GSA TodAy | 2012 JAnuAry
Alycia L. Stigall, Dept. of Geological Sciences and Ohio Center
for Ecology and Evolutionary Studies, Ohio University, Athens,
Ohio 45701, USA, email@example.com
The Late Devonian (Frasnian-Famennian) interval includes one
of the most dramatic intervals of biotic turnover in the
Phanerozoic. Statistical evaluation of diversity change reveals that
the primary cause of biodiversity decline was reduced speciation
during the crisis interval, not elevated extinction rates. Although
various hypotheses have been proposed to explain extinction
increase during the Late Devonian, potential causes for reduced
speciation have previously been largely unaddressed. Recent
analyses focusing on biogeographic and phylogenetic patterns of
species in shallow marine ecosystems of Laurentia indicate that a
dramatic increase in interbasinal species invasions, facilitated by
transgressive pulses, fundamentally affected biodiversity by
enabling range expansion of ecological generalists and eliminating
vicariance, the primary pathway by which new species typically
form. Modern species invasions may result in similar speciation
loss, exacerbating the current biodiversity crisis.
A dramatic interval of biodiversity loss and ecosystem
reorganization occurred at the boundary between the Frasnian
and Famennian stages of the Late Devonian Period (ca. 375 Ma).
This event was originally considered to rank among the “Big Five”
mass extinction events in the Phanerozoic (Raup and Sepkoski,
1982), and it is still listed as the “Frasnian-Famennian Mass
Extinction” in most introductory and historical geology
textbooks. The designation of “mass extinction,” however, is
misleading because the Frasnian extinction rate was neither
elevated relative to the Middle Devonian nor statistically higher
than the background rate of extinction throughout the
Phanerozoic (Bambach et al., 2004; Alroy, 2008). Rather, an
anomalously low rate of speciation, the origination of new species,
was the primary cause of this decline in biodiversity (Bambach et
Global standing biodiversity is controlled equally by the
number of new species forming and the number of species
becoming extinct during an interval. All episodes of biodiversity
loss require that extinction rate exceeds speciation rate. For an
event to be classified as bona fide mass extinction, however, the
extinction rate of the crisis interval must statistically exceed both
the background extinction rate of the Phanerozoic and be elevated
above that of the adjacent stages. Biodiversity crises occur when
speciation rates have a statistically significant decline compared to
the background rate while extinction rates remain within the
limits of statistical normal. Reduced speciation rate combined
with slightly elevated extinction levels can result in a dramatic
biodiversity crisis, and this is what transpired during the Late
Devonian. The Frasnian-Famennian event is, therefore, better
termed a “biodiversity crisis” than a “mass extinction.”
The shift in status of the Frasnian-Famennian event from a
“mass extinction” to a “biodiversity crisis” does not imply a
reduction in the severity of the effects on global ecosystems. In
fact, the level of marine ecosystem reorganization that occurred
during the Late Devonian, including a fundamental collapse of
the reef ecosystem, is second only to the Permo-Triassic mass
extinction (McGhee et al., 2004). The Middle Devonian included
the most geographically widespread metazoan reef ecosystem in
Earth’s history, but its extent was reduced by a factor of 5000
following the crisis interval (Copper, 1994). Other biotic changes
included the spread of cosmopolitan species facilitated by
rampant species invasions documented across many clades
(reviewed in McGhee, 1996).
A series of local and global environmental changes occurred
coincident with biotic overturn. These included changes related to
the development of complex forest ecosystems on land, such as
eutrophication and alteration of terrestrial weathering patterns
(Algeo and Scheckler, 1998), high frequency sea-level changes (ver
Straeten et al., 2011), widespread anoxia events (Buggisch and
Joachimski, 2006), overall warming of the global oceans (van
Geldern et al., 2006), and pulses of enhanced carbon burial that
resulted in rapid cooling events at the Frasnian-Famennian
boundary (van Geldern et al., 2006; also see reviews in McGhee,
1996, 2005; Racki, 2005). Most of these environmental factors
(and various combinations of them) have been proposed as
drivers for the “mass extinction.” Theoretically, abrupt or even
gradual changes in environmental conditions could result in
increased extinction of species because extinction occurs when
members of a species can no longer cope with changing
environmental conditions (abiotic or biotic) and population size
decreases to zero. These environmental factors are undoubtedly
involved with ecosystem degradation and certainly contributed to
the observed elevation of extinction levels.
None of these abiotic changes, however, supply a satisfactory
explanation for speciation collapse because they do not directly
impact the speciation process. In order to better understand the
ecological crisis during the Late Devonian, the mechanisms of
speciation decline must be examined. A speciation event is a
unique episode in geologic time that transpired at a discrete
geographic location within a specific lineage of organisms.
Identifying causal factors for speciation decline requires both a
detailed temporal and geographic framework and robust hypotheses
of ancestor-descendant relationships (Benton and Pearson, 2001).
This type of detail is only available in clades for which species-
level phylogenetic hypotheses have been generated. Therefore,
speciation analysis requires a fundamentally different and more
Speciation collapse and invasive species dynamics
during the Late Devonian “Mass Extinction”
GSA Today, v. 22, no. 1, doi: 10.1130/G128A.1.
GSA TodAy | 2012 JAnuAry
detailed dataset than analyses of potential extinction mechanisms.
Although the significance of speciation reduction during this
interval has been known for more than 25 years (McGhee, 1984),
sufficiently resolved phylogenetic data have only recently become
available to assess changes in speciation mode across the Middle
to Late Devonian interval. Significantly, recent analyses discussed
herein indicate that the widespread extra-basinal migrations of
species (analogous to modern invasive species) during the Late
Devonian facilitated speciation decline by preventing geographic
isolation, the primary process by which new species arise.
ORIGINATION AND SPECIATION RATE ANALYSES
The importance of reduced origination in driving Frasnian
diversity decline was first recognized by McGhee (1984) from
analyses of genera of articulate brachiopods from Catskill delta
complex of eastern North America and from the Ural Mountains.
Subsequent analyses (e.g., Foote, 1994; Bambach et al., 2004;
Alroy, 2008) using stratigraphic range data for marine invertebrate
families and genera culled from global compendia confirmed that
reduced origination was the primary driver of biodiversity
collapse (Fig. 1A). Certainly, patterns of biodiversity change were
not congruent across all clades. Some previously prolific clades,
such as the atrypid brachiopods, experienced high extinction
rates, while other clades, including the crinoids, radiated to effect
a pronounced change in post-crisis shallow marine ecosystems
(see discussion in Racki, 2005). However, the general pattern of
depressed origination—but only moderately elevated extinction
rates—documented in cross-faunal database analyses is robust to
variations in sampling procedures, rate metric used, taxonomic
level (family vs. genus) analyzed, or database employed (Foote,
1994; Bambach et al., 2004; Alroy, 2008). This supports depressed
origination as a primary driver of Late Devonian biodiversity loss.
From a biological standpoint, the most appropriate taxonomic
level to assess reduced origination is the species level. Species are
biological entities that are defined by attributes related to
reproductive cohesion in both time and space (deQueiroz, 2007).
Therefore, analysis of origination at the species level equates to
examination of actual biological processes, whereas generic and
familial analyses are increasingly distant proxies. Species-level
phylogenetic hypotheses, which include an evolutionary
framework to constrain timing of speciation events, are necessary
to calculate the most accurate speciation rates (Smith, 1994).
Unfortunately, very few species-level phylogenies have been
published with substage temporal resolution for Late Devonian
clades (e.g., Rode, 2004; Stigall Rode, 2005).
Stigall (2010a) utilized recently published species-level
phylogenetic hypotheses of Rode (2004) and Stigall Rode (2005)
for three Late Devonian clades (two articulated brachiopod genera
and one bivalve subgenus), primarily from North America, to
examine whether reduced origination was also significant at the
species level. These clades serve as a reasonable proxy for shallow
marine biota of Laurentia because these monophyletic lineages
had excellent preservation potential, include common members of
the shallow marine benthos, and their combined fifty species
inhabited the full suite of nearshore to offshore marine
environments. Results are consistent with the earlier analyses
based on higher taxa (Fig. 1B). Overall biodiversity plummeted
during the Frasnian crisis interval, and this change was driven
primarily by speciation loss (Stigall, 2010a). Late Frasnian
extinction rates, while moderately elevated, do not exceed pre-
crisis levels for any clade and are not statistically higher during the
crisis interval than the average value for each rate over the
duration of the clade (Stigall, 2010a).
The combination of species, generic, and family-level analyses
firmly establishes the loss of speciation as a fundamental driver
for biodiversity loss during the Late Devonian, at least among
shallow marine taxa where the crisis was most pronounced.
Examining the process of speciation and the factors that promote
or hinder that process is, therefore, required to identify causal
factors for the crisis.
SPECIATION MODE ANALYSIS
Investigating the cause of speciation collapse during the Late
Devonian first requires determining which speciation
mechanisms were compromised during the crisis interval.
Speciation requires a group of organisms to become
reproductively isolated from its ancestral population in order to
establish a new biological entity. This isolation typically occurs via
Figure 1. Comparison of extinction versus origination/speciation across the
Middle to Late Devonian interval. Late Devonian Biodiversity Crisis interval is
indicated in yellow. (A) Proportion of generic extinction or origination per
interval. Modified from Bambach et al. (2004). (B) Instantaneous rates of species
extinction and speciation for two brachiopod genera (Schizophoria and Floweria),
one bivalve genus (Leiopteria), and all three clades combined. Modified from
Stigall (2010a). Similar patterns occur at both taxonomic levels: Origination/
speciation rates are substantially reduced during the crisis interval, but extinction
rates are lower than during the Middle Devonian background interval.
GSA TodAy | 2012 JAnuAry
either geographic separation of the incipient species from the
ancestral population (allopatric speciation) or via shifts in
reproductive timing or chromosomal count within the same
geographic space as the ancestor (sympatric speciation) (Mayr,
1963). Sympatric speciation is commonly undetectable in the
fossil record, but allopatric speciation is perceptible because it
typically results in morphological shifts as incipient species adapt
to environmental conditions that differ from those of the
ancestral range. Allopatric speciation occurs via two primary
mechanisms: vicariance and dispersal, which are characterized by
discrete biogeographic patterns related to the geographic range of
daughter species relative to the ancestral population (Fig. 2)
(Wiley and Mayden, 1985). Thus, it is possible to identify
speciation events of each type in fossil taxa where evolutionary
relationships are known and ancestral ranges can be inferred
(Lieberman, 2000) (Fig. 3).
To assess speciation mode during the Late Devonian, Stigall
(2010a) conducted a biogeographic analysis on species-level
phylogenetic hypotheses of four common groups of Devonian
marine organisms: an order of predatory crustaceans, one bivalve
genus, and two brachiopod genera, published in Rode and
Lieberman (2002), Rode (2004), and Stigall Rode (2005),
respectively. This cross-phyla analysis included common taxa
within both the sessile benthos and pelagic predator guilds and,
thus, is a reasonable proxy for faunal dynamics in shallow marine
environments of Laurentia. Speciation by vicariance was limited
relative to speciation by dispersal in each of these clades, ranging
from only 12% to 50% of quantifiable speciation events, for a
combined rate of 28% speciation by vicariance versus 72%
speciation by dispersal. Similar analyses of the modern biota have
demonstrated vicariance to be the dominant form of speciation by
a factor of almost 3 to 1 (Brooks and McLennan, 2002), and
analyses of speciation mode conducted for other Paleozoic
intervals (reviewed in Stigall, 2010a) have always recovered higher
frequencies of speciation by vicariance versus dispersal (Table 1).
Speciation mode during the Late Devonian is evidently different
from the typical pattern in Earth history. This incongruity
provides the framework for a mechanistic explanation for
speciation decline during the crisis interval. During the Late
Devonian, vicariance, normally the most prevalent style of
speciation, was essentially extinguished. In fact, each of the few
vicariance events present in the clades analyzed precede the late
Frasnian crisis interval (Stigall, 2010a). Speciation by dispersal,
although still operational during the crisis interval, typically
occurred at a lower rate and accordingly resulted in few Late
Devonian speciation events. Therefore, elimination of the
dominant mode of speciation led to the dramatic reduction in
speciation rate, and consequently biodiversity, at the end of the
The differential loss of speciation type provides a foundation
against which to analyze causes of biodiversity decline.
Satisfactory causes for biodiversity collapse must be able to
explain both the lack of vicariant speciation and the slight
Figure 2. Geography of allopatric speciation modes. In vicariance, the ancestral
population (Species A) is passively divided by a geographic barrier. Incipient
species (Species A' and A") form during geographic isolation and later diverge
to become new species (Species B and C). In dispersal, a subpopulation of the
ancestral species (Species A) actively migrates across a geographic barrier to
form an incipient species (Species A'), which later diverges to become a new
species (Species B).
Figure 3. Biogeographic areas mapped onto a strato-cladogram for the
brachiopod genus Floweria. Speciation events where daughter species occupy a
subset of the ancestral range are interpreted as vicariance events. Speciation
events where daughter species occupy areas additional to the ancestral areas
are interpreted as dispersal events. Modified from Stigall (2010a).
Table 1. Comparison of speciation mode through geologic time
Geologic interval % Speciation by
28% (sd 16%)
54% (sd 16%)
74% (sd 35%)
% Speciation by
72% (sd 16%)
46% (sd 16%)
26% (sd 34%)
Cambrian to Middle Devonian
Late Devonian clades exhibit substantially limited speciation by
vicariance when compared to taxa in other geologic intervals. Modified
from Stigall (2010a); sd—standard deviation.
GSA TodAy | 2012 JAnuAry
elevation in extinction rates. Abiotic explanations alone, such as
global cooling or basin anoxia, do not provide adequate
explanations for the differential reduction in vicariance compared
to speciation by dispersal; however, biotic factors, such as the
spread of invasive species, potentially could.
INVASIVE SPECIES DURING THE LATE DEVONIAN
Extensive interbasinal species migrations have been
documented in many clades during the Frasnian (reviewed in
McGhee, 1996). These migrations are characterized by the
dispersal of a species native to one tectonic basin into a second
tectonic basin outside its original geographic range (Fig. 4).
Because these species establish secondary populations in
ecosystems in which they did not evolve, they are analogous to
modern invasive species (Vermeij, 2005; Stigall, 2010b). Species
migrations have occurred throughout geologic time; however,
most episodes of biotic exchange are limited to a localized
dispersal pathway (e.g., Great American Biotic Interchange; see
review in Vermeij, 2005). During the Late Devonian, species
introductions were rampant on a global scale.
The impact of these Late Devonian invasions was quantified by
Rode and Lieberman (2004) using Geographic Information
Systems–based analyses to calculate geographic ranges and map
invasion events in more than 300 Middle and Late Devonian
articulate brachiopod and bivalve species of Laurentia. Species
ranges were mapped at conodont zone resolution to produce a
high-precision temporal framework for identifying invasion
events (Fig. 5). A substantial increase in invasion intensity
occurred coincident with the decline in speciation during the
Frasnian; 68% of all identified invasion events occurred in the
Frasnian. Rapid transgressions provided pathways for species
dispersal; 65% of Frasnian invasion events correlate with
transgressive events. Additionally, the mean size of both native
and invasive species’ geographic ranges increased during the
Frasnian (Rode and Lieberman, 2004). Moreover, species with
larger geographic ranges, an episode of interbasinal invasion in
their history, and/or expansion of their geographic ranges during
the late Frasnian survived into the Famennian at statistically
higher rates than non-invasive species with narrow geographic
ranges (Rode and Lieberman, 2004; Stigall Rode and Lieberman,
These species invasions, facilitated by sea-level changes, could
have caused the observed reduction in speciation coupled with
moderately elevated extinction. The combination of overall range
expansion and frequent invasive events would have prohibited
sustained geographic isolation, thereby impeding the primary
requirement for vicariant speciation, as well as hindered the
successful development of migrant populations into new species,
thus restricting speciation by dispersal. The preferential
extinction of species with small geographic ranges could have
produced the observed elevation of extinction levels.
SYNTHESIZING INVASIVE SPECIES EFFECTS, ECOLOGY,
The results of speciation and biogeographic analyses provide a
framework in which to examine the mechanisms that reduced
speciation and slightly elevated extinction rates during the Late
Devonian Biodiversity Crisis. In particular, three features—
differential extinction of narrowly ranging species, impact of
invaders on native species, and macroevolutionary differences
between ecological generalist and specialist species—are critical
for explaining biodiversity decline.
A striking feature of the Late Devonian biogeographic pattern is
the differential survival of species with large geographic ranges.
Species with larger geographic ranges tend, on average, to have
broader ecological tolerances than those with small ranges
(Jackson, 1974; Fernández and Vrba, 2005). Ecological specialists
Figure 5. Interbasinal invasion intensity during the Middle and Late Devonian.
The spread of invasive species was facilitated by sea-level rises, indicated by
blue arrows. Modified from Rode and Lieberman (2004).
Figure 4. Interbasinal invasions of Late Devonian brachiopod, Pseudatrypa devoniana. (A) During the early Frasnian Stage, P. devoniana occupied its ancestral
basin in the New York region. (B) This species invaded the Iowa basin during the middle Frasnian and (C) subsequently invaded the New Mexico basin during
the late Frasnian. Invasions correspond to sea-level rises indicated by the second and third arrow in Figure 5. Modified from Rode and Lieberman (2004).
GSA TodAy | 2012 JAnuAry
are confined in terms of both their habitat preferences and the
geographic region where those conditions occur (Stanley, 1979).
Conversely, ecological generalists can successfully utilize a wider
set of environmental conditions, which typically allows them to
occupy larger geographic areas. Thus, although Middle Devonian
biotas included both ecological specialists and generalists, most
native species that survived the crisis had large geographic ranges
(Rode and Lieberman, 2004) and were presumably ecological
Furthermore, Devonian invaders were dominantly, if not
exclusively, ecological generalists. Modern invasive species are
characterized by broad environmental tolerances, which
contribute to their ability to survive during both the transport
and establishment phases of invasion (Lockwood et al., 2007).
Devonian invaders were likely similar, because ecological niches
of Devonian invaders must have been sufficiently broad to allow
colonization of both the invasion pathway and the new tectonic
basin. Consequently, the arrival of the Devonian invaders into
new tectonic basins effectively resulted in an influx of new
ecological generalists into the ecosystem. Studies of modern and
Cenozoic invasive species have demonstrated that invader species
regularly displace native species through higher resource
efficiency (Johansson, 2007) or competitive ability (Vermeij,
2005). Similar processes operating during the Late Devonian
would have caused differential extinction of narrowly ranging
ecological specialist species. This resulted in elevated extinction
rates and a proportional increase of broad ranging ecological
generalists versus geographically restricted specialists in the biota.
Clades of ecological generalists tend to have lower speciation
rates and contain fewer species relative to specialist lineages (Vrba,
1987; Eldredge, 1989). This discrepancy relates to the mechanics
of the allopatric speciation process. If a group of specialists
undergoes vicariance, it will likely be exposed to environmental
conditions that differ in some way from their ancestral range, and
the population must either adapt to those conditions or become
extinct. On the other hand, generalists are more likely to be pre-
adapted via their broad ecological niche to the new set of
conditions encountered so that no adaptive change is required.
Consequently, specialist lineages experience both higher
speciation and extinction rates than ecological generalists.
The differential extinction of native specialist species during
the Late Devonian reduced the potential ancestral species pool
from which new specialist species could evolve, which resulted in
speciation depression. Furthermore, native and invasive generalist
lineages would have had few opportunities to speciate as
expansion of geographic ranges facilitated by sea-level rise
prevented effective long-term vicariance from ancestral
populations. Rather, incipient generalist species were more likely
to be subsumed as a geographic extension of the expanding
ancestral species than to develop into new species. This
combination of preferential extinction of specialist species and
expansion of the geographic ranges of generalist species (native
and invasive) facilitated the dramatic speciation reduction of the
This pattern of differential survival, range expansion, and
speciation occurred within the most common components of the
Late Devonian shallow marine ecosystem but may not be
transferable to all marine clades or other environments. Central to
this argument is the frequency of range expansion among native
generalist species and the introduction of invaders resulting in
competitive interactions on the seafloor. Continental ecosystems,
including both terrestrial and freshwater habitats, and marine
taxa potentially less amenable to these processes did not
experience the same level of biodiversity loss during the Late
Devonian (reviewed in McGhee, 1996).
The Late Devonian Biodiversity Crisis was one of the most
significant intervals of biodiversity loss and faunal overturn during
the Phanerozoic. Unlike “true” mass-extinction events, such as the
Late Permian and End Cretaceous events, the primary driver of
biodiversity loss was a severe reduction in speciation rate, not
substantially elevated levels of extinction. Purely abiotic explanations
for biodiversity loss during the Late Devonian fail to provide a
complete explanation for the biodiversity crisis. Shifts in Late
Devonian biogeographic patterns were driven by range expansion of
generalist taxa within basins and rampant species invasions between
basins associated with transgressive events. These shifts provide a
mechanistic explanation for the reduction in speciation during the
crisis interval, particularly speciation by vicariance.
The central role of invasive species in mediating biodiversity
decline during that Late Devonian Biodiversity Crisis parallels
aspects of modern biodiversity crisis affecting our planet. The
primary drivers of the current biodiversity crisis are habitat
destruction, climate change, and the spread of invasive species
(Thuiller, 2007). The current rate of biodiversity loss is as high as
or higher than during any interval in the Phanerozoic (Barnosky
et al., 2011). The impacts of habitat degradation and climatic
change have long been analyzed within the context of geologic
time and are known to cause substantial elevation of extinction
rates. Comparison with the Late Devonian interval suggests that
the modern influx of invasive species will result in substantially
reduced speciation rates. The modern combination of habitat
destruction coupled with species introductions is, therefore, likely
to result in total biodiversity loss that may be even greater than
that experienced during the Late Permian coupled with an
extensive recovery interval due to speciation depression. These
implications highlight the need for conservation efforts to target
specialist taxa for protection in addition to preventing species
introductions and preserving habitat.
Thanks to Peter Harries, Bernard Housen, and three
anonymous reviewers for constructive reviews and to Damian
Nance for the invitation to submit this paper to GSA Today. This
research was supported by U.S. National Science Foundation
grant EAR-0922067 and a grant from donors to the American
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Manuscript received 18 May 2011; accepted 3 Nov. 2011.