ISSN 0001-4370, Oceanology, 2016, Vol. 56, No. 6, pp. 863–875. © Pleiades Publishing, Inc., 2016.

Original Russian Text © M.S. Barash, 2016, published in Okeanologiya, 2016, Vol. 56, No. 6, pp. 946–958.

MARINE GEOLOGY

Causes of the Great Mass Extinction of Marine Organisms

in the Late Devonian
M. S. Barash
Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia
e-mail: barashms@yandex.ru
Received June 9, 2015

Abstract―The second of the five great mass extinctions of the Phanerozoic occurred in the Late Devonian.
The number of species decreased by 70–82%. Major crises occurred at the Frasnian–Famennian and Devo-
nian–Carboniferous boundary. The lithological and geochemical compositions of sediments, volcanic
deposits, impactites, carbon and oxygen isotope ratios, evidence of climate variability, and sea level changes
reflect the processes that led the critical conditions. Critical intervals are marked by layers of black shales,
which were deposited in euxinic or anoxic environments. These conditions were the main direct causes of the
extinctions. The Late Devonian mass extinction was determined by a combination of impact events and
extensive volcanism. They produced similar effects: emissions of harmful chemical compounds and aerosols
to cause greenhouse warming; darkening of the atmosphere, which prevented photosynthesis; and stagnation
of oceans and development of anoxia. Food chains collapsed and biological productivity decreased. As a
result, all vital processes were disturbed and a large portion of the biota became extinct.

DOI: 10.1134/S0001437016050015

The Devonian Period (419.2 ± 3.2‒358.9 ±
0.4 Ma ago) (from here on, the age of stratigraphic
boundaries after [30]) was marked by the second of
five Phanerozoic great mass extinctions, when over
75% of organisms became extinct in relatively short
time intervals. For the Late Devonian, the reduction
in marine organism diversity is estimated as 13‒38%
at the family level, 55‒60% at the generic level, and
70‒82% at the species level [42]. The terrestrial com-
munity suffered as well: 43‒53% of plant taxa disap-
peared. Most extinctions of individual taxa of marine
organisms in the Late Devonian correspond to crises
near the Frasnian–Famennian boundary (FFB, 372.2
Ma ago) and the terminal Devonian near its boundary
with Carboniferous Period (DCB, 358.9 Ma ago)
(Fig. 1). The first of these crises (FFB) was stronger.
The latter is termed as the great mass extinction.
During the FFB crisis, 33 families of Devonian
brachiopods, which represented a diverse and abun-
dant constituent of benthic communities, became
extinct; representatives of 30 of them were tropical
dwellers. The extinction was most significant among
brachiopods of the middle and outer parts of the shelf.
In the Devonian, bryozoans represented a thriving
group that numbered over 1000 species. They became
extinct prior to the extinction of most other organisms,
i.e., near the boundary between the Givetian and
Frasnian stages: 43% of their families, 64% of genera,
and 69% of species disappeared at that time [28]. Like
brachiopods, bryozoans survived the DCB extinction
event. In the Late Devonian, 31% of bivalves also
became extinct.
Rugose corals disappeared almost completely. At
the FFB, 97% of their shallow-water and 60% of deep-
water species disappeared [48]. Tabulate corals, which
populated reefs, lost 80% of their genera during this
crisis, although they survived the crisis at the Devo-
nian‒Carboniferous transition. Stromatoporoidea
were main reef-building organisms in the Late Paleo-
zoic forming the largest buildups in the Frasnian Age.
Nevertheless, 50% of their families disappeared by the
end of this age and role of main reef-builders passed to
calcareous algae. Among Stromatoporoidea, only
benthic communities composed of cold-resistant taxa
continued existing. The end of the Frasnian Age was
marked by extinction of their last six families. Reef cal-
careous algae also disappeared at that time.
For trilobites, the extinction at the end of the
Famennian Age was catastrophic. Ostracods during
the FFB crisis experienced strong ecological stress,
which is evident from the disappearance of most spe-
cies (70%). This is particularly true of detritovore
ostracod taxa, while species capable of living in oxy-
gen-deficient environments and planktonic forms suf-
fered less. Ammonoids were also subjected to mass
extinction: of 26 families, only one survived [29]. In
the Late Devonian, the crisis involved all Chordata
representatives except for sharks: the diversity of cono-
donts was drastically reduced, 9 Agnatha families dis-
appeared, as well as 63 of 70 Arthrodira genera among

863
864 BARASH

Pelagic organisms
Givetian Frasnian Famennian Tournaisian

Devonian Carboniferous
Agnatans

Placoderms

Actinopterygians

Ammonoids

Conodonts

Cricoconarids

Ostracoda

Benthic organisms

Brachiopods

Bryozoans

Foraminifera
?
Rugose corals
?

Stromatoporoids

Trilobites

Ostracods

Fig. 1. Variations in diversity of pelagic and benthic organisms in Late Devonian‒initial Carboniferous, after [27]; age of stage
boundaries, after [30].

Placoderms. Freshwater Placoderms taxa overcame
the crisis with less significant losses.
Paleontological evidence of the crises is synchro-
nous with lithological, geochemical, and isotopic
indicators of environmental changes (see review in
[46]). The corresponding stratigraphic levels are
marked by black shales, volcanogenic deposits,
impactites, carbon and oxygen isotopic excursions,
and features reflecting climate changes, sea level fluc-
tuations, and other events. The main reasons for
extinction are considered tectonic movements, fluctu-

OCEANOLOGY Vol. 56 No. 6 2016

 CAUSES OF THE GREAT MASS EXTINCTION OF MARINE ORGANISMS
Panthalassa
Ocean
La
Rheic Ocean
Fig. 2. Late Devonian paleogeography, 370 Ma ago (after [53], modified). Plates: (Sib) Siberian, (La) Laurentia, (Bal) Baltica,
(Gond) Gondwana.
ations in climate and sea level, volcanism, the collision
of Earth with cosmic bodies (impact events). These
abiotic factors influenced the biota of the ocean
through the hydrological structure and hydrothermal
composition of water: its evolution, distribution areas
and sizes of organisms, biodiversity, and bioproductiv-
ity. Unfavorable combinations of these factors resulted
in the mass extinction of organisms.
The mass extinction occurred against the back-
ground of Gondwana’s position near the South Pole
and development of the main Acadian‒Eovariscan
fold belt (Fig. 2). Injurious environmental distur-
bances could be determined by intensified silicate
weathering on elevated land domains and decreased
greenhouse CO2 concentrations in the atmosphere.
They could have also resulted from intense develop-
ment of terrestrial vegetation and mass burial of
organic carbon. It is believed that primary production
increased due to the intensified influx of nutrients
from continents.
The repeated global development of black shale
facies, enrichment of sediments with trace elements,
increase in δ34S of pyrite, positive shifts of δ13C, and
dominant development of benthic species tolerant to
oxygen deficiency in water represent the characteristic
features of the extremely unfavorable conditions that
existed during the last two Late Devonian extinctions
(FFB and CDB). Two models are used to explain the
elevated burial of organic carbon in sediments: better
preservation of organic matter in stagnant oceans and
increase in primary production, which resulted in its
intensified influx to sediments. The latter model
implies increased transport of biogenic elements from
land during regressions. According to calculations,
OCEANOLOGY Vol. 56 No. 6 2016
865
Prototethys
Sib Ocean
Paleotethys
Ocean
Bal
Gond
this process could represent the main factor responsi-
ble for the formation of two +3‰ δ13C excursions in
the late Famennian Age. The main causes were evi-
dently drops in sea level [13]. In the Late Devonian,
black shales enriched in organic matter were deposited
due to the high influx of organic matter into sediments
and favorable oxygen-deficient conditions for its pres-
ervation at the water‒bottom interface.
The main impulse of extinction in Upper Devonian
sections near the FBB is termed the Kellwasser Event,
which is reflected in the presence of two black shale
members. Both Kellwasser members have been
recorded in Laurentia, Gondwana, Siberia, and
southern China. Organic-molecular indicators from
marine sedimentary sequences of Belgium in the FFB
interval point to a wide distribution of forest fires, soil
erosion, and oceanic hypoxia, which was responsible
for the mass extinction of marine organisms. The
magnetic susceptibility data imply the prevalence of an
arid climate, which stimulated forest fires [33].
According to isotopic investigations of sulfur, the
FFB interval in southern China and Poland is charac-
terized by frequent disturbances in its cycle [15]. The
authors explain this phenomenon by sea level and cli-
mate fluctuations, which were responsible for changes
in the nutrient influx from continents, primary pro-
duction volumes, and organic matter burial. The
increase in nutrient influx during the a drop in sea
level could significantly increase bacterial sulfate
reduction. During a rapid rise sea level at the begin-
ning of the Famennian Age, this was followed by the
formation of long-term euxinic environments. The
temporal coincidence of two extinction phases with
geochemical reorganizations and euxinization of the
866 BARASH
photic zone confirms the assumption that these events
played an important role in the biotic crisis.
The Kellwasser Event affected mostly shallow-
water thermophilic organisms (systems of coral reefs,
brachiopods, trilobites, ammonites, conodonts,
acritarchs, etc.). The FFB interval in two significantly
remote areas (Belgium and China) exhibits positive
shifts and high δ13Ccarb values. Some organic matter
components near the FBB in both sections demon-
strate similar peaks. These data indicate the global dis-
tribution of anoxic or euxinic environments, which
should have affected the biota [57].
The difference in the geochemical characteristics
of two Kellwasser black shale members in the Harz
Mountains (Germany) gave grounds to assume of
their different depositional environments [51]. The
formation of the lower member resulted from an
increased influx of nutrients from land, increased pri-
mary production, and development of anoxia on sub-
merged platforms. The upper member formed due to
the development of oxygen-deficient water in deep
parts of the ocean from intermittent stratification of
the water column during the maximum of the Fras-
nian transgression. Oxygen-deficient water could have
spread over shoals during this transgression.
An insufficient oxygen content in water was not
widely considered responsible for the disappearance of
reefs due to unavailability of data on the shallowest
Kellwasser facies. The intermittent occurrence of
anoxic conditions on the very shallow-water carbonate
platform during the Upper Kellwasser Event was
proved by a geochemical study in Alberta, Canada
[12]. In the Late Devonian, this area was located in the
equatorial zone. Transgressions were accompanied
there by distribution of euxinic or anoxic waters,
which is evident from the enrichment of sediments in
organic carbon and uranium.
In China, the Upper Devonian Hongguleleng For-
mation includes the Upper Kellwasser Member [14].
Unlike previous researchers who argued that this event
was characteristic of shallow-water marginal epiconti-
nental basins of the closing Rheic Ocean, the authors
of [14] show that the latter is also reflected on the
shoals of the open part of Paleotethys Ocean.
The terminal extinction pulse recorded immedi-
ately below the DCB is also reflected in the lithology
as Hangenberg black shales, which have been docu-
mented in Europe, North Africa, North America, and
China [45]. Their accumulation coincided with the
mass extinction and glacioeustatic sea level fluctua-
tions. The accurate U‒Pb zircon dates obtained for
volcanic ash beds which bound this black shale mem-
ber made it possible to specify the age of this event
(from 358.97 ± 0.11 to 358.89 ± 0.20 Ma ago) and its
duration (~50–100 ka). The glacioeustatic climate
fluctuations likely reflect Milankovitch cycles. The
terminal Devonian extinction is synchronous with
volcanic eruptions, climate changes, sea level fluctua-
tions, and the development of anoxia and probably
hypercapnia. The DCB readily correlates at the sub-
global scale owing to the positive δ13C peak [16, 35, 46]
(Fig. 3). This peak is considered synchronous with the
Hangenberg black shale member.
The hydrochemical changes in the ocean during the
Devonian‒Carboniferous transition from euxinic to
aerobic conditions was able to be determined by
replacement of vertical haline circulation with thermo-
haline circulation. In the Devonian, deep parts of the
ocean were filled with transformed dense hypersaline
water that formed in epicontinental and shelf seas at low
latitudes, while cooling resulted in its formation at high
latitudes from the sinking of cold, well-aerated water.
Another probable cause of these changes could be a low
oxygen concentration in the Devonian atmosphere and
an increase in it by the end of this period [5].
Unlike in most DCB sections represented by black
shales with high concentrations of organic matter, the
section located in the eastern Alps (Austria) is com-
posed of carbonate facies. The last section was sub-
jected to high-resolution geochemical studies [11].
The concentrations of organic matter and δ13C, δ18O,
δ34S isotopes reveal a shift to higher values, which
reflects cooling, intensification of oceanic circulation,
and elevated bioproductivity.
The Hangenberg black shales that accumulated on
the Late Devonian shelf in its Poland part are evidence
that they were deposited under alternatively varying
(anoxic to well-aerated) conditions in the bottom
water layer. U/Th values exceeding 1.25, Ni/Co values
exceeding 7, a total content of organic matter above
10%, and other geochemical features in the lower part
of the Hangenberg member indicate anoxic or euxinic
environments. The formation of the upper part of the
black shale member was also accompanied by inter-
mittently occurring anoxic conditions. All the exam-
ined samples contained small pyrite framboids and
biomarkers confirming euxinic conditions in the pho-
tic zone [41]. The middle part of the Hangenberg shale
member in the same section encloses a bed of volcano-
genic sediments with geochemical characteristics typ-
ical of oceanic basalts. Proceeding from these data, the
authors believe that volcanism could have resulted in a
reduction in biodiversity.
The Kellwasser and Hangenberg black shale mem-
bers enclose bacterial mats similar to their counter-
parts formed by recent cyanobacteria [36]. Cyanobac-
teria are tolerant to low-sulfide conditions. They char-
acterize conditions in Late Devonian epicontinental
seas as euxinic and enriched in H2S, i.e., unfavorable
to most benthic organisms. It is conceivable that bot-
tom conditions during the Late Devonian varied from
well-aerated to anoxic depending on the rates of oxy-
gen production by cyanobacteria, intensity of H2S
emission from decomposed mats, and plankton pro-
duction rates.
OCEANOLOGY Vol. 56 No. 6 2016
 Temperature, °С δ13C, ‰ Impact events Volcanism Extinction
20 25 30 2 4

OCEANOLOGY
Viluy trap

ferous
Carboni-
DCB
“Hangenberg
Charlevoix Flinn Creek
Aorounga event”

Vol. 56
Siljan
Woodleigh

No. 6
Black
shale

Famennian

2016
Alamo

?
FFB “Kellwaasser

Late Devonian
event”
Ilyinets
Kaluga

Frasnian
Alamo

1 2 3
– +
Sea level

Givetian
Mid. Devonian
Fig. 3. Correlation between abiotic factors and mass extinctions of organisms in Late Devonian. Environmental conditions: surface water temperature in low latitudes, after [31];
CAUSES OF THE GREAT MASS EXTINCTION OF MARINE ORGANISMS

Frasnian‒Famennian boundary (FFB); sea level: (“+”) transgression, (“‒“ regression, after [32]; carbon isotope ratio δ13C, after [13]; impact events, crater diameters:
(1) >100 km, (2) 100‒10 km, (3) <10 km; volcanism of Viluy trap province (periods of intensification are indicated). Rectangles show stratigraphic position of black shale mem-
bers and Kellwaasser and Hangenberg mass extinction events. Dashed lines show assumed intervals of age estimates.
867
868
Transgression
Fig. 4. Extinctions of marine organisms and sea level fluctuations in Late Devonian (after [52], modified). The position of the
Alamo impact event and glacioeustatic rise in sea levels during the Famennian interglacial are indicated.
The Frasnian Age was marked by a global rise in sea
level (Fig. 4). Mass extinction occurred against the
background of its sharp drop at the Franian/Famen-
nian transition due to the onset of glaciation in the
Southern Hemisphere. These processes resulted in
reduction in the main ecological niche for marine
organisms and was accompanied by accumulation of
Kellwasser black shales. The Famennian Age was
characterized by a global drop in sea level, which was
interrupted by its several glacioeustatic rises and cul-
minated in the less significant Hangenberg mass
extinction during another sharp drop in sea level,
which was, in turn, determined by the maximum gla-
ciation on Gondwana [52].
The authors of [52] believe that catastrophic
eustatic rises in sea level and subsequent mass extinc-
tion in the late Frasnian are most likely explained by a
series of comet showers. They substantiate this
assumption by many known and presumable impacts,
such as the Siljan and Flinn Creek events. The major
rise in sea level after the Alamo impact event is consid-
ered as an argument in favor of relations between cat-
astrophic rise in sea levels and comet showers. The rise
in sea levels induced by the Alamo impact event and
subsequent comet showers weakened the ecosystem
BARASH
D/C extinction
Famennian Stage
Interglacials
F/F extinction
Frasnian
Alamo impact event
Givetian
Regression
and strengthened the injurious effect of the sea level
fall at the end of Frasnian Age marked by mass extinc-
tion.
Climate changes such as both warming and cooling
are also considered as being responsible for mass
extinction. Warmings associate with emanation of
greenhouse gases during volcanic eruptions and sub-
sequent development of anoxia [10, 23, and others].
Coolings associate with glaciation in the Southern
Hemisphere and drop in sea level. The Devonian
cooling could be caused by outburst of terrestrial
plants, which consumed carbon dioxide from the
atmosphere, intensified weathering, anoxia develop-
ment, and burial of organic carbon [4].
Cooling at the Frasnian‒Famennian transition is
substantiated by the dominant extinction of reef-
dwelling and other low-latitudinal organisms, survival
of cold-resistant organisms during and after crisis
periods, evidence of Gondwana glaciation, and rapid
sea level fluctuations. Cooling in the low-latitude
zone could have resulted from the convergence
between the Gondwana and Laurentia plates in the
Frasnian Age, blocked equatorial circulation, and
deviation of cold high-latitude currents to the tropical
zone [18].
OCEANOLOGY Vol. 56 No. 6 2016
 CAUSES OF THE GREAT MASS EXTINCTION OF MARINE ORGANISMS
Number of conodont
δ13Ccarb, ‰ PDB Δ13C
0 1 2 3
10
m
8
0 FFB
–2 Kellwasser
–4
–6
–8
Fig. 5. Correlation between variations in carbon isotope ratio, evolution of conodonts, and sea level fluctuations near FBB in
Upper Devonian section of southern China (after [56], modified).
Available publications contain many paleontologi-
cal and lithological data on the terminal Frasnian
cooling related to Gondwana glaciation and indicating
that the latter marked the onset of the Late Paleozoic
glacial epoch [43]. Its first and second impulses took
place approximately 376 Ma ago (lower Kellwasser)
and 375 Ma ago (upper Kellwasser), respectively. Both
impulses are reflected in the reduction in biodiversity.
The second stage of the Late Paleozoic glacial epoch
corresponded to the Famennian cooling event. Glaci-
ation in Gondwana reached its maximum at that time.
Coastal sections (Kentucky, United States) contain
glacial detritus including boulders. The minimum size
of the Famennian ice sheet in western Gondwana at
the second stage is estimated as 16 × 106 km2.
Many continents yield evidence of a rapid eustatic
drop in sea levels in the Late Devonian: carstification
and brecciation of Frasnian and Famennian carbonate
rocks determined by their subaerial exposition, strati-
graphic hiatuses, evaporitic and black shale formations.
Cooling and Famennian glaciations in southern
Gondwana can also be explained by tectonic factors
[7]. The simultaneous formation of the main Aca-
dian‒Eovariscan fold belt gives reasons to explain
relations between environmental perturbations and
intense continental rise through two mechanisms
responsible for the decrease of CO2 concentrations in
the atmosphere: (1) intensification of silicate weather-
ing on continents, which is evident from the signifi-
cant increase in the 87Sr/86Sr value of seawater at the
OCEANOLOGY Vol. 56 No. 6 2016
869
Sea level
species – +
4 12
Frasnian‒Famennian transition; (2) mass burial of
organic carbon (Kellwasser Event) in semiclosed
basins due to the reduction in equatorial interoceanic
connections between the Panthalassa and Paleotethys
oceans. It is also assumed that primary production
increased owing to the elevated influx of nutrients
from continents.
The sea level changes documented in deep-sea sec-
tions, on submarine rises, slopes, outer and inner parts
of the shelf [52], as well as the global positive δ13Ccarb
excursion, confirm the correlation of the mass extinc-
tion near the FBB with the drop in sea level and
changes in the carbon cycle [56].
To determine CO2 concentrations, Δ δ13C (= δ13Ccarb ‒
δ org) is used [20], since the difference between the
13C
carbonate and organic components is determined by
fractionation during photosynthesis, which depends
partly on the CO2 concentrations in seawater. In order
to reveal correlations between variations in CO2 con-
centrations in the atmosphere and extinctions in the
FFB interval, paired analyses of inorganic (δ13Ccarb)
and organic (δ13Corg) carbon isotopes were performed
for two Upper Devonian sections in southern China
[56] (Fig. 5). A more significant fall in the share of
organic isotope as compared with its carbonate coun-
terpart indicates increased CO2 concentration in the
atmosphere. This decrease was preceded by regres-
sion, which corresponded to the beginning of mass
conodont extinction. The drop in temperature deter-
870 BARASH
mined by the decreased greenhouse effect of atmo-
spheric CO2 could have contributed to the mass
extinction.
According to [16], which investigated marine sec-
tions in the United States (Illinois), the δ13Ccarb values
increased from ‒1.0 to +5.0‒7.0‰ (Hangenberg
excursion) at the Famennian‒Tournaisian transition,
indicating that the greenhouse climate was replaced by
a glacial one [16]. High-resolution investigations of
changes in depositional environments in the DCB
interval carried out in eastern Anti-Atlas (Morocco)
[34] revealed high-frequency sea level fluctuations,
including two global transgressive and anoxic events
corresponding the accumulation of Hangenberg
(DCB) and Lower Alum black shales (Tournaisian
Stage of the Lower Carboniferous). In this region, both
events coincide with mass extinctions. Correlation with
glacial phases in southern Gondwana implies the gla-
cioeustatic factor as responsible for sea level fluctua-
tions and an overall drop by approximately 100 m, cor-
responding to the main regressive phase of the Han-
genberg Event. These results are consistent with
assumptions of a mild humid climate for the terminal
Devonian, which stimulated expansion of glaciation in
high latitudes and elevated accumulation of detrital
sediments in the middle latitudes of Northwest Africa.
Apatite of conodonts abundant in Paleozoic sections
was used for quantitative reconstruction of Devonian
paleotemperatures [31]. The oxygen isotope ratio was
measured in 639 samples from sections of North Amer-
ica, Europe, and Australia. In the tropical belt, paleo-
temperatures were as high as 30‒32°C in the Early
Devonian; in the Middle Devonian (397‒385 Ma ago)
they decreased to 22‒25°C; in the Frasnian Age, they
increased again to reach maximal values (30‒32°C) at
the Frasnian‒Famennian transition. These recon-
structed paleotemperatures indicate that coral‒stro-
matoporoidea reefs developed under relatively mild cli-
mate conditions, while reefs formed by microbial com-
munities were characteristic of very warm environments
of the Early and Late Devonian. In the middle‒late
Famennian, moderate cooling was crowned by brief
glaciation at the Devonian‒Carboniferous transition,
which is reflected in the positive δ18O shift in apatite of
conodonts. Judging from these paleoreconstructions,
two Devonian extinction events (at the Give-
tian‒Frasnian and, in particular, Frasnian‒Famen-
nian transitions) corresponded to temperature growth
episodes.
The Kowala section in Poland reflects climatic
fluctuations determined by astronomical rhythms
[21]. Sediments of its lower part were deposited under
conditions of a hot greenhouse climate, while overly-
ing sediments imply transition to glacial environ-
ments. The lithological variations in the Fras-
nian‒Famennian section correspond to eccentricity
rhythms lasting 405 and 100 ka. This interpretation is
confirmed by high-resolution measurements of stable
isotopes. According to the developed age model,
events accompanied by accumulation of lower and
upper Kellwasser black shales are separated by a period
of 800 ka. The pyrite distribution and Th/U dates
showed that during the late Kellwasser Event, the oxy-
gen content remained low during 400 kyr. According
to δ13Corg measurements, the carbon cycle after its shift
by +3‰ became stable 600 kyr later.
The Famennian‒Tournaisian (DCB) interval
demonstrates lithological variations corresponding to
100-ka-long astronomical cycles. In the opinion of the
above-mentioned authors [21], black shale members
including that reflecting the Hangenberg Event accu-
mulated during the transgression, when the extremely
high eccentricity stimulated destruction of small con-
tinental ice sheets in southern latitudes of western
Gondwana.
Volcanic eruptions, including the eruption of Viluy
traps, which was the largest in the Devonian,
undoubtedly had a significant influence on the extinc-
tions. The Viluy (Yakutsk) trap province developed in
the Lena, Markha, and Viluy river valleys along rifto-
genic normal faults and on slopes of the Anabar and
Aldan shields. Intrusive rocks are represented by dikes,
sills, and bedded basaltic breccias. Basalts are interca-
lated by ashes and tuffs. The Late Devonian‒Early
Carboniferous volcanics and sediments rest upon Pro-
terozoic and Lower Paleozoic rocks. The Viluy aula-
cogen extends northeast for 600 km to dip under the
Verkhoyansk fold belt. The largest part of the Viluy
trap province is eroded, and its western part is overlain
by Siberian Permian‒Triassic traps. At the same time,
only the aulacogen proper contains approximately
300000 km3 of igneous rocks [39].
Eruptions represented a multistage process. The
thickness of the sedimentary sequence is 3‒7 km in the
western part of the rift. The total volume of igneous
rocks in the Viluy rift is estimated to be 1000000 km3.
The 40Ar/39Ar age averaged for several samples is
373.4 ± 0.7 Ma, which corresponds to the FFB age,
accepted as 372.2 ± 1.6 Ma [19]. Other dates obtained
for samples from the Viluy rift constitute two clusters
with mean weighted values of 364.4 ± 1.7 and 376.7 ±
1.7 Ma. The first value is close to the DCB age, some
ahead of it. The authors believe that these data support
the hypothesis on relationships between extinctions
and volcanism [50]. The application of the new
40Ar/39Ar technique for dating basaltic eruptions tak-
ing into account their alteration during weathering
allowed the author of [8] to draw a conclusion on the
sufficient dating accuracy and argue that eruption of
Viluy traps corresponds in age to the FBB, i.e., the
main mass extinction of the Late Devonian.
There are also other large regions of Late Devonian
magmatism in addition to the Vilyui trap province.
Two stages of basaltic magmatism are recorded in the
South Urals: in the last third of the Frasnian Age and
more powerful corresponding to the FBB [54]. A large
OCEANOLOGY Vol. 56 No. 6 2016
 CAUSES OF THE GREAT MASS EXTINCTION OF MARINE ORGANISMS
region of alkaline magmatism 100000 km2 in size,
which includes 25 magmatic centers, is known in the
East European Platform [55]. The Pripyat–Dnieper–
Donets rift zone extends for over 800 km [38]. The
volcanic zone in northern Iran was likely a constituent
of the large Late Devonian volcanic province in the
peripheral part of Paleotethys [24]. Reflections of Late
Devonian magmatic processes are also known in
northern and southern China. The impact of eruptions
on the biota includes several probable mechanisms:
ejections of large volumes of CO2 and SO2 into the
atmosphere, global darkening, acid rains, warming,
rise in sea level, formation of high-saline deep water,
relatively strong vertical stratification, and develop-
ment of anoxia in the ocean. The ratios between REE,
Fe, Mn, Al, and other elements in limestones of south-
ern China indicate the development of hydrothermal
processes at that time, which could have influenced
oxidation of seawater, its pollution by heavy metals,
and eutrophication, and could have resulted in distur-
bance of the neritic ecosystem [58].
Upper Devonian sections demonstrate reflections
of intense bombardment of the Earth by cosmic bodies
(asteroids and/or comets), which should have injuri-
ously affected biodiversity. They are represented by
craters and areas with elevated iridium concentrations,
shocked quartz, and microspherules. (Estimates of
ages of impact events obtained by different methods
are characterized by the significant scatter.) One such
manifestation, the Woodleigh crater, is near the west-
ern coast of Australia (26°03′ S, 114°40′ E). The crater
is estimated to be Late Devonian in age: 364 ± 8 Ma
after [22] and 369 ± 4 Ma after [44]. Its diameter is
120 km (probably, up to 160 km). This crater is fourth
in size among similar known structures over all the
Earth. It is overlain by sediments; nevertheless, its
structure has been revealed by gravimetric measure-
ments, and that of oceanic rocks, by drilling. The cra-
ter bears distinct features of an astrobleme. The central
uplift of the granitoid basement is approximately
20‒25 km across. It is located at a depth of 171 m, i.e.,
1800 m above the regional basement. The uplift is sur-
rounded by a ring trough. The sediments contain
shocked quartz and other mineralogical indicators of
the impact crater. The size of the impact body (aster-
oid, comet) is estimated to have been 5‒12 km across.
The Siljan crater is located in central Sweden
(61°2′ N, 14°52′ E; diameter 50‒60 km). Its age is
estimated as either 361.0 ± 1.1 or 368 ± 1 Ma [44] (in
any case, Late Devonian). This impact structure
resulted from a granite massif 1700 Ma old being
struck by a cosmic body, which stimulated the forma-
tion of a melted granite layer ~200 m thick, overheated
to 2000‒2200°C, and a spacious system of fractures in
the granite massif extending to a depth of 5‒6 km.
Several structures (Khibiny, Lovozero, and Kontoz-
ero) are confined to a single NE-trending line with the
undoubted Siljan astrobleme, which likely implies the
OCEANOLOGY Vol. 56 No. 6 2016
871
same formation mechanism [3]. Glassy spherules with
morphological and chemical properties similar to the
properties of tectites and microtectites have been
found at the stratigraphic level of the FFB in sections
of Belgium [17]. It is assumed that these spherules rep-
resent products of ejection during the impact event,
most likely related to the formation of the Siljan crater.
The Flinn Ceek crater is in Tennessee (United
States) (36°17′ N, 85°40′ W; diameter 3.6 km). It is
estimated as 360.0 ± 20.0 Ma old [22, 44]. This struc-
ture resulted from an asteroid hitting Ordovician car-
bonate deposits. The crater is filled with basal breccia
and Upper Devonian black shales.
The Aorounga impact crater (19°06′ N, 19°15′ E),
discovered in northern Chad (Africa) in 1992, is 12.5–
18.0 km across (according to different estimates) and
less than 345 Ma old. The crater is deeply eroded. Its
structure is represented by two ring swells approxi-
mately 100 m high located up to 7 and 11 km from the
center. The swells are separated by a ring depression
with a hummocky area in the central part approxi-
mately 1.5 km wide. The section includes fine-grained
breccia composed of particles with fluidal texture [9].
The maximum age estimate of the impact derived
from target rocks corresponds to the Late Devonian.
Subsequently, the sizes of the structure and its ele-
ments were specified and new mineralogical features
indicating its impact nature were found [37]. In recent
catalogs, the age of the impact event is indicated as
<345 Ma, i.e., Carboniferous. Subsequently, a chain
of four craters was established (10.9–12.0 km across
and located 14‒20 km from each other), which
extends northeast. The age of three of them was deter-
mined as ~360 Ma; i.e., this impact event occurred
immediately prior to the Devonian‒Carboniferous
transition.
The giant Alamo structure (37°30′ N, 116°30′ W;
diameter up to 190 km) is described in North America
(Nevada and neighboring states). Its age is estimated
as 367 Ma (some authors consider it to be 382.1 ±
3.0 Ma old [44]). This region, ~25000 km2 in size,
hosts Alamo impact breccia, representing a member of
shallow-water marine limestone 50‒100 m thick that
disintegrated into large blocks and was deformed.
There are geological, petrographical, and geochemical
features indicating that the Alamo Event reflects the
impact of a large cosmic body (~5 km across) hitting a
relatively deep-sea area beyond the carbonate plat-
form on the tectonically active margin of the Pan-
thalassa Ocean or on the shelf margin. The differenti-
ated crater, 50–75 km in diameter, buried under sedi-
ments is located at a water depth of 1.5 km [47].
Over 20 mountainous areas surrounding the crater
exhibit outcrops of allochthonous and semiautoch-
thonous breccias, impact ejection products, and seis-
mically disturbed layers. The lower part of the breccia
sequence contains sediments deposited several sec-
onds after the impact, as well as tsunamites and fine-
872 BARASH
Tectonics Mantle plumes
Volcanism
oscillations
Climate Emission of CO2, SO2, Cl, F, CH4
Greenhouse effect
fluctuations
Stagnation of the ocean
Sea level
Anoxia
Mass extinction
Fig. 6. Schematic model of abiotic factors responsible for great mass extinction event in Late Devonian.
grained sediments deposited subsequently over sev-
eral hours or a longer period. An iridium anomaly
(~70 ppt), grains of shocked quartz, carbonate accre-
tionary lapillis, products of impact-induced meta-
morphism, and different ejection products, including
glassy spherical grains, confirm the impact nature of
this structure. Alamo breccias are found at a distance
of at least 100 km away from the intended point of
impact, and distal quartz grains emissions, at dis-
tances up to 250 km.
Several other astroblemes are also attributed to the
Late Devonian: Charlevoix (Quebec, Canada, 47°32′ N,
70°18′ W, diameter 54 km), Ilyinets (Ukraine, 49°08′ N,
29°11′ E, diameter 4.5 km), Kaluga (Russia, 54°30′ N,
36°15′ E, diameter 15 km), Elbow (Canada, 50°59′ N,
106°43′ W, diameter 8 km). Figure 3 presents assumed
ages of these events.
In southern China, uppermost Frasnian sediments
contain microspherules of presumably impact origin
[40] and peak iridium concentration at the FBB,
which is interpreted by different researchers as either
reflecting the impact event or resulting from reducing
processes. The moderate iridium anomaly exceeding
its background values approximately 20 times was dis-
covered near the FBB in Western Australia [49]. This
anomaly may be related to development of the stro-
matolite layer with cyanobacterium Frutexites, fibers
of which concentrate some elements such as Ir, Pt,
Co, As, Mn, Fe, and others exceeding their back-
ground values by two to five times. Thus, the iridium
Asteroids Comets
Impact events
Ash, aerosols
Darkening of atmosphere,
reduction in UV radiation
Reduction of photosynthesis
and bioproductivity
Destruction of food chains
anomaly in question may be related to other processes,
not an impact event.
A special investigation aimed at discovering the
features indicating impact events that could have been
responsible for the biotic crisis were undertaken in the
FFB stratotype of southern France [26]. In contrast to
previous investigations of the same section, no high Ir
concentrations, Ni spinel, or microtectites were found
by these authors. A twofold excess of Ir concentration
found in some samples is explained by these authors as
the accumulation of common cosmic dust during
periods of low sedimentation rates. Nevertheless,
there is convincing evidence that Late Devonian
extinctions correspond to many (including major)
impact events, which should have been responsible for
environmental deteriorations.
All of this indicates that mass extinction in the Late
Devonian was determined by a combination of impact
events and extensive volcanism. They could have acted
during a continuous (longer or shorter) time interval,
as has been noted for almost all great mass extinctions
[2]. Their joint action was the most catastrophic. Vol-
canism and an impact event result in similar conse-
quences for environments (Fig. 6): ejection of harmful
chemical elements and aerosols into the atmosphere,
the greenhouse effect, warming, darkening of the
atmosphere preventing penetration of ultraviolet light
and photosynthesis, stagnation of oceans, and devel-
opment of anoxia. These processes are accompanied
by a reduction in bioproductivity and destruction of
food chains. Mass extinctions are confined to time
OCEANOLOGY Vol. 56 No. 6 2016
 CAUSES OF THE GREAT MASS EXTINCTION OF MARINE ORGANISMS
intervals in which the combination of harmful condi-
tions reached a level impossible for the further exis-
tence of different groups of organisms: disturbances of
vital processes led to mass extinction of the biota.
Such a conclusion broadens the views on the possi-
ble abiotic factors responsible for great mass extinc-
tions, which have been discussed for many decades.
Climate changes, volcanic eruptions, tectonic move-
ments, sea level fluctuations, and impact events,
which have been considered as such factors, represent
only links in chain processes resulting in mass extinc-
tions. It seems that the succession of all these pro-
cesses is primarily determined by sinusoidal oscilla-
tions of the solar system with respect to the galaxy and
crossing of its arms during orbital motion around the
center [25]. According to these authors, reiteration of
extinctions at the same points of different spiral arms
confirm the main (galactic) cause of these events,
which reflects its influence on terrestrial and extrater-
restrial processes.
Some authors doubt this hypothesis or even ignore
it due to temporal inconsistencies between volcanism,
impact events, and extinctions. In my opinion [1, 2],
these inconsistencies are insignificant, since cosmic
factors are responsible for intervals of unfavorable
conditions millions of years long, which determine
such stepwise extinctions as, e.g., the Late Devonian
and Triassic‒Jurassic events. Mass extinctions occur
within intervals in which environmental conditions
become extremely unfavorable.
Recently, several researchers reached the same
conclusion discussed in this work: mass extinctions
were determined by the combination of impact events
and volcanism. Analysis of extinctions at the generic
level for 73 stratigraphic units of the Mesozoic and
Cenozoic revealed that most significant extinctions
took place only during simultaneous action of volca-
nism and asteroid impacts [6]. Volcanism, climate
changes, and sea level fluctuations exert pressure on
ecosystems to change their structures. Impact events,
as well as oceanic anoxia, are such sudden catastrophic
events that determine mass extinctions.
ACKNOWLEDGMENTS
This work was supported by project no. 0149-2014-
0027 of the Shirshov Institute of Oceanology (govern-
ment contract of the Federal Agency of Scientific
Administration) and by the Presidium of the Russian
Academy of Sciences (program no. 28 “Problems of
the Origin of Life and Development of the Biosphere).
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Translated by I. Basov