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School of Zoology, Tasmanian Aquaculture and Fisheries Institute, University of Tasmania, Private Bag 5, Hobart, Tas 7001,
Australia, email g.edgar@utas.edu.au
Tasmanian Aquaculture and Fisheries Institute, University of Tasmania, Nubeena Crescent, Taroona Tas 7053, Australia
Abstract: We used Tasmania as a case example to question the consensus that few marine species have
recently become extinct or are approaching extinction. Threats to marine and estuarine speciesprimarily
in the form of climate change, invasive species, fishing, and catchment dischargesare accelerating, fully
encompass species ranges, and are of sufficient magnitude to cause extinction. Our ignorance of declining
biodiversity in the marine environment largely results from an almost complete lack of systematic broad-scale
sampling and an overreliance on physicochemical data to monitor environmental trends. Population declines
for marine species approaching extinction will generally go unnoticed because of the hidden nature of their
environment and lack of quantitative data.
Key Words: climate change, effects of fishing, global warming, introduced species, marine protected area,
marine reserve, siltation, threatened species
Extinci
on de Especies en el Medio Marino: Tasmania como Ejemplo Regional de Perdidas de Biodiversidad Ignoradas
Resumen: Usamos a Tasmania como ejemplo para cuestionar el consenso de que pocas especies mari-
nas se han extinguido o est an cerca de la extinci on recientemente. Las amenazas a especies marinas y
estuarinasprincipalmente el cambio clim atico, las especies invasoras, la pesca y las descargas en cuencas
est
an acelerando, abarcan completamente el a rea de distribucion de especies y tienen la suficiente magnitud
para causar extincion. Nuestra ignorancia de la declinacion de la biodiversidad en el ambiente marino resulta
en gran medida de una carencia casi total de muestreos sistem aticos a gran escala y una confianza excesiva
en datos fisicoqumicos para monitorear tendencias ambientales. Las declinaciones poblacionales de especies
marinas cercanas a la extincion generalmente pasar an inadvertidas por la naturaleza oculta de su ambiente
y por la ausencia de datos cuantitativos.
Palabras Clave: area marina protegida, calentamiento global, cambio climatico, efectos de la pesca, especies
amenazadas, especies introducidas, reserva marina, sedimentaci
on
Paper submitted January 29, 2004; revised manuscript accepted August 18, 2004.
1294
Conservation Biology 12941300
C 2005 Society for Conservation Biology
DOI: 10.1111/j.1523-1739.2005.00159.x
Edgar et al. Loss of Marine Biodiversity 1295
Table 1. The total number of terrestrial species and marine species (in parentheses) listed under the World Conservation Union (IUCN) Red Lista
and Tasmanian Threatened Species Protection Actb on 23 January 2004.
animal species are marine (Table 1), and only a single ma- number of terrestrial species, whereas negligible num-
rine plant is listed (0.01% of total). No obligate marine bers of marine invertebrates and plants are included.
fish, one marine plant, and only four marine invertebrate Thus, numerous air-breathing marine species (whales,
species (all gastropods) are listed as extinct (IUCN 2001). sea lions, albatross, petrels, and turtles) are recognized
Regardless, given our rudimentary knowledge of the as threatened, whereas only eight fully aquatic species
distribution and abundance of most marine species and are listed (21% of total).
recognition that endangered species listings are biased Several hypotheses explain this trend: (1) higher verte-
by paucity of information, we need to critically assess the brates produce few offspring per year, making them dis-
possibility that our consensus largely reflects ignorance proportionately vulnerable to episodic impacts because
(Roberts & Hawkins 1999). We reviewed available infor- of slow population recovery; (2) higher vertebrates pos-
mation on threats to marine biodiversity for one region sess relatively large body mass, making them preferred
Tasmaniaand the ways managers and scientists assess hunting targets for humans; (3) species with both ter-
those threats. The Tasmanian region should not be con- restrial and marine habitat requirements are dispropor-
sidered unusual, other than perhaps its recent European tionately vulnerable to threats because they must cope
colonization (1803) and low human population density with hazards in both realms; (4) the charismatic nature of
(450,000 people inhabiting 67,000 km2 ), which make higher vertebrates enhances public and scientific interest,
it less affected by human activity than most temperate increasing the likelihood that a researcher will investigate
coasts. a species and generate data suitable for successful threat-
The Tasmanian marine environment is publicly per- ened species listing; or (5) air-breathing taxa are moni-
ceived to be in reasonable condition, aside from contami- tored relatively easily at the sea surface or on land, pro-
nation by heavy metals and sewage discharge that affects viding some long-term data on population trends. These
some estuaries, localized infestations of invasive species, hypotheses are not mutually exclusive; all are likely to be
and high rates of sediment and nutrient runoff (DPIWE true in some cases. Nevertheless, on their own, hypothe-
1997). For Australia in general, water quality and habitat ses 1, 2, and 3 do not fully account for observed patterns.
loss are considered the two major marine environmen- For example, the majority of shark species produce fewer
tal issues (Australian State of the Environment Committee offspring annually than sea turtles. Many fish and kelp
2001). The Australian State of the Marine Environment Re- species grow to a larger mass than seabirds. Numerous
port considered that the nations ecosystems were gener- cetacean species are listed despite lacking terrestrial con-
ally fair to excellent, with the exception of estuaries, nections.
seagrass beds, and open sandy beaches in developed ar- In reality, the almost complete absence of baseline pop-
eas, which were reported as poor to fair (Zann 1995). ulation data for marine species other than those commer-
cially exploited or visible at the sea surface makes it vir-
Endangered Species Listings tually impossible to successfully propose marine species
for listing under the Tasmanian Threatened Species Pro-
Few marine species are listed under the Tasmanian Threat- tection Act. Consequently, for example, only one mol-
ened Species Protection Act as extinct, endangered, lusk species (Gazameda gunnii ) has been listed as
vulnerable, or rarethe four official categories of threatened, despite the majority of the >1000 Tasma-
threat (Table 1). Inspection of listed species nevertheless nian mollusk species (May & Macpherson 1958) not hav-
reveals a clear taxonomic trend. Among the higher verte- ing been sighted or collected alive during the past two
brates, the number of marine species listed exceeds the decades.
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Volume 19, No. 4, August 2005
1296 Loss of Marine Biodiversity Edgar et al.
Moreover, regardless of magnitude of threat, few ma- or may not progress to extinction. If threatening pro-
rine invertebrates and plant species are likely to be listed cesses act within a subset of the range of a species, then
in the foreseeable future because of the near absence of local extinction is possible, whereas total extinction is
specialist researchers. The majority of biologists (50) unlikely. Conversely, a wide species distribution provides
and biologically trained managers (30) dealing with little insurance against extinction if the scale of a threat-
marine subjects in Tasmanian government institutions ening process fully encompasses that range.
and universities are concerned with fisheries issues. No Most marine species possess a widely dispersing life-
macroalgal botanist is currently employed in Tasmania, cycle phasegenerally planktonic larvaeand are dis-
and only five ecologists (no taxonomists) predominantly tributed over thousands of kilometers of coast. Others
study unexploited marine invertebrates. develop directly from demersal eggs or by live birth (e.g.,
all amphipods and isopods, some fishes, mollusks, echin-
oderms, and polychaetes) and can be highly localized
Available Evidence on Population Declines in distribution (e.g., the live-bearing seastar Patiriella
vivipara, which is restricted to <1 km2 of intertidal shore
Although no population data are available for most Tasma-
near Hobart, Tasmania; Prestedge 1998). Regardless of
nian marine species, limited inferences on interdecadal
mode of dispersal, few species possess refuges from the
trends can be made for some groups by using (1) catch
major threats to inshore biodiversity: climate change, in-
statistics for commercially exploited fishes and inver-
troduced species; fishing; and siltation, nitrification, and
tebrates, (2) historical aerial photographs for canopy-
other catchment effects. These threats also interact, often
forming marine plants, and (3) dated sediment cores for
in unpredictable ways.
mollusk assemblages. All data sets indicate major pop-
ulation declines for the majority of species, supporting
Climate Change
results from studies elsewhere (Pauly et al. 2000; Jackson
2001; Jackson et al. 2001) that show widespread histori- Mean surface sea temperature off the Tasmanian east
cal changes to inshore ecosystems. These declines gener- coast has increased by >1 C since the 1940s (Crawford
ally have gone unnoticed with changes between human et al. 2000). This change has been accompanied by local
generationsthe so-called sliding baseline syndrome transformation of habitat types, both in terms of popula-
(Dayton et al. 1998). tion gain for warm-temperate biota such as the barrens-
Catch statistics indicate that populations of most major forming sea urchin (Centrostephanus rodgersii) and pop-
fisheries species (most notably native oysters, commer- ulation loss for cool-temperate species such as the giant
cial scallops, southern rock lobster [ Jasus edwardsii], kelp.
orange roughy [Hoplostethus atlanticus], eastern gem- If global warming contributes another 12 C rise over
fish [Rexea solandri], barracouta [Thyrsites atun], south- the next century, an average change predicted by current
ern bluefin tuna [Thunnus maccoyii], jack mackerel [Tra- models (Houghton et al. 2001), suites of cool-temperate
churus declivis], school shark [Galeorhinus galeus], and organisms are likely to disappear, including a number
trumpeter [Latridopsis forsteri]) have declined by >50% of endemic species restricted to southeastern Tasma-
over three generationsthe IUCN criterion for endan- nia (Dartnall 1974). As with oceanic islands, Tasmanian
gered statusand many have declined by >80% (IUCN ecosystems are particularly susceptible to temperature
critically endangered; Harries & Croome 1989; Kailoa rise because the southward range extension of species
et al. 1993; Edgar & Samson 2004). Aerial photographs is prevented by a deepwater barrier. By contrast, species
indicate that giant Macrocystis pyrifera (L). C. Ag. kelp with ranges currently centered on mainland Australia are
beds, once sufficiently large to be commercially har- potentially able to migrate south to maintain residence in
vested, have declined by approximately half along the Tas- preferred temperature bands and avoid competition with
manian east coast since 1944 (Edyvane 2003). According species better adapted to warmer conditions. Sea tem-
to one estimate ( Rees 1993), seagrass beds have declined perature rise is an accelerating global threat that extends
in area by approximately 25% since the 1950s. Inshore beyond the range of Tasmanian species.
mollusk biodiversity has decreased from an average of 21
species per 5-cm slice for sediment cores dated at the start Introduced Species
of the twentieth century to 7 species per slice in 1990
The threat of introduced species is similarly global, largely
(Edgar & Samson 2004). No kelp, seagrass, or commer-
uncontrolled, and accelerating. Introduced species
cially exploited fish species is listed under the Tasmanian
threaten native species and the functioning of ecosys-
Threatened Species Protection Act.
tems through habitat modification, competition, preda-
tion, disease, and poisoning (e.g., Carlton 1989; Carlton
Scale of Threats & Geller 1993).
Species at all trophic levels were introduced dur-
Catastrophic population declines, as many Tasmanian ma- ing the past century into Tasmanian coastal waters
rine species have experienced over the past century, may and are now proliferating rapidly. Among the most
Conservation Biology
Volume 19, No. 4, August 2005
Edgar et al. Loss of Marine Biodiversity 1297
Fishing
Targeted fish catches will rarely cause extinction of a
species because the cost of capture of overharvested
populations will eventually exceed economic returns
(i.e., commercial extinction) and effort will transfer
to other species. Indirect effects of fishing, however
habitat damage, bycatch, and trophic cascadesare not
closely linked to economics and can potentially lead to
extinction.
Fishing has probably caused extinction of species in
the Tasmanian region through trawling of deepwater
seamounts that rise 300600 m from the continental slope
in water depths of 10002000 m. Large schools of or-
ange roughy were discovered aggregating on the approx-
imately 70 seamounts off southern Tasmania in 1989, and
a trawl fishery blossomed over the subsequent 5-year pe-
Figure 1. Ratio of introduced species to total species riod until more stringent management controls were en-
for mollusk shell fragments in 5-cm-thick sediment acted. During the initial gold rush period, all shallow
slices for 13 dated sediment cores collected from (<1000 m depth) seamounts were extensively trawled
shallow depths (414 m) across the southeastern with heavy bottom gear, some more than 3000 times
Tasmanian region (C.R.S. & G.J.E., unpublished data; (Koslow & Gowlett-Holmes 1998; Koslow et al. 2001).
for methodology see Edgar & Samson [2004]). The typical basal diameter of seamounts is 2 km.
Conservation Biology
Volume 19, No. 4, August 2005
1298 Loss of Marine Biodiversity Edgar et al.
Conservation Biology
Volume 19, No. 4, August 2005
Edgar et al. Loss of Marine Biodiversity 1299
10/km2 (Edgar & Barrett 2000). Estuaries isolated from despite considerable expense in remedial management
human activity possess sandy sediments and a predomi- actions remains unknown.
nance of epifaunal species, whereas estuaries with popu- Rather than ecological efficacy, SOE indicators were
lated catchments possess muddy sediments and a major- selected primarily for reasons of low perceived cost (i.e.,
ity of infaunal animals. Downstream runoff from catch- able to be statistically compiled without significant ad-
ments, primarily in the form of increased sediment loads ditional cost for data gathering). Biological condition
associated with land clearance, is probably responsible indicators are reliant on ad hoc rather than systematically
for this broad-scale habitat transformation. Other poten- directed research and are thereby confounded by chang-
tially important catchment impacts that affect estuarine ing effort, debasing their long-term value. Extent and
biota include freshwater diversion and increased nutrient condition of aquatic habitats is widely used as a condi-
loadings that can lead to die-off of seagrass beds. tion indicator (Ward et al. 1998); however, marine habitat
Although Tasmania is unusual in a global sense in that classes are primarily physical (e.g., hard sand, low-profile
it possesses numerous estuaries with uninhabited catch- reef ) and remotely sensed and thus unlikely to change
ments that are reserved as national parks, such estuaries greatly over time (seagrass beds are an exception). Al-
are largely confined to the western and southern coasts though aerial and satellite habitat mapping is extremely
and have remained uninhabited because of poor soils, effective for monitoring large-scale changes in terrestrial
which in turn generates low aquatic productivity and vegetation types, it is much less useful in the marine en-
depauperate estuarine biota (Edgar et al. 1999). Almost vironment.
all estuaries on the northern and eastern coasts, where Overall, changes in existing SOE indicators should be
the majority of Tasmanian estuarine species are located recognized as providing little insight into the current state
(Edgar et al. 1999), have been badly degraded by catch- or trends in marine ecosystems. Complete transformation
ment activity (Edgar et al. 2000). of communities could occur with, for example, the arrival
of invasive species or climate change, and no indicator
Pollution, Shore Reconstruction, and Other Local Threats would detect this trend. To be effective, the systematic
collection of biological data is unavoidable. No long-term
Oil spills, fish farms, land reclamation, foreshore develop- surveys of marine communities are currently being un-
ment, sewage effluent, heavy metal discharge, and chem- dertaken in Tasmania, other than an MPA monitoring pro-
ical outfalls frequently generate intense impacts on the gram that relies on ad hoc funding.
marine environment, including complete loss of flora
and fauna in extreme cases; however, such effects are
generally highly localized (Crawford et al. 2000). Thus,
in contrast to public perceptions that pollution poses Conclusion
the greatest hazard to marine life (L. Dropkin, www.
compassonline.org/activities/summary.html), pollution Our main goal is to question the widespread assumption
is unlikely to cause species extinction, other perhaps than that an absence of unequivocal proof implies that extinc-
for taxa with extremely restricted intertidal or shallow tion is an extreme rarity in the marine environment. Proof
subtidal ranges. of extinction is unattainable, and the hidden nature of
the marine environment and lack of baseline data means
Assessment of Threats to Marine Biodiversity that population declines for almost all species approach-
ing extinction will go unnoticed. Rather than assuming
Trends in the health of Tasmanian marine ecosystems are population persistence, a key question is, How do rare
monitored using a defined set of environmental indicators species manage to find mates and survive in the face of
that are reported every 5 years in the Tasmanian and Aus- multiple interacting and ubiquitous threats and increasing
tralian State of the Environment Reports (DPIWE 1997; population fragmentation?
Australian State of the Environment Committee 2001). De- Adequate comprehension of change in the marine en-
spite this coverage, little information is available on the vironment requires reversal of the burden of proof
changing biological state of the marine environment. Vir- (Gerrodette et al. 2002) and the systematic collection of
tually all State of the Environment (SOE) indicators relate broad-scale data sets. As scientists, we continually be-
to human pressure or management response, or are moan difficulties in assessing human impacts in the en-
physicochemical rather than biological metrics. Thus, for vironment in the absence of baseline data, yet rarely un-
example, considerable information is available on chang- dertake quantitative sampling over large spatial scales to
ing levels of total phosphate input into estuaries, but neg- provide useful comparative information for the future.
ligible information is available on the biological response Until current funding to conserve marine biodiversity is
of ecosystems to changing loadings. Whether greater re- partially applied to a systematic quantitative global sur-
ductions in phosphates are required to have a significant vey, we will continue to grope blindly with unrealistic
ecosystem effect or whether phosphates are unimportant models when assessing and addressing threats.
Conservation Biology
Volume 19, No. 4, August 2005
1300 Loss of Marine Biodiversity Edgar et al.
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