Silurian and Early Devonian, but themetazoan reef ecosystem could not be sustainedin the 21 Myr long cool Famennian, the preludeto the Late Palaeozoic icehouse. Reef corals dis-appeared ¢rst, but even the aragonite stromatop-oroids did not survive the Late Famennian, de-spite a Late stromatoporoid scordatura in the¢nal CZ.The Mid-Palaeozoic reef database con¢rms the

basic global pattern that most metazoan reefs intropical low latitudes disappeared within or belowthe rhenana CZ, one CZ below the F/F boundary.Reefs ceased not in a single event, but duringprotracted stepdown episodes probably lastingmore than 1.0^1.5 Myr, with regional responsebeing variable. Relatively few reefs are known inthe latest Frasnian linguiformis CZ. There is nocatastrophic reef ‘kill horizon’ at the F/F bound-ary known anywhere in the world, with reefsoverlain by anomalous organic-rich facies carry-ing Pt/Ir signatures, or impactites (Grieve et al.,1995). The Late Devonian mass extinctions andreef decline rank second only behind the Permianfor their severity. The highly diverse Mid-Palaeo-zoic benthic reef consortium with its more than200 calcitic tabulate and rugose coral genera andsome 60 genera of aragonitic stromatoporoidswas eliminated, never to return in a signi¢cantway. The 21 Myr long Famennian saw no basicrecovery of the metazoan coral^sponge reef eco-system, except for rare and minor stromatoporoidor lithistid patch reefs: many carbonate sequencesare condensed, and Famennian carbonate produc-tion was only a fraction of that in the Frasnian.Stromatoporoids became completely extinct dur-ing the praesulcata CZ at the close of the Famen-nian, also marked by a regressive phase, as is theF/F boundary. Calcimicrobes, usually the samegroups that had been prominent from the EarlyCambrian onwards, were the dominant reef for-

PALAEO 2824 31-5-02


mers in the Famennian, persisting and expandingfrom the Frasnian without a break, often formingstromatolite-like mounds. True stromatolites ap-pear to be no more common in the immediatepost-extinction phase than before.Cooling climates and lowered sealevels, a¡ect-

ing both marine and terrestrial environments,oceans and atmospheres (Veevers and Powell,1987; Handford and Loucks, 1993; Isaacson,1997; Grader and Isaacson, 1997; Caplan andBustin, 1999), accompanied by rapid overturningof ocean water masses in the Late Devonian, nowappear to be favored by the bulk of the evidenceas the causes for the F/F reef demise and Famen-nian changes. They can best explain the 14 fea-tures cited above for reef decline, and faunal^£o-ral turnovers. Strong supporting evidence comesfrom Late Devonian temperature declines basedon Ca/Mg ratios (Yasamanov, 1981), Late Devo-nian glaciation in South America (Caputo, 1985;Martinez and Isaacson, 1996), sealevel curves(Johnson and Klapper, 1992), and rising NO18 sta-ble isotope ratios (Berner, 1999a). Are there com-parable climate and reef signatures in the Ceno-zoic? The Cenozoic shifted Earth into a broad‘icehouse’ climate mode in the latest Cretaceousand Early Palaeocene, following the Mesozoicgreenhouse (Berner, 1999a). With a warming in-terruption during the Early Eocene, progressivedecline, and then sharp cooling into the EarlyOligocene, warming in the Early and Middle Mio-cene, and then renewed cooling from the LateMiocene (Tortonian) into Plio-Pleistocene, equa-torial climates were strongly a¡ected (Diester-Haas and Zahn, 1996). Early and mid-Mioceneaverage SST temperatures and reef abundance ex-ceeded those of the Oligocene and Holocene, andreefs declined in the Late Miocene (Isern et al.,1996), accompanied by losses in coral diversity.Late Miocene reefs of the Mediterranean shiftedinto low diversity Porites, rhodalgal and stroma-tolite phases, as climates cooled (Esteban, 1996).Budd (2000) has shown that origination and ex-tinction rates of Cenozoic coral genera and spe-cies match these climate trends, and ‘‘the mostintense peak in generic extinction occurred duringthe Plio-Pleistocene, as climates deteriorated inresponse to the onset of northern hemisphere gla-

ciation’’. This de¢nes the closure of the isthmus ofPanama, kicking in the Gulf Stream. Thismatches the pattern seen in the Late Frasnianand Famennian with the ¢nal closure of the Rheicocean and formation of Pangea, except that gla-ciation was only in the southern hemisphericGondwana continent (Copper, 1984).The most reasonable explanation for the demise

of the Mid-Palaeozoic coral^sponge reef ecosys-tem appears to be the double blow of coolingclimates and major sealevel lowstands duringcooling and glacial episodes that dominated theFamennian, punctuated by interglacial high-stands. Mid-Devonian and Frasnian calcite reefsand reef corals appear to have been relatively tol-erant to high temperatures, as noted for somePersian Gulf corals today (Kinsman, 1968). Thehighest modern coral biodiversity today is in theSulu sea near Sulawesi, which also has the highestaverage seasurface temperatures of the modernocean (Linsley, 1996). High Frasnian coral toler-ance to raised SSTs may have made them moresusceptible to cooling events. Since many areaslack any evidence for F/F anoxic events, andFrasnian back and o¡-reef biotas were welladapted to hypoxia through their Devonian his-tory, multiple global hypoxic or anoxic ‘burps’seem unlikely causes for reef losses in shallowmarine, carbonate platform settings: in additionanoxia cannot account for terrestrial change-oversin biota at the F/F boundary.

Acknowledgements

This reef database is based on my own reefcompilations over the past two decades, withadditions from the Canadian Reef Inventory(CSPG 1998), the Russian reef databases (e.g.Sokolov, 1986; Belenitskaya and Zadoroshnaya,1990), and the PaleoReefs database of Erik Flu«gelat Erlangen University (vide Kiessling et al., 1999).The Natural Sciences and Engineering ResearchCouncil of Canada are thanked for long termsupport of field, museum, and library workworldwide, and the many colleagues who havekindly provided tips and clues into the buriedpaleo-reef literature are also thanked. Jim Sorauf

PALAEO 2824 31-5-02


and Wolfgang Kiessling are thanked for thoroughreviews that improved the manuscript.

Appendix 1. Appendix of Frasnian reef localities

For a more complete discussion of these reefsand their distribution and signi¢cance, consultCopper (2002).

Northwestern Europe (western Baltica plate)

UK: Devon, only Early Frasnian reef mud-mounds (Scrutton, 1977).Belgium: limited Early Frasnian reefs, maximal

mid-Frasnian, absent latest Frasnian, mostlymudmounds 30^150 m thick (Lecompte, 1936,1970; Tsien, 1976; Monty et al., 1982; Dreesenet al., 1985a,b, 1986; Monty et al., 1988; Boul-vain et al., 1988; De Jonghe and Mamet, 1988;Boulvain, 1993, 2001; Casier, 1987, 1992; Casierand Lethiers, 1997; Hilali et al., 1998). Reefs cov-ered by Late Frasnian Matagne shales.France: Boulonnais, Massif Armoricain, Mon-

tagne Noire, reefs and mudmounds absent (Elloy,1972; Mistiaen and Poncet, 1989; Bourrouilh andBourque, 1995).Germany: patch reefs, mudmounds, atolls (Jux,

1960, Krebs, 1967, 1971, 1974; Franke, 1973;Palme, 1977; Eder and Franke, 1982, 1983; Kasigand Wilder, 1983; May, 1987, 1995; Ziegler,1988; Weller, 1989; Wilder, 1989; Fuchs, 1990;Gischler et al., 1991; Gischler, 1992, 1995; Schind-ler, 1993). Reefs ceased rhenana CZ, karsted, con-densed cap.Poland: microbial mounds prominent, also

patch reefs, reef banks, mostly Early to mid-Fras-nian (Szulczewski, 1971; Szulczewski and Racki,1981; Racki et al., 1985, 1993; Racki, 1988, 1992,1998b; Narkiewicz, 1988; Narkiewicz and Ho¡-man, 1989; Racki and Balin‹ski, 1998; Racki andSobon‹-Podgo¤rska, 1992; Sandberg et al., 1992;Szulczewski et al., 1996). Reefs generally ceasedin lower rhenana zone, karsted, condensed cap,hiatus.

Southern and central Europe (terranes, tectonicplates)

Czech Republic: reef growth limited? or absent(Chlupac, 1988, 1998).Slovakia: general reef decline to Late Frasnian,

with reefs terminating earlier to north (Zukalovaand Skocek, 1979; Dvorak, 1980; Zukalova andChlupac, 1982; Dvorak, 1986; Hladil, 1986;Galle et al., 1995). Some Late Frasnian reefscapped by stromatolites.Sardinia: no Givetian^Frasnian reefs known

(Gnoli et al., 1981).Spain and French Pyrenees: reduced small

Frasnian patch reefs into linguiformis zone (Reij-ers, 1980; Van Loevezijn, 1987, 1989; Mendez-Badia and Soto, 1984; Van Loevezijn and Raven,1984; Joseph et al., 1980; Fernandez et al., 1996).Carnic Alps: reefs ended in rhenana (gigas) CZ,

F/F boundary phosphatic (Scho«nlaub, 1979,1998; Vai, 1963, 1967, 1998; Bandel, 1972).

Morocco^Algeria (northern Gondwana plate)

Reefs ceased by Late Givetian, but one possibleFrasnian patch reef site in Atlas Mts (Hollard,1967; Gendrot, 1973; Wendt, 1985, 1988; Corne¤eet al., 1990; Becker, 1993b).

Russia (eastern part of Baltica plate, RussianPlatform)

Pechora: thick reefs said to end at F/F bound-ary (Menner et al., 1996).Novaya Zemlya: deep-water carbonates, but no

reef facies (Andreeva et al., 1979).Vaigach: small patch reefs (Shuiskii, 1980).Beloruss^Donets graben: coral^stromatoporoid

patch reefs (Makhnach et al., 1986), Late Fras-nian only bryozoans and calcimicrobes.SW Russian Platform: small, 6 2 m thick cor-

al^stromatoporoid, ‘algal’ patch reefs (Sorokin,1978), emergent late Frasnian.W. slope Urals^Volga: mid- and Late Frasnian

patch, some barrier and atoll reefs, many micro-bial ; some areas only patch reefs (Ulmishek,

PALAEO 2824 31-5-02


1988; Bogoyavlenskaya and Lobanov, 1995; An-toshkina, 1997, 1998).Ural foredeeps: Semiluki coral^stromatoporoid

patch reefs (Ulmishek, 1988).S. Urals : Bashkir and Skhapova platforms with

thick patch, atoll and barrier reefs (Ulmishek,1988; Chuvashov and Shuiiski, 1990; Rusetskayaand Yaroshenko, 1990; Dronov, 1993).

Siberia

W. active margin Siberia: no Frasnian reefs,only Givetian and older (Stepanov, 1990; Bo-goyavlenskaya and Lobanov, 1995).Kuznetsk Basin: Early Frasnian rugose coral

patch reef, barrier reef complex, 80 and 150 kmlong (Belskaya, 1960; Ivanova, 1983; Krasnov etal., 1986).NE Russia with patch reef belts (Ioganson,

1990a,b,c; Ioganson and Bazanov, 1990; Belya-eva and Ioganson, 1990; Ganelinand Ioganson,1990; Khaiznikova, 1989).

Kazakhstan

SS Balkhash^Dzungar: reefs from Frasnian toTournaisian?, Early Frasnian patch reefs in vol-canic setting (Berg et al., 1980; Zadoroshnaya etal., 1990a).Caspian Basin: 400 km2 Tengiz Frasnian reef

atoll, extending into Visean (Pavlov et al., 1988).S. Tyan-Shan: mid-Frasnian stromatoporoid

patch reefs (Kim and Erina, 1984).

Afghanistan, Pakistan, Iran

Afghanistian: Early and mid-Frasnian fringingand platform coral^stromatoporoid reef tract,coral patch reefs 40^50 m thick; Late Frasnianbryozoan and ‘algal’ reefs (Brice, 1977; Mistiaen,1985).Pakistan: Givetian, but no Frasnian reefs (Gae-

tani, 1968).Iran: patch reefs and biostromes in Givetian

and Frasnian (Wendt et al., 1997); biostromes

and small? patch reefs from the jamieae Zone,ending in rhenana Zone near Isfahan, were re-placed by bryozoan and stromatolite communities(Huckriede et al., 1972; Dastanpour, 1996;Wendt et al., 1997; Mistiaen et al., 2000).

Laurentia

W. Canada sedimentary basin: 1500 km longgiant barrier, fringing and patch reef complexes,primarily in mid-Frasnian, with major declines inLate Frasnian (Davies, 1975; Burrowes andKrause, 1987; Norris et al., 1982; Moore, 1988;Reinson et al., 1993; Weissenberger, 1994; Ed-wards and Brown, 1999).N. Alberta, BC: deeper slope reefs (MacKenzie,

1967; Mountjoy and Riding, 1981; Pratt andWeissenberger, 1989; Hemphill et al., 1970).N. Alberta, BC subsurface, Rockies : reef-

rimmed platforms towards Peace river arch(McLaren, 1963; Andrichuk, 1958; Belyea,1960; Belyea and McLaren, 1962; McLaren andMountjoy, 1962; Mountjoy, 1965, 1980; McCa-mis and Gri⁄th, 1967; Langton and Chin, 1968;Mackenzie, 1969; Mountjoy and MacKenzie,1973; Coppold, 1976; Anderson and Machel,1989; Mountjoy, 1989; Fischbuch, 1968; Green-lee and Lehmann, 1993; McLean and Mountjoy,1993; Ville¤ger, 1996; McLean and Klapper, 1998;Whalen et al., 2000); Hay River area mid-Fras-nian patch reefs (Jamieson, 1971).Yukon-Mackenzie: during Frasnian deeper

water black limestones/shales (Morrow, 1999).NWT-NE British Columbia: mid- to Late Fras-

nian coral patch and platform reefs extendingsouth to Alberta, some with restricted micro-bial^stromatoporoid community (Belyea andMcLaren, 1962; Wendte et al., 1992; Norris,1985; Hedinger and Workum, 1989; McLeanand Klapper, 1998).Mexico, southern USA: Early, mid-Frasnian

coral patch reefs, serpulid reefs, Arizona (Stoya-now, 1936; Huddle and Dobrovolny, 1952; Tei-chert, 1965; Beus, 1980a,b), mid-Frasnian patchreefs Idaho (Isaacson and Dorobek, 1988; Isaac-son et al., 1989).Nevada, California: carbonate platform but no

PALAEO 2824 31-5-02


reefs (Warme and Sandberg, 1996), but possiblepatch reefs near Alamo Breccia (J.E. Sorauf, per-sonal communication; Johnson et al., 1991).W. Arctic, Banks Island: three early-mid-Fras-

nian reef levels, total ca. 150 m thick, 100 km longbarrier, patch and back-reef tract, corals^stroma-toporoids (Thorsteinsson and Tozer, 1962; Embryand Klovan, 1971, 1976; Miall, 1976; Embry,1991); covered by siliciclastics Late Frasnian.Central and eastern arctic : no Frasnian reefs,

carbonate platform smothered by giant siliciclas-tic delta complex from Greenland Caledonides(McLaren, 1963; Kerr, 1967; Trettin 1978,1991; Embry, 1991; De Freitas and Mayr, 1998).Central and E. North America: Hudson Bay,

Michigan through Missouri, no Frasnian reefs(Stumm, 1969).

Australia

Lennard Shelf^Canning Basin: passive margin,prograded Frasnian coral^stromatoporoid^calci-microbial reef tract in cyclical stages, especiallymid-Frasnian (Playford, 1980; Begg, 1987; Play-ford and Cockbain, 1989; Playford et al., 1989;Southgate et al., 1993; Brownlow et al., 1996;Wood, 1998, 1999, 2000a); top of Frasnianmarked by karst (Holmes and Christie-Blick,1993).Carnarvon Basin: patch reefs (Cockbain and

Playford, 1988).E. Australia: no post-Emsian reefs preserved

on active margin (Conaghan et al., 1976).

SE Asia: China, Vietnam, Thailand

Guizhou, Guangxi : coral^stromatoporoid^cal-cimicrobial patch reefs to atoll, reef-rimmedbanks, lagoonal facies with receptaculitid mounds(Yu and Wu, 1988; Bai et al., 1994; Yu and Shen,1998; Yu et al., 1999). Rhenana CZ marked bysealevel lowstand, linguiformis CZ sealevel high-stand (Muchez et al., 1996).Hunan: tabulate coral^stromatoporoid patch

reefs (Dai, 1982; Liu, 1986; Zeng et al., 1992;Liu and Yang, 1997).

Laos, Cambodia: no Frasnian reefs, but back-reef stromatoporoid facies present (Thanh et al.,1988).Vietnam: Famennian stromatoporoid patch

reefs in Vietnam (Nguyen and Mistiaen, 1998).North China: no reefs (potential reefs in NW

China, Dzungar Basin and Tyan-Shan ranges asextensions of Russian reef provinces) (Suetenko etal., 1977; Su, 1988).Thailand: Middle Devonian reefs, NE Thai-

land, none in Frasnian (Fontaine and Suteethorn,1994).

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Documents

famennian reefs

metazoan reefs

frasnianfamennian ff

carbonate platforms

reefs extinctions ocean

myr long famennian

end frasnian

lithistid sponge patch