Survival and Extinction Decon

Survival and Extinction: Deconstruction, Extinction
Studies, Paleontology
Luke Donahue
Abstract What is extinction? What is the difference between

death and extinction? Between evolution (surviving as other) and
extinction (not surviving at all)? This article begins by considering
the articulation of extinction in Extinction Studies. Combining rig-
orous philosophical insights with cutting-edge scientific research,
Extinction Studies offers one of the best understandings we have:
the extinct is not “what it is” (absolutely gone), but remains (in
both senses of the word). The extinct’s absence survives, calling
to us from beyond the grave. However, I argue almost the exact
opposite in this article: extinction is remainderless. And extinction
itself is going extinct.
Key Words Deconstruction; Derrida; ecocriticism; evolution;

extinction; Extinction Studies; Multispecies Studies; paleontology;
remainderless; survival; trace; Van Dooren
Introduction: Survival
Whether it began with the Holocene some 12,000 years ago, with
the Industrial Revolution, or with the Great Acceleration following
the Second World War, an extinction event has overtaken the globe.
Biodiversity is plummeting with an acceleration unprecedented
outside of the Earth’s major mass extinctions. The word “extinction,”
today, cannot be heard without the faint echoes of the Sixth Extinction
pounding on the well-insulated doors of our everyday consciousness.
Like the phenomenon itself, research on extinction has been under-
going a resurgence, and not only in the biological and paleontolog-
ical sciences, but in the interdisciplinary humanities, too. In Imagining
Extinction: The Cultural Meanings of Endangered Species, for example,
Ursula Heise argues that extinction must be understood as a culture
object—a sign used to resolve contradictions, reinforce identities, and
articulate sociopolitical critiques. In fact, she argues, cultural narratives
not only exploit the scientific concept of extinction but also institute
the very values that shape scientific inquiry. From an entirely different
Theory & Event Vol. 24, No. 4, 922–950 © 2021 Johns Hopkins University Press
Donahue | Deconstruction, Extinction Studies, Paleontology 923
angle, Claire Colebrook’s writings teach us that anthropogenic extinc-

tion (as well as our own auto-extinction) is not a mere accident in the
history of humanity but is part and parcel of humanity’s specular
and speculating functions.1 Finally, a whole discourse in the human-
ities explores the peculiar conceit of the Anthropocene: at the moment
humanity realizes it is as powerful as tectonic plates at shaping Earth
systems, humanity announces its own default and extinction, only to
then imagine some future (presumably non-human) geographer who
will archive it in the calendar of geological time.2
Within the vast landscape of ecocriticism, science studies, and new
realisms, a subgenre of “Extinction Studies” is emerging, announced
with three recent collections of essays: Extinction Studies: Stories of
Time, Death, and Generations; After Extinction; and Arts of Living on a
Damaged Planet. A diversity of perspectives rules in Extinction Studies,
from stories that immerse themselves in the cacophony of human and
nonhuman voices involved in local extinction crises, to theorizations
of how the Sixth Extinction transforms ethics, aesthetics, and politics;
from research on the biopolitics of zoos and wildlife management,
to imaginative creations of knowledge and action in a world of mass
death.
Despite this diversity of perspectives, a particular (though not
always explicit) concept of extinction emerges in many of the arti-
cles in this nascent field. Scholars associated with Extinction Studies
routinely complicate (but do not quite reject) the definition of extinc-
tion standard in the biological sciences. After all, extinction, today,
exceeds biology: our current Sixth Extinction cannot be extricated from
the destructive power of human narrative and rhetoric, economics and
politics, science and technology, and phenomenology and ideology.
However, extinction’s complexity is more fundamental and applies
even prior to the advent of anything like homo sapiens. Extinction is
complex because species are complex. As Thom van Dooren writes in
Flight Ways: Life and Loss at the Edge of Extinction:
To allow the term “extinction” to stand for only the death of the
last of a kind is to think within an impoverished notion of “spe-
cies,” a notion that reduces species to specimens, reified represen-
tations of a type in a museum of life, and in so doing ignores the
entangled relations that are a particular form of life.3
In other words, when we understand extinction as the simple passage

from the presence of a species to its absence, we erroneously posit species
as abstract kinds that are arithmetically added to or subtracted from
the world. Van Dooren is expressing a sentiment common throughout
Extinction Studies, by way of Multispecies Studies: species are not
isolated and discrete realities, but emerge in and as their entanglement
924 Theory & Event
with others.4 Before distinct species, their relations: enfoldings and

envelopments, avoidances and indifferences, mutualisms and parasit-
isms, as well as all sorts of other symbiotic entwinements wherein to
be is to be other—classically illustrated by the bacteria that humans
are (not), or by the semiotic exchanges and creations between bee and
flower.
In contrast to the traditional account of species as historical
lineages that have congealed into heterogenous kinds based on their
inability to produce viable offspring, many of the scholars associated
with Extinction Studies privilege sympoetic and biosemiotic braids
and fabrics. They insist that partial linkages among species (among
their genetic sequences, internal cellular and bacterial life, develop-
mental processes, morphological and behavioral traits, epigenetic and
cultural inheritances, and modes of reading, registering, responding
and recreating) convene to hold a species together while guaran-
teeing that that species is not a bounded identity but an entangled and
disseminated knot. Species are not there, isolate kinds, that then form
relationships of co-becoming: the relations and differences are within
species as much as between them. For an elk to be (or be categorized
as) an elk, it must first be the bacterial and cellular life composing it,
the technologies and rituals it constitutively requires, and the ecolog-
ical webs that not only surround it but intersect it. This sympoetic or
symbiogenetic understanding of species should not be confused with
a reactionary and wholesale rejection of the evolutionary account: no
one doubts that a ponderosa pine seed cannot grow into a manza-
nita, but the coherence and replicability of ponderosas as ponderosas
require whole arrays and networks of bacteria and plants and animals.
Far from our popular notion of individuated species competing in a
zero-sum evolutionary game, life is a messy, multispecies affair.
This insight has profound consequences for what we mean by
“extinction.” As the editors of Arts of Living on a Damaged Planet put it,
the concern is no longer “the loss of individual species but of assem-
blages.”5 They continue: “We often tally the plants and animals at risk
of extinction one by one on lists of endangered species. But single
species are not the best units through which to see extinction—because
they are not the units of life.”6 If species’ “[i]rresistible attraction
toward enfolding each other is the vital motor of living and dying on
earth”—if species are their involution with other species before they
are separated out as distinct (which is to say, if there aren’t species per
se)—then that which goes extinct cannot be simple species, but in-defi-
nite and borderless entanglements.7
Putting the complexity of extinction in more temporal than spatial
terms, Van Dooren notes that extinction
is the loss not of a single fixed “kind,” but of a potentially limit-

less set of emergent and branching flight ways from the present
into the diversity of the future. Each species is ultimately a flight
way beyond itself […] And so what is lost in extinction is not “just”
the current manifestation of a flight-way—a fixed population of
organisms—but all that this species has been, as well as all that its
past and present might have enabled it to one day become.8
What is lost is not only what was, but its past and future. Van Dooren,
though, is careful not to totalize the loss. For, if a species is its spatial
and temporal entanglements, then its extinction will echo, following
lines of conduction and osmosis, as it were. In this sense, the extinct
remains, “continu[ing] long afterward, haunting future possibilities for
a host of living beings.”9 Speaking of the extinct-in-the-wild Hawaiian
crow, ‘alalˉa, he argues:
their spectral presence is inscribed in the forest landscape. Plants

call out to ‘alalˉa, their fruiting and flowering bodies shaped by
past attractions and associations […] An appreciation of these
kinds of entanglements makes it easier to understand why a spe-
cies like ‘alalˉa cannot be neatly excised from our living world.10
The bird may be gone, but it survives as a ghost, an absent presence

inscribed in ecological and evolutionary systems.
Van Dooren, here, is giving voice to one of the most penetrating
insights (sometimes implicit, sometimes explicit) reverberating
throughout much of Extinction Studies: that which goes extinct para-
doxically survives. The extinct remains by cascading, affecting after the
fact other species entangled and coevolving with “it.” Since species are
assemblages that constitutively exceed their boundaries, assembling
with other assemblages, they cannot be completely and finally wiped
out. Otherwise, a single extinction would be a total one, the extinction
of all life. Instead of a simple end, the extinction of a species (which
was “itself” a knot of disseminated traces) leaves more traces behind.
As the editors of Extinction Studies put it: “the unraveling of forms of
life” is at stake (italics mine).11 And the editors of Arts of Living on a
Damaged Planet boldly assert in a subtitle: “Extinction Leaves Traces.”
They continue: “As life-enhancing entanglements disappear from our
landscape, ghosts take their place […] showing us how living land-
scapes are imbued with earlier tracks and traces.”12 Finally, the editor
of After Extinction explains that the volume was motivated by the
imperative to “think the event of extinction not as destructive or final,
but as generative.”13
Accordingly, it no longer suffices to understand extinction as the
clean cut of a phylogenetic line or as the death of the last organism of
a species. If we refuse to neatly differentiate a species from its traces
926 Theory & Event
and entanglements, then an extinction can never be a simple absence.

Rather, the extinct paradoxically survives, its absence inscribed,
remaining as ghostly echoes. In this vein, Extinction Studies refuses
the discourse of apocalyptic ends. As Cary Wolfe puts it, extinction is
haunted by “retentions from an evolutionary past and protentions of
an evolutionary future.”14 Extinction is the traces of (the) extinction (of
what survived as entangled traces).
This argumentative thread, which runs throughout much but
certainly not all of Extinction Studies, emerges in the context of the
vanguard ecological sciences that highlight symbiosis, sympoetics,
and biosemiotics over neo-Darwinian competition. But it also emerges
in the context of twentieth-century poststructuralism, which asserts
that to be is “to be” a ghost (which do not quite have being). Whatever
the unit in question (concept or thing, self or community, race or sex),
“it” survives from the first as other than itself, as the trace of a consti-
tutively erased ipse. “It” survives only and ever as remains, in excess
to an original nature, essential being, or self-coincident identity that
never existed in the first place. In twentieth-century poststructuralism,
survival replaces Being. Arguably, this receives its most radical and
influential formulation in the writings of Jacques Derrida. In his final
interview he says: “All the concepts that have helped me in my work,
and notably that of the trace or of the spectral, were relating to this
‘surviving’ as a structural and rigorously originary dimension,” a
dimension prior to the difference between life and death.15
Survival, for Derrida, “is not to be added on to living, and dying. It is
originary: life is living on, life is survival [la vie est survie]. To survive
in the usual sense of the term means to continue to live, but also to live
after death.”16 Not only in order to live, but in order to die, “it” first
survives. In all its unforgiving irreversibility and absoluteness, death
is nonetheless predicated upon survival. What died, in order to die,
was retroactively inscribed, transcribed, incorporated, iterated. The
supplemental inscription supplants—and thereby constitutes—what
it supplemented, which thereby survives as its own ghostly remains.
Speaking of ecology as generalized writing, Vicki Kirby conceives of a
play with neither “beginning nor end—an enfolded genesis wherein
every ending is an inventive reiteration, a beginning.”17
Extinction Studies dwells in this intellectual climate. Once species
are deconstructed and refigured as ghostly knots in fields of dissem-
inated traces, extinction cannot but become a movement of survival.
Blending the most rigorous philosophical insights with the most
cutting-edge scientific research, Extinction Studies offers the best
understanding we have of extinction today: the extinct is not “what it
is” (viz. extinct), but remains (in both senses of the word).
However, when extinction is understood as the unraveling of a
knot, as the inscriptions of the absent, do we fail to specify it? Do we
fail to differentiate extinction from death more generally? By extending

extinction via a logic of survival, do we make it a little too thinkable,
a little too familiar, a little too congruent with our poststructural
paradigms? And when we bring the extinct to life, hearing the voices
of the departed, do we limit our definition of extinction to extinc-
tion-among-humans, for humans? Finally, do we risk passing over the
radicality and nuances that might already be present in the traditional,
evolutionary conception of extinction?
A perusal of Extinction Studies reveals a routine slippage from the
language of extinction to the language of death. The editors of Extinction
Studies, for example, name and define the eponymous subfield as “a
particular focus on understanding and responding to processes of
collective death, where not just individual organisms, but entire ways
and forms of life, are at stake.”18 Note the shift in terminology. The
refiguration of species as “ways and forms of life” dictates that we can
no longer speak of extinction, but only of “collective death.”19 Once
the species concept has been deconstructed, extinction can no longer
be defined as the extinction of a species; it becomes a certain quantity
of death, an extension of death as “collective,” rather than a unique
type of event with a distinct quality. And once the species concept has
been deconstructed, incorporated in and as death, the extinct survives.
Extinction Studies’ rigor forces it, despite itself, to give up on the tradi-
tional, scientific conception of extinction; to give up on the difference
between collective death and extinction; to give up on the specificity of
extinction. If entanglements rather than discrete species are the units
of life, how can we maintain a concept of extinction that is irreducible
to death and destruction?
Later in Extinction Studies, James Hatley perceptively questions
whether “extinction” is the right word when discussing the extinct
ˉ
Japanese wolf, Okami: “Indeed, one is called on to witness that Oka-ˉ
mi’s disappearance not only is to be questioned but already is a ques-
tioning, uncannily interrogating we who remain behind. Is ‘extinction’
even the right word to turn to…?”20 If we—bioculturally entangled with
ˉOkami—remain afterwards to witness it and inscribe it and respond to
it, then perhaps the concept of extinction won’t do; for, Okamiˉ is not
entirely gone. Insofar as Extinction Studies tends to have its eye on
extinction today—on extinction as we register it and thereby bring it
back into the play of life and death—we need, Hatley suggests, a new
term. “Extinction is a necessary thought, but it is not the best thought
ˉ
in regard to Okami here and now. […] [M]y very rendering of Okamiˉ
as ‘extinct’ finds that it is already undermined by the ongoing naming
ˉ
of Okami.” 21
By extending extinction beyond its paleontological and
evolutionary domain and into a cultural one, Hatley finds himself
in need of a different concept—one that falls between extinction and
memory, between the loss of genetic memory and the life-support of
928 Theory & Event
cultural memory, one that can simultaneously countenance the scien-

tific extinction concept as well as its deconstruction.
I mean to highlight that insofar as Hatley deconstructs extinction
(in order to make extinction matter, to heed its complexity, to respond
ethically), he struggles to maintain the concept. I mean to highlight,
that is, something that Extinction Studies seldom comes out and says:
insofar as we deconstruct extinction, it’s not entirely obvious what
we’re left with, what extinction means anymore. The very reality of
extinction comes into crisis, as does the truth and utility of the term.
My reader, no doubt, may very well ask: “Isn’t extinction, like any and
every process or event, both its reality and a complexity that refuses
to congeal into an ontic simplicity, both its concept and the disartic-
ulation of that concept? Isn’t extinction both irreducible to collective
death and a form of eradication that cannot rigorously be kept separate
from collective death?”
I cannot disagree: there’s no reason that we cannot maintain the
traditionally-received fact of extinction and deconstruct this fact;
no reason we cannot say that extinction names the death of the last
member of a species and an eradication that refuses such delimitation.
But what would be the fate of the concept of extinction here? Would it
be relegated to a technical scientific term that, in the context of global
unwindings and untanglings, has marginal importance? Would mass
unraveling displace the old concept of mass extinction? Perhaps it
ought. Perhaps, as Richard Grusin puts it, “the concept’s theological
origins still inflect current understandings of extinction […and] the
very idea of extinction bear[s] traces of an ontology that is alien to
natural, social, and human scientists in the twenty-first century.”22
Nonetheless, I will argue in this paper that the deconstruction of
extinction risks bypassing too quickly the potential insights that might
be waiting within the more traditional concept of extinction in pale-
ontological theory. I will argue that the threat of extinction precisely
exceeds—indeed, attacks—the generative play of life and death, of
life-death or survivance, as Derrida would have it. The traditional
concept of extinction (the death of the last organism of a species), once
unpacked, puts pressure precisely on our most rigorous paradigms of
survival.
My gamble is that if we are going to affirm and explore the differ-
ence between death and extinction, then we have to do so from within
the notion of species. After all, what separates extinction from death is
that it happens to species. No matter how complex the species ques-
tion may be, an investigation of extinction should start with the tradi-
tional phylogenetic conception of species, for that is the horizon of our
inherited meaning of “extinction.”23 If the event of extinction is distinct
from collective death, it is so insofar as it happens to species. As such,
to understand extinction today—and to understand how the history
of anthropogenic extinction may scramble its “natural” manifestation

and thus transform (what we have taken to be) its concept or reality—
we first have to unpack the basic, scientific definition.
I ask, in the old language of definitions: What is the difference
between collective death and extinction? What do we mean by, and
what is, “the extinction of a species”? More emphatically, what is the
difference between evolution and extinction? That is, what does it
mean to no longer survive? What does it mean to no longer evolve, to
no longer survive as other, as mutant?
The challenge that extinction poses, I will argue, is perhaps greater
than we imagined. For it challenges even and especially our most
nuanced, anti-metaphysical thinking. Instead of using the deconstruc-
tion of life and death to rearticulate extinction as a form of survival, I
will argue in this paper that extinction undermines our very under-
standing of traces, remainders, constitutive entanglements, surviv-
ance, etc. It figures the remainderless—an almost unintelligible notion
(any child can tell you that destructions leave remainders).
How to label the strategies of reading and argument that follow?
Rather than investigating the social, material, and epistemological
formation of the fact of extinction (à la Latour), the cultural responses
to and guiding frames of this fact (à la Heise), the specular-specula-
tive programming of human auto-extinction (à la Colebrook), or the
articulation of extinction as a form of survival (à la Van Dooren and
other researchers in Extinction Studies), I try to immerse myself in the
empirical findings and deductive explanations of paleontological and
evolutionary literature. By treating this literature as a text with its own
unthought assumptions/insights, I tease out an underlying conceptu-
alization of extinction as remainderless. I then ask: how do biologists
and paleontologists differentiate between evolution and extinction,
between surviving-as-other and complete eradication? My method-
ological strategy will be somewhere between scientific explication,
philosophical argumentation, and an almost literary reading of scien-
tific research.
With no small degree of apprehension, I will be more doggedly
traditionalist than contemporary sciences and will completely bracket
anthropogenic mass extinction. I do this because, before looking at
today’s extinctions, we need to pay heed to the general concept of
extinction, however inflated, idealized, and artefactual its objectivity
and generality may be.
Extinction is Remainderless
In the biological and paleontological sciences, extinction is the end
of a species or of a clade (i.e., a group of species on the tree of life
930 Theory & Event
with a single ancestor). I will use the term “taxon” to refer to both.
Extinction occurs the moment the last living organism of a taxon dies.
Paleontologist Paul Taylor writes: “Extinction is quite simply the
‘death’ of a taxon. The extinction of a species occurs when the last indi-
vidual of that species dies.”24 Such is the dominant definition in the
scientific literature today.25 Of course, things will prove a little more
complicated.
In one of the better attempts to bridge paleontology (the study
of the history of life) with biology (the study of the structure of life),
Jeffrey Levinton concedes that extinction “is the province of the pale-
ontologist.”26 After all, what is is not extinct. Indeed, our scientific
understanding of extinction comes not from empirical observations of
the present world, but from the fossil record of the deep past. Without
fossils of nonextant species, our understanding of extinction would
be founded upon hearsay (or, at the very least, we would only have
knowledge of extinction as a contemporary phenomenon, as an effect
of human history, and not of extinction in general).
But while the fossil record proves the fact of extinction, it fails
to archive the event of extinction. The fossil record does not archive
the defining moment when “the last individual of [a] species dies.”
In paleontological literature, the “Signor-Lipps effect” insists that the
real moment of extinction and its archived date do not coincide. As
Taylor explains: since the fossil record is overwhelmingly incomplete,
“[t]he last appearance of a species (or genus, family, etc.) in the fossil
record seldom coincides with the time of its extinction. Instead, the
last appearance will always precede the true time of extinction.”27
Benton and Harper are more emphatic: “you never find the last fossil
of a species” (my italics).28 Although it is technically possible that the
last living organism will, by a magnificently rare chance, happen to
fossilize, scientists’ confidence is not unjustified: the likelihood that
we will find the fossil of the very last member of a taxon is analo-
gous to the likelihood that life will be eradicated tomorrow. Indeed,
the absence of the moment of extinction in the archive (i.e., the fossil
of the last individual) does not result from our inability to access the
entirety of the fossil record; rather, the fossil record is itself the tiniest,
faintest trace of past life.29 Even if we were to survey Earth’s fossils
from an inhuman and entirely omniscient perspective, there would
most probably still be no fossils of the last members of taxa. (Again, I
am excepting today’s extinction crisis, where we can witness the death
of the last member of a taxon.)30
Because of the contingencies of fossilization, extinction is remain-
derless in fact. More fundamentally, though, it is remainderless in prin-
ciple. Even if the last surviving member of a taxon happened to leave
a fossil behind, the extinction would still be uninscribed, unarchived.
For, on its own, a fossil only archives the past life of an individual
organism. Fossils, that is, are not the signs or remains of extinct taxa,
but of deceased individuals. The fossils of dinosaurs, for example, are
the fossils of bodies, not of a taxon or its extinction (as a taxon, dino-
saurs are not extinct). Extinction happens to entire taxa, and unlike the
bodies of organisms, taxa do not leave fossils behind.
That extinction is absent from the fossil record (which comprises
only of the bones or imprints of bodies) is perhaps too obvious,
which is why, perhaps, the scientific research says it without giving
it much thought. Nevertheless, according to paleontological research,
the moment of extinction (the death of the last organism of a taxon)
is uninscribed. Both in fact and in principle, it is not inscribed in the
spatial layers of rock that make up the basis of paleontological study.
In the only archive (i.e., the fossil record) where we might locate extinc-
tion, the event of extinction leaves no remainder, no fossil. Extinction
is a temporal event that does not leave behind a spatially inscribed
remainder. An archive of extinction would require the record of an
absence that cannot be recorded through what it leaves behind. As
such, to aestheticize extinction as its fossil remainders is to misun-
derstand it entirely. To figure extinction through its traces or remains,
through the bones of the extinct, is to assimilate extinction to survival
and confuse extinction with death.
We’ve seen that, because of the incompleteness of the fossil record,
extinction occurs after the last extant fossil. Less intuitively, there is
a sense in which extinction occurs before the death of the last living
organism of a taxon. Scientists use the phrases “dead clade walking”
and “extinction debt” to name “species committed to eventual extinc-
tion following a forcing event,” such as geochemical upheaval, sea
level rise, or bolide impact.31 Of course, the eradication of a taxon’s
conditions of survival cannot program extinction in a deterministic
manner. An extinction-bound taxon can always happen to develop a
beneficial adaptation; or, conditions can change to its benefit. When
scientists speak of extinction debt and dead clade walking, they are
speaking of probabilities rather than biological certainties.
However, once there is enough discord between a taxon’s phys-
iology and its external conditions—and once the taxon has been
reduced to a “minimal viable population”—it makes sense to say that
the taxon goes extinct before the event. The phenomenon of “dead
clade walking” is best illustrated in miniature: if only one organism of
a sexually reproducing species remains, then the species has already
gone extinct in principle (it would be ludicrous to imagine that the
last member of a sexually reproducing species suddenly develops the
means to reproduce asexually). When the final remaining member
(bereft of a sexual partner, of its conditions of reproduction) dies, the
taxon has already, for all intents and purposes, been extinct. Extinction
happens in principle before the death of the last remaining organism.
932 Theory & Event
Indeed, and interestingly, taxa do not grow old and die. They do
not irrevocably and necessarily deteriorate from internal senescence
as individual organisms do. The universally cited study here is Van
Valen’s 1973 finding that the age of a taxon has no bearing on its like-
lihood of extinction.32 Rather, extinction occurs when the biotic and
abiotic structures on which a taxon relies transform too fast for it to
adapt. Most unlike death, which can happen by external accident but
would otherwise and inevitably happen by internal decay, extinction is
not a function of aging. Instead of going extinct because they march
through time, taxa go extinct when their conditions of survival suddenly
change. At a moment (but at no moment in particular) prior to the death
of the last surviving organism, taxa go extinct in advance, following a
structural transformation of their requisite conditions.
Because it is remainderless, extinction can only be registered some-
where after the last remaining fossil; and because it strikes a taxon’s
conditions of the survival rather than being necessitated by the taxon’s
aging, the death of the last member is the delayed “effect” of an extinc-
tion that has already occurred, as it were, at some unspecific prior
moment. These temporal peculiarities certainly complicate Taylor’s
definition (“Extinction is quite simply the ‘death’ of a taxon. The
extinction of a species occurs when the last individual of that species
dies”). But there’s a more fundamental way in which the basic paleon-
tological and evolutionary literature undermines its own definition of
extinction, thereby pointing us toward its own insights.
Obviously, what goes extinct does not evolve, and what evolves
is not extinct. The less-than-intuitive twist is that a species can be
completely wiped out—each and every member dead—but not be
extinct. In phylogenetic evolution, a species can branch into two new
species. If the parent species disappears but not the daughter species,
extinction has not occurred. Even if the parent species is no longer
extant—even if the last member of the parent species has died—the
species is not extinct: it has evolved into two new species. This evolu-
tionary eradication of a species is termed “pseudo-extinction,” which
has to do with speciation, the very opposite of extinction.33 The death
of the last member of a taxon is not necessarily the extinction of that
taxon. It might be its evolution.
The eradication of a taxon counts as an extinction if and only if it
is also the eradication of every phylogenetic mutation of that taxon.
Extinction occurs if the last member of a taxon dies and if that taxon
leaves behind no daughter species. For a taxon to be extinct, it is not
enough that it does not exist: no population (however small) that
might have evolved into a different species, whether a minute or a
millennium before, and perhaps on the other side of the world, can
survive. Extinction precludes not only surviving, but also and espe-
cially surviving as other, as mutated or evolved. At stake is the eradi-
cation not simply of a taxon, but of its evolution, of its ability to survive
as new species, as ghosts of itself.
The figuration of extinction I am teasing out is quite different than
that which dominates in Extinction Studies. Instead of viewing extinc-
tion as a death that haunts, the buried arguments of the biological and
paleontological literature view it as a remainderless eradication of the
constitutive condition of surviving as other.
Evolution as Survival, Remaining as Remains

Evolution is about survival; extinction about a strange cessation of
survival. It is thus necessary to juxtapose the two. In this section I
will provide a reading of evolution and draw out what I take to be
an uncontroversial idea: insofar as something evolves, it does not live
as itself, but only and ever as a mutant of “itself,” as the remainder of
its own death. Following the work of Catherine Malabou, Francisco
Vitale, and others on deconstruction and evolution, I mean to suggest
that natural selection, in its most rigorous formulation, can be read as
another name for the différant play of survival prior to life and death.34
I will ultimately argue that if extinction is the extinction of a taxon’s
evolution, then it has to be understood as the eradication of the very
mode of survival that constitutes life as afterlife. The disappearance of
ghosts, not their production or evolution, is at stake.
A picture of evolution begins with genes (delimited sequences of
nucleic acids in a DNA strand). Genes are transcribed as RNA, which
are translated into proteins, which in turn are expressed as pheno-
typic variations (observable chemical, morphological, physiolog-
ical, or behavioral features—big leaves or small ones, for example).
Phenotypic variations are selected or “compete.” If, differentially, indi-
viduals with certain amalgamations of phenotypes reproduce more
successfully and die less often than individuals with others, then those
amalgamations are “selected.”
But, beginning with the transition from gene to phenotype, things
are a little more complicated. The phenotypic expression of a gene also
depends upon: the extra-genetic material enveloping the gene (e.g.,
methylation); the gene’s combination with other variations of that gene
(e.g., homozygous or heterozygous); the interaction of the proteins
coded by a gene with the quantities, densities, frequencies, and combi-
nations of proteins coded by different genes (e.g., multiple loci inter-
action); and the chemical and ecological inputs from the external envi-
ronment that can determine whether a phenotype will be expressed at
all and thus make possible the very expression of phenotypic traits (the
gene-environment interaction). Like the unit of communication (the
signifier’s expression of a signified), the unit of evolution (the pheno-
typic expression of a gene) does not have its essence within itself, but
934 Theory & Event
in and as the networks from which it emanates. Which is to say that

there is no simple code through which a gene translates into a pheno-
type. Which is to say that the expression of a gene is not quite what it
is. The phenotypic expression of a gene is its exteriority, its contextual
field of differences and interactions.
And yet, while the relationship between a gene and its phenotype
is entirely dependent upon and dispersed in its singular context, this
relationship nonetheless repeats outside of that context. In infrequent
but illustrative circumstances, genes can be removed from a body,
inserted into a member of a different species, and produce the very
same phenotype. If genes didn’t have this ability to follow a simple
code regardless of context, then genetic code would be a nonrepeat-
able, noncodified mush (a private language, which in principle is no
language at all). Indeed, for genetic code to be a code, it must first and
foremost be repeatable, able to be passed on from generation to gener-
ation. To appreciate the importance of the machinic replication of this
code, one only need look at the marvelous proofreading mechanisms
on the molecular level. In fact, evolutionary life probably evolved not
with the emergence of repeatable code, but with the emergence of a
certain degree of repetition’s machinelike accuracy: less-than-precise
self-replicating RNA doubled and evolved into a structure far more
conducive to accurate replication: DNA.
But while machinic repetition is the basis of genetic code, so is
random mutation. Without being able to err—to repeat with a differ-
ence, as a variation of itself—genetic code would remain the same in
any environmental context. Without being able to err, life would have
ceased with the first environmental changes. As such, the ceaseless
exposure to random error paradoxically makes the replication of genetic
code and thus the code itself possible in the first place. Evolution, then,
simultaneously requires that replication be as unfailing as possible and
that this replication fail. The failure of the code necessary for life is
necessary for life—which is to say that life becomes life only when it
has already suffered the duplicity of its origin.
Genetic code (the play between molecular context and the ability
to repeat out of context; the play between machinic accuracy and
mutant erring) is one condition of natural selection. The other is the
environmental context. But just as code is not quite what it is (i.e., a
simple code regulating the passage from genotype to phenotype),
neither is environmental context. If context were static, populations
would eventually become perfectly harmonious with their environ-
ments, thus bringing evolutionary life to an end. Rather than a back-
ground to be teleologically adapted to, environmental context is a
transforming complex of interpenetrating yet discontinuous chemical,
genetic, ecological, and geological levels.
At the same time, the environment context must be static enough

that the selection of one phenotype over another amounts to direc-
tionality, however local and short. For, if the environment were not
sufficiently stable, variations would not have the time to be privi-
leged, deleted, or disseminated—which is to say that there would be
no adaptation. As the genetic code is doubled between the machinic
repetition and random error, the environment, too, is doubled: it must
simultaneously be conceived of as the background anchoring selection
and also as a complexity that cannot and does not coagulate into a
mere background.
Natural selection occurs with the meeting of doubled code and
doubled context. When, in redoubled environmental circumstances,
individuals with one randomly produced phenotype (big leaves, say)
happen to survive longer and reproduce more frequently or abun-
dantly than individuals with another (small ones), that phenotype is
selected, and the population as a whole evolves. As I’ve emphasized,
phenotypic traits are inscribed, repeated, and disseminated not based
on their internal value, but based on how well they replicate when
statistically and differentially compared to other variations of that
trait in contingent conditions. Such is natural selection: the differential
dissemination of one variation over others at the dynamic collision of
code and context. Nothing adapts, if “to adapt” means the autogene-
rated unification of a population to its context. There is only a play of
differences.
From the meeting of redoubled code and redoubled context, the
very possibility of life arises: the phenotype. A phenotype is a peculiar
thing. Every phenotype is a “phenotypic variation.” It “is” its differ-
ence from other variations. Otherwise, it would be a character (the fact
of having a leaf, say, of being leaved) and not a phenotype (big leaves
or small). “Phenotype” names the variation of a trait rather than a trait
itself. It is a virtual thing. Indeed, as an emergent property irreducible
to code or context, it is ghostly in its reality: the virtualization of the
encounter of a repeatable genetic variation with its contingent envi-
ronment, the virtualization of what it is not (neither gene nor environ-
ment, code nor context). Moreover, phenotypes are both selected by
their environment (already there, genetically coded) and created by
that environment (without selection in an environment, a phenotype
is singular error).
Genes and phenotypes, codes and contexts, survive as their own
erasure, traces of the impossibility of being what they are. To take
natural selection seriously is to insist that, scientifically, the scientific
terms don’t hold. Indeed, if natural selection occurs with the meeting
of genetic code and environmental context, it simultaneously scram-
bles this very dichotomy. While it may be intuitive to think of genetic
code as internal and environment as exterior, we can just as well
936 Theory & Event
think of genetic code as an externality disseminated throughout the

biosphere and environmental context as the internal space and time of
a single vector. Likewise, while it is intuitive to think of genetic code
as a repetitive machine and its encounter with its environment as a
singular event, we can just as well think of random genetic mutations
as events and the ecological context as mechanically selecting those
mutations. Natural selection names both the institution and the disar-
ticulation of the difference between the twohanded genetic system and
the redoubled environmental one.
Life, then, emerges from and as the meeting of two dynamic
systems that are, strictly speaking, neither living nor evolving. The
environmental context of a population is not, qua context, evolu-
tionary life; and genetic code without an environment is merely a
nonevolving machinics. Evolution, in this sense, is a precipitate of the
meeting of dead code and dead context. It is a precipitate, moreover,
that retroactively opens the difference between them (for, without the
selection of code in a context, there is no difference between the two,
and thus no code and no context). Evolution opens the space and time
of life: “space” amounts to the constitutive supplementation of dead
code by dead context (or vice versa) and thus temporal deferral, and
“time” amounts to the incorporation of error and thus spatial inscrip-
tion. According to evolutionary thought, to live is “to be” the uncanny
ghost of the play of death.
It is worth remembering that lives—the breathing or photosynthe-
sizing beings surrounding us—do not evolve (only populations do).
Lives, in this sense, are the “effect” of the movement of life, which
cannot thereby be posited as some transcendental over and above
those lives. A living being survives (as) its severance from the move-
ment of life. Things become even more complicated when we move
from single beings and populations to entire taxa, but I will cut myself
short. In this section of my essay, I mean only to have read evolution,
in the most scientific terms, as a movement of différant survivance. So
much for evolution. What, then, of extinction?
Evolution and Extinction

If to evolve is to survive as a ghost, what does it mean for a ghost
to “die’? If to evolve is to survive as remains, what does it mean to
be eradicated without remainder? How does the scientific literature
confront the problem of extinction vis-à-vis evolution?
No matter how prevalent it is in the media, extinction is an ill-un-
derstood event. Paleontologist Donald Prothero writes: “Despite its
importance to life’s history, there has been surprisingly little research
on extinction. Biologists have spent decades and written hundreds
of books on how species evolve, but not on how they disappear […]
Although this has changed since 1980 […] extinction theory is still
not a major part of evolutionary biology.”35 According to Levinton,
“we know very little about the extinction of species, expect by fairly
obvious mechanisms such as hunting.”36 The recent Princeton Guide to
Evolution, for example, contains nine chapters on speciation and one
on extinction.37 We know so little about extinction, I would suggest,
precisely because of extinction’s peculiar evolutionary status: remain-
derless eradication does not sit well with the theory of surviving as
remains.
Unsurprisingly, the questions driving contemporary extinc-
tion research are defined by this very evolutionary-nonevolutionary
divide. To what extent should we imagine extinction as triggered by
shocks that interrupt the normal conditions of evolutionary competi-
tion? Conversely, to what extent should we imagine extinction as the
accumulation of this competition?38 How should we measure extinc-
tion? By the gross loss of species in disregard to evolutionary history,
or by the amount of evolutionary history that is lost?39 Is the global
decrease of extinction rates due to evolutionary patterns or abiotic
contingencies?40
During most of the twentieth century, biologists had not seen in
extinction a great challenge to evolutionary theory. Although natural
selection properly pertains to the differential reproduction and death
of organismal phenotypes, biologists were and still are keen to extend
it: just as organisms die and reproduce, species go extinct and speciate
(split into or “give birth” to two species). Such was Darwin’s thought:
“as new species in the course of time are formed through natural selec-
tion, others will become rarer and rarer, and finally extinct. The forms
which stand in closest competition with those undergoing modification
and improvement, will naturally suffer the most.”41 When two species
or taxa, like two individuals, are competing for the same resources,
one of them will eventually develop a beneficial mutation and outper-
form the other, slowly driving it to extinction. According to this line of
reasoning, extinction seems perfectly consistent with natural selection.
Such is the Red Queen hypothesis, where species constantly run in
place: as one species outperforms a competitor through a beneficial
mutation, the competitor does the same, leading to an arms race of
adaptation. If extinction results, then it does so through natural selec-
tion, through the failure to adapt.42
But while this might be theoretically and analogically convincing,
it has been persistently disputed. One of the twentieth century’s most
prominent scholars of extinction, paleontologist David Raup, high-
lights the difference between extinction and the evolutionary system
of traces as follows:
938 Theory & Event
The notion that slow, gentle pressure produces extinction is part of

the Darwinian paradigm. In The Origin of Species, Darwin used the
metaphor of a log of wood with many wedges driven into its sur-
face. Newly driven wedges were the newly evolved species. With
the crowding of wedges (species), each new wedge displaced and
expelled old ones from the log. The clear implication is that gentle
pressure exerted by new—and better-adapted—species leads to
the extinction of one or more incumbent species. This is appealing
and has been learned by generations of biology students. But its
verification from actual field data is negligible.43
In contrast to evolutionary biology’s explanation of extinction, pale-

ontologists have questioned whether a beneficial adaptation in one
species is the mechanism that leads to another’s extinction. According
to the data, they assert, a species may have to adapt, exploit new
resources, or migrate when outcompeted for a limited resource, but
it will not usually go extinct. Outperformed taxa prove surprisingly
resilient and plastic.44 As Prothero puts it, “Many of the classic stories
of competition and extinction in the fossil record do not hold up under
close scrutiny.”45 In The Princeton Encyclopedia of Evolution, we read
that “new species can almost always be added to natural communities
without resulting in extinction.”46
For paleontology, taxa do not go extinct because they fail to adapt
to the slow and constant transformations of their environment, nor
because they are differentially outperformed by their competitors’
adaptations. Rather, sudden disturbances in, say, the ocean’s oxygen
content randomly kill off taxa without regard to their evolutionary
or adaptive success. Paleontology thus explains extinction through
discrete events that act on—rather than continuously result from—
evolutionary competition and adaptation. Extinction “requires an
environmental shock (physical or biological) which is not normally
encountered during the geological life spans of such species or
groups, and the shock must be applied rapidly enough over a broad
geographic area to prevent adaptation by natural selection or escape
by migration.”47 It does not matter how well a taxon was differentially
performing in its local environment: the rapid and structural recon-
figuration of that environment indiscriminately kills off the most and
least successful. Being adaptively successful is immaterial. Extinction
comes, as it were, from above and outside natural selection.48 Such is
the Court Jester model, where a wild card is thrown into the mix, inter-
rupting and indifferent to running queens.49
But even if paleontologists have challenged the understanding of
extinction as a properly evolutionary phenomenon,50 they have none-
theless retained the evolutionary paradigm (evolution is, after all, the
unifying principle of genesis and structure in the life sciences). While
paleontologists generally refuse to view extinction as the effect of the

microevolutionary selection of individuals or species in a local context,
they proffer it as an effect of macroevolutionary taxon selection.51 Taxa
possess characteristics (e.g., geographic range) that cannot be reduced to
the cumulative effect of phenotypes (e.g., distance of a single organism’s
travel): the organisms of a taxon may barely travel, but the taxon’s
geographic range may be extensive. Now, taxa inhabiting a wide range
will, compared to taxa with a restricted one, speciate more frequently
and go extinct less often. At stake is not how well taxa perform in their
local context, but whether they possess characteristics (geographic
range, clade size and density, turnover rate, latitude, trophic position,
endemism, average body size, evolvability, and specialization) that
make them, in the statistical context of the totality of taxa across the
planet, more resistant or susceptible to shocks. Accordingly, so long as
evolution pertains not only to the differential selection of organismal
phenotypes in a local environment, but also to the differential selec-
tion of taxic characteristics in the global context, then extinction would
seem to prove no problem for evolution.
There is no question that both microevolutionary and macroevo-
lutionary selection count as evolution. Yet, the difference in prefix is
revealing: whereas the death of individual organisms (in microevolu-
tion) is a condition of possibility of survival, such cannot be said of
the extinction of taxa (in macroevolution). Only microevolution (the
differential erasure and iteration of virtual phenotypes through the
interfacing of code and context) is a condition of possibility of life—of
remaining as remains. In contrast, the differential speciation and extinc-
tion of taxic characteristics is not a constitutive element of life. It is
entirely imaginable that life could originate and diversify, continually
filling finer and finer ecospaces, without extinction. As Raup puts it,
there is not “any known reason why a species could not live forever…
one can imagine an evolutionary system organized without extinc-
tion—and this may exist on planets elsewhere.”52 While speciation is
an inherent consequence and necessary condition of life, extinction,
as far as we know, is not. As such, even if extinction is meaningless if
abstracted from evolution (for only the phylogenetically evolving can
go extinct—otherwise, extinction would be little more than collective
death), it is not comprehensible through the différant movement of
life/death.
Unlike death, extinction does not make survival possible in the
first place. For unlike death, it is not an absence constitutively virtual-
ized by and incorporated into the play of code and context, genotype
and phenotype, descent and modification. Although a supplement of
evolution—and although this supplement may shape the history of
life—extinction is not, in our paleontological knowledge, a constitutive
one.
940 Theory & Event
Like many biologists, the most astute thinkers of extinction in the

humanities have missed this strange relationship between evolution
and extinction. Too often, extinction is read as a continuation or part
of evolution. In the introduction to After Extinction, Grusin conceptu-
alizes extinction “not as an aberration but as part and parcel of the
process of natural selection.”53 Kirby reads the phylogenetic branches
cut off by extinction as the very process of survival, using Derrida’s
“nothing outside the text” to insist that there is no outside of evolu-
tionary branching:
Darwin finds life in death here, wherein broken and dead branch-
es are internal to processes of viability […] Darwin further under-
lines this sense that there is no outside branching […] In sum,
what Darwin emphasizes is that what follows from branching is
yet more branching, just as Derrida finds that the root of things is
strangely ubiquitous.54
For Kirby, extinction, the cut of the phylogenetic branch, amounts to

the very engine of evolution. In the deconstructive and evolutionary
picture alike, surviving as remains the paradigm.
But in contrast to the evolutionary and deconstructive arguments,
paleontologists implore that extinction exceeds the evolutionary
movement of life/death; it exceeds the law of surviving as other,
wherein life’s supplemental virtuality, mutation, and death supplant
its essence, thereby bringing life alive. Exceeding the constitutive and
retroactive movement of remaining as remains, extinction is precisely
the remainderless excision of a species’ différantial movement.
Extinction eradicates a taxon’s constitutive doubling and undoing
(without thereby doubling what it undoes, without thereby keeping it
alive in the process).
Conclusion: Extinction
I began this essay by considering the work of Thom van Dooren and
other contributors to the blossoming field of Extinction Studies. I
found there a deconstruction of the species concept and, as a conse-
quence, the articulation of extinction as a play of survival. I then asked
whether this deconstruction has been too quick—too quick to bypass
the traditional, scientific extinction concept, too quick to bypass the
difference between death and extinction. In the course of my argu-
ment I similarly read the evolution of species as the deconstruction of
species, wherein to evolve is to survive as a ghost. But by juxtaposing
evolutionary theory with the paleontological literature, I have arrived
at an almost opposite conclusion from Extinction Studies. The decon-
struction of the species concept does not imply the spectral survival of
the extinct; rather, extinction poses a threat to this very form survival.
In this manner, I have endeavored to distinguish between evolu-

tion and extinction, death and extinction. In the biological context,
death is the motor of life, incorporated in and as the evolution/survival
of a population. But extinction is different. It names the “death” of
death. It names the “death” of a taxon’s life/death. My argument is not
without precedent. In fact, it is made by one of the inaugural thinkers
of Extinction Studies, Deborah Bird Rose, who offers the notion of
“double death”:
So many extinctions that the process of evolution is unable to keep

up. More species die than are coming into being. The death of evo-
lution itself […] The death of temporal, fleshy, metabolic relation-
ships across generations and species.
The destruction of the future of one’s own death, which starts to

collapse along with the future of flourishing others and ecosys-
tems.
Double death is a despoiler. It smashes the relationship between

life and death, fracturing a compact that has been integral to life
on earth. The despoliation of death throws the lives of earth crea-
tures into a barren place with no future and with a rapidly unrav-
eling past.55
But while Bird Rose puts her finger on what I have been trying to put
mine on, her aim is to say “no” to extinction, to hear the call of ethics
and thus to hear the call of the extinct from beyond its buried grave.
In other words, at the very moment she registers extinction as double
death, she refuses her own insight: she reads the extinct as its remains
and implorings, as its survival. She cares too much to entertain the
impossibility of witness. Bird Rose articulates extinction as the “death”
of the play of death, but her ethical commitment of hearing the call
of the other prevents her from appreciating her own insight that the
extinct no longer remains to call. She says that extinction is the “death
of evolution itself,” proffers the thought, but then turns away from her
own radicality.
I mean to think the “death” of death, to take the unsettling pale-
ontological conceptualization of extinction seriously. Insofar as
extinction eradicates that which makes the extinct and extinction
possible—namely, evolution, non-self-coincidence, iteration as other—
it somehow refuses to fold back into life/death, where it might be kept
alive. The extinct is not structurally contingent upon the remainders it
leaves behind or upon the possibility of witness. As a general concept,
extinction is remainderless (only recently—only in today’s human
landscape—does extinction leave behind a remainder that is archived,
brought back into the play of life/death). As a general concept, extinc-
942 Theory & Event
tion is not the material’s death-by-virtualization (natural selection), nor

the virtual’s death-by-materiality (fossil remainders), but the absolute
absenting of the extinct’s constitutive excesses, mutations, and inscrip-
tions. At play is not the interruption of a system, but the short-circu-
iting of a system predicated upon its own interruptions.
The paleontological and catachrestic figure of extinction thus chal-
lenges the foundations and limits—or anti-foundations and de-limita-
tions—of poststructural and evolutionary thought. The overwhelming
axiomatic in poststructuralism, evolutionary theory, and Extinction
Studies is that every being or event is retroactively constituted by its
remains, by its survivance or evolution, even and especially the event
of death. Just like the origin, no destruction is absolute, no end final.
This has almost become something of an a priori. Ghosts cannot die
without leaving more ghosts behind and ahead: the erasure of a trace
(under erasure) leaves behind traces (under erasure). Accordingly, and
by necessity, that which has been destroyed or has disappeared will
have repeated with a difference—registered, incorporated, or tran-
scribed by a reading machine (phenomenological or semiotic, biolog-
ical or chemical). A ghost, Derrida remarks, “never dies, it remains
always to come and to come-back.”56 Martin Hägglund puts it more
forcefully, “any moment always must be recorded in order to be.”57
“There is thus no exception to the law of survival.”58 The deconstructive
articulation—in alignment with evolutionary theory and Extinction
Studies—denies the possibility of extinction.
There is truth to this denial, truth to this assertion of survival as
prior to life and death. There is truth to the argument that extinction
must be understood as a movement of survival. After all, when a group
of organisms (taxon or not) dies and leaves an empty ecospace behind,
the traces of the group’s absence remain, surviving as its effects,
affecting the ecosystem. While this mode of survival is not specific to
extinction and does not distinguish extinction from collective death,
it certainly applies to extinction. Such cannot be denied. And from
this perspective—wherein the deferrals of evolution interface with the
dispersals of ecology—the extinct survives.
In a sense, the issue is a question of where to begin. Scholars asso-
ciated with Extinction Studies tend to begin with sympoetic systems
of ecological entanglement, and they tend to begin with the context of
anthropogenic extinctions and the entwinement of nature and culture.
In contrast to the ecological entanglement of species, I’ve started with
the phylogenetic species concept. And in contrast to the entwinement
of extinction and cultural networks, I’ve started with an idealized
notion of extinction in disregard to humanity. From these two starting
points, two radically divergent conclusions are reached: survival on
the one hand and remainderless eradication on the other.
Importantly, there’s no reason we can’t have it both ways. I would
insist, though, that we would thereby have to have it both ways.
We would not only have to open extinction up to its survivance (as

Extinction Studies would have it); we would also have to conceive of
it as a radical end. Extinction as its deconstruction, as a movement of
evolution and remains/remaining; but also extinction as the remain-
derless. Extinction as its deconstruction, as a form of ghosting; but also
extinction as a threat to that deconstruction.
To say we can have it both ways is not, however, to say that these
two ways are congruent or harmonious. For, to differentiate collective
death from extinction—to understand extinction as a remainderless
eradication of that which survives as remains—is inevitably to fall afoul
of our most rigorous, anti-metaphysical conception of survivance. To
understand extinction as remainderless is inevitably to challenge our
poststructural predilection that eradications leave retroactively consti-
tutive traces. This is not an easy task. After all, how can we think the
remainderless eradication of that which constitutively spreads beyond
any spatial or temporal border? How can we think the extinction of a
taxon if we cannot think of a taxon as an ontologically congealed type
susceptible to eradication? To think extinction as remainderless would
require a figuration that evolutionary, poststructural, and everyday
thinking prohibits (again, any child can tell you that destruction leaves
remainders).
Perhaps, though, such a figuration is necessary when our contem-
porary thinking remains so impotent in the face of its own cataclysmic
effects. Maybe it’s worth giving credence to the insights of the pale-
ontological thinking of extinction, no matter how inconceivable. For
maybe they address the unthinkability of extinction for contempo-
rary thought. Maybe extinction compels us—if not to simply reject its
deconstruction—to differentiate it from death. If there is extinction, it
leaves traces (no denying it), but (I’ve argued) it leaves none.
A final question. A question I’ve been hinting at, preparing for. I
have thus far ignored contemporary extinctions. But what would the
figure of the remainderless teach us about extinctions today—today,
when we can bear extinction with us, carrying it along into the deathly
archives that keep it alive, witnessing it? Can we, as Hatley asked,
call contemporary extinctions “extinctions”? The current extinction
crisis is marked not only by the acceleration of extinction, but also by
the disappearance of extinction, by the appropriation of extinction as
survival. Extinction is itself going extinct. Perhaps, then, extinction,
today, is at risk—in addition to survival.
Notes
1. See, for instance, Claire Colebrook, Death of the PostHuman: Essays on
Extinction, Vol. 1 (Ann Arbor, MI: Open Humanities Press with Michigan
Publishing, 2014).
2. The literature is extensive. See, for example, Jason W. Moore, ed.,
Anthropocene or Capitalocene? Nature, History, and the Crisis of Capitalism
944 Theory & Event
(Oakland, CA: PM Press, 2016); Dipesh Chakrabarty, “Anthropocene Time,”

History and Theory 57, no. 1 (2018): 5–32; Timothy Clark, Ecocriticism at the
Edge: The Anthropocene as a Threshold Concept (London: Bloomsbury, 2015);
Tom Cohen, Claire Colebrook, and J. Hillis Miller, Twilight of Anthropocene
Idols (London: Open Humanities Press, 2016).
3. Thom Van Dooren, Flight Ways: Life and Loss at the Edge of Extinction (New
York: Columbia University Press, 2014), 58.
4. See Donna Haraway, Staying with the Trouble: Making Kin in the Chthulucene
(Durham, NC, and London: Duke University Press, 2016); Eben Kirksey,
ed., The Multispecies Salon (Durham, NC, and London: Duke University
Press, 2014); Van Dooren, Eben Kirksey, and Ursula Münster, “Multispecies
Studies: Cultivating an Arts of Attentiveness,” Environmental Humanities
8, no. 1 (2016): 1–23; Scott F. Gilbert, Jan Sapp, and Alfred I. Tauber, “A
Symbiotic View of Life: We Have Never Been Individuals,” The Quarterly
Review of Biology 87, no. 4 (2012): 325–341.
5. Anna Tsing, Heather Swanson, Elaine Gan, and Nils Bubandt, eds., Arts of
Living on a Damaged Planet (Minneapolis, London: University of Minnesota
Press, 2017), G4.
6. Tsing, Swanson, Gan, and Bubandt, Arts of Living on a Damaged Planet, M141.
7. Haraway, Staying with the Trouble, 59.
8. Van Dooren, Flight Ways, 38–9.
9. Van Dooren, Flight Ways, 46.
10. Van Dooren, “Spectral Crows in Hawai’i: Conservation and The Work of
Inheritance,” in Extinction Studies: Stories of Time, Death, and Generations,
eds. Deborah Bird Rose, Thom van Dooren, and Matthew Chrulew (New
York: Columbia University Press, 2017), 190–93.
11. Rose, van Dooren, and Chrulew, Introduction to Extinction Studies, 8.
12. Tsing, Swanson, Gan, and Bubandt, Arts of Living on a Damaged Planet,
G4–6.
13. Richard Grusin, Introduction to After Extinction (Minneapolis, London:
University of Minnesota Press, 2018), xi.
14. Cary Wolfe, “Condors at the End of the World,” in After Extinction, ed.
Grusin, 117.
15. Jacques Derrida, Learning to Live Finally, trans. Pascale-Anne Brault and
Michael Naas (Hoboken, NJ: Melville House, 2007), 26.
16. Jacques Derrida, Learning to Live Finally, 26.
17. Vicki Kirby, “Un/Limited Ecologies,” in Eco-Deconstruction: Derrida and
Environmental Philosophy, eds. Matthias Fritsch, Philippe Lynes, and David
Wood (New York: Fordham University Press, 2018), 129.
18. Bird, van Dooren, and Chrulew, Extinction Studies, 5.
19. Without supplying a barrage of evidence, I would suggest that the
tendency to slip from the language of extinction to the language of death
is pervasive. The difference between extinction and death, in other words,
remains surprisingly unarticulated, as if it were not even a question. See
Cary Wolfe’s contributions to Extinction Studies and After Extinction as
well as Philippe Lynes’s Futures of Life Death on Earth: Derrida’s General

Ecology (London, New York: Rowman and Littlefield, 2018).
ˉ
20. Hatley, “Walking with Okami, the Large-Mouthed Pure God,” in Extinction
Studies, 26.
ˉ
21. Hatley, “Walking with Okami,” 28, 35.
22. Grusin, After Extinction, x.
23. For a starting place into the species question, see Robert A. Wilson, ed.,
Species: New Interdisciplinary Essays (Cambridge, MA: MIT Press, 1999).
24. Paul Taylor, “Extinction and the Fossil Record,” in Extinctions in the
History of Life, ed. Paul Taylor (Cambridge, UK: Cambridge University
Press, 2004), 7.
25. Ecologists sometimes use “extinction” to refer to the disappearance of a
population, but such use is qualified as “local extinction.”
26. Jeffrey S. Levinton, Genetics, Paleontology, and Macroevolution, second ed.
(Cambridge, UK: Cambridge University Press, 2001) 369. See also Purvis,
“Phylogenetic Approaches to the Study of Extinction,” Annual Review of
Ecology, Evolution, and Systematics 39 (2008): 301–319.
27. Taylor, “Extinction and the Fossil Record,” in Extinctions in the History of
Life, 8.
28. Michael J. Benton and David A. T. Harper, Introduction to Paleobiology and
the Fossil Record (Oxford: Wiley-Blackwell, 2009), 27.
29. 99% of species have left behind no fossils at all, and this is to say nothing
of the last surviving organism of those species. For the relationship
between the fossil record and extinction, see Andrew B. Smith, “Large-
scale Heterogeneity of the Fossil Record,” Philosophical Transactions:
Biological Sciences 356, no. 1407 (March 2001): 351–367; P.W. Signor, “Real
and Apparent Trends in Species Richness Through Time,” in Phanerozoic
Diversity Patterns, ed. Valentine (Princeton, NJ: Princeton University
Press, 1985); and the second chapter from John Lawton and Robert
May, eds., Extinction Rates, (Oxford, UK: Oxford University Press, 1995).
Historically relevant for the debates surrounding the fossil record is: J.
John Sepkoski, Jr, Richard K. Bambach, David M. Raup, and James W.
Valentine, “Phanerozoic Marine Diversity and the Fossil Record,” Nature
293 (October 1981): 435–437.
30. The possibility of witnessing the moment of extinction is entirely unprece-
dented in life history, as is the “possibility” of de-extinction. In this sense,
the Anthropocene would not simply witness an increase in extinction
rates, but a transformation of extinction itself.
31. Stephen T. Jackson and Dov F. Sax, “Balancing Biodiversity in a Changing
Environment: Extinction Debt, Immigration and Species Turnover,” Trends
in Ecology and Evolution 25, no. 3 (Mar. 2010), 153. See also David Tilman, et
al., “Habitat Destruction and the Extinction Debt,” Nature 371 (September
1994): 65–66; and David Jablonski, “Survival Without Recovery After
Mass Extinctions,” Proceedings of the National Academy of the Sciences 99.12
(Jun. 2002): 8139–8144. “Extinction debt” generally names a probability
based on population dynamics; “dead clade walking” generally names
particular taxa in the Phanerozoic fossil record.
946 Theory & Event
32. Leigh van Valen, “A New Evolutionary Law,” Evolutionary Theory 1 (1973):
1–30. Younger taxa, though, are generally at greater risk for extinction. See
George E. Boyajian, “Taxon Age and Selectivity of Extinction,” Paleobiology
17, no. 1(Winter 1991): 49–57; and Seth Finnegan, Jonathan L. Payne, and
Stephen C. Wang, “The Red Queen Revisited: Reevaluating the Age
Selectivity of Phanerozoic Marine Genus Extinctions,” Paleobiology 34, no.
3 (Summer 2008): 318–341.
33. See Michael J. Benton, “Causes and Consequences of Extinction,” in
The Princeton Guide to Evolution, ed. Jonathan B. Losos (Princeton, NJ:
Princeton University Press, 2017), 582; Taylor, “Extinction and the Fossil
Record,” in Extinctions in the History of Life, 9–10.
34. In addition to Francesco Vitale’s Biodeconstruction: Jacques Derrida and
the Life Sciences, trans. Mauro Senatore (Albany, NY: SUNY Press, 2018)
and Catherine Malabou’s “Whither Materialism? Althusser/Derrida,”
in Plastic Materialities: Politics, Legality, and Metamorphosis in the Works of
Catherine Malabou (Durham, NC, and London: Duke University Press,
2014), see Colin Milburn, “Monsters in Eden: Darwin and Derrida,”
MLN 118, no. 3 (2003): 603–621 and Colm J. Kelly, “Trace, Dissemination,
“‘Survivre’: Derrida’s ‘Biology’ and the ‘Socius,”” The International Journal
of the Humanities: Annual Review 5.12 (2008): 191–200.
35. Donald R. Prothero, Bringing Fossils to Life: An Introduction to Paleobiology,
second ed. (New York: McGraw-Hill, 2004), 83–4.
36. Levinton, Genetics, Paleontology, and Macroevolution, 368.
37. Most paleontology and evolution textbooks are not much better. In some
cases, extinction is restricted to the area where we have the most knowl-
edge, the few instances of mass extinction. In other cases, extinction is
restricted to the anthropocentric perspective of the Sixth Extinction. When
extinction in general is considered, it is typically approached through
birth-death models borrowed from population ecology awkwardly fused
with evolutionary theory. In the best of treatments, textbooks present a
hodgepodge of established facts and accepted hypotheses (99% of all life
that has graced Earth is extinct; mass extinctions account for only 4% of
extinctions; extinction rates have decreased throughout the Phanerozoic;
the age of a species has no bearing on the likelihood of extinction; some
characteristics of taxa make them more susceptible to extinction, etc.). See
the sections on extinction in: Carl Zimmer and Douglas J. Emlen, Evolution:
Making Sense of Life. second ed. (New York: Freeman, 2016); Jon C. Herron
and Scott Freeman, Evolutionary Analysis, fifth ed. (London: Pearson, 2014);
Douglas J. Futuyma, Evolution (Sunderland, MA: Sinauer, 2005); Mark
Ridley, Evolution, third ed. (Oxford: Blackwell, 2004); Prothero, Bringing
Fossils to Life; Benton and Harper, Introduction to Paleobiology and the Fossil
Record; Michael Foote and Arnold I. Miller, Principles of Paleontology,
third ed., revision of Raup and Stanley’s version (New York: Freeman,
2006); Derek E. G. Briggs and Peter R. Crowther, Paleobiology II (Oxford:
Blackwell, 2001).
38. This is perhaps the central question, and the major research deserves to be
cited. See David M. Raup and John J. Sepkoski, “Mass Extinctions in the
Marine Fossil Record,” Science, New Series 215, no. 4539 (Mar. 1982): 1501–
1503; A. Hallam and P. B. Wignall, Mass Extinctions and Their Aftermaths

(Oxford, UK: Oxford University Press, 1997); David Jablonski, “Mass
Extinction and Macroevolution,” Paleobiology 31, no. 2 (2005): 192–210;
David M. Raup, “The Role of Extinction in Evolution,” Proceedings of the
National Academy of Sciences 91, no. 15 (Jul. 1994): 6758–6763; A. J. Boucot,
Evolutionary Paleontology (Amsterdam: Elsevier, 1990), 538; Richard K.
Bambach, Andrew H. Knoll, and Steven C. Wang, “Origination, Extinction,
and Mass Depletions of Marine Diversity,” Paleobiology 30, no. 4 (2004):
522–542; Wang, “On the Continuity of Background and Mass Extinction,”
Paleobiology 29.4 (Autumn 2003): 455–467; K. W. Flessa, et al., “Causes and
Consequences of Extinction: Group Report,” in Patterns and Processes in the
History of Life, ed. David M. Raup and David Jablonski (Berlin: Springer-
Verlag, 1986).
39. According to one measurement, extinction events are measured in disre-
gard to evolution by the number of species that go extinct (E) over the
total number of species (D, diversity) over the timespan of the extinction
event (T). But according to another measurement, extinction is measured
as the amount of hard-won evolutionary morphologies that are lost.
Imagine three morphologically unique clades, each with five species. If
four species from each clade go extinct (twelve species total), then no
damage is done to the evolutionary history of life, as all three unique
morphologies survive. But if one whole clade goes extinct (only five
species total) and the other two survive entirely intact, then the extinction
is far greater in terms of evolutionary history. See Sean Nee and Robert
M. May, “Extinction and the Loss of Evolution History,” Science 278, no.
5338 (Oct. 1997): 692–694; Douglas H. Erwin, “Extinction as the Loss of
Evolutionary History,” in In the Light of Evolution: Volume II: Biodiversity
and Extinction, ed. John C. Avise, Stephen P. Hubbell, and Francisco Ayala
(Washington, DC: National Academies Press, 2008).
40. See Herron and Freeman, Evolutionary Analysis, 711–12.
41. Darwin, The Origin of Species, chapter 4, “Extinction caused by Natural
Selection,” The Origin of Species by Means of Natural Selection, sixth ed.
(Project Gutenberg, 2009, http://www.gutenberg.org/files/2009/2009-
h/2009-h.htm).
42. See Michael Brockhurst, et al., “Running with the Red Queen: The Role of
Biotic Conflicts in Evolution,” Proceedings: Biological Sciences 281, no. 1797
(Dec. 2014): 1–9; Levi T. Morran, et al., “Running with the Red Queen:
Host-Parasite Coevolution Selects for Biparental Sex,” Science, New
Series 333, no. 6039 (Jul. 2011): 216–218. Related is Gause’s competitive
exclusion principle and the notion of “carrying capacity”—see Robert
H. MacArthur and Edward O. Wilson’s seminal The Theory of Island
Biogeography (Princeton, NJ: Princeton University Press, 1967, 2001);
Garrett Hardin, “The Competitive Exclusion Principle,” Science, New
Series 131, no. 3409 (Apr. 1960): 1292–1297; Robert MacArthur and Richard
Levins, “Competition, Habitat Selection, and Character Displacement,”
Proceedings of the National Academy of the Sciences 51, no. 6 (1964): 1207–
1210; Robert G. Jager, “Competitive Exclusion: Comments on Survival
and Extinction of Species,” BioScience 24, no. 1 (Jan. 1974): 33–39; Sepkoski,
948 Theory & Event
“Competition in Macroevolution: The Double Wedge Revisited,” in

Evolutionary Paleobiology, ed. Jablonski, Erwin, and Lipps (Chicago:
University of Chicago Press, 1996), 211–255. The argument has also been
presented that even mass extinction results from local, internal system
disturbances—see Kim Sneppen, et al., “Evolution as a Self-Organized
Critical Phenomenon,” Proceedings of the National Academy of the Sciences 92,
no. 11 (May 1995): 5209–5213, and Richard Solé, et al., “Self-Similarity of
Extinction Statistics in the Fossil Record, Nature 388 (Aug. 1997): 764–767.
This claim has received quite a bit of doubt (see John Alroy, “Dynamics
of Origination and Extinction in the Marine Fossil Record,” in In the Light
of Evolution: Volume II: Biodiversity and Extinction, ed. Avise, Hubbell, and
Ayala, 207–225.) An interesting and related hypothesis is that extinction
can result not from adaptive failure, but adaptive success—see E. H. Day,
X. Hua, and L. Bromham, “Is Specialization an Evolutionary Dead End?
Testing for Differences in Speciation, Extinction and Trait Transition Rates
Across Diverse Phylogenies of Specialists and Generalists,” Journal of
Evolution Biology 29 (2016): 1257–1267.
43. David M. Raup, Extinction: Bad Genes or Good Luck? (New York, London:
Norton, 1991), 185.
44. See Haley Lindsey, et al., “Evolutionary Rescue from Extinction is
Contingent on a Lower Rate of Environmental Change,” Nature 494 (Feb.
2013): 463–467.
45. Prothero, Bringing Fossils to Life, 84.
46. Luke Harmon, “Macroevolutionary Rates,” in Princeton Guide of Evolution,
ed. Losos, 572.
47. Raup, “The Role of Extinction,” Proceedings of the National Academy of
Sciences 91, no. 15 (Jul. 1994): 6762.
48. The notion of a “tipping point” is more nuanced than Raup’s shocks,
but while established in Earth sciences and discourses around climate
change, its use in extinction research is underdeveloped. The idea is that
slow, incremental changes in an ecological or biospheric system pass a
threshold of instability and produce positive feedback loops such that
that system abruptly and irreversibly reorganizes into a different state.
While taxa are ordinarily capable of surviving environmental transfor-
mations, they cannot survive a tipping point’s restructuration. See Roy
E. Plotnick and Michael L. McKinney, “Ecosystem Organization and
Extinction Dynamics,” PALAIOS 8.2 (1993): 202–212; Carlos Botero, et al.,
“Evolutionary Tipping Points in the Capacity to Adapt to Environmental
Change,” Proceedings of the National Academy of Sciences 112, no. 1 (Jan
2005): 184–189.
49. See Anthony D. Barnosky, “Distinguishing the Effects of the Red Queen
and the Court Jester on Miocene Mammal Evolution in the Northern
Rocky Mountains,” Journal of Vertebrate Paleontology 21, no. 1 (Mar. 2001):
172–185; Geerat J. Vermeij and Peter D. Roopnarine, “Reining in the
Red Queen: The Dynamics of Adaptation and Extinction Reexamined,”
Paleobiology 39, no. 4 (Fall 2013): 560–575; Thomas H. G. Ezard, Tracy
Aze, Paul H. Pearson, and Andy Purvis, “Interplay Between Changing
Climate and Species’ Ecology Drives Macroevolutionary Dynamics,”

Science, New Series 332, no. 6027 (April 2011): 349–351; Michael J. Benton,
“The Red Queen and the Court Jester: Species Diversity and the Role of
Biotic and Abiotic Factors through Time,” Science, New Series 323, no.
5915. (February 2009): 728–732; Foote, “Pulsed Origination and Extinction
in the Marine Realm,” Paleobiology 31, no. 1 (Winter 2005): 6–20. For
critical assessments of the competitive exclusion principle, see Robert
A. Armstrong and Richard McGehee, “Competitive Exclusion,” The
American Naturalist 115, no. 2 (February 1980): 151–170; Pieter J. den Boer,
“The Present Status of the Competitive Exclusion Principle,” Trends in
Ecology and Evolution 1.1 (1986): 25–28; Benton, “On the Nonprevalence of
Competitive Replacement,” in Evolutionary Paleobiology, ed. by Jablonski,
Erwin, and Lipps, 185–210; Timothy D. Walker and James W. Valentine,
“Equilibrium Models of Evolutionary Species Diversity and the Number
of Empty Niches,” The American Naturalist 124, no. 6 (December 1984):
887–899.
50. More precisely, as paleontologists admit, we do not know if natural selec-
tion explains extinction. The fossil record seldom provides sufficiently
high-resolution images for determining whether extinction is a product of
anevolutionary shock or the pressures of natural selection—or, the over-
lapping patterns and processes and degrees of each. As such, we have
to observe the contemporary world to learn about extinction up-close.
In the contemporary world, though, extinction is no longer caused by
natural selection, but almost exclusively by the anevolutionary excres-
cences of human consumption and waste. McKinney writes: “Human
impacts have been so profound that not a single case of nonanthropogenic
species extinction can be documented in the last 8000 years” (“Extinction
Vulnerability and Selectivity: Combining Ecological and Paleontological
Views,” Annual Review of Ecology and Systematics 28 [1997]: 495–516).
51. Macroevolution is a field of its own, but for important articles, see Elisabeth
S. Vrba and Stephen Jay Gould, “The Hierarchical Expansion of Sorting
and Selection,” Paleobiology 12, no. 2 (Spring 1986): 217–228; Jablonski,
“Species Selection: Theory and Data,” Annual Review of Ecology, Evolution,
and Systematics 39 (2008): 501–524; McKinney, “Extinction Vulnerability
and Selectivity”; Andy Purvis, John L. Gittleman, Guy Cowlishaw, and
Georgina M. Mace. “Predicting Extinction Risk in Declining Species,”
Proceedings: Biological Sciences 267, no. 1456 (October 2000): 1947–1952;
Stephen Jay Gould, chapter eight of Structure of Evolution Theory
(Cambridge, MA: Belknap Press of Harvard University Press, 2002).
52. Raup, Extinction: Bad Genes or Good Luck (New York, London: Norton,
1991), 6, 19.
53. Grusin, After Extinction, viii.
54. Kirby, “Un/Limited Ecologies,” in Eco-Deconstruction, 131–2.
55. Deborah Bird Rose, “Double Death,” The Multispecies Salon, WordPress,
accessed August 5, 2020, www.multispecies-salon.org/double-death/.
56. Derrida, Specters of Marx, trans. Peggy Kamuf (New York: Routledge,
1994), 123.
950 Theory & Event
57. Martin Hägglund, Radical Atheism: Derrida and the Time of Life (Stanford,
CA: Stanford University Press, 2008), 27 (italics in original).
58. Martin Hägglund, Radical Atheism, 122 (italics in original). It is not
entirely fair to suggest that remainderless eradication has not received
consideration in poststructural thought. An entire if under-appreciated
thread emerges in response to the Nazi death camps. Holocaust survi-
vors have remarked on the horror of the inability to bear witness to the
horror: to have suffered the death camps is to have “experienced” the
traumatic shattering of experience, the shattering of the ability to testify
to the experience. It is in this context that Derrida develops the figure of
ash, which is arguably a figure for the remainderless eradication of what
survives under erasure. He writes: “Ash, this is also the name of what
annihilates or threatens to destroy even the possibility of bearing witness
to the annihilation. Ash is the figure of annihilation without remainder,
without memory, or without readable or decipherable archive” (“Poetics
and Politics of Witnessing,” in Sovereignties in Question: The Poetics of Paul
Celan, trans. By Rachel Bowlby and ed. Thomas Dutoit and Outi Pasanen
[New York: Fordham University Press, 2005], 68). But despite flirting with
this idea, Derrida drops the term “ash” after 2000. Moreover, when he did
engage it, he usually could not but assimilate ash to trace, conflate extinc-
tion with survival: ash, he writes, “renders better what I meant to say with
the name of trace, namely, something that remains without remaining,
which is neither present nor absent” (“‘There is No One Narcissism’
(Autobiophotographies),” in Points… Interviews, 1974–1994, trans. Peggy
Kamuf and ed. Elisabeth Weber [Stanford, CA: Stanford University Press,
1995], 208). It is at this moment, buried deep in a footnote at the final end
of this article, that I want to give my thanks to Cathy Caruth, whose work
on trauma and erasure provided the impetus for this essay on the most
fundamental level.
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Survival and Extinction: Deconstruction, Extinction

Abstract What is extinction? What is the difference between

Key Words Deconstruction; Derrida; ecocriticism; evolution;

angle, Claire Colebrook’s writings teach us that anthropogenic extinc-

In other words, when we understand extinction as the simple passage

with others.4 Before distinct species, their relations: enfoldings and

is the loss not of a single fixed “kind,” but of a potentially limit-

their spectral presence is inscribed in the forest landscape. Plants

The bird may be gone, but it survives as a ghost, an absent presence

and entanglements, then an extinction can never be a simple absence.

fail to differentiate extinction from death more generally? By extending

cultural memory, one that can simultaneously countenance the scien-

of anthropogenic extinction may scramble its “natural” manifestation

Evolution as Survival, Remaining as Remains

in and as the networks from which it emanates. Which is to say that

At the same time, the environment context must be static enough

think of genetic code as an externality disseminated throughout the

Evolution and Extinction

The notion that slow, gentle pressure produces extinction is part of

In contrast to evolutionary biology’s explanation of extinction, pale-

paleontologists generally refuse to view extinction as the effect of the

Like many biologists, the most astute thinkers of extinction in the

For Kirby, extinction, the cut of the phylogenetic branch, amounts to

In this manner, I have endeavored to distinguish between evolu-

So many extinctions that the process of evolution is unable to keep

The destruction of the future of one’s own death, which starts to

Double death is a despoiler. It smashes the relationship between

tion is not the material’s death-by-virtualization (natural selection), nor

We would not only have to open extinction up to its survivance (as

(Oakland, CA: PM Press, 2016); Dipesh Chakrabarty, “Anthropocene Time,”

well as Philippe Lynes’s Futures of Life Death on Earth: Derrida’s General

1503; A. Hallam and P. B. Wignall, Mass Extinctions and Their Aftermaths

“Competition in Macroevolution: The Double Wedge Revisited,” in

Climate and Species’ Ecology Drives Macroevolutionary Dynamics,”

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