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Frasnian/Famennian mass extinction and cold-water oceans

Paul Copper Department of Geology, Laurentian University, Sudbury, Ontario P3E 2C6, Canada

ABSTRACT
Suturing of Laurussia and Africa, elimination of easterly tropical currents, and as well as elimination of earlier floras via
consequent equatorial diversion of cold and possibly dysaerobic waters are suggested as competition (Knoll, 1984). Nonmarine
prime causes of the Frasnian/Famennian (Late Devonian) extinction. These are explained by bivalves appeared in the Late Devonian,
using a new paleogeographic model. Evidence for stepdown extinctions that took place from though pulmonate land snails, aquatic and
Givetian through Frasnian time is taken particularly from atrypoid brachiopods, one of the terrestrial oligochaete worms, nematodes, ter-
most abundant representatives of the global tropical shelly marine benthos during the Middle restrial scorpions, millipedes, centipedes, and
and Late Devonian. Only 5 of 17 Frasnian genera or subgenera of atrypoids are known to be onychophorans followed later in the Early
present in late Frasnian sediments, and possibly only a single genus, Spinatrypa, was present Carboniferous (Rolfe, 1980).
at the Frasnian/Famennian boundary. Evidence for stepdown extinction from other Nearly all the evidence collected to date to
invertebrate and vertebrate groups in general corroborates that provided by brachiopods. support or deny a "sudden" mass extinction
has been collated at the family, superfamily,
INTRODUCTION the cool Malvinokaffric region from Givetian or ordinal level. Relatively little evidence has
At least four major extinctions appear to through Famennian time. been accumulated at the species or genus
have occurred in the early Phanerozoic interval The Famennian is generally characterized level, with the exception of comprehensive
from latest Precambrian through Devonian by relatively widespread regressive phases work on rugose corals (Pedder, 1982) and
time. The first of these extinction crises elimi- (Johnson et al., 1985), progradational coastal some brachiopod genera (Boucot, 1975). The
nated the Ediacaran fauna about 630 Ma (Seil- development of clastics (e.g., sandstone, silt- evidence has largely been muted because it is
acher, 1984). A second brought on the demise stone, mudstone, lagoonal to brackish water, standard practice to extend the known range
of the early micromorph invertebrate (Tommo- organic black shale), and a widespread ex- of Devonian genera to the end of the Fras-
tian) and archaeocyathid reef-building fauna at pansion of terrestrial nonmarine environ- nian or the end of the Givetian. The general
the end of the Early Cambrian (Lenian, ca. 530 ments that included fish, amphibians, terres- appearance of the fossil record, as a result,
Ma; Hill, 1972). A third was a major turnover trial invertebrates, and plants. Tropical reef has been to show an apparent catastrophic
of faunas within or toward the end of the Late habitats were extremely scarce, and reefs decline at the end of the Frasnian and an
Ordovician (Ashgill, ca. 445 Ma; review in were very small and patchy, impoverished or abrupt synchronous termination of many taxa
Brenchley and Newall, 1984). The fourth was depleted "mud-mound" faunas being known at a critical boundary. It leaves the impres-
the Late Devonian or Frasnian/Famennian from only a few places in the world. Famen- sion that one should find a bed full of Fras-
mass extinction about 360 Ma (McLaren, nian reefs are completely unknown in North nian fossils below the boundary, covered by a
1970; Copper, 1966,1977). For early Phanero- America. Where thin carbonate sequences of nearly barren bed of Famennian age, with a
zoic extinctions, the extinction event (or series Famennian age exist, such as in Poland, the "Carboniferous" fauna (McLaren, 1983). In
of events) occurred below the standard chron- Caucasus, and south China, there was a reap- complete, uninterrupted, marine Frasnian/
ostratigraphic boundary. For example, the pearance of stromatoporoids resembling Late Famennian boundary sections known to date,
Frasnian/Famennian extinction crisis occurred Ordovician taxa (Dong, 1964). In the Can- there are no known sharp faunal transitions
well before (possibly as much as 10 m.y. be- ning Basin of Australia, Famennian reef like this. The possible exception to this may
fore) the end of the Devonian. Similarly, Late niches were vacated by invertebrates and be in the Canning Basin, where the Frasnian/
Ordovician extinction(s) took place during reoccupied by skeletal, normal marine poro- Famennian boundary in the fore-reef and
Ashgillian time (below Conodont Zone 13, the stromatolites such as Sphaerocodium, and pos- reef-flat environment is drawn at the appear-
Gamachignathus zone), probably at the end of sible encrusting calcareous forams (Playford, ance of porostromatolites (e.g., Sphaeroco-
the Richmondian, and not at the end of the 1976). The Famennian marine faunas were dium), bacterial Frutexites, and possible en-
Ordovician. The archaeocyathid fauna became unlike their earlier Devonian counterparts in crusting foraminiferal horizons (Playford,
extinct at the end of the Early Cambrian, and the Givetian and Frasnian; they had a decid- 1976; Playford et al., 1984). However, there
the Ediacaran extinction occurred well before edly "Carboniferous" aspect; i.e., represented are no indications at this horizon of mass
the beginning of the Tommotian, the first stage by families and orders of invertebrates com- mortality of a thriving reef community, nor is
of the Cambrian. mon in the Carboniferous and rare or absent there a marked unconformity.
in the Devonian.
FRASNIAN/FAMENNIAN SETTING The Frasnian/Famennian was a "noisy" ATRYPOID BRACHIOPODS
The Late Devonian lasted about 21 m.y., time interval, both biologically and tectoni- One group commonly cited to have under-
the Frasnian taking about 9.7 m.y. and cally. The earliest tetrapods, the amphibians, gone "catastrophic world-wide extinction at the
the Famennian about 11.2 m.y. (Ziegler, appeared on land and in shallow marine end of Frasnian time" is the atrypoid brachio-
1978). The Frasnian was a period of cos- coastal or estuarine regions (Young, 1981; pods (Copper, 1966), one of the most abundant
mopolitanism for nearly all marine taxa; Turner and Tarling, 1982). Insects and spi- benthic marine components in the Frasnian.
areas today as far apart as eastern and west- ders showed their first radiation (Rolfe, Plotting their extinction at the family level
ern North America, western Europe, and the 1980), probably in response to the first de- shows that the Palaferellidae disappeared in late
northern Urals show a common marine shelf velopment of soils, forests, arborescent trees Givetian time, and Lissatrypidae and Atrypidae
fauna. There was widespread marine trans- with trunks, and the evolutionary introduc- at the end of Frasnian time (Fig. 1, left). This
gression at this time and widespread reef tion of seed habits (Chaloner and Sheerin, thereby corroborates the picture of extinction at
development in many areas. In contrast, no 1979; Edwards, 1980). There was rapid floral the close of the Frasnian as compiled by Sep-
shelly or coral marine faunas are known from biomass expansion of terrestrial communities, koski (1982). However, recalculation of these

GEOLOGY, v. 14, p. 835-839, October 1986 835

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data at the subfamily level changes this pic- Biernat (1970). A check of these records shows appear to occur near the top of the succession
ture—two subfamilies disappear during middle that none of these specimens were colic cted in but are extremely rare (Deh6e, 1929; Drever-
Givetian time, two at the end of the Givetian, situ from rocks of definitive Famenniar. or mann, 1901) . In North America, a small Spina-
and three at or near the end of the Frasnian younger age. Some include reworked s< ali- trypa is the youngest occurrence in the Missis-
(Fig. 1, right). It is clear that the finer the taxo- ments; other strata have been redated as Fras- sippi Valley (Weller, 1906), and in New
nomic refinement of data, the more accurate nian or older, or the specimens have been lost. Mexico, Spinatrypa compacta occurs near the
the picture becomes (cf. Fig. 2). There are thus far also no records of atiypoid top of the Contadero Formation of late Fras-
Were there any holdover Famennian atry- brachiopods occurring immediately below nian age (Cooper and Dutro, 1982). In Iowa,
poids? This is not clearly established at present. (i.e., within 3 m) the Frasnian/Famennian the Owen Member of the Lime Creek Forma-
There are only isolated reports of Famennian boundary. tion in the uppermost Frasnian has abundant
or Early Carboniferous atrypoids. These in- What were the youngest known atrypoids lowatrypa and rare Pseudoatrypa, but Spina-
clude reports by Herrick (1887), Drevermann from the latest Frasnian rocks? In northwest trypa is missing here, though it is abundant in
(1901), Weller (1906), Mansuy (1912), Dehee Europe, both a small Spinatrypa and the the members directly below. In the Soviet
(1929), Veevers (1959), Gunia (1968), arid smooth atrypoid "Glassia " [= Cryptatrypal] Union, Spinatrypina [= ? Spinatrypa] and
Pseudoatrypa are the youngest known genera
(Lyashenko, 1959; Rzhonsnitskaya, 1975).
family subfamily Thus, contrary to general views, the genus
Atrypa did not become extinct at the end of the
Frasnian (McLaren, 1983), but well before the
end of the Frasnian. The subgenus Atrypa (Kyr-
I 1 tatrypa), whose last occurrences were in Austra-
Figure 1. Middle and Late lia (Grey, 1978), became extinct before the end
Devonian range chart for I 5 of the Frasnian, within the lower Palmatolepis
families and subfamilies
of atrypoid brachiopods
gigas conodont zone (Fig. 2). Elsewhere, this
showing that significance subgenus is unknown higher in the stratigraphic
of disappearance of atry- column except in Givetian rocks; thus, Austra-
poids in Late Devonian is lia may have represented an anomalous refu-
masked by taxonomic
gium for the genus Atrypa in Frasnian time.
data.
= e- i
= | ! S & What do the ranges of other atrypoid genera
within the late Middle Devonian (Givetian)
I 1 through Frasnian tell us? Precise stratigraphic
!m m•
o a. information at the species level is available for
few Upper Devonian sections at present (except
Norris and Uyeno, 1981; Norris, 1985; Lya-
shenko, 1959; Balinski, 1979). The conven-
tional practice of extending the range of a
species through the complete thickness of a
stratigraphic unit precludes a bed-by-bed as-
signment of precise occurrences. Genus range
charts show that the disappearance of genera
was not a terminal Frasnian event, but was
phased out from the Givetian through late
Frasnian in stepdown fashion.
§ £ f «a
~ Atrypoid brachiopods were tropical to sub-
tropical, carbonate to clastic shelf inhabitants of
up
£ Devonian seas (Copper, 1966). They were ab-
sent in the Southern Hemisphere, cold climate,
Malvinokaffric faunal province during the
Devonian (Copper, 1977). The decline in these
« brachiopods is; paralleled by similar Givetian
i I
through Frasnian progressive extinction rates in
other groups (Fig. 3), with the possible excep-
up
tion of the rugose corals, which appear to dis-
play a different pattern, if all taxa, not just
hermatypic genera, are taken into account
(Pedder, 1982). No data are available on the
survival of colonial vs. solitary rugosans in the
Figure 2. Provisional genus and subgenus range chart of atrypoid brachiopods during Mliddle
Givetian through Famennian. The successful,
and Late Devonian; showing stepdown extinction patterns. For Frasnian, informal terms "mid-
dle," "lower," and "upper" have been based on conodont zones using brachiopod abundance surviving, and new Famennian rugose coral
data: lower Frasnian designates lower to upper Polygnathus asymmetricus Zones, middle genera were usually solitary, deep-water types
Frasnian for Ancyrognathus triangularis through Lower Palmatolepis gigas Zones, and upper (Sorauf, 1984, oral commun.): a sharp extinc-
Frasnian for Upper gigas through Middle Palmatolepis triangularis Zones. Atrypoid brachio- tion apparently occurs at the Frasnian/Famen-
pods had mostly disappeared by end of gigas Zone. Exact distribution of atrypoids within
zones is approximated because of present lack of precise stratigraphic information. Absolute
nian boundary in the rugose corals. This tends
age dates are extrapolated from Ziegler (1978). to support McGhee's (1982) observation that

836 GEOLOGY, October 1986

 Downloaded from geology.gsapubs.org on January 4, 2015
the cold, deep-water, hexactinellid glass
sponges did particularly well in the Famennian;
they were uncommon in the preceding Fras-
nian. The stromatoporoid record is ambiguous
and not well documented at present, and it
shows an apparent rise in the Frasnian and a
sharp decline in the Famennian.
Of the 12 groups shown in Figure 3, 7 had
greater extinctions at the end of the Givetian
than during the Frasnian. Ammonoids and
phyllocarids declined progressively through the
Frasnian; the Givetian extinction was less se-
vere. Information on tabulate corals is incom-
plete because there is no documentation of last
appearances in the Frasnian and Famennian
(Hill, 1981).
POSSIBLE CAUSES FOR THE LATE
DEVONIAN EXTINCTIONS
The Frasnian/Famennian was a geologically
active period. It was the time of the Antler oro-
geny in the western United States, the Ellesmer-
ian orogeny in northeastern Canada and Green-
land, and late phases of the Acadian orogeny
in central and northwestern Europe (closing
of the Scottish Highlands, translational faulting;
Ziegler, 1982). Strong sinistral displacement
occurred between the adjacent continents of
Laurentia and Baltica. Extensive volcanism,
folding, and faulting occurred in the central Eu-
ropean fold belt (Germany, Poland, Czechoslo-
vakia), and there was local gentle folding and
faulting in northwest Gondwana (South Amer-
Figure 3. Extinction rate chart of major Devonian macroflora and invertebrate groups calcu-
lated on basis of numbers of species of phyllocarids; genera of macroflora, nautiloids, ammo-
noids, gastropods, stromatoporoids, rugosans, and tabulates; subfamilies of trilobites; and fam-
ilies of ostracodes and echinoderms. Bryozoans are based on percentage of taxa that became
extinct. Data taken from House et al. (1979). Numbers shown are number of taxa that became
extinct per time unit.
837 GEOLOGY, October 1986
ica and North Africa). There was, however, a
marked downturn in the granitic emplacement
cycle (Engel and Engel in Fischer, 1981).
Devonian eustatic fluctuations were particu-
larly cyclical and more sharply defined in the
Givetian through Famennian time intervals
(Lenz, 1982; House, 1983; Sutton and
McGhee, 1985). The Famennian was charac-
terized by widespread regressive phases (John-
son et al., 1985), periods of nondeposition,
and/or periods of progradational coastal devel-
opment with silts, sands, and conglomerates.
Restricted marine to lagoonal, brackish, or
stagnant-water and nearshore, organic-rich,
black mud environments with plants and an
intermittent, impoverished, species-poor, re-
stricted benthic marine fauna were widespread
in North America. They characterize the New
Albany and Chattanooga black shales of the
north-central and east-central United States and
the Antrim, Kettle Point, and Long Rapids
black shales of the Michigan Basin, southern
Ontario, and the Hudson Bay region to the
north. Eroded reef tops are recorded in post-
Frasnian outcrops or subcrops of western Can-
ada. Localized Frasnian disconformities and
karst development are present in the midconti-
nent region of the United States and in Bel-
gium, Germany, and the Canning Basin of Aus-
tralia. Sandstones containing plant remains, fish
scales, and the primitive nonvascular plant Pro-
tosalvinia were common in the Famennian suc-
cessions of the Amazon, Parnaiba, and Parana
basins of Brazil (Caputo and Crowell, 1985).
Sandstones and conglomerates are also a con-
spicuous component in many tropical parts of
the world, frequently overlying normal marine
carbonate or shale successions of Frasnian age.
In both deep basinal and shallow marine shelf
areas of Europe (e.g., the Montagne Noire in
France, and in Czechoslovakia), localized
phosphates were developed, a feature virtually
unknown in the Middle Devonian and Fras-
nian and possibly associated with cold waters
upwelling onto carbonate platforms in equator-
ial belts (Zukalova and Skocek, 1979).
Middle Devonian and probably early Fras-
nian climates are recognized to have been ex-
ceptionally warm in equatorial belts (Fischer,
1981). Reef habitats were widespread, and
large barrier reef tracts more than twice the
length of the Great Barrier Reef of today are
known to have existed in the Old World faunal
province (Copper, 1974). Global sea levels
were generally high, and transgression was
widespread. No terrestrial glacial deposits are
known in Middle Devonian through Frasnian
time in the polar regions.
At the close of the Devonian a shift from the
global greenhouse climatic mode to the global
icehouse of the Carboniferous has been sug-
gested (Fischer and Arthur, 1977; Fischer,
1984). A Famennian ice age is documented
in Brazil by the presence of diamictites with
striated pebbles, dropstones, rhythmites, erra-
tics, striated pavements, and ice-pushed sedi-
ments (Caputo, 1985; Caputo and Crowell,
1985).
At present, four ideas generally prevail as
immediate or direct explanations for the Late
Devonian extinction phase: (1) asteroid or bo-
lide impact (e.g., McLaren, 1970,1983); (2)
climatic change—e.g., oceanic cooling (Copper,
1977; Zukalova and Skocek, 1979; Stanley,
1984) or glaciation (Caputo and Crowell,
1985); (3) oceanic carbon dioxide venting
(Fischer and Arthur, 1977; Berry and Wilde,
1978; Wilde and Berry, 1984); and (4) rapid
sea-level fluctuations, including either drowning
(House, 1983) or regression (Johnson et al.,
1985), or a cycle of both.
Little attention has thus far been directly
paid to global paleogeography and its climatic
implications, specifically during the Frasnian/
Famennian intervals. Though several recon-
structions are available for the Late Devonian
in general (Bambach et al., 1980), or the Fras-
nian (Boucot and Gray, 1983; Young 1981;
Turner and Tarling, 1982), only one Famen-
nian reconstruction appears to have been at-
tempted (Scotese, 1984). The last contains two
radically different options, one with a fused
Pangea, the other with a 2000-km-wide south
equatorial ocean separating Laurussia and
Gondwana; paleomagnetic data are ambiguous.
Boucot and Gray's (1983) reconstruction, per-
haps the most convincing because it draws on
 Downloaded from geology.gsapubs.org on January 4, 2015
wide-ranging sedimentological and stratigraphic
data, shows a South African south pole and a
Southern Hemisphere location for all of North
America and Eurasia. However, it is impossible
to crowd so much continental plate mass into
the Southern Hemisphere unless the Devonian
Earth were about 30% larger. North America
must straddle the equator or be located north of
the equator, as shown by Scotese (1984). Bou-
cot and Gray (1983) also showed a single fused
Eurasian plate, a model that few others have
favored because of substantial evidence for
major Paleozoic and Mesozoic suturing in
China, southeast Asia, Iran, Turkey, and Ka-
zakhstan. Newer plate reconstructions of China
show that the north and south China block;,
may have been part of Gondwana and that
substantial clockwise rotation of Gondwana.
may have occurred during the Paleozoic (Lin
et al., 1985). Most specialists working on the
Pangea model for the Permian-Carboniferous
have paid little attention to the generally more
widely accepted archipelagic model for the
Devonian, and vice versa.
It is generally agreed that during the Devo-
nian, the South American, African, Indian,
Antarctic, and Australian plates formed a uni-
fied region known as Gondwana. The Malvin-
okaffric marine shelf faunas of this area are
spread over a wide area, and there is some mix-
ing at the northern and eastern boundaries.
There is also general agreement that North
America, Greenland, Spitsbergen, and north-
western Europe represented a unified land area
(Laurussia) fringed by common continental
shelves in the Late Devonian. There is no
agreement where the Siberia, Kazakhstan,
Figure 4. Tentative model ol Frasnian and Famennian paleogeography based on marine in-
vertebrate distribution. Cosmopolitanism in Frasnian time suggests open marine connec-
tions from east to west. During Famennian time, rare reef formation occurred only on east-
ern side of sutured Gondwana-Laurussia, though there were sporadic connections between
eastern North America and western Europe. Onset of Famennian glaciation, accompanied
by sea-level drawdown and large-scale regressions, may have introduced cold waters into
tropical shelf areas.
838
China, or Sunda segments were dispersed.
Vertebrate paleontologists, looking at Devonian
fresh- and brackish-water terrestrial fishes and
primitive amphibians, suggest that collision
of Laurussia and northwestern Gondwana
(i.e., North Africa) occurred during the Lite
Devonian "at or near the Frasnian-Famennian
boundary" (Young, 1981; Turner and Tabling,
1982). This appears to make sense not only for
vertebrate distribution but also to account for
Frasnian marine cosmopolitanism and Famen-
nian restrictions and extinctions of benthic
marine faunas.
A Frasnian/Famennian collision of Laurus-
sia and Gondwana would create a large conti-
nental plate straddling the equator (Fig. 4).
Easterly equatorial currents would be diverted
and would thus bring cold waters into equator-
ial belts along the western margins of conti-
nents. The westerly drift of the southern
subpolar gyre would spill cold and poorlj
oxygenated waters over warmer, subtropical
shelves of Laurussia. Restricted circulation
would produce euxinic basins on the flanlcs of
the land areas adjacent to the paleo-Tethys
(e.g., west of the Appalachians), though these
shelves would remain warmer because of the
impingement of the eastwardly flowing tropical
currents. This could account for the unusual
presence of small Famennian and earliest Car-
boniferous mud-mound reefs that have relict
faunas in Poland, the Caucasus, and China, and
of the relict reef province in the Canning Basin
of Australia. Recovery of a fairly cosmopolitan
Famennian brachiopod shelf fauna of rhyncho-
nellids, spiriferids, athyrids, and productids, all
with distinct "Carboniferous" taxonomic affini-
ties, may have l)een the result of periodic flood-
ing of continental areas and rapid recoloniza-
tion of vacant niches. Possibly this was due to
episodic raised sea-level intervals from inter-
glacial cycles melting ice caps covering South
America at the time (Caputo and Crowell,
1985). However, Famennian time would, over-
all, remain a period of cool oceans and cool
shelf habitats.
CONCLUSIONS
1. Atrypoid brachiopod distribution during
Frasnian time suggests a progressive stepdown
extinction of the tropical marine shelly fauna,
final disappearance being below or near the
Frasnian/Famennian boundary. Most atrypoids
appear to have vanished by the end of the gigas
conodont zone on a worldwide basis. The last
surviving genera were Spinatrypa and, possibly,
the smooth atrypoid "Cryptatrypa. "
2. Tropical marine carbonate shelf organ-
isms were more strongly affected by the Frasni-
an/Famennian extinction than those that lived
in cool or deeper waters or in clastic environ-
ments. Small, solitary, deep-water rugose corals
and cold, deep-water siliceous sponges survived
the extinction, possibly representing "Lazarus"
taxa that repopulated Famennian shelf habitats.
3. Late Devonian terrestrial vertebrates were
not strongly affected by the Frasnian/Famen-
nian extinction; indeed, some groups show di-
versification at this time. There is no evidence
of any mass extinction of land plants, which
were diversifying, evolving into new directions
and becoming increasingly abundant.
4. Late Devonian climatic change, via oce-
anic cooling and overturning of the water mass,
may have been triggered by closure of the
ocean between Laurussia and Gondwana,
which resulted in the diversion of ocean cur-
rents at the equator. Collision may have further
exaggerated mountain building and increased
land elevation in the Appalachian-Caledonian-
Hercynian mountains. Thus, paleogeographic
configuration may have stimulated seasonal
extremes that, in turn, initiated glaciation in
South America. Extinctions were exacerbated
by major downdrops in sea level, which in-
itiated a cycle of regressions at the Frasnian/
Famennian boundary.
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ACKNOWLEDGMENTS
I thank Steve Stanley, J. G. Johnson, and an
anonymous reviewer for their comments on an ear-
lier version of this manuscript. Research was sup-
ported by the Natural Sciences and Engineering Re-
search Council of Canada. This paper was based on a
talk presented at the Geological Society of America
Northeastern Section Meeting in Providence, Rhode
Island, in March 1984.
Manuscript received March 17, 1986
Revised manuscript received June 11, 1986
Manuscript accepted June 23, 1986
839
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Geology

Frasnian/Famennian mass extinction and cold-water oceans

Paul Copper

Geology 1986;14;835-839
doi: 10.1130/0091-7613(1986)14<835:FMEACO>2.0.CO;2

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