Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65

www.elsevier.com/locate/palaeo

Reef development at the Frasnian/Famennian

mass 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 reef
episode had relatively low diversity compared to Middle Devonian highs (with over 200 genera of calcitic rugose and
tabulate corals). Following an initial early rise after Late Givetian coral and stromatoporoid extinctions, reefs
expanded 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 were
wiped out by the end Frasnian: some Famennian reefs, while partly retaining the structure of the underlying
carbonate 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 thamnasterioid
modules) 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 21
Myr long Famennian, also a time of very low platform carbonate production. Rare, small, isolated stromatoporoid
sponge, and lithistid sponge patch reefs returned episodically during the Famennian in North America, western
Europe, Australia and China: the aragonitic stromatoporoids became extinct at the end of the Famennian. During a
Late Devonian tectonically very active, collisional Caledonian mountain-building phase, oceanic and atmospheric
cooling, accompanied by sealevel lowstand systems, exposed most carbonate platforms, accelerating coastal erosion
and karsting. This increased the amount of clastics in the shelf-slope setting, in the last 1^3 Myr prior to the F/F
boundary, 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 link
between black, organic-rich horizons and reef demise at or close to the F/F boundary. The F/F boundary not also
marks the largest change from widespread flooded Early and Mid-Paleozoic continental cratons to narrow, distal
shelves, 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 tropical
coastal 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
* Fax: +1-705-673-6508.
E-mail address: pcopper@nickel.laurentian.ca (P. Copper). 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

PALAEO 2824 31-5-02

28 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
nerozoic acme of metazoan reef and carbonate
platform productivity (Copper, 1994; Kiessling
et 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 reefs
reached their maximum known geographic extent,
with barrier reef complexes far exceeding in size
those of today’s interglacial Holocene. Reefs were
periodically projected well into latitudes 45‡S and
possibly 60‡N, as seasurface temperatures were
10^15‡ higher than today’s interglacial 16‡C glob-
al average. Exceptional global warming was
coupled to atmospheric CO2 levels 14^24U great-
er than today, and more than double the CO2
levels of the warm Mesozoic (Berner, 1994,
1998, 1999b). High rates of weathering of terres-
trial Ca/Mg silicates in the presence of CO2 -rich
atmospheres ultimately meant a long term trans-
fer of atmospheric CO2 to oceanic-stored CaCO3
during the Early and Mid-Palaeozoic. Sealevel
highstands £ooded vast continental interiors,
and epicontinental seas greatly exceeded in size
their modern tropical analogues, the relatively
small South China, Java and Arafura seas. This
£ooding created ample accommodation space for
shallow-water, ‘skeleton framework reef facto-
ries’, and coral^stromatoporoid sponge carpets
and meadows, suitable for accelerated calcite
and aragonite production under warm tropical
climates (e.g. as seen for the mid-Frasnian, Fig.
1). Prime examples of such carbonate factories
existed on the areas fringing and covering Lau-
rentia during the mid-Devonian : more than half
of the present 10 million km2 land area of Canada
was covered in tropical seas at that time, as North
America 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 by
tabulate and rugose corals, and cyanobacterian
calcimicrobes possessing calcite skeletons, and
by the aragonitic stromatoporoid sponges. Oceans
were in a calcite mode, in contrast to the modern
Cenozoic 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-
PALAEO 2824 31-5-02
nite saturation system, triggered by rising O2 and
cooler temperatures. The Famennian^Tournaisian
was also marked by a major radiation of the sili-
ceous planktic radiolarians, identi¢ed by a rise
and 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 such
tree trunks are found preserved in mid-Frasnian
reefs and inter-reef strata of Banks Island (Embry
and Klovan, 1971), the ¢rst time this is seen in the
geologic record. Dramatic changes in spore sizes
of the global land £ora at the Frasnian/Famen-
nian (F/F) boundary are related to new strategies
for plant reproduction, e.g. seeds and pre-pollen
(Streel and Loboziak, 1995). These ¢rst rainfor-
ests a¡ected soil development and the storage of
carbon by soil microbiota, chemical rock weath-
ering, and coastal sediment transport (Wilder,
1989; Algeo et al., 1995; Berner, 1997). Fairly
stable levels of 15^20% O2 prevailed in the Cam-
brian through Mid-Ordovician, probably with O2
generated primarily by marine phytoplankton.
Levels of O2 then rose during the Late Ordovi-
cian, as CO2 declined (Berner, 1998), with the ¢rst
arrival of very simple small plant cover of bryo-
phytes (Retallack, 1985). Oxygen levels then shot
up dramatically to elevated levels, possibly near
35% 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 Late
Ordovician and Late Devonian oceanic cooling
episodes were marked by a reduction of CO2
and 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- and
salt-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-
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65 29

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 were
poorly developed during the ¢nal Frasnian linguiformis CZ. Famennian reefs were dominantly calcimicrobial consortia, with rare
stromatoporoid, or lithistid patch reefs, e.g. in the Early Famennian of the Bonaparte-Canning basins, western Canada carbonate
platform. Following repeated regressive cycles and cooling events, mid- to late Famennian carbonate platforms declined, followed
by transgressive intervals of isolated mudmounds to sponge patch reefs (e.g. Belgium, Poland small patch reefs in the marginifera
and 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).

rolepids all became extinct during the Frasnian,
with placoderms and lobe-¢nned ¢sh dying out in
the Famennian (Long, 1995). This is the largest
extinction and diversity loss of ¢sh known in the
Phanerozoic. The only survivors appear to have
been 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 slime
cocooning and burrowing. This strongly suggests
that coastal lowland, terrestrial habitats were
equally as much a¡ected as shallow marine trop-
ical successions in the Late Devonian, ruling out
marine anoxia as a prime cause for extinctions. In
turn, changing land conditions (e.g. the rise of the
¢rst forests, rise of oxygen via photosynthesis,

PALAEO 2824 31-5-02

30 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
reduced atmospheric CO2 , weathering and erosion
pro¢les) had a run-o¡ impact on neighboring
tropical marine systems.
The Siluro-Devonian coral^sponge reef ecosys-
tem had a coral biodiversity of more than 200
genera (Scrutton, 1998), 30% greater than that
of the scleractinian corals for the Cenozoic (Ve-
ron, 1995; Budd, 2000). Complementing this were
more than 60 genera of aragonitic stromatoporoid
sponges adapted to open, illuminated, fore- and
back-reef settings (Stearn et al., 1999), compared
to only a handful of primarily cryptic coralline
sponges today. Metazoans were accompanied by
diverse coralline algae and green algae. Frasnian
reefs were dominated by alveolitid and thamno-
porid tabulates, and giant phaceloid to aphroid
rugosan corals, but with a diversity much lower
than seen in the mid-Devonian (showing that
£ourishing reef growth was not necessarily corre-
lated with diversity). Many were capped by very
large, meter-sized, platy to domal stromatopor-
oids in the fore-reef and reef £at areas (Fager-
strom, 1994). Calcimicrobes played a signi¢cant
cementing, encrusting and cavity-¢lling role, as
from the Miocene to today (Riding et al., 1991;
Reid and Macintyre, 1992; Esteban, 1996). From
the close of the Frasnian, the metazoan, shallow
tropical reef ecosystem never fully recovered in
size or diversity in the Famennian, nor Late Pa-
laeozoic. For tropical marine biota, the Late De-
vonian was the second largest mass extinction in
the Phanerozoic, exceeding severity of Ordovician
events for the metazoan reef ecosystem (wiping
out more than 70% of the primary reef dwellers
and builders and shallow carbonate platform bio-
tas, as well as losing the Mid-Palaeozoic metazo-
an, coral^sponge reefs). Famennian microbial
reefs covered 6 10^20% of the shelf space seen
in the Frasnian.
F/F boundary events still remain controversial.
Controversy relates to (a) the timing of the events
within conodont zones (CZs), whether sudden
(Claeys and Casier, 1994), or staggered, gradual
or 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
PALAEO 2824 31-5-02
of these, and (c) whether only marine environ-
ments were a¡ected (House, 1985; Becker,
1993a,b), or whether both terrestrial and marine
systems were disturbed (Wilder, 1989; Algeo et
al., 1995; Copper, 1998). Debate also swirls
around 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 and
cycles, etc.). When examining causal e¡ects, the
question 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 or
cooling trigger anoxia?, nutrient £ux?, sealevel
change), or whether there were multi-causal or
cascading threshold factors (were sealevel draw-
down, cooling and anoxia inter-linked, and in
what order?). Can the same extinction factor
have multiple, opposing explanations? (e.g. was
anoxia due to stagnant, sluggish ‘hot’ oceans, or
the product of rapid overturn by sea surface tem-
perature (SST) cooling, bringing up deep CO2 -
rich waters and nutrients?). For those favoring
anoxia as a primary cause, were black shales or
black limestones the result of high oceanic surface
productivity under well-oxygenated bottoms, or
the result of bottom anoxia and burial factors?
(i.e. were black shales the cause, or the e¡ect, or
organic burial events unrelated to mass extinc-
tions?). Were pelagic events coupled to shelf
events, or not? Were atmospheric events linked
to oceanic events? Were some events simply
symptoms of change, or the real agents of mass
extinction, e.g. were sealevel lowstands just re-
£ecting glacial cooling events, or did eustatic sea-
level drain shelf areas, removing habitable space
and biotas (e.g. Johnson, 1974)? Was the F/F ex-
tinction due to anomalous nutrient (biogeochem-
ical) cycling and eutrophication (e.g. Murphy et
al., 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/F
boundary, at ca. 376 Ma (Tucker et al., 1998;
Okulitch, 1999), the oceanic crust that might
have given us distinctive platinum^iridium signa-
tures, has largely been lost by subduction. Despite
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
exhaustive search, no de¢nitive impact crater, nor
boundary clay horizon, nor iridium anomalies
have been detected at the F/F boundary, though
anomalies have been noted well above and below
the boundary in both shallow and deeper water
sections.
It is generally agreed that the equatorial coastal
marine ecosystems, particularly shallow-water
reefs, from interior £ooded cratons to shelf mar-
gins, were most strongly a¡ected by the F/F
events. During the Middle Devonian (Eifelian^
Givetian), metazoan reefs reached latitudes of
up to 60‡N (if the Mongolia plate is correctly
plotted), 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 reefal
to peri-reefal atrypid and pentamerid brachiopods
declined, with the loss of many families and even
a whole suborder (Copper, 1998; Godefroid and
Helsen, 1998). The terminal Frasnian extinctions
in these groups were less than the end-Givetian
biodiversity 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 their
greatest extent of the Phanerozoic, using four fac-
tors: (1) overall size of reef tracts (cumulative
thickness and areal extent), (2) pervasive occupa-
tion of widely £ooded, tropical and subtropical
continental interiors and platform margins to
high latitudes, (3) high coral and coralline sponge
diversity, with more than 200 genera of tabulate
and rugose corals (Scrutton, 1997, 1998), and 60
Devonian stromatoporoid genera (Stearn et al.,
1999) and (4) peripheral invasion by deep-water
slope, and high-latitude, cool shallow-water mud-
mounds. A number of mid-Devonian reef provin-
ces were at least three to four times longer, and
several times wider, than the modern Great Bar-
rier Reef (e.g. the Emsian^Givetian carbonate reef
platform extending from Nevada into Ellesmere
Island, some 5000 km along the northwestern
margin of Laurentia). In the Frasnian this reef
belt disappeared almost totally from the Cana-
da^Greenland arctic, shrinking by nearly 2000
km to only a 150 km belt along Banks island at
the western margin.
PALAEO 2824 31-5-02
31
The Late Devonian encompassed two phases of
reef evolution, a restricted Frasnian episode of
coral and stromatoporoid reefs, and a Famennian
episode of catastrophic metazoan reef collapse,
and replacement by microbial reefs. During the
Frasnian, the taxa that survived the severe coral
biodiversity losses of the Late Givetian (norrisi
CZ), probably triggered by a phase of cooling,
global sealevel drawdown, and exposure of the
shallow tropical reef setting to subaerial karst ero-
sion, had regrouped. Surviving tabulate corals
and colonial rugosans formed reef tracts which
were regionally extensive, such as in the western
Canada sedimentary basin and the Urals. In the
high 40^50‡S latitudes, for example NW Africa
(Morocco and Algeria), and even in lower latitude
areas such as the Montagne Noire and Sardinia,
reefs completely vanished during the Frasnian.
Early Frasnian reefs were usually smaller, with
peri-reefal settings of lower diversities: reefs in-
cluded more mudmound reef types than seen ear-
lier. By mid-Frasnian time, reefs had temporarily
recovered to a considerable extent in size and
areal extent, with cosmopolitan coral taxa,
though with reduced diversity compared to the
Givetian. Late Frasnian reefs were diminished in
abundance and geographic distribution, and these
appear to have become even more restricted to-
wards the F/F boundary, with few localities
known where reefs extended to the ¢nal extinction
events at the close of the linguiformis CZ (last
zone below the F/F boundary). Commonly, latest
Frasnian reefs were replaced by bryozoan bio-
stromes or stromatolitic units in the rhenana
through linguiformis zones. The Famennian was
marked by major, worldwide, sealevel drawdown
pulses, interrupted by rapid transgressions: meta-
zoan reefs were replaced by calcimicrobial, Renal-
cis-dominated reefs. Famennian coral reefs are
unknown, but a few patch reefs with stromatop-
oroid and lithistid sponges occurred regionally
(Fig. 1). Though some stromatoporoids survived
as 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 loss
of the Mid-Palaeozoic reef ecosystem from the
32 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
Frasnian through Famennian. To date no paper
has yet been devoted exclusively to looking at
what happened to reefs worldwide at the F/F
boundary. Problems remain with the lack of pre-
cise dating of reef terminations in a number of
areas. Since conodonts are scarce in reef settings,
the precise age of reef cores may not be fully
determinable, though the reef base and top may
be better dated. Data from the early 1990s sug-
gested that the Frasnian was a relatively lengthy
episode lasting ca. 10 Myr, with the Famennian
about 4^5 Myr long. More recently, Okulitch
(1999) has compiled radiometric data suggesting
the converse, a 5^6 Myr long Frasnian followed
by a much longer Famennian lasting 21 Myr. This
suggests that the end-Givetian through Frasnian
extinction and diversity losses in reefs lasted 6 6
Myr, but that the post-extinction phase was
nearly four times longer than previously sus-
pected. It also changes the general picture of
what happened to reefs, with a considerably
shorter period of Frasnian reef expansion, and a
much more protracted phase of reef absence and
recovery. In addition, the accumulation rates of
pre- and post-F/F carbonate platforms must be
re-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 the
mid-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, slightly
better than those of the Holocene. Because many
Frasnian reefs are petroleum reservoirs (e.g. in
western Canada and Timan^Pechora), these are
often better studied than Middle Devonian exam-
ples. Broadly summarized, there were three global
Frasnian reef trends, probably related to major
sealevel cycles plotted by Johnson and Klapper
(1992) and to climate change. Regressive Late
Givetian trends were succeeded by transgressive
PALAEO 2824 31-5-02
episodes with reef development controlled partly
by local tectonics, or limited by siliciclastic delta
development (Canadian arctic, Catskill delta of
the Appalachians), but with overall reef develop-
ment remarkably coordinated worldwide. Reefs
are 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 the
mid-Frasnian (MNCZ 5^11: approximately has-
si-lower rhenana CZ of others) saw maximal reef
development. The Late Frasnian (MNCZ 12^13:
approximately upper rhenana-linguiformis CZ of
others) saw the broad decline of reefs, with small-
er sizes and generally more restricted faunas, ter-
minating in regressions, or regional black shale
events. A typical Middle Frasnian highstand car-
bonate systems tract saw a broad passive margin
with barrier and platform reefs and isolated large
reef mounds up to several kilometers in diameter
(Fig. 2). The Frasnian interval represents the ‘dy-
ing’ episodes and close of the Middle Palaeozoic
metazoan reef ecosystem, with reefs losing more
than 50% of their tabulate and rugose coral ge-
neric diversity by the end Givetian. Only 39% of
the Devonian coral suborders and superfamilies
survived into the Carboniferous (Scrutton,
1997). The stromatoporoid sponges, the other ma-
jor reef builders, saw four orders surviving into
Famennian (Labechiida, Clathrodictyida, Stroma-
toporida and Amphiporida), and with one possi-
ble syringostromatid (Stearn, 1987). However, 10
genera extended into the Famennian (with seven
others more doubtfully ranging into the Famen-
nian: Stearn et al., 1999). Total diversity of the
stromatoporoids in the Middle Devonian was ca.
64 genera, with 38 recorded from the Frasnian
(including four new genera), so that extinction
of stromatoporoid genera by the end Givetian
was ca. 47%, and by the end Frasnian 74% of
the remaining genera were lost. There are no
post-Famennian stromatoporoids, with Mesozoic
spiculate forms probably independently derived.
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65 33

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 that
expanded carbonate platform margins seawards, and back-stepped reefs inland. Note the coral^stromatoporoid complexes that
dominated the reefs. O¡-reef settings featured organic-rich carbonates and shales. Coastal lowlands had the ¢rst pteridophyte
rainforests.

The Frasnian reefs of western Europe grew on
carbonate platform terrain that covered less than
a million km2 , but reef outcrops form only a
fraction of this southern border of the Old Red
continent, 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 this
belt, 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 the
classic sections around Frasne and Famenne (Le-
compte, 1936), limited Early Frasnian reef
growth, was followed by maximal expansion in
the 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 growth
in Belgium and the Aachen area of Germany.
Boulvain and Herbosch (1996) demonstrated
that Frasnian mudmounds of Belgium were in£u-
enced in their growth by bathymetry, as the mid-
Frasnian rimmed shelf changed to a ramp by the
Late Frasnian, signifying a slowdown in carbon-
ate production. Reefs disappeared in Belgium at
the top of the F2j level, roughly at the base of the
Matagne organic-rich shales, well below the F/F
boundary (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 Neuville
Fm., 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% loss
of benthic ostracodes at the F/F boundary. In
Germany, reefs were best developed east of the
Rhine, in part as shelf or ramp carbonates, and
in part as isolated atoll-like, commonly ‘mud-

PALAEO 2824 31-5-02

34 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
mound’ complexes in a tectonically active, vol-
canic belt of the Rhenish Schiefergebirge (Jux,
1960; Krebs, 1967, 1971, 1974). Rhenish reefs
were best developed in the Early and Middle
Frasnian, commonly over Givetian highs, but
metazoan reef and microbial mudmound growth
ceased in the Late Frasnian (Weller, 1989; Gisch-
ler, 1995). This has been related to regressive
cycles of siliciclastics in shelf areas, capping layers
marking restricted facies (shallow water?), or de-
position of hypoxic, lower Kellwasser black lime-
stones in the deeper water facies of the Rhenish
trough (Krebs, 1974; Fuchs, 1990).
In Poland, a 600 km long Frasnian carbonate
shelf, stretched along the eastern margin of the
Old Red Continent, is typi¢ed by extensive coral
and 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 mounds
appear to have locally become increasingly impor-
tant upwards in the Frasnian succession of
Poland (Racki, 1992). Reef growth halted in the
lower rhenana CZ, when condensation and
depositional hiatuses, with basin-wide facies re-
structuring, and local karsting took over. Synse-
dimentary tectonics complicated carbonate plat-
form foundering, and sealevel drawdown, with
shallowing sealevel signatures even in basinal
sections 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 and
central 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 the
former Yugoslavia were situated between Baltica
and Gondwana in south subequatorial latitudes.
Frasnian reef development was very limited
here or locally wiped out, or possibly con¢ned
to low diversity sponge and microbial mud-
mounds, with other areas showing small coral^
stromatoporoid patch reefs. In Moravia some
reef growth continued to the F/F boundary, while
in the Montagne Noire and Prague Basin Fras-
nian reefs were absent. Reef growth had ceased in
western North Africa (Morocco, Algeria) by the
end Givetian (Wendt, 1988), though possible ex-
PALAEO 2824 31-5-02
ceptions are noted in the Atlas Mountains (Gen-
drot, 1973).
Frasnian reef development was relatively vigo-
rous on the eastern side of the Russian Platform
or eastern Baltica (western slopes Urals from Vai-
gach Island in the north through the south Volga
region). Reefs are known both from outcrop and
subsurface, and many are petroleum-bearing
(Menner et al., 1996; Antoshkina, 1997). In
southern Timan, Frasnian stromatoporoid^coral
reefs (Sirakhoi, Vezhavozh, Sedyu, Bolshoii Ke-
ran reefal limestones) extended into the lower rhe-
nana CZ, but highest Frasnian linguiformis strata
are 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 Frasnian
barrier and patch reef complexes are located in
the Kuznetsk Basin (Belskaya, 1960; Ivanova,
1983). Stromatoporoid^coral reefs of the Glubo-
kaya complex, associated with solenoporids and
oncolites, range into the rhenana CZ, but are lack-
ing in higher strata (Rzhonsnitskaya et al., 1992).
The Kazakhstan Dzungar and Tyan-Shan had
small, lower Frasnian coral patch reefs, linked
to volcanic suites (Kim and Erina, 1984; Zado-
roshnaya et al., 1990a). Neighbouring Afghanistan
had a coral^stromatoporoid tract about 100 km
long (Mistiaen, 1985), and in Iran biostromes and
small coral^stromatoporoid patch reefs occurred
in the upper Givetian through mid-Frasnian to
the top of the jamieae CZ (Wendt et al., 1997;
Mistiaen et al., 2000). These identify the shelf
margin of a separate plate positioned adjacent
to 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 and
the Yukon); many of these formed as rimmed
platform and isolated reef complexes. Canadian
reef studies have placed emphasis on di¡erent as-
pects, e.g. oil and gas resources (Davies, 1975;
Reinson et al., 1993), tectonics, basement and
geography (Moore, 1988), platform-basin se-
quence stratigraphy (Whalen et al., 2000), and
coral distribution correlated with conodonts with-
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
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 reef
episodes are de¢ned for the early, middle and lat-
est Frasnian. Reefs particularly £ourished in the
Middle Frasnian, succeeded by major declines in
the Late Frasnian (CZ 12^13: Klapper, 1997).
During the Frasnian, reefs vanished completely
from the central and eastern arctic (Victoria
through Ellesmere islands, over 1500 km of shelf),
as a result of giant delta construction. In the west-
ern Canada sedimentary basin, Frasnian reefs
were developed as barriers, banks, platforms, lin-
ear trends, or large patch reef complexes many
kilometers in diameter, or as arcuate fringing reefs
around land areas. Deep seismic surveys indicate
that Upper Devonian reefs of Alberta partly in-
herited their regional distribution from underlying
Precambrian and Cambrian basement highs, but
some reef trends show no correlation with preced-
ing platform topography (Edwards and Brown,
1999). Some rare Frasnian deeper water, slope
reefs with renalcids or receptaculitid green algae
sat in troughs or embayments between carbonate
platforms : these were probably still within the
photic zone 6 100 m deep (MacKenzie, 1967;
Mountjoy and Riding, 1981; Pratt and Weissen-
berger, 1989). Carbonate reef mega-breccias, reef
olisthostromes and reef margin debris £ows also
reveal evidence of nearby shallower water reefs,
where adjacent reefs themselves are not evident
in 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 and
Northwest Territories (NWT). If distal outcrops
of carbonate platform with reefs are added, this
belt 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, from
the disparilis CZ through Frasnian CZ 4, a marine
PALAEO 2824 31-5-02
35
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 Lake
and Beaverhill carbonate platforms to central
north Alberta (Andrichuk, 1958), and fringing
reefs around the Peace River Arch to the north-
west. Many of these reefs were terminated at the
end of cycle 5 (within CZ 11, the rhenana zone).
Small reefs continued growth around the West
Pembina area (Nisku reefs) during more regres-
sive phases of succeeding Winterburn cycle 6 in
the 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 of
the Frasnian (Geldsetzer and Morrow, 1992), pro-
viding the source for petroleum trapped in the
reefs. These Frasnian black limestones, compara-
ble to the Givetian^Frasnian Domanik facies of
the 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 Banks
Island shows the northernmost Frasnian reefs in
Canada, developed as a ca. 150 m thick carbonate
barrier bank on the distal margins of a giant delta
complex s that stretched west from Greenland
(Thorsteinsson and Tozer, 1962). Reefs became
scarce in the uppermost Frasnian (CZ 13, lingui-
formis zone), largely disappearing with regressive
facies, or absent due to crustal £exing and deep-
ening into open marine facies in the west and
north (NWT and Yukon). Uppermost Frasnian
reefs of the District of Mackenzie and NE British
Columbia, belonging to the Kakisa Formation,
were open marine, small coral patch reefs with
an abundant and diverse rugose coral fauna, re-
markably the most diverse Frasnian coral fauna
in western Canada with some 32 species (McLean
and Klapper, 1998). Uppermost Frasnian rugose
coral patch reefs up to 30 m thick also outcrop in
the Rocky Mountains, and succeeding Famennian
strata shallow into intertidal facies.
36 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
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 Barrier
Reef came during the mid-Frasnian. Such reefs
featured a £ourishing coral^stromatoporoid and
calcimicrobial community (Wood, 1998, 1999).
Wood (2000a) suggested that the back-reef area
of the Canning Basin was dominated by a micro-
bial community that also included branching and
very large domed stromatoporoids, the latter up
to 5 m in diameter. Precise expansion and con-
traction of the various Australian barrier, atoll,
platform, patch and pinnacle reefs is not yet
known 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 Basin
in many respects, with the growth of coral^stro-
matoporoid-rimmed banks, barriers and patch
reefs curtailed by Famennian intertidal or re-
stricted marine facies.
3. The F/F boundary and succeeding Famennian
calcimicrobial reefs
The long Famennian time interval eliminated
coral^sponge reef habitats on a worldwide basis,
with a protracted ‘winter’ of condensed shelf car-
bonate productivity. The Famennian must have
been a 21 myr long period of continued stress in
the tropical carbonate setting. In terms of CaCO3
production (e.g. Timan^Pechora carbonate plat-
form), the total thickness for the Frasnian was
350 m, but only 160 m for the Famennian (Men-
ner et al., 1996), which indicates that carbonate
sediment production dropped from ca. 70 m per
Myr to 8 m per Myr after the F/F extinction, a
drop of nearly 90%. This di¡erence is comparable
to average tropical carbonate production on the
Great Barrier Reef versus the temperate cool-
water Great Australian Bight, and greater than
the decline of platform reefs in the Late Miocene
due to temperature drops (Isern et al., 1996).
Although other areas have not been compared,
PALAEO 2824 31-5-02
this suggests that at the F/F boundary, carbonate
production fell at rates as great as reef losses, and
that the systems were coupled (but contrast Kiess-
ling et al., 2000). Area available for carbonate
platforms also shrank as sealevel lowstands were
favored (Fig. 3). Dramatic global biodiversity de-
clines and extinctions, and absence of innovation
of new species of reef faunas for the Late Fras-
nian (McGhee, 1982) were the second factor at
work. 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- and
reef-building invertebrate phyla, and of equatorial
taxa in general, primarily the corals and stroma-
toporoid sponges. Famennian rugose corals were
mostly solitary, with only rare colonial forms
(Sorauf, 1989, 1992; Poty, 1999). Very few tabu-
late corals are prominent (Smirnova in Simakov
et al., 1983; Scrutton, 1998), and neither coral
groups were reef builders. Stromatoporoids were
stragglers that limped through the Early Famen-
nian, had a minor resurgence in the Late Famen-
nian Strunian substage, and disappeared by the
close of the Famennian (Smirnova in Simakov
et 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, low
diversity, Lazarus metazoans, such as labechiid
stromatoporoids and lithistid sponges. Most
were mudmound reefs, or occurred as microbial
reef caps in open marine, shelf settings. Though
local alpha diversity for lithistid sponges and cal-
cimicrobes in some reefs was moderate in the
Canning Basin (Wood, 2000b), this paled by com-
parison to similar reefs in the Frasnian, and corals
were absent in these reefs. The ‘crisis-progenitor
model’ of Kau¡man and Harries (1996), e.g. that
here 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¢ed
by calcimicrobes (Fig. 3), along with solenoporid
red 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-
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65 37

Fig. 3. Schematic setting for global, Early Famennian, lowstand setting, with calcimicrobial reef development, following the F/F
extinction 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.

crops were formerly dated as Carboniferous). Cal-
cimicrobes and algae radiated rapidly into many
new genera in a not surprising burst of evolution,
countering the pattern of extinction in nearly all
other 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, and
the loss of accommodation space due to global
regressive phases, and changing cooler climates
shifting 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 down
into 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 over
800 000 km2 , extending from the southern NWT,
through northern Alberta, over the Peace River
arch, 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 lowstands
began as early as the rhenana zone and persisted
through the F/F boundary in shallow shelf set-
tings: reefs were karsti¢ed, and developed ¢ssures
which were in¢lled. In the central and eastern
arctic, 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 Banks
Island to the far west (Goodbody, 1988; Embry,
1991). No Late Frasnian, nor Famennian reefs
were developed along a coastal lowland system
over 1500 km long, where reefs had once been

PALAEO 2824 31-5-02

38 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65

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 the
southern polar regions. Ocean currents were thus equatorially blocked, except by narrow straits, producing divergent warm-water
masses: 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-Famennian
cooling episodes, with data mainly derived from localities in Belgium and Poland, e.g. isolated build-ups from the marginifera
and postera-praesulcata zones. Vacated and karsted reef platforms were partially recovered by calcimicrobial consortia present
during 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.

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 of
former marine platforms, and renewed river inci-
sion inland. The F/F boundary here can be de-
¢ned only within terrestrial sandstone sequences
containing miospores and plant remains (Embry
and Klovan, 1976).
3.1. Western Europe (W. Baltica plate)
During the Famennian, the north European
platform was located in the 15^20‡S latitudes,
£anking the Old Red continent Laurussia (Fig.
4). In general, Europe featured a major regressive
megasequence in the Famennian, interrupted by
shorter, periodic transgressive events, triggered
by regional epeirogenic movements as seen in dis-
conformities, erosional gaps, evaporites, black
shales, extensive sandstones, and paleosols (Dree-
sen et al., 1985a, 1988). Coral^sponge reefs of the
Belgian Ardennes shelf progressively died out
within the rhenana CZ, below the linguiformis
CZ, and the F/F boundary (Dreesen et al.,
1985b). Black shales mark the terminal Famen-
nian Hangenberg event (praesulcata CZ), but
ironstones, ooids, and hardgrounds record ero-
sional gaps, and turbulent, cyclic oceanic events
throughout the Famennian (Dreesen et al., 1988).
Mid-Famennian mudmounds up to 100 m thick,
formed by crinoids, dasyclad green algae, and
sponges cemented by calcimicrobes, in an open
marine, marginal ramp setting, and possibly emer-

PALAEO 2824 31-5-02

 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
gent in later stages, are described from the mar-
ginifera CZ in Belgium and Germany (Kasig and
Wilder, 1983; Dreesen and Flajs, 1984; Dreesen
et al., 1985a, 1991; Dreesen, 1989). The ¢nal stro-
matoporoid patch reefs of Belgium occur within
the Etroeungt (Strunian) praesulcata CZ, directly
below the Carboniferous boundary (Dreesen,
1989). Stromatoporoids occurred more as thickets
or biostromes, such as in the Epinette and
Etroeungt formations of Belgium, rather than as
prominent reefs (Paproth et al., 1986). Rare small
coral patch reefs, 6 1^2 m thick, also occur in the
Strunian of Belgium (Poty, 1999).
During the latest Frasnian and Famennian,
some of the Frasnian reefs, e.g. the Iberg atoll
complex of Germany, built on a volcanic pile,
were eroded, karsted, pierced by neptunian dykes,
and depressions were in¢lled or covered with
black 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-water
limestones. The subsiding atoll thus became a sea-
mount, covered by deep-water coral banks during
the marginifera CZ, but emergent for most of the
remaining Famennian (Gischler, 1996). He also
remarked 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 the
western Paci¢c ocean today, part of the sinking
hotspot system that a¡ects seamounts as they ei-
ther become inactive volcanic piles, or move out
of the equatorial belt (Wheeler and Aharon, 1991;
Wilson et al., 1998). However, ancient platform
‘drowning’ (i.e. loss or reduction of carbonate
production) may be a direct response to cooling
climates, and have nothing to do with relative
sealevel 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 Frasnian
rhenana CZ, except in deeper water facies (Szulc-
zewski, 1986; Narkiewicz, 1988). Casier et al.
(2000) suggested that normal marine carbonate
PALAEO 2824 31-5-02
39
facies, under a sealevel drop of 10 m, at least
locally switched the platform to semi-restricted
back shoals and erosional surfaces at the F/F
boundary. 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 calcimicrobial
mounds, or small, shallow-water stromatoporoid
patch reefs were present in higher rhomboidea to
postera 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). In
the Prague basin, reefs had disappeared by the
beginning of the Frasnian (Zukalova and Skocek,
1979; Zukalova and Chlupac, 1982). For Mora-
via, carbonate production declined from the Early
Frasnian 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 earlier
in the north, forming emergent karst highs in the
Famennian, or replaced by shoaling foraminiferal
shelf facies, algal laminites and evaporites. Fa-
mennian basinal facies in Moravia were marked
by cherts with planktic radiolarians and benthic
sponges (Chlupac, 1988).
The stratotype F/F boundary is set in the Mon-
tagne Noire area of southern France, in deeper
water, ammonoid and conodont-bearing, pelagic
facies, lacking reefs or reefal mudmounds at the
boundary (Klapper et al., 1993). Reefs ceased here
in the Late Emsian. In the Montagne Noire, the
Famennian ostracode survivors were suggested to
have come from oxygen oases in shallow waters,
without reefs or mudmounds (Lethiers and Ca-
sier, 1994). Spain and the French Pyrenees were
devoid of reefs by the Famennian, as platforms
were converted to deeper water, black shale^lime-
stone facies. In the Carnic Alps, F/F boundary
strata are phosphatic (up to 90%), and no Famen-
nian mudmounds or reefs are known (Vai, 1963,
1967; Bandel, 1972). In Morocco, the Famennian
lacked metazoan reefs and mudmounds, with se-
quences dominated by siliciclastics, turbidites, or
40 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
erosional gaps (Gendrot, 1973; Wendt, 1988;
Benbouziane et al., 1993).
3.2. Central and eastern Baltica (including Russian
Platform, Urals)
In the Timan^Pechora region, stromatoporoid^
coral reefs ceased after the Frasnian and were
replaced by ‘algal bioherms and mud hills’ (Men-
ner et al., 1996). In the northern Pechora Urals a
diverse assemblage of calcimicrobes such as Re-
nalcis, Epiphyton, Izhella and Shuguria formed
framework in Famennian patch reefs, and were
accompanied by inter-reef or lagoonal facies con-
taining calcispheres (Belyaeva, 1986; Antoshkina,
1997). Some of these Famennian Pechora reefs
occur 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, from
drill-core data, were said to continue on tops of
highs created by underlying Frasnian reefs in the
southwestern Urals, and to be present in strata
possibly 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 to
shelf limestones at the F/F boundary (Ulmishek,
1988). In the Urals, some reefs were mudmounds
with an impoverished bryozoan, sponge or un-
known microbial contribution, and minor skeletal
remains, and in others, Famennian calcimicrobial
patch reefs with distinctive fabrics (Shuiskii,
1986). Such reefs were also said to continue into
the Early Tournaisian (Mirchink, 1974). Famen-
nian reefs disappeared from the shallow platform
to the west, and were con¢ned to calcimicrobial
build-ups along the new shelf edge margin, S and
E 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 et
al. (1986) ¢gured Early Famennian microbial (‘al-
gal’) reefs, from the Pripyat High or ‘Beloruss’
shelf, in a 250 km wide tract, covered by Late
PALAEO 2824 31-5-02
Famennian dolostones and evaporites. No Fa-
mennian black shales are reported as £ooding
this platform, but microbial (‘algal’) reefs were
repeated in the Carboniferous in the same, albeit
reduced, area of Beloruss and Ukraine (Makh-
nach et al., 1986). In the Precaspian region of
the SE Russian Platform, Famennian (and Tour-
naisian) microbial reefs, 6 10 m thick, are present
near 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 the
N margin of the Caspian Sea, 400^500 km E of the
Volga delta (NW Kazakhstan, Karaton), identify
subsurface reefal limestones of Famennian and
Early Carboniferous age. These are of unde-
scribed composition, in a succession some 80^90
m thick (Rusetskaya and Yaroshenko, 1990). In
the Caucasus to the south, a collisional continen-
tal margin facies suppressed Famennian carbon-
ates, which are frequently interrupted by erosional
disconformities, slumped units, and sandstones.
Much of the sequence is regressional, reef activity
having 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 (condensed
Famennian carbonate section = 50^100 m), they
extend discontinuously for about 150 km; Tour-
naisian build-ups are thicker. In the northern
Tyan-Shan Fold Belt (Karatau range, S. Kazakh-
stan), Famennian reef belts stretch about 500 km
SE towards the Akshirak^Moldotau range, west
of Lake Issuk Kul. These appear to be dolomi-
tized, Famennian rimmed, ‘algal’ reef complexes,
covering an area about 10 km2 but ‘algal’ textures
are diagenetic relicts (the Famenne section here is
about 500 m thick). The carbonates are near the
top of a 1^5 km thick, rapidly subsiding sequen-
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
ces of molasse and volcanics, probably derived
from an actively uplifting area during the Devo-
nian and Carboniferous (Zadoroshnaya et al.,
1990b). From Bolshoi Karatau, the same general
area, Cook et al. (1994) ¢gured Famennian
‘Waulsortian’-type reef mounds, dominated by
crinoids, bryozoans and algae, up to 100 m thick
and 2 km long. Tournaisian reefs continue this
north Tyan-Shan section. In the Caspian Basin
of southern Kazakhstan, seismic modeling has
discovered a very large, horseshoe-shaped reef
atoll (the Tengiz structure, 400 km2 ), initiated in
the Frasnian, interrupted by probable sealevel
lowstands but continuing into the Famennian
and up to the Visean (Pavlov et al., 1988). Be-
cause of its great depth at 4^5 km, the more
than 1 km thick Tengiz reefal units, with six
stratigraphic breaks, are not precisely dated, and
the nature of the Frasnian and Famennian com-
ponents of the reef are not known. In the south-
ern Tyan-Shan ranges of Zeravshan, Famennian
strata of the Yatavluk Fm., some 500^550 m
thick, contain dolomitized stromatolitic mounds
associated with lagoonal or peri-tidal micrites
(Kim et al., 1984). Such Famennian microbial
reefs may also be present in the Chinese Tyan-
Shan ranges, but remain to be discovered. No
Famennian reefs are known from Iran, Pakistan
or 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 in
the rhenana CZ by bryozoan and stromatolite
communities (Mistiaen et al., 2000).
3.4. Siberia plate
Lower Famennian calcimicrobial (‘algal’) reefs,
several meters thick, and locally associated with
bryozoans, superceded Frasnian coral reefs along
the 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 two
Famennian ‘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
PALAEO 2824 31-5-02
41
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 in
the Kuznetsk Basin, close to Kemerovo, stretch-
ing over a distance of nearly 100 km. Famennian
and Early Carboniferous ‘algal’ reef massifs also
occur in the Alai Ranges near Archaltur (Dronov
and Natalin, 1990). Yolkin et al. (1994) pointed
out that ‘powerful volcanism’ within the Altai
mountains, the fold belt £anking the Kuznetsk
Basin in the Late Devonian, may have disrupted
some of the reef development in the region.
3.5. Mongolia (Tuva), North China and NE
Russia plates
Famennian-age reefs are thus far unknown in
the far-eastern parts of Russia (Kolyma^Chukot)
and in Mongolia (Tuva). Though a major reef
belt nearly 2000 km long on the Mongolian plate
extended 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 microbial
reefs, 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 of
active tectonism, but it is also possible that the
whole continent had simply moved northwards
out of the tropical reef belt, which was already
at 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 shedding
no 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 metazoan
reef, or microbial mounds (Simakov et al., 1983).
Similarly, for the North China plate no Middle
Devonian nor Frasnian^Famennian reefs have
been described to date. These areas without reef
growth suggest possible drift of these plates out of
the tropics, or unsuitable conditions for reef
growth in the Famennian.
42 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
3.6. Laurentia
The western Canada sedimentary basin, which
had the world’s largest Frasnian reef tracts, lost
its metazoan coral^sponge reefs by the Famen-
nian. Calcimicrobial reefs are also not known,
possibly due to shallow to emergent conditions
over much of the carbonate shelf. In Alberta,
the weathered, karsted surface of the latest Fras-
nian Southesk Formation, e.g. at Jasper Park in
the central Rockies, is typically overlain by the
primarily intertidal dolomitic siltstones of the Fa-
mennian Sassenach Fm., with reefs ending in the
mid-Frasnian Mount Hawk Fm. (MacKenzie,
1969). Reinson et al. (1993) identify the F/F break
at the top of their cycle C6, the Winterburn
Group, which is succeeded by cycle C7, the Wa-
bamun Group. Stromatolites in the Wabamun
sediments suggest highly restrictive and stressed
conditions on the western side of Laurentia. The
F/F disconformity is followed in other areas by
the ‘Graminia Silt’, a regressive unit recording a
thickening siliciclastic wedge, the base of which
marks the F/F boundary locally (Reinson et al.,
1993). McLean and Klapper (1998), however,
dated the regressive lower Graminia Silt as latest
Frasnian : this suggests that uppermost marine
strata 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, are
small stromatoporoid patch reefs of mid- to late
Famennian age recorded from Alberta borehole
data (Stearn et al., 1987; Nishida, 1987; Stearn,
1988; Halim-Dihardja and Mountjoy, 1988), and
some small outcropping mounds along Humming-
bird Creek, probably in the crepida CZ (Stearn,
1987). In basinal sequences, a regressive event is
also recorded in western Canada (Van Buchem et
al., 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 N34 S values at the F/F
boundary, a feature not con¢rmed elsewhere.
However, this level also has icriodid conodonts,
indicating shallowing. In a 1200 km long belt
from Nevada to Montana, the terminal Famennian
(praesulcata CZ) is featured by vast beds of Girva-
PALAEO 2824 31-5-02
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 Late
Frasnian shale ¢ssure in¢lls containing brachio-
pods (Day, 1998). Frasnian reefs were absent in
eastern North America (Fagerstrom, 1983). The
Famennian in southern and south central Lauren-
tia (e.g. Michigan Basin, Ontario, New York,
Pennsylvania) was marked by shallow, hypoxic
to anoxic waters, pyrite and organic-rich, silici-
clastic mud deposition, or local deepening (Gut-
schick and Sandberg, 1991), partly as a response
of crustal £exing and the expansion of the Catskill
delta. Even bryozoan, or microbial mounds are
unknown. The Famennian black shales (the An-
trim, Kettle Point formations, etc.) were rich in
woody trunks, leaves, and fresh-water molluscs
and insects, deposited in delta-distant, coastal
lowlands under fresh water, or estuarine to partly
marine conditions. Widespread local marine black
shale horizons, precisely dated by conodonts and
bentonites, occur below, at and above the F/F
boundary in eastern North America, and thus
Kellwasser-type ‘anoxia’ here appears to mark lo-
cal sedimentary anoxic or hypertrophic events of
di¡erent ages (Over, 2000).
3.7. South China
Wang (1985) made no mention of the Chinese
Famennian as an unusual episode in reef history,
marked by regressions and loss of reef biotas.
South China is assumed to have been located
proximal to the Australian plate in the low lati-
tudes 6 30‡S of the equator (Fig. 4). In south
China, Famennian (Xiwangshanian) strata were
most commonly restricted intertidal to supratidal
facies, 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 margin
and slope, calcimicrobial reefs were dominated
virtually exclusively by massive Renalcis and Epi-
phyton (labeled as ‘algal’ mounds) on the platform
margins, such as those in Guangxi, eastern
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
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 thick
and ca. 50^180 m diameter calcimicrobial ‘Renal-
cis^Epiphyton’ mounds of the upper Yongshien
Fm. near Miaomen and Zhaijiang, west of Guilin
(Guangxi province) have their surfaces intersected
by sediment-in¢lled ¢ssures showing contempora-
neous erosion (Yu and Shen, 1998). Inter-reef
sediments with gastropods, ostracodes and forams
indicate a restricted subtidal to peri-tidal setting
for these reefs. The famous Baisa reef in Guangxi
is a Famennian calcimicrobial mound (Tsien et
al., 1988; Yu and Shen, 1998).
The South China calcimicrobial reefs replaced
stromatoporoid reefs of the Late Frasnian. Fa-
mennian stromatoporoid reefs are scarce, though
endemic stromatoporoids, including labechiids
were common to abundant, albeit not generally
reef-building. Rare, small and 6 5 m thick, upper
Famennian stromatoporoid patch reefs are re-
ported from the Dongcun Fm., at the village of
Etaoucun, 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, solitary
forms that appear to have been con¢ned to deeper
waters (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 deeper
water, o¡-reef sections of Maanshan in Guangxi
province, the top of the Frasnian is marked by a
brachiopod bed of ‘‘shell-beach facies under high
dynamic conditions [sic]’’, de¢ning a short shal-
lowing event directly below the boundary. There
is no record of any within-Famennian faunal ex-
tinction in forams or microbes, nor any sedimen-
tologic carbonate platform cessation in South
China, e.g. in the crepida CZ, as suggested by
Wang et al. (1994). Wang (1992) discovered glassy
microspherules, below a siderophile anomaly,
within the crepida CZ from the northern margin
of the South China carbonate platform, and sug-
gested Taihu Lake near Shanghai as the impact
site : neither reefs, nor diversity losses, are re-
ported in these strata. Muchez et al. (1996) com-
PALAEO 2824 31-5-02
43
pared the South China and Belgian sections of the
F/F boundary and suggested that the same eu-
static sealevel fall occurred in both, coinciding
with the base of the triangularis CZ. In Vietnam,
small stromatoporoid patch reefs are reported
from the latest Famennian (Strunian) by Nguyen
and Mistiaen (1998).
3.8. Australia
There is an erosional gap at the top of the
Frasnian, and possibly the earliest Famennian,
in much of the Canning Basin, which made the
shallow carbonate platform emergent (Playford,
1984; Cockbain and Playford, 1988; Playford et
al., 1989). Famennian reefs of the Nullara and
Windjana limestones were dominated by back-
stepping cycles with the cyanobacteria Renalcis,
Shuguria, Girvanella and Rothpletzella in shallow
waters less than 45 m deep on both the platform
and shallow slope (Playford and Cockbain, 1969;
Begg, 1987). Some calcimicrobial reefs were prob-
ably at the limit of the photic zone at 100 m, and
lithistid sponge reefs existed in still deeper waters
(Playford and Cockbain, 1969). Reef architecture
was largely inherited from the pre-existing car-
bonate platform, and modi¢ed by regressive
pulses. Neptunian fracturing and platform col-
lapse may be partly related to rapid sealevel £uc-
tuations at this time (Southgate et al., 1993). It
was suggested by Wood (2000b) that a local novel
reef 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 the
reef ecosystem had been fully re-established.
Nevertheless, corals played no role in the reefs,
as they did in the Frasnian, and stromatoporoids
were minor or absent.
The iron-concentrating cyanobacterium Frutex-
ites marks a hypoxic condensation surface, and is
found on some reef and reef slope surfaces within
the Famennian crepida CZ, at a level post-dating
the F/F extinction by possibly 0.5^1.5 Myr or
more. This con¢rms that the iridium anomaly is
not associated with any metazoan reef or faunal
extinction here, contradicting Wang et al. (1994).
No black shales are known from the carbonate
44 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
platform. Carbonate platform and calcimicrobial
reef growth seems to have persisted through the
Late 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-Famennian
platform margin collapse during a sealevel high-
stand of the Canning Basin, marked by dislodged
reef blocks, and this was suggested to be due to
over-steepening and/or tectonic activity. On the
tectonically active eastern margins of Australia
and New Zealand, no post-Emsian reefs are
known.
4. Temperate to cool Gondwana (South America,
southern Africa)
This region was located within the south polar
cold to cool temperate climate regime, as seen
from the dominance of siliciclastics, total absence
of carbonates, and declining abundance of marine
shelf 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 favositid
corals had vanished. Reefs and microbial mud-
mounds were completely absent in this realm
throughout 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 striated
pebbles, 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/F
boundary
Deep-water settings during the F/F boundary
PALAEO 2824 31-5-02
interval generally record continuous sedimenta-
tion, and little biotic disturbance of their benthic
faunas: the zonal stratigraphic scheme is exclu-
sively based on this environment. Sorauf and Ped-
der (1986) demonstrated that deep-water solitary
rugose coral faunas persisted in the Famennian,
and were almost undisturbed. There were virtually
no Famennian colonial rugosans, though some
returned in the Strunian (Poty, 1999). For the
ostracodes, although shallow-water taxa were
strongly a¡ected by F/F extinctions, deep-water
ostracodes were ‘almost untouched’ (Lethiers
and Casier, 1996, 1999a,b). Famennian deep-
water mudmounds appear to have been very
rare to absent, though the glass sponges and ra-
diolarians saw a radiation in North American and
European slope to basin sections (McGhee, 1982;
Racki, 1999; Racki and Cordey, 2000). The Mon-
tagne Noire section at Coumiac, with condensed
dark gray to black, carbonate^shale facies was
deposited 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, small
epiplanktic bivalves, palynomorphs, acritarchs,
chitinozoans, microforams, and pelagic to benthic
ostracodes 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 dropped
dramatically from about 10 000 elements per kg of
rock to less than 100 per kg, about 1 m below the
extinction 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 the
Late Frasnian pelagic system, as also seen in east-
ern North America and Brazil at this time. In
contrast, chitinozoan plankton were relatively
low in abundance in the Late Frasnian, but rose
to 19 000 specimens per g of rock in the Early
Famennian, 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 were
remnants of detritivores (egg cases, resting
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
stages?), feeding on the enriched planktic detritus
derived from the ¢nal collapse at the F/F bound-
ary (possibly a disaster biota). Paris et al. (1998)
suggest basal Famennian chitinozoan abundance
was due to a ‘‘temporary lowering of ocean tem-
peratureT and very low activity of the usual pred-
ators’’. Girard and Albarede (1996), who looked
at 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 ecosystem
collapse ?
The global nature of the ‘catastrophic’ F/F ex-
tinction was ¢rst suggested by Copper (1966), and
expanded by McLaren (1970), who proposed an
extra-terrestrial impact, associated with giant tsu-
namis. Most of the theories regarding the F/F
extinction have been succinctly surveyed by
McGhee (1996), but no papers have focused on
what happened globally to carbonate platforms
and reefs at the time. To determine the causes
of 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 two
CZs, 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/F
boundary, 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, and
the overlying Famennian is initially also usually
regressive, with the break usually marked by dis-
conformities, erosional boundaries and sharp
changes 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 and
Pedder, 1986; Oliver and Pedder, 1994).
(6) Deeper and shallower water stromatopor-
PALAEO 2824 31-5-02
45
oids declined markedly in the latest Frasnian, in
the rhenana CZ, below the F/F boundary (Stearn,
1975), and survived the F/F extinction, only to
perish 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 only
surviving 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 close
of 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 the
F/F boundary have been proposed (Copper,
1998; Racki, 1998a for brachiopod losses). Some
suggested global warming for reef decline
(Thompson and Newton, 1988; Brand, 1989).
Nutrient (phosphorus) poisoning was a theme in
other explanations (Schlager, 1981; Hallock and
Schlager, 1986; Wood, 1993; Eliuk, 1998). Theo-
ries for direct causes presently seem to revolve
mainly around two schools of thought: (1) one
or two cycles of worldwide oceanic anoxia asso-
ciated with the lower and upper Kellwasser lime-
stones (Walliser, 1984, 1996) and supported by
others (e.g. Joachimski and Buggisch, 1993;
May, 1995; Murphy et al., 2000), or (2) global
climatic cooling and sealevel drawdown, a¡ecting
46 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
both sea and land, ¢rst proposed by Copper
(1975, 1977). A corollary of these two e¡ects
might 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/R
cycles (Buggisch, 1991). The database for F/F
reefs is far from complete, and new discoveries
are 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 nektic
taxa in key sections or drill-core, leading to a
postulated gradual or stepdown interpretation,
when, in fact, all organisms may have died at
the 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 or
boreholes. They also represent an entire commun-
ity raised above the sea£oor, usually preserved as
massive, or thickly bedded stratigraphic units,
highly unlikely to erode completely in outcrop,
even if diagenetically altered. Thus the reef record
is as close to complete for any ecosystem, and
much more complete than the geologic record
for terrestrial rainforests.
6.1. Anoxia
How would anoxia, explained by water column
strati¢cation during transgressive pulses, force
reef demise and the loss of marine biotas on the
tropical, shallow shelves ? Brongersma-Sanders
(1966) suggested that anoxia could be created by
upwelling, of nutrient-rich and O2 -depleted waters
via strong winds, but considered this a local, not
global model. The main dilemma is how giant
‘burps’ of deep water from below the carbonate
compensation depth (CCD) could kill o¡ reefs on
a worldwide basis, and what would trigger such
burps? And do black shale events have the same
timing as reef demise ? Wignall and Hallam (1992)
and Wignall and Twitchett (1996) favored the
idea of anoxia as the prime killer for the end
Permian extinctions, though Hallam and Wignall
(1997) also added regression as a factor, perhaps
even over-riding anoxia as the main cause. The
end Permian mass extinctions coincide with well-
PALAEO 2824 31-5-02
dated £ood basalt events in NW Siberia, the larg-
est basalt traps of the Phanerozoic, and possible
sources for atmospheric CO2 or H2 S enrichment:
no such large scale end Devonian £ood basalts
are known at present. Vogt (1989) suggested in
a general model that volcanism drove anoxia
and reef death, but there is limited evidence for
this on the huge passive margin of Laurentia,
though bentonites are common on the eastern
side (Tucker et al., 1998). Kellwasser events
were related to transgressions by Walliser (1984)
and Joachimski et al. (2001). Black shales and
limestones have traditionally been viewed as
markers for high rates of carbon burial (hence
the development of suitable petroleum source
rocks), and strongly thermally layered, relatively
stagnant oceans with stable water column strati¢-
cation, e.g. a deep CCD. Black shales thus have
been regarded as proxies for transgressive sealevel
highstands and global warming. Nevertheless, this
is contradicted by new models for organic-rich
black shales, which include those of the Black
Sea, the western coast of Peru and Ecuador in
South America, the Gulf of California, the Nor-
wegian coast, and the west coast of Africa. Mod-
ern counterparts for black muds occur mostly in
temperate areas, and on the cool western shelves
of continents, marked by upwelling and mixing,
and high nutrient spill-over on the shallower shelf
or embayments. This led Pedersen and Calvert
(1990), Calvert and Pedersen (1992) and Calvert
et al. (1992) to suggest that organic-rich shales in
shallow and deep areas could best be explained by
high planktic productivity, stimulated by shelf
and slope upwelling, and cooling. Murphy et al.
(2000) suggested, to the contrary, that marine C,
N and P biogeochemical cycles were decoupled
from mass extinction : they preferred an anoxic
eutrophication model, tied to global cooling, for
the mass extinction.
How would reefs be a¡ected by global anoxia,
and is worldwide, simultaneous anoxia of the
tropical shallow shelf feasible? A major problem
with the anoxia hypothesis is that it is di⁄cult to
imagine how ‘giant megaburps’ of CO2 (and
SO2 )-enriched waters, brought up from below
the CCD, could simultaneously spill over all the
world’s tropical shelf areas, where reefs were
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
growing. No mechanism is known for this via
warming or transgression. At best, rapid oceanic
overturn seems possible only during major diver-
sions in ocean currents and deep sea water
masses, accompanied by cooling and loss of shelf
areas by regressions, as it did in the modern ice-
house during the Pleistocene. Most reefs today
appear to be reasonably well adapted to living
on the margins of ecosystems which face the
problem of being ‘shot in the back’ by a stagnant,
eutrophic lagoon or back-reef zone (Neumann
and Macintyre, 1985). Throughout the Middle
and Late Devonian, black shales and limestones
were widespread in inter-reef basins, and in back-
reef zones. Many patch reefs and their associated
corals such as thamnoporids, and the delicate,
stick-like amphiporid and stachyodid sponges,
and speci¢c brachiopods (leiorhynchids, atrypids,
productids, etc.) were adapted to oxygen-starved
settings. Indeed, they form a distinctive Eifelian
through 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 other
areas, there is no evidence for anoxia and black
shales or limestones at the F/F boundary, e.g. in
the Canning reef complex, yet the metazoan reefs
still died out (Becker and House, 1997). In South
China, black shales are also missing at the F/F
boundary, yet metazoan reefs gave way to calci-
microbial reefs (Yu and Shen, 1998). In Nevada,
black shale events pre-date the F/F boundary, and
are 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 (Embry
and Klovan, 1971; Embry, 1991). There is no
evident, direct relationship between black shale
horizons and reef disappearances in any sections,
as the timing of these do not coincide. There may
be no direct link between black shales and bottom
anoxia, except as indicators of high surface pro-
ductivity in an icehouse climate setting. Black
shale horizons are widespread in the Late Fras-
PALAEO 2824 31-5-02
47
nian, Famennian and Early Carboniferous, and
best tie in to widespread climatic cooling, and
vigorous 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 since
high nutrient input may stimulate the growth of
soft algal carpets, smothering reef metazoans.
Hallock and Schlager (1986) proposed a nutrient
kill model for reef demise, including the Late
Devonian. Quinn and Johnson (1996), and others,
however, have noted relatively strong tolerance
for some modern reefs in marginal settings to sea-
sonal high nutrient loading. What was the side
e¡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 low
diversity and low growth (Quinn and Johnson,
1996; Coles, 1997). Is there evidence for phos-
phorites or silica-producing biotas at the F/F
boundary, 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 siliceous
sponges 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 feature
elsewhere in the Devonian, particularly on passive
margin, stable carbonate platforms, nor in the
giant epicontinental seas of the Late Devonian.
6.2. Impacts
The impact origin for the F/F mass extinctions
was initially promulgated by McLaren (1970),
who favored instant mass killing and destruction
of ecosystems at the boundary by giant tsunamis.
No evidence has yet been found for the tsunami
concept. The impact theory has been stressed by a
number of other workers, but the geochemical
anomalies (Geldsetzer et al., 1987; Goodfellow
48 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
et al., 1988; McLaren and Goodfellow, 1990;
Nicoll and Playford, 1993; Wang et al., 1991,
1994, 1996), and spherules (Wang, 1992) occur
in the Famennian, well above the F/F boundary,
and mostly within the crepida CZ, or below the
F/F boundary in the rhenana CZ. Geochemical
anomalies within the two Frasnian Kellwasser ho-
rizons also suggest an internal oceanic source for
anomalous values of N13 C (Joachimski, 1997). A
mid-Famennian spherule horizon comes from
black shales of eastern Germany (Marini et al.,
1997). Belgian examples of ‘spherules’ at or below
the F/F boundary were contaminated by road
marker glass beads (Claeys et al., 1992, 1996).
Re-examination of the boundary stratotype in
the Montagne Noire for Pt/Ir has failed to con-
¢rm earlier anomalies suggested by Geldsetzer
(Girard et al., 1997). The only postulated Late
Devonian impact structure, the ‘Alamo Breccia’
(Warme and Sandberg, 1996), has been dated as
Early Frasnian, probably 3^4 Myr prior to the F/
F, and this had no known e¡ect on the carbonate
platform on which it landed in Nevada, nor on
global or regional diversity (J.E. Sorauf, personal
communication, regards these as slope-toe plat-
form breccias, as they occur along strike for
more than 150 km). Claeys and Casier (1994)
and Claeys et al. (1996) have suggested the 55-
km diameter Siljans crater in Sweden as a Late
Devonian impact site, but the crater has not been
reliably dated (McGhee, 1996), and no proximal^
distal e¡ects are identi¢ed. The Flynn crater of
eastern Laurentia is too small ( 6 5 km) to have
been e¡ective in extinction or reef demise. Wang
(1992) suggested Lake Taihu in China, near
Shanghai, as a possible Late Devonian impact
site, but the crater age is not con¢rmed. Thus
several possible impacts are known from the 26
Myr long Late Devonian, none of which had an
e¡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 reef
carbonate accretion is high in healthy reefs (Neu-
mann and Macintyre, 1985), and there is every
PALAEO 2824 31-5-02
indication that Devonian reef accretion rates
were 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 and
Walker (1992) suggested that reefs were net ex-
porters of CO2 to the atmosphere, because of
the CaCO3 equation, which dictates that CO2 is
released, as CaCO3 is precipitated : this should
theoretically enhance atmospheric CO2 and global
warming. 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 a
fossil record already in the Neoproterozoic. Ware
et al. (1992) concluded that modern reefs were net
sources of CO2 to the atmosphere, not sinks, even
with symbionts. But this is certainly not clear for
the Mid-Palaeozoic calcite greenhouse system
(Copper, 1997). The demise of metazoan reefs at
the F/F boundary coincides with a rapid rise of
O2 in the atmosphere, but it seems unlikely that
Late 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, Kleypas
et al. (1999) proposed that rapid increase in mod-
ern CO2 was a major threat to reefs because of
increased solubility of aragonite in more acid
oceans, leading to a drop in CaCO3 production.
However, they noted also that such CO2 increases
would favor calcite-secretors, or the growth of
non-skeletal organisms. The largest reefs of the
Phanerozoic occurred at a time of maximal Pha-
nerozoic CO2 levels and calcite-dominated oceans,
suggesting that at this time reefs, dominated by
coral calcite precipitators (rugosans and tabu-
lates), were capable of tolerating high CO2 , and
that globally warm shelf seas over-rode the factor
of oceanic CO2 concentrations. The three most
important factors favoring the growth of modern
coral reefs, in order of importance, are temper-
ature, light and carbonate saturation (Buddemeier
and Fautin, 1996). The fact the Siluro-Devonian
metazoan reefs showed high-temperature equato-
rial and high-latitude distribution, high growth
rates similar to those seen in modern reef biotas
(Gao and Copper, 1997), and shallow-water pref-
erence, indicates that the same three factors were
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
important in the Palaeozoic. There is little evi-
dence that Mid-Palaeozoic reefs su¡ered during
greenhouse conditions with much higher SSTs
than today, as seen in low diversity Panamian
reefs on the west coast of the Americas (Glynn,
1988; Glynn and D’Croz, 1990). Carbonate plat-
form development and reefs in the Coral Sea, the
northern extent of the Great Barrier reef, shrank
by 50% when Late Miocene SSTs dropped to
averages of 18^20‡C (Isern et al., 1996), showing
that for modern reefs temperature is also the
prime control. A parallel decline is seen in the
Late Miocene tropical Porites^Tarbellastraea reefs
of the semi-enclosed Mediterranean (Esteban,
1996).
7. Conclusions
The Mid-Palaeozoic calcite-dominated coral
reef ecosystem was adapted to tropical, carbonate
platform SSTs well above the Holocene intergla-
cial icehouse norm. Calcite secretors are bu¡ered
against excessively hot tropical shallow waters, in
contrast to those which secrete aragonite, and can
thus 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 to
retreat to small tropical oases (refugia), devolve a
skeleton, or to become extinct, leaving a gap in
the fossil record, as seen in the Famennian. High
equatorial 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). The
main driving mechanism today for the production
of coral reefs and thick carbonate platforms at the
equator, and up to 30‡N and S, is warm, tropical
temperatures (Buddemeier and Fautin, 1996). The
distribution pattern of tropical SSTs and coral
reefs, as plotted by satellite data, and concomitant
carbonate saturation, is identical. The second fac-
tor is light, therefore the requirement of shallow
waters for reef-dwelling symbionts that assisted in
high rates of CaCO3 precipitation : Mid-Palaeozo-
ic reefs favored shallow carbonate platforms. The
distribution of Palaeozoic reefs matched and ex-
ceeded these two main requirements, as tropical
PALAEO 2824 31-5-02
49
temperatures were present at much higher lati-
tudes than in the Cenozoic. The Mid-Palaeozoic
coral^sponge reef ecosystem evolved from the
Late Ordovician into the Late Devonian, for
over 100 Myr, and became strongly adapted to
the global greenhouse. Reefs survived short-lived
glaciation events ( 6 1 Myr) in the ¢nal stage of
the Ordovician, and repeated sealevel changes in
the Late Silurian and Early Devonian, but the
metazoan reef ecosystem could not be sustained
in the 21 Myr long cool Famennian, the prelude
to 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 in
tropical low latitudes disappeared within or below
the rhenana CZ, one CZ below the F/F boundary.
Reefs ceased not in a single event, but during
protracted stepdown episodes probably lasting
more than 1.0^1.5 Myr, with regional response
being variable. Relatively few reefs are known in
the latest Frasnian linguiformis CZ. There is no
catastrophic reef ‘kill horizon’ at the F/F bound-
ary known anywhere in the world, with reefs
overlain by anomalous organic-rich facies carry-
ing Pt/Ir signatures, or impactites (Grieve et al.,
1995). The Late Devonian mass extinctions and
reef decline rank second only behind the Permian
for their severity. The highly diverse Mid-Palaeo-
zoic benthic reef consortium with its more than
200 calcitic tabulate and rugose coral genera and
some 60 genera of aragonitic stromatoporoids
was eliminated, never to return in a signi¢cant
way. The 21 Myr long Famennian saw no basic
recovery of the metazoan coral^sponge reef eco-
system, except for rare and minor stromatoporoid
or lithistid patch reefs: many carbonate sequences
are 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 the
F/F boundary. Calcimicrobes, usually the same
groups that had been prominent from the Early
Cambrian onwards, were the dominant reef for-
50 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
mers in the Famennian, persisting and expanding
from the Frasnian without a break, often forming
stromatolite-like mounds. True stromatolites ap-
pear to be no more common in the immediate
post-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 and
Bustin, 1999), accompanied by rapid overturning
of ocean water masses in the Late Devonian, now
appear to be favored by the bulk of the evidence
as 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 comes
from Late Devonian temperature declines based
on 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 Cretaceous
and Early Palaeocene, following the Mesozoic
greenhouse (Berner, 1999a). With a warming in-
terruption during the Early Eocene, progressive
decline, and then sharp cooling into the Early
Oligocene, warming in the Early and Middle Mio-
cene, and then renewed cooling from the Late
Miocene (Tortonian) into Plio-Pleistocene, equa-
torial climates were strongly a¡ected (Diester-
Haas and Zahn, 1996). Early and mid-Miocene
average SST temperatures and reef abundance ex-
ceeded those of the Oligocene and Holocene, and
reefs declined in the Late Miocene (Isern et al.,
1996), accompanied by losses in coral diversity.
Late Miocene reefs of the Mediterranean shifted
into 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 most
intense peak in generic extinction occurred during
the Plio-Pleistocene, as climates deteriorated in
response to the onset of northern hemisphere gla-
PALAEO 2824 31-5-02
ciation’’. This de¢nes the closure of the isthmus of
Panama, kicking in the Gulf Stream. This
matches the pattern seen in the Late Frasnian
and Famennian with the ¢nal closure of the Rheic
ocean and formation of Pangea, except that gla-
ciation was only in the southern hemispheric
Gondwana 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 cooling
climates and major sealevel lowstands during
cooling and glacial episodes that dominated the
Famennian, punctuated by interglacial high-
stands. Mid-Devonian and Frasnian calcite reefs
and reef corals appear to have been relatively tol-
erant to high temperatures, as noted for some
Persian Gulf corals today (Kinsman, 1968). The
highest modern coral biodiversity today is in the
Sulu sea near Sulawesi, which also has the highest
average seasurface temperatures of the modern
ocean (Linsley, 1996). High Frasnian coral toler-
ance to raised SSTs may have made them more
susceptible to cooling events. Since many areas
lack any evidence for F/F anoxic events, and
Frasnian back and o¡-reef biotas were well
adapted to hypoxia through their Devonian his-
tory, multiple global hypoxic or anoxic ‘burps’
seem unlikely causes for reef losses in shallow
marine, carbonate platform settings: in addition
anoxia cannot account for terrestrial change-overs
in biota at the F/F boundary.
Acknowledgements
This reef database is based on my own reef
compilations over the past two decades, with
additions 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«gel
at Erlangen University (vide Kiessling et al., 1999).
The Natural Sciences and Engineering Research
Council of Canada are thanked for long term
support of field, museum, and library work
worldwide, and the many colleagues who have
kindly provided tips and clues into the buried
paleo-reef literature are also thanked. Jim Sorauf
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
and Wolfgang Kiessling are thanked for thorough
reviews that improved the manuscript.
Appendix 1. Appendix of Frasnian reef localities
For a more complete discussion of these reefs
and their distribution and signi¢cance, consult
Copper (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, mostly
mudmounds 30^150 m thick (Lecompte, 1936,
1970; Tsien, 1976; Monty et al., 1982; Dreesen
et al., 1985a,b, 1986; Monty et al., 1988; Boul-
vain et al., 1988; De Jonghe and Mamet, 1988;
Boulvain, 1993, 2001; Casier, 1987, 1992; Casier
and 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 and
Bourque, 1995).
Germany: patch reefs, mudmounds, atolls (Jux,
1960, Krebs, 1967, 1971, 1974; Franke, 1973;
Palme, 1977; Eder and Franke, 1982, 1983; Kasig
and 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 and
Sobon‹-Podgo¤rska, 1992; Sandberg et al., 1992;
Szulczewski et al., 1996). Reefs generally ceased
in lower rhenana zone, karsted, condensed cap,
hiatus.
PALAEO 2824 31-5-02
51
Southern and central Europe (terranes, tectonic
plates)
Czech Republic: reef growth limited ? or absent
(Chlupac, 1988, 1998).
Slovakia: general reef decline to Late Frasnian,
with reefs terminating earlier to north (Zukalova
and Skocek, 1979; Dvorak, 1980; Zukalova and
Chlupac, 1982; Dvorak, 1986; Hladil, 1986;
Galle et al., 1995). Some Late Frasnian reefs
capped 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 possible
Frasnian patch reef site in Atlas Mts (Hollard,
1967; Gendrot, 1973; Wendt, 1985, 1988; Corne¤e
et al., 1990; Becker, 1993b).
Russia (eastern part of Baltica plate, Russian
Platform)
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,
52 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
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; Rusetskaya
and 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 km
long (Belskaya, 1960; Ivanova, 1983; Krasnov et
al., 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 to
Tournaisian?, Early Frasnian patch reefs in vol-
canic setting (Berg et al., 1980; Zadoroshnaya et
al., 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 fringing
and platform coral^stromatoporoid reef tract,
coral patch reefs 40^50 m thick; Late Frasnian
bryozoan 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
PALAEO 2824 31-5-02
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 long
giant barrier, fringing and patch reef complexes,
primarily in mid-Frasnian, with major declines in
Late Frasnian (Davies, 1975; Burrowes and
Krause, 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 and
Weissenberger, 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 and
Mountjoy, 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 extending
south to Alberta, some with restricted micro-
bial^stromatoporoid community (Belyea and
McLaren, 1962; Wendte et al., 1992; Norris,
1985; Hedinger and Workum, 1989; McLean
and 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 patch
reefs Idaho (Isaacson and Dorobek, 1988; Isaac-
son et al., 1989).
Nevada, California: carbonate platform but no
 P. Copper / Palaeogeography, Palaeoclimatology, Palaeoecology 181 (2002) 27^65
reefs (Warme and Sandberg, 1996), but possible
patch 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 long
barrier, patch and back-reef tract, corals^stroma-
toporoids (Thorsteinsson and Tozer, 1962; Embry
and 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, especially
mid-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 Frasnian
marked 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-rimmed
banks, lagoonal facies with receptaculitid mounds
(Yu and Wu, 1988; Bai et al., 1994; Yu and Shen,
1998; Yu et al., 1999). Rhenana CZ marked by
sealevel 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).
PALAEO 2824 31-5-02
53
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 as
extensions of Russian reef provinces) (Suetenko et
al., 1977; Su, 1988).
Thailand : Middle Devonian reefs, NE Thai-
land, none in Frasnian (Fontaine and Suteethorn,
1994).
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