ArticlePDF Available

Abstract and Figures

The International Union for Conservation of Nature (IUCN) is the premier global biodiversity conservation organization. Its Red List is a rigorous vehicle for assessing the conservation status of plant and animal species. However, although all animal and bird species recognized by IUCN have been evaluated , only a tiny fraction of invertebrates have been evaluated. As a measure of the numbers of extinct species (since around the year 1500) the Red List is probably quite accurate for birds and mammals, but severely underestimates the numbers for invertebrates. Nonetheless, molluscs stand out as the major group most severely impacted by extinction, with 297 of the 744 animal species listed as extinct in the third issue of the 2016 Red List. Here we review efforts to obtain a more realistic , albeit less rigorous, assessment of the numbers of extinct mollusk species. Our approach has been based on biblio-graphic research and consultation with experts, rather than following the highly detailed but restrictive IUCN Categories and Criteria. In 2009, this led to an assessment that 533 mol-lusk species were extinct, far more than the number on the Red List. In the present study we revisited this approach and here list 638 species as extinct, 380 as possibly extinct, and 14 as extinct in the wild, a total of 1,032 species in these combined categories, and more than twice as many as listed by IUCN in these categories. However, this approach only considers species for which information is available; it is therefore biased. In a study published in 2015 we developed an alternative approach, based on a random global sample of land snails, and estimated that 3,000–5,100 mollusk species have gone extinct. We review the main reasons for these extinctions: habitat destruction, impacts of introduced species, exploitation and collecting, and, potentially, climate change, and discuss relevant case studies.
Content may be subject to copyright.
Measuring the Sixth Extinction: what do mollusks tell us?
Robert H. Cowie
Pacific Biosciences Research Center
University of Hawaii
Honolulu, HI 96822 USA
cowie@hawaii.edu
Claire Re
´gnier
Benoı
ˆt Fontaine
Philippe Bouchet
Muse
´um national d’Histoire naturelle
Paris 75005, FRANCE
ABSTRACT
The International Union for Conservation of Nature (IUCN) is
the premier global biodiversity conservation organization. Its
Red List is a rigorous vehicle for assessing the conservation
status of plant and animal species. However, although all ani-
mal and bird species recognized by IUCN have been evalu-
ated, only a tiny fraction of invertebrates have been evaluated.
As a measure of the numbers of extinct species (since around
the year 1500) the Red List is probably quite accurate for birds
and mammals, but severely underestimates the numbers for
invertebrates. Nonetheless, molluscs stand out as the major
group most severely impacted by extinction, with 297 of the
744 animal species listed as extinct in the third issue of the
2016 Red List. Here we review efforts to obtain a more realis-
tic, albeit less rigorous, assessment of the numbers of extinct
mollusk species. Our approach has been based on biblio-
graphic research and consultation with experts, rather than
following the highly detailed but restrictive IUCN Categories
and Criteria. In 2009, this led to an assessment that 533 mol-
lusk species were extinct, far more than the number on the
Red List. In the present study we revisited this approach and
here list 638 species as extinct, 380 as possibly extinct, and
14 as extinct in the wild, a total of 1,032 species in these
combined categories, and more than twice as many as listed by
IUCN in these categories. However, this approach only con-
siders species for which information is available; it is therefore
biased. In a study published in 2015 we developed an alterna-
tive approach, based on a random global sample of land snails,
and estimated that 3,000–5,100 mollusk species have gone
extinct. We review the main reasons for these extinctions: hab-
itat destruction, impacts of introduced species, exploitation and
collecting, and, potentially, climate change, and discuss rele-
vant case studies. Oceanic island land snails, especially those of
Pacific islands, have suffered the greatest proportion of the
extinctions, with some species having gone extinct before being
discovered and described scientifically. The Amastridae, an
endemic Hawaiian family of 325 recognized species, may have
lost all but 18 species. We outline the phases in this catastro-
phe: 1) pre-human and/or prehistoric extinction, either natural
or anthropogenic, with species known only as fossils/subfossils;
2) extinction due to habitat destruction and introduction of a
number of alien species by Pacific island people as they settled
the islands; 3) extinction due to extensive habitat destruction
and introduction of highly destructive invasive alien species
following colonization by Westerners; 4) extinction following
the advent of large-scale agriculture at the end of the 19th
Century, at the time of a major increase in the land snail
extinction rate globally; 5) extinction due to increased military
activity, tourism, commerce, urbanization and the concomittant
rapidly increasing introduction of invasive species after the
Second World War. Extrapolating from our assessments of
mollusks, we estimate that approximately 7.5–13% of all spe-
cies have gone extinct since around year 1500. This is orders of
magnitude greater than the 860 (0.04% of 2 million) listed as
extinct by IUCN (2016). The biodiversity crisis is real.
Additional Keywords: Amastridae, biodiversity crisis, bivalves,
Euglandina, Gambier Islands, Hawaii, IUCN, Melanopsis,
Mollusca, non-marine, Powelliphanta,Rhachistia aldabrae,
Red List, snails
INTRODUCTION
Over a decade ago, Lydeard et al. (2004) published a key
paper outlining the decline of non-marine mollusks, the
threats they face, and the high level of extinction com-
pared with other major animal groups that had been
documented as of 2002 by the International Union for
Conservation of Nature on its Red List. The Red List
program was initiated in 1964 and mollusks were first
included in it in 1983, when 28 species were listed as
extinct (Wells et al., 1983). The Red List only considers
extinctions in modern historical times, from around the
year 1500. Following the realization that an ill-conceived
biological control program had caused the extinction in
the wild of the entire fauna of partulid tree snails on the
island of Moorea in French Polynesia (Murray et al.,
1988), more effort was put into documenting mollusk
extinctions on the Red List. A Moorean partulid appeared
on the front cover of the 1990 Red List (IUCN, 1990), and
when the 1994 Red List (Groombridge, 1994) was pub-
lished, 255 species were listed as extinct. The number has
gradually increased and the most recent Red List (IUCN,
2016) lists 297 mollusks as Extinct out of a total of 860
extinct species listed.
If we accept a figure of 2 million described species
(Chapman (2009) estimated 1.9 million, and IISE (2017)
documents a current yearly increment of around 18,000
newly described species; the Red List accepts 1,736,081
species), this means that between one and two species
THE NAUTILUS 131(1):3–41, 2017 Page 3
have gone extinct per year since 1500, the year from
which IUCN starts counting, or about 0.8 species extinc-
tions per million species years (E/MSY). The background
rate, based on the fossil record, is around 0.1–2.0 E/MSY
(Ceballos et al., 2015). That the rate documented by
IUCN is within the estimated range of the background
rate has provided support for the suggestion by environ-
mental skeptics (e.g., Lomborg, 2001) that there is no
“Biodiversity Crisis”, despite the views of many scientists
and the media publicity surrounding the notion of the
“Sixth Extinction”, caused by human activities (Novacek,
2001; Leakey and Lewin, 1996). So the question becomes,
is there really a crisis or is it a false or exaggerated claim
by environmental activists and scientists with an arguably
political agenda?
The key question to ask in trying to resolve this con-
flict is: how accurate really are the IUCN extinction
data? This review summarizes the approaches that have
been developed since the review of Lydeard et al. (2004)
and that have attempted to begin to answer this question
(Re
´gnier et al., 2009, 2015a). It updates the assessments
of Re
´gnier et al. (2009) and reviews a case study of a
Hawaiian land snail family, the Amastridae, that used
these new approaches to obtain a realistic assessment
of extinction (Re
´gnier et al., 2015b).
IS THE IUCN RED LIST APPROPRIATE FOR
ASSESSING EXTINCTION RATE?
The IUCN has assessed 85,604 species (IUCN, 2016).
This represents a huge amount of detailed work by ded-
icated biologists, but nonetheless represents only 4.3%
of the total 2 million animal and plant species. Although
a small sample of overall biodiversity, if it were a random
sample, some confidence could perhaps be placed in
its assessment of extinction rate. However, it is not a
random but a highly biased sample.
IUCN (2016) estimated that there are 5,567 known
mammal species and 11,121 known bird species, total
16,688, although Chapman (2009) estimated 5,487 and
9,990, total 15,477, probably a result of differing taxo-
nomic treatments and estimation protocols. IUCN
(2016) has assessed all mammal and bird species that it
recognizes. Of these, only 849 (5%) were placed in the
IUCN category “data deficient”, that is, they lacked suf-
ficient information to assess their conservation status
according to the IUCN Red List Categories and Criteria
(IUCN, 2012). Thus, the number of extinctions (239)
listed for mammals and birds by IUCN (2016) is proba-
bly quite accurate.
However, the situation is very different for inverte-
brates, which constitute >95% of described animal
diversity, about 1.31 million (IUCN, 2016) or 1.5 million
species (1.36 million of Chapman (2009) extrapolated
by an annual increment of 1%). Only 18,609 of these
species have been assessed, 1.2% of the total, with 7,205
(39%) of these deemed data deficient. Why is this? There
are two main reasons: 1) taxonomic bias, and
2) the related relative difficulty of obtaining adequate
data to assess the conservation status of invertebrates
compared to vertebrates according to the IUCN criteria.
TAXONOMIC BIAS
There are on average many specialists able to assess the
conservation status of each mammal or bird species. As
most mammal and bird species have been discovered
and described, these specialists are primarily field biolo-
gists working on ecology, population biology, behavior,
etc. In contrast, most invertebrate specialists are taxono-
mists or systematists (in the broad sense including those
studying biogeography, phylogenetics, and the assess-
ment of biodiversity), and most of these systematists
each deal with tens to hundreds of species. There are
roughly equal numbers of specialists focused on verte-
brates, on plants and on invertebrates, yet plant species
are roughly ten times, and invertebrates a hundred
times, more numerous than vertebrates (Gaston and
May, 1992; May, 2011).
This bias and the 100-fold greater relative number of
vertebrate specialists compared to invertebrate special-
ists is reflected in the numbers of IUCN Species Survival
Commission Specialist Groups focused on particular
taxa: 73 for vertebrates and only 12 for invertebrates,
with only one for the entire phylum Mollusca (Table 1).
In contrast, many of the vertebrate Specialist Groups
are focused on just one or a few species (e.g., African
elephant, hyaenas, vultures, pelicans, etc.).
IUCN CRITERIA
The IUCN criteria are detailed and complex. They
include precise quantitative determinations of remaining
numbers of individuals, life history details, area occupied,
Table 1. Number of IUCN Specialist Groups for animals.
Vertebrates No. of Groups Invertebrates No. of Groups
Mammals 35 Insects 4
Birds 16 Other arthropods 3
Reptiles/amphibians 12 Coral 1
Fish 10 Mollusks 1
Geography/habitat 3
Total 73 Total 12
Page 4 THE NAUTILUS, Vol. 131, No. 1
trends in abundances and range and many other param-
eters that are precisely defined. This detail and precision
was developed by IUCN in response to criticism that its
assessments were too qualitative and subjective, indeed
secretive (e.g., Mrosovsky, 1997). All mammal and bird
species have been evaluated based on these stringent
criteria, with very few species considered Data Defi-
cient. This has been possible because, as explained
above, there are many specialists in the field generating
the kind of data that are required.
In contrast, for the great majority of invertebrate
species, few data relevant to the IUCN criteria exist
other than what are available in the original descriptions
(type localities and little else) and perhaps a small number
of subsequent publications. Most of the field research of
any relevance is undertaken as part of biodiversity inven-
tories, the discovery and subsequent description of the
vast number of species as yet unknown to science, or
indeed to humanity.
MOLLUSKS ASSESSED BY IUCN
The most careful estimate of the number of described
mollusk species (Rosenberg, 2014) suggested that there
are 70,000–76,000, although IUCN (2016) estimated
85,000, following Chapman (2009). Compared to other
invertebrate groups, a relatively high proportion of mol-
lusk species has been assessed: 7,276 species (IUCN,
2016), or roughly 8.5–10%. However, in contrast to mam-
mals and birds, a high proportion of these species was
assessed as Data Deficient (2,463 species, 34%), for lack
of adequate information addressing the IUCN criteria.
The Red List (IUCN, 2016) lists 860 species
(744 animals, 116 plants) as Extinct, including 297 mollusk
species. Mollusks, despite the small proportion of them
that has been assessed, thus represent 35% of all species
extinctions and 40% of animal extinctions, as reported
by IUCN.
ALTERNATIVE APPROACHES TO ASSESSING
MOLLUSK EXTINCTIONS
RE
´GNIER ET AL. (2009)
Given the shortcomings of the Red List in assessing the
level of extinction of invertebrates overall, alternative
approaches have been sought. Re
´gnier et al. (2009)
re-evaluated mollusk species listed as Extinct on the
Red List of 2007 based on a review of the literature and
by asking a cadre of biologists with expert knowledge to
provide their opinion on the veracity of the Red List
assessments. This literature review and gathering of
expert knowledge also identified additional species not
on the Red List but either documented in the literature
as extinct or simply known to the experts as extinct. Some
of the species listed as extinct were considered in fact
not to be so, either because they had been found alive
since being listed, or because they had been synonymized
with extant species; and two species were excluded as
being nomina dubia. Despite this reduction, from 302
species listed to 269 considered in fact to be extinct,
overall the number of species considered extinct (both
listed and not) increased to 533, including those consid-
ered Extinct in the Wild (13) and those considered
“possibly extinct” (71) (Re
´gnier et al., 2009: Supporting
Information online), roughly twice as many as correctly
considered Extinct on the Red List.Re
´gnier et al. (2009)
also listed 33 subspecies (including 5 extinct in the wild
and 1 possibly extinct), for a total of 566 taxa.
UPDATE OF RE
´GNIER ET AL. (2009)
Following the same approach as that of Re
´gnier et al.
(2009), i.e., literature search and expert consultation (see
the Acknowledgements for the names of the experts
who provided information), we have updated the list of
species considered extinct. We took the most recent
evaluation of each species as representing its current
status, which was either the most recent IUCN evalua-
tion as listed in the Red List (IUCN, 2016), Re
´gnier et al.
(2009), or our own literature/expert consultation. We
excluded species listed as Extinct, Possibly Extinct, or
Extinct in the Wild on the Red List and/or by Re
´gnier
et al. (2009) if they are now thought to be extant
(Appendix Table A1). We also did not consider subspe-
cies, neither those recognized in the Red List nor those
listed by Re
´gnier et al. (2009). Subspecies and synonyms
that have been recognized in the literature subsequently
but that are still retained as valid species on the Red List,
as well as undescribed species listed with provisional
names on the Red List, were also excluded (Appendix
Table A2).
Of the 297 species listed as Extinct on the Red List,
we considered six as now only Possibly Extinct, three
as Extinct in the Wild, 20 as extant, and 11 that have now
been considered synonyms, subspecies, nomina dubia,
or unrankable. Of the 124 listed as “Critically Endan-
gered (Possibly Extinct)” on the Red List, three are
known to be extant, six are considered as now extinct,
with a further five (Galapagos Bulimulus species)
undescribed, five synonymized, and five unrankable.
And of the 14 listed as extinct in the wild, we considered
one to be extant in the wild and five to now be extinct.
Thus, excluding undescribed species and species still
listed as valid on the Red List but that have been synon-
ymized, reduced to subspecies, considered unrankable,
or are now thought to be extant, the current Red List
(IUCN, 2016), in our view correctly lists 386 valid spe-
cies in the combined categories of Extinct, Critically
Endangered (Possibly Extinct), and Extinct in the Wild,
which is 44 species fewer than the 430 actually listed by
IUCN (Table 2).
The additional information derived from the literature
search and expert consultation allowed us to estimate
that in fact 638 species are extinct, 380 possibly extinct
(EX?), and 14 extinct in the wild, a total of 1,032 species
in the combined categories (Appendix Tables A3–5).
R.H. Cowie et al., 2017 Page 5
Of these, 47 are known only as “fossil” or “subfossil” but
in many of these cases it was not possible to say when
they went extinct, perhaps in some cases from natural
causes such as non-anthropogenic climate change. For
comparability with the approach officially taken by
IUCN of focusing on species that have gone extinct since
around the year 1500 (though 11 fossil/subfossil species
are included in the Red List), we might conservatively
exclude the 47 fossil/subfossil species in our list and then
would consider 591 species as extinct, 380 as possibly
extinct, and 14 as extinct in the wild, total 985 species.
Even so, our numbers of extinct and possibly extinct
species both greatly exceed those of the Red List,by
more than two and three times respectively. This total is
also approximately double the number listed by Re
´gnier
et al. (2009).
Of the 1,032 species (Appendix Tables A3–5), 803 are
land snails, from 52 families but dominated by species of
four Pacific island families, the Amastridae (307 species),
which is a Hawaiian endemic family (see below),
Endodontidae (92 species), Partulidae (52 species) and
Achatinellidae (44 species), as well as one more wide-
spread family (though with greatest diversity in the
Pacific), the Charopidae (54 species). Freshwater
snails are represented by 177 species from 22 families,
with all but two families (Hydrobiidae, 61 species;
Pleuroceridae, 32 species) represented by 12 or fewer
species. Freshwater bivalves are represented by
46 species from five families, with only the Unionidae
(40 species) having more than one or two species. Six
marine gastropods are listed.
RE
´GNIER ET AL. (2015)
Re
´gnier et al. (2009) and the updated assessment pro-
vided above, have only dealt with species already assessed
by IUCN and those additional species that were known to
be extinct, possibly extinct, or extinct in the wild, both
documented in the literature and as known to biologists
with expert knowledge. These species were therefore not
a random sample. A more realistic estimate of the true
number of mollusk extinctions would only be provided by
assessing a random sample of mollusk species.
Therefore, Re
´gnier et al. (2015a), focusing on land
snails, generated a rigorously random sample of 200 spe-
cies from a wide representation of localities across the
globe. They evaluated these species based on the IUCN
categories and criteria (IUCN, 2012) by reviewing the
literature as well as major museum collections. For com-
parison with this IUCN-based evaluation, they also sent
the list of 200 species to numerous land snail experts,
asking them to evaluate whether those species for which
they had personal knowledge and experience were
extinct. For species for which no expert was available,
Re
´gnier et al. (2015a) made their own assessment based
on collection records. In addition, Re
´gnier et al. (2015a)
developed a mathematical probabilistic model, based on
collection dates as documented in major museum mala-
cological collections. This model evaluated the probabil-
ity of extinction for each of the 200 species, and thereby
offered an independent means of corroboration (or not)
of the expert evaluation.
Based on the IUCN categories and criteria, Re
´gnier
et al. (2015a) were only able to evaluate 31 of the 200 spe-
cies, the other 169 being categorized as Data Deficient.
Of the 31, three (1.5% of the 200, but 10% of the evalu-
ated 31) were evaluated as extinct. Under the assumptions
that the 200 land snail species in the random sample are
representative of the described non-marine molluscan
diversity—roughly 30,000 species (Rosenberg, 2014)
and that marine molluscan extinction is negligible com-
pared to non-marine extinction (e.g. Carlton, 1993; but
see Peters et al., 2013), extrapolation leads to an estimate
of 3,000 extinct mollusk species.
In contrast, the experts were able to evaluate 118 of the
200, the remaining 82 being “Impossible to Assess”.
Twenty (10% of the 200, but 17% of the evaluated 118)
were evaluated as extinct. Note that Re
´gnier et al. (2015a)
used slightly different terminology from the IUCN
categories in order to draw attention to the differences
between the two approaches. The probabilistic model
broadly corroborated the expert evaluations in terms of
the proportion of species considered extinct.
Of the 76,000 described mollusk species (Rosenberg,
2014), about 46,000 are marine (WoRMS, 2017) and
roughly 30,000 non-marine. Therefore, as 10–17% of
the 200 land snail species were considered extinct,
extrapolation suggests that in fact around 3,000–5,100
mollusk species are extinct, far more than the 297 on the
Red List (IUCN, 2016), the 532 estimated by Re
´gnier
et al. (2009), the 1,032 estimated above updating Re
´gnier
et al. (2009), but in the same region as the 3,000 extrapo-
lated from the assessments of Re
´gnier et al. (2015a) based
on the IUCN categories and criteria. This estimate of
3,000–5,100 mollusk extinctions, even taking into account
that it is based on a small sample, is shocking. And many
are going extinct before they have been discovered and
described (e.g., Richling and Bouchet, 2013; Sartori et al.,
2013; 2014).
Table 2. Numbers of mollusk species considered extinct
(EX), critically endangered (possibly extinct) (CR(PE)), and
extinct in the wild (EW) in the Red List (IUCN, 2016), with
species on the Red List re-assessed herein, and the results of
the present study based on additional literature search and
expert consultation.
IUCN category Red List
1
Red List
re-assessed This study
2
EX 297 268 646
CR(PE) / EX?
3
119 107 373
EW 14 11 14
Total 430 386 1,032
1
Includes 11 fossil/subfossil species listed as extinct
2
Includes 46 fossil/subfossil species listed as extinct
3
CR(PE) in the Red List, EX? in this study
Page 6 THE NAUTILUS, Vol. 131, No. 1
WHY ARE NON-MARINE MOLLUSKS
GOING EXTINCT?
There are at least four possible causes of non-marine
mollusk extinction, which are, for the most part, the same
causes of the extinction of non-marine species in general:
habitat destruction, impacts of introduced species, exploi-
tation and collecting, and, potentially, climate change.
HABITAT DESTRUCTION
Urbanization, deforestation, agricultural expansion and
exploitation of natural resources have all had impacts
on mollusks. Three examples serve to illustrate some of
these threats.
Gambier Island Land Snails: Based on collections
made by the Bishop Museum (Honolulu) Mangarevan
Expedition in 1934 and by the Muse
´um national
d’Histoire naturelle (Paris) in 1997, 46 endemic species
have been recorded from the Gambier Islands in the
families Euconulidae, Endodontidae, Assimineidae, and
Helicinidae (Abdou and Bouchet, 2000; Bouchet and
Abdou, 2001; 2003; Richling and Bouchet, 2013). Only
three of these species were still extant; the remainder
were described from empty shells collected from the
shell bank of the soil.
The cause of the extinction of almost this entire fauna
was deforestation (Richling and Bouchet, 2013). Defor-
estation began with the first arrival of Polynesian settlers
around 1,000 years ago and reached a peak in the 17th
and 18th centuries with the total destruction of the
native flora (Conte and Kirch, 2008), no doubt exacer-
bated after the arrival of Europeans in the early 19th
Century. A few of the snail species were still extant in
the 1840s–1860s, but no living specimens of all but the
three known to be extant have been collected since the
19th Century (Richling and Bouchet, 2013). Similar sce-
narios have played out across the islands of the Pacific.
Melanopsis parreyssii in Romania: This freshwater
species was listed as Critically Endangered on the Red
List in 2013 (Fehe
´r, 2013). It was deemed Extinct in
the Wild in 2016 (Sı
ˆrbu and Benedek, 2016). It was
extremely narrowly endemic in Romania but had also
been introduced to Hungary and Bulgaria. However, by
2010 these introduced populations had vanished (Fehe
´r,
2013; Sı
ˆrbu et al., 2013). The Romanian locality was part
of a system sustained by a geothermal aquifer that was
declared a nature reserve and a Natura 2000 Site of
Community Importance. There were lakes and creeks
fed by thermal springs, forming the only habitat of
Melanopsis parreyssii. However, rapidly increasing recent
development of the geothermal waters, especially for tour-
ism, led to the springs becoming clogged and the natural
thermal lakes diminished, up to the point where the only
natural, but shrinking, lake that remained was Pet¸ea
(referred to as Ba
˘ile Episcopiei by Fehe
´r, 2013). By 2011
the spring serving Pet¸ea Lake ceased activity and by 2015
the lake had become little more than a puddle supporting
no mollusks except an invasive bivalve (Sı
ˆrbu et al., 2013;
ˆrbu and Benedek, 2016). Captive breeding efforts have
met with little success (Sı
ˆrbu and Benedek, 2016).Human
greed and disregard for the environment, including laws
supposedly protecting it, had led to the destruction of
the habitat of this narrowly endemic species and thus
its extinction.
Powelliphanta augusta in New Zealand:
Powelliphanta species are large predatory, worm-eating
land snails endemic to New Zealand and most have very
small ranges, making them highly vulnerable to habitat
destruction (Walker et al., 2008; Boyer et al., 2013). A
species of Powelliphanta, first collected in 1996 but not
recognized as a possible new species until 2003, and
confirmed as such by Trewick (2005), was discovered on
Mount Augustus, a peak on the Stockton Plateau in
New Zealand’s South Island and the site of a large open
cast coal mine (Trewick et al., 2008). By 2003, much of
the snails’ habitat had been destroyed, with the entire
remaining 8.5 ha of ridge-top habitat under severe threat
from the mining. With this imminent threat, and follow-
ing legal action (see Walker et al., 2008; Boyer et al.,
2013), all snails and eggs that could be found were col-
lected and brought into captivity, beginning in 2006.
Soon thereafter, all but a tiny piece of snail habitat was
destroyed (Walker et al., 2008). Many of the snails were
transferred back to the wild at three sites with sup-
posedly similar habitat, but the mortality rate in these
populations was such that they were unlikely to survive
(Morris, 2010). One of these sites was created by trans-
ferring entire habitat from the original site to an area not
slated to be mined, but the large trees did not survive
well and the habitat was invaded by weedy species
(Morris, 2010). The captive snails exhibit slower growth
and higher hatchling mortality than estimated in the orig-
inal wild population (James et al., 2013). Furthermore, a
large proportion of the captive snails died following an
electrical malfunction in their temperature-controlled
facility (James et al., 2013). The species was described as
Powelliphanta augusta in 2008 (Walker et al., 2008).
Although P. au g us t a is not yet extinct, the destruction of
its entire habitat by coal mining has left it on the brink.
IMPACTS OF INTRODUCED SPECIES
It is generally difficult to demonstrate definitively
that an invasive species has caused the extinction of
another species. For example, following the zebra mussel
(Dreissena polymorpha) invasion of North America
beginning around 1985, many of the native freshwater
mussels (Unionoida) were thought to be doomed
(Ricciardi et al., 1998). At localities with high densities
of D. polymorpha, local populations of native mussels
were being extirpated and some of the native species
were in steep decline or becoming regionally extinct.
Over 60 species were thought to be in danger of global
extinction from the combined effects of zebra mussels
and habitat degredation (Ricciardi et al., 1998). However,
R.H. Cowie et al., 2017 Page 7
a decade later, Strayer and Malcom (2007), focusing on
four species in the Hudson River, showed that although
they had declined steeply following zebra mussel inva-
sion, by 2000–2004 populations of these species had
stabilized at 4–22% of their pre-invasion densities, offering
a slender hope that the native mussels might be able to
co-exist with the invaders, albeit at much lower densities
(Strayer and Malcom, 2007).
In contrast, the prime example of an invasive species
causing extinction of mollusk species is the introduction
of the predatory snail Euglandina rosea to the islands of
the Pacific, notably to the Hawaiian Islands and the
Society Islands of French Polynesia but also elsewhere
(e.g., Cowie and Cook, 2001), in poorly considered
efforts to control the invasive giant African snail,
Achatina fulica (Hadfield, 1986; Murray et al., 1988).
The clearest evidence of a direct impact was that as
E. rosea spread across the island of Moorea, the endemic
Partula tree snail species vanished in its wake; it did not
control A. fulica (Murray et al., 1988; Cowie, 2001). On
the other islands of the Society group the same story
played out (Coote and Loe
`ve, 2003; Gerlach, 2016).
In Hawaii, the combination of E. rosea and invasive
rats, following on from habitat destruction, has caused
the decline of endemic achatinelline tree snails (Hadfield
et al., 1993), and another introduced predatory snail,
Oxychilus alliarius, may yet impact endemic Hawaiian
species, notably the single species in the endemic mono-
typic helicarionid genus Kaala (Curry et al., 2016). The
invasive predatory flatworm Platydemus manokwari has
caused the extinction of endemic Pacific island snails,
notably in the Ogasawara Islands (Chiba and Cowie,
2016). Competition between invasive and native snails
may also be important, but no definitive instances of this
have been documented (Cowie, 2005).
The impacts of invasive species are often inextricably
linked to those of habitat destruction or modification, as
invasive species, such as rats (e.g., Athens, 2009), may
drastically alter habitat, and habitat alteration may facil-
itate the spread of invasive species (Didham et al., 2007).
As such, they can be at least the partial cause of extinc-
tion. However, invasive species may act in concert with
or consecutively with habitat alteration, making it diffi-
cult, with some clear exceptions, to say that invasive
species, per se, have been the cause of specific mollusk
species extinctions.
EXPLOITATION AND COLLECTING
Numerous non-marine mollusk species are exploited for
human consumption. In Europe, and especially in coun-
tries bordering the Mediterranean, various of the larger
species of land snails are collected and eaten, most nota-
bly Helix pomatia, the “escargot de Bourgogne”, and
Cornu aspersum, the “petit gris”, both of which are read-
ily available in most French markets, but also more
widely. However, although C. aspersum remains abun-
dant and widespread in western Europe, H. pomatia has
declined, notably in France, and efforts are increasingly
being made to culture it for export, especially in eastern
Europe (Ligaszewski et al., 2007). Nonetheless, H. pomatia
is listed as of Least Concern on the Red List (IUCN, 2016).
Various other species are eaten around the Mediterranean
(Yildirim et al., 2004) but none seems to have attracted
conservation concern. The collection in the wild for the
restaurant trade, in combination with habitat loss and
alien species, has endangered the endemic “bulimes”
(genus Placostylus) of New Caledonia (Brescia et al.,
2008; Neubert et al., 2009). In Asia, various species of
Ampullariidae, Viviparidae, and Pachychilidae in partic-
ular are eaten, as are a number of clams and mussels
(e.g., Ko
¨hler et al., 2012), and Achatinidae are eaten in
West Africa (e.g., Nyoagbe et al., 2016); but none of
these species has attracted great concern because of this.
There are a few records of land snails being used for
medicinal purposes, e.g., Theba pisana (Benı
´tez, 2011)
and Achatina fulica (Cowie and D.G. Robinson, 2003),
and religious purposes, e.g., Achatina fulica (Neto et al.,
2012), and they may be a significant part of local rural
economies (Osemeobo, 1991); they may also be intro-
duced beyond their native range for such religious pur-
poses (Va
´zquez et al., 2016). But there is no evidence
that these usages have led to the decline and certainly
not extinction of these species.
In the 19th Century, freshwater mussels (Unionida)
were commercially harvested for their pearls, notably in
the United States; over-harvesting led to decline of
the populations and the fishery was largely abandoned
(Neves, 1999; Anthony and Downing, 2001). However,
soon thereafter, the demand for shells of freshwater
mussels for the button industry burgeoned, causing fur-
ther declines and adding to the already serious and
increasing threats from habitat degredation; but this
industry essentially died out with the advent of plastics
(Neves, 1999; Anthony and Downing, 2001; Strayer
et al., 2004), although it persists in other parts of the
world (Beasley, 2001). However, the discovery in Japan
that mussel shell material could act as nuclei for the
production of cultured pearls, resulted in a further
phase of exploitation of mussels in the United States for
export, although demand declined drastically by the late
1990s (Neves, 1999) Although habitat degredation has
been generally considered the primary cause of mussel
decline, over-exploitation has also been important (Strayer
et al., 2004).
The hobby of shell-collecting is generally more
focused on marine than on non-marine species, with
some marine taxa, for instance in the genus Conus,
threatened as a result (Peters et al., 2013). Nonetheless,
among non-marine species there are a few notable
instances in which shell collecting and ornamental use
may have been at least in part responsible for the decline
and perhaps extinction of certain species (Cowie, 2004).
Most notably, collecting of snails by late 19th and early
20th Century shell collectors quite possibly had an
important impact on some of the larger and more colorful
Hawaiian species, primarily but by no means exclusively
the beautifully colored and patterned Achatinellinae
Page 8 THE NAUTILUS, Vol. 131, No. 1
(Hadfield, 1986). Compared to marine species, there is a
much more limited collectors’ trade in shells of non-
marine species, which nonetheless may lead to endanger-
ment. However, the legal instruments of control (notably
the Convention on International Trade in Endangered
Species (CITES)) list only three non-marine gastropod
genera: the genus Achatinella, with 39 species listed as
Extinct or Critically Endangered by IUCN (2016), the
genus Polymita, with no species listed, and Papustyla
pulcherrima, the Manus Island (off the north coast of
New Guinea) green tree snail, which is listed as Near
Threatened by IUCN (2016) (see also Whitmore, 2016).
The collection of the 10,000 or so shells of the partulid
tree snail Eua zebrina that once made up the chandeliers
in the lobby of American Samoa’s then main hotel surely
must have significantly reduced at least some populations
of that species (Cowie, 1993).
Overall, therefore, exploitation and collecting have not
been a major cause of mollusk extinction, with a number
of notable exceptions.
CLIMATE CHANGE
Gerlach (2007) declared that Rhachistia aldabrae,an
endemic cerastid from Aldabra Atoll that was still locally
abundant in the 1970s (Gerlach, 2009), had gone extinct
in the late 1990s as a result of declining rainfall. It was
therefore placed on the Red List as Extinct (Gerlach,
2009). This is the only instance of a mollusk being
reported as having gone extinct as a result of climate
change. However, in 2014 it was discovered alive
(Battarbee, 2014). Nonetheless, there is only one tiny
population (J. Gerlach, pers. comm.) and it seems likely
that with ongoing climate change it may yet succumb.
The Red List (IUCN, 2016) has not yet been updated.
Baur and Baur (1993) concluded that the local extir-
pation of the widespread European land snail Arianta
arbustorum at sites around the city of Basel, Switzerland,
had resulted from climate warming in otherwise suitable
areas close to extensive urban development, and that
sites from which A. arbustorum had disappeared had
higher surface temperatures than sites where it remained.
The same authors (Baur and Baur, 2013) compared
historical records from 1916–1917 with survey results
from 2011–2012 on nine mountain slopes in Switzerland.
They found that the upper elevational limit for A.
arbustorum had risen 164 m in the 95 year period, during
which mean annual temperature in the area had risen
1.6 C. Although only a local impact, this study demon-
strated the potential for climate change to affect popula-
tions of land snails.
Similarly, Pearce and Paustian (2013) undertook
extensive elevational surveys in Pennsylvania, USA, to
assess whether, with climate warming, species forced
ever upward would eventually have nowhere further to
retreat to (cf. similar studies on arthropods: Meyer et al.,
2015). Of the 69 species recorded, five appeared espe-
cially susceptible. This kind of susceptibility is of partic-
ular concern on oceanic islands, especially in the Pacific.
On many Pacific islands, habitat destruction and the
introduction of invasive species at lower elevations has
resulted in most of the remaining endemic land snail
species being confined to higher elevation refugia
(Durkan et al., 2013), either because their lower eleva-
tion populations have been extirpated or because they
are evolutionarily adapted to the lower temperatures
at these higher elevations and historically only ever
occurred there. As such, with limited opportunity to
move to higher elevations as the climate warms, they
face extinction.
Thus, there is no evidence that climate change has
caused the extinction of any non-marine mollusk species.
However, continued warming will probably have more
serious effects in the future, and ocean acidification
resulting from raised carbon dioxide levels may impact
marine species (Peters et al., 2015).
EXTINCTION ON PACIFIC ISLANDS:
A CASE STUDY
Among the species assessed as extinct by Re
´gnier et al.
(2009), more than 70% were from oceanic islands and
most of these were from the Hawaiian Islands, French
Polynesia and the Mascarene Islands. Previous authors
have suggested that 65–90% of the Hawaiian land snail
species have gone extinct (Solem, 1990; Cowie and A.C.
Robinson, 2003; Lydeard et al., 2004). The proportion
differs among families, but three groups (Achatinellinae,
Amastridae, Endodontidae) appear to have suffered
“catastrophic extinction” (Solem, 1990; and see above).
To begin to get a more accurate assessment of the level
of extinction in Hawaiian land snails, Re
´gnier et al.
(2015b) focused on the Amastridae, a family endemic to
the Hawaiian Islands and with 325 known valid species
(Cowie et al., 1995). Rather than using the rigid IUCN
categories and criteria (IUCN, 2012), they took a less
rigorous approach similar to that taken by Re
´gnier et al.
(2015a). They based their assessments on a comparison
of information from historical collections and archived
field notes in the Bishop Museum, with data from
modern extensive surveys undertaken throughout the
Hawaiian Islands by K.A. Hayes, N.W. Yeung, and col-
laborators between 2004 and 2013. They also consulted a
diversity of experts with experience in the Hawaiian land
snail fauna. A species was considered extinct if it had not
been found since 2004 at any recently surveyed location
where it had formerly been recorded. In addition, and
again taking a similar approach to that adopted by
Re
´gnier et al. (2015a) in order to provide independent
corroboration, Re
´gnier et al. (2015b) undertook a statis-
tical assessment of extinction probabilities, based on
collection years and using the methods of Thompson
et al. (2013) and Lee (2014).
Of the 325 species, 131 were assessed as extinct; there
was insufficient evidence of extinction for 179, although
most of these can probably be considered extinct (and
were considered possibly extinct in the updated analysis
R.H. Cowie et al., 2017 Page 9
of global extinctions, above); but only 15 were considered
definitively extant (three subsequently found extant;
N.W. Yeung and K.A. Hayes, pers. comm.). Thus, a
minimum of 131 (40%) and maximum 310 (95%) were
considered extinct. The probabilistic assessment was
consistent with the expert assessment: the probabilities
of being extant for those species assessed as extinct
was <0.01 (111 species), <0.1 (16 species) and 0.1–0.3
(4 species); and for those species assessed as extant it was
0.38–0.93 (15 species); the species for which there was
insufficient evidence of extinction were not assessed
probabilistically. The Red List (IUCN, 2016) lists only
33 amastrid species (10%) as extinct.
There have been five phases of amastrid extinction,
discussed in more detail by Re
´gnier et al. (2015b),
as follows.
1) Description as fossils or subfossils and not known to
be extant, but it is not possible currently to determine
when they went extinct, i.e., prior to or after human
colonization of the Hawaiian Islands, or prior to or
after around the year 1500.
2) Subsequent to the first colonization of the islands
by Polynesians, 800–1000 years ago, which led to con-
siderable habitat destruction and introduction of a
number of alien species.
3) Following European colonization, when extensive
additional habitat destruction took place and highly
destructive invasive alien species were introduced.
4) Following the advent of large-scale agriculture at the
end of the 19th Century, at the time of a major
increase in land snail extinction rate globally, identi-
fied as around 1895 by Re
´gnier et al. (2015a).
5) After 1945 and the end of the Second World War, with
the increased military activity, tourism, commerce,
urbanization and rapidly increasing introduction of
invasive species, including snails (Cowie, 1998).
If the extinction rate were constant over this roughly
1,000 year period, it would have been between roughly
0.4 and 1.0% of the fauna per decade, given the extremes
of 131 and 310 of the 325 amastrid species having gone
extinct. However the rate was certainly not constant but
probably has increased in a roughly exponential and
step-wise manner over time. Re
´gnier et al. (2015b)
therefore modeled a number of scenarios reflecting this
increasing rate. The overall rates obtained ranged from
0.4% of the amastrid fauna per decade (131 species
extinct, beginning in the year 1000, as above) to 14%
per decade (310 species extinct, beginning in 1945).
These scenarios are certainly over-simplistic, but none-
theless provide a framework for discussion.
DISCUSSION
The most conservative estimate of 0.4% per decade for
the Hawaiian amastrid extinction rate is similar to the
global biodiversity extinction rate of <1% per decade
estimated by Costello et al. (2013). However, this extinc-
tion rate is probably not reflective of the true rate of
amastrid extinction, as not only is it based on a very
conservative estimate of the number of species that have
gone extinct (131) but it assumes a constant rate since
the year 1000. Undoubtedly, the rate has increased over
the millenium and Re
´gnier et al. (2015b) suggested
that a rate of around 5% per decade over the last 150–
200 years would be more realistic, indeed still rather
conservative, given the maximum rate they estimated
of 14% per decade. A rate of 5% would lead to a loss of
>50% of a fauna within 150 years (Costello et al., 2013).
Indeed, for the amastrids, with only 18 of 325 species
known to be extant (i.e., including the three species
discovered alive since Re
´gnier et al., 2015b; see above),
this scenario seems to have already played itself out.
This rate (5% per decade) is much higher than the
global estimate of the loss over the last 500 years or so
of 3,000–5,100 (10–17%) of the 30,000 known land snail
species, as estimated by Re
´gnier et al. (2015a) and out-
lined above. The amastrids, however, may be an extreme
case, although land snail groups from other Pacific
islands have suffered similar fates, notably the
Endodontidae (Solem, 1976; Zimmerman et al., 2009;
Sartori et al., 2013; 2014) and Partulidae (Coote et al.,
2003; Gerlach, 2016), and many extinct species continue
to be found, as empty shells, even before their scientific
description (e.g., Richling and Bouchet, 2013). Oceanic
island biotas are in general much more susceptible
to extinction than more buffered continental faunas
(Triantis et al., 2010). And some taxa may be more sus-
ceptible than others. Therefore it may be dangerous to
base generalizations regarding extinction rates on global
estimates, though even these, such as the loss of 10–17%
of land snail species in 500 years described here, are
cause for great concern. While it is crucial to increase
awareness of the high level of global extinction, subsum-
ing more local extinction rates, especially of particularly
susceptible faunas such as those of oceanic islands, or of
particularly susceptible taxa such as the Amastridae,
under global rates will doom those plants and animals
to extinction as their special vulnerability and need
for conservation will be overlooked, or at least not
adequately appreciated.
If we assume that 1) the 200 land snail species
sampled by Re
´gnier et al. (2015a) are representative
of known non-marine invertebrate diversity and their
extinction rate, 2) three-quarters of species are non-
marine (Mora et al., 2011), and 3) marine extinctions
are negligible compared with non-marine extinctions
(only four marine mollusks are considered as extinct;
IUCN (2016), and see above), then approximately 7.5–
13% of all species have gone extinct since around 1500.
This is orders of magnitude greater than the 860 (0.05%
of 2 million) listed as extinct by IUCN (2016). The bio-
diversity crisis is real.
But what of the IUCN? The studies reviewed herein
have shown that it is inappropriate to use the IUCN Red
List as a source of data on global extinction rates (except
for mammals and birds), and more generally that
Page 10 THE NAUTILUS, Vol. 131, No. 1
assessing overall levels of threat to all biodiversity based
on the species listed by IUCN seriously downplays that
threat, notably because the great majority of biodiversity
(invertebrates) has not been evaluated. A similar critique
was voiced by Possingham et al. (2002), who argued that
threatened species lists (such as the IUCN Red List)
should not be used to indicate the overall status of bio-
diversity and changes in it, largely because of uneven
taxonomic treatment and variation in observational effort
(as described above for vertebrates compared to inverte-
brates). Nonetheless, IUCN is the premier and most
influential global conservation organization. But its goal
is to “highlight taxa threatened with extinction, and
thereby promote their conservation” (IUCN, 2016);
documenting extinction is incidental to this goal as once
extinct a species cannot be conserved. For terrestrial
vertebrates IUCN is well on the way to achieving its goal,
but invertebrates present a daunting challenge both
because of their immense diversity and because of the
lack of adequate data to apply the IUCN criteria for the
vast majority of them.
Major focused efforts by IUCN continue to be made
to evaluate additional mollusk species (e.g., Seddon,
2011; 2014; 2015; Pippard, 2012; Peters et al., 2013;
Seddon et al., 2014; Allen et al., 2016; Bo
¨hm and
Allcock, 2016; Neubert et al., 2017). These efforts have
focused on particular taxa, habitats and geographic loca-
tions that were deemed a priori as especially in need of
attention, i.e., to address the IUCN goals of highlighting
taxa in need of conservation (above), and for which
funding could be obtained. Nonetheless, at the current
rate it will be many years before all mollusks, or even all
non-marine mollusks, have been assessed. The approach
we have taken in the two studies reviewed herein
(Re
´gnier et al., 2015a, b), as well as our update of the
analysis of Re
´gnier et al. (2009) based on new informa-
tion, is an attempt to speed up the evaluation process
and to develop a method that allows global trends to be
more realistically determined. Admittedly, our approach
is less rigorous than the process of applying the IUCN
criteria to assign species to the IUCN categories, with
peer review required (when at best only one specialist
has any knowledge of the fauna), but is considerably
quicker and more cost-effective. While there is a chance
that our approach might incorrectly list some species as
extinct and thereby cut them off from conservation
attention, it has the potential to identify many more
species in need of conservation, species that would be
listed as Data Deficient by IUCN and therefore also
ignored. Our approach also has the advantage that it
can provide a much more realistic overview of the bio-
diversity crisis than can the Red List, especially regard-
ing the extraordinary levels of extinction, which was
our immediate focus in the studies reviewed above.
Nonetheless, IUCN remains the preeminent global con-
servation agency with a crucial role in identifying con-
servation needs and developing global conservation
strategies. Our efforts do not in any way compromise
those roles.
ACKNOWLEDGMENTS
We thank our collaborators Amaury Lambert and
Guillaume Achaz (Re
´gnier et al., 2015a) and Ken Hayes,
Norine Yeung, Carl Christensen, and Daniel Chung
(Re
´gnier et al., 2015b). We also thank all the experts who
contributed information and assessments for those two
studies, the numerous students and others who par-
ticipated in the Hawaiian survey work, and others who
provided assistance, as acknowledged in those two
publications. This paper is based on a presentation
given by Robert Cowie at the Mollusks in Peril Forum
(Bailey-Matthews National Shell Museum, Sanibel
Island, Florida, May 2016). We thank the organizers of
the Forum, notably Jose
´H. Leal and Dorrie Hipschman,
respectively Science Director and Executive Director
of the Bailey-Matthews National Shell Museum, and
Museum benefactors Smoky and Stephanie Payson, for
the invitation to participate and for funding in support of
that participation. Additional information was provided
by M.R. Alonso, Gary Barker, Ru¨ diger Bieler, Fred
Brook, Robert Cameron, Satoshi Chiba, Carl Christensen,
Zoltan Fehe
´r, Justin Gerlach, Jozef Grego, Brenden
Holland, Yasunori Kano, Miguel Iban
˜ez, Ben Rowson,
Rebecca Rundell, Menno Schilthuizen, Steve Trewick,
and Norine Yeung. We thank Chuck Lydeard for
reviewing the manuscript. This contribution is partly
based on research supported by the French National
Research Agency Losers Project Grant ANR-09-PEXT-
007, an Ars Cuttoli Foundation grant, NSF grant DEB-
1120906 and by grants from the U.S. Department of
Agriculture (CAPS program) and the Oahu Army Natural
Resources Program. Contribution number 9977 of the
University of Hawaii School of Ocean and Earth Sciences.
LITERATURE CITED
Abdou A. and P. Bouchet. 2000. Nouveaux gaste
´ropodes
Endodontidae et Punctidae (Mollusca, Pulmonata)
re
´cemment e
´teints de l’archipel des Gambier (Polyne
´sie).
Zoosystema 22: 689–707.
Allen, D., E. Neubert, and M.[B.] Seddon. 2016. Final stage
of the European Red List of terrestrial molluscs starts.
Tentacle 24: 55–56.
Anthony, J.L. and J.A. Downing. 2001. Exploitation trajectory
of a declining fauna: a century of freshwater mussel
fisheries in North America. Canadian Journal of Fisheries
and Aquatic Sciences 58: 2071–2090.
Athens, J.S. 2009. Rattus exulans and the catastrophic dis-
appearance of Hawai’i’s native lowland forest. Biological
Invasions 11: 1489–1501.
Battarbee, R.W. 2014. The rediscovery of the Aldabra banded
snail, Rhachistia aldabrae. Biology Letters 10: 20140771.
http://dx.doi.org/10.1098/rsbl.2014.0771
Baur, B. and A. Baur. 1993. Climatic warming due to thermal
radiation from an urban area as possible cause for the
local extinction of a land snail. Journal of Applied Ecology
30: 333–340.
Baur, B. and A. Baur. 2013. Snails keep the pace: shift in upper
elevation limit on mountain slopes as a response to climate
warming. Canadian Journal of Zoology 91: 596–599.
R.H. Cowie et al., 2017 Page 11
Beasley, C.R. 2001. The impact of exploitation on freshwater
mussels (Bivalvia: Hyriidae) in the Tocantins River, Brazil.
Studies on Neotropical Fauna and Environment 36: 159–165.
Benı
´tez, G. 2011. Animals used for medicinal and magico-
religious purposes in western Granada Province, Andalusia
(Spain). Journal of Ethnopharmacology 137: 1113–1123.
Bo
¨hm, M. and L. Allcock. 2016. New cephalopod additions
to the IUCN Red List. Tentacle 24: 54–55.
Bouchet, P. and A. Abdou. 2001. Recent extinct land snails
(Euconulidae) from the Gambier Islands with remarkable
apertural barriers. Pacific Science 55: 121–127.
Bouchet, P. and A. Abdou. 2003. Endemic land snails from
the Pacific islands and the museum record: documenting
and dating the extinction of the terrestrial Assimineidae
of the Gambier Islands. Journal of Molluscan Studies
69: 165–170.
Boyer, S., S.D. Wratten, A. Holyoake, J. Abdelkrim, and R.H.
Cruickshank. 2013. Using next generation sequencing
to analyse the diet of a highly endangered land snail
(Powelliphanta augusta) feeding on endemic earthworms.
PloS ONE 8(9): e75962.
Brescia, F.M., C.M. Po
¨llabauer, M.A. Potter and A.W. Robertson.
2008. A review of the ecology and conservation of
Placostylus (Mollusca: Gastropoda: Bulimulidae) in New
Caledonia. Molluscan Research 28: 111–122.
Carlton, J.T. 1993. Neoextinctions of marine invertebrates.
American Zoologist 33: 499–509.
Ceballos, G., P.R. Ehrlich, A.D. Barnosky, A. Garcı
´a, R.M.
Pringle, and T.M. Palmer. 2015. Accelerated modern
human-induced species losses: entering the sixth mass
extinction. Science Advances 1(5): e1400253.
Chapman, A.D. 2009. Number of Living Species in Australia
and the World. Second edition. Department of the
Environment, Water, Heritage and the Arts, Australian
Government, Canberra, 80 pp.
Chiba, S. and R.H. Cowie. 2016. Evolution and extinction of
land snails on oceanic islands. Annual Review of Ecology,
Evolution, and Systematics 47: 123–141.
Conte, E. and P.V. Kirch. 2008. One thousand years of human
environmental transformation in the Gambier Islands
(French Polynesia). In: G. Clark, F. Leach and S.
O’Connor (eds.) Islands of Inquiry (Terra Australis 29).
Colonization, Seafaring and the Archaeology of Maritime
Landscapes. Australian National University Press, Acton,
pp. 253–264.
Coote, T. and E
´. Loe
`ve. 2003. From 61 species to five: endemic
tree snails of the Society Islands fall prey to an ill-judged
biological control programme. Oryx 37: 91–96.
Costello, M.J., R.M. May and N.E. Stork. 2013. Can we
name Earth’s species before they go extinct? Science
339: 413–416.
Cowie, R.H. 1993. Why tree snails are becoming scarce
in Samoa. Hawaiian Shell News 41(3): 1, 9.
Cowie, R.H. 1998. Patterns of introduction of non-indigenous
non-marine snails and slugs in the Hawaiian Islands.
Biodiversity and Conservation 7: 349–368.
Cowie, R.H. 2001. Can snails ever be effective and safe bio-
control agents? International Journal of Pest Management
47: 23–40.
Cowie, R.H. 2004. Disappearing snails and alien invasions: the
biodiversity/conservation interface in the Pacific. Journal
of Conchology Special Publications 3: 23–37.
Cowie, R.H. 2005. Alien non-marine mollusks in the islands of
the tropical and subtropical Pacific: a review. American
Malacological Bulletin 20: 95–103.
Cowie, R.H. and R.P. Cook. 2001. Extinction or survival:
partulid tree snails in American Samoa. Biodiversity and
Conservation 10: 143–159.
Cowie, R.H. and A.C. Robinson. 2003. The decline of native
Pacific island faunas: changes in status of the land snails of
Samoa through the 20th century. Biological Conservation
110: 55–65.
Cowie, R.H. and D. G. Robinson. 2003. Pathways of introduc-
tion of nonindigenous land and freshwater snails and
slugs. In: G. Ruiz and J.T. Carlton (eds.) Invasive Species:
Vectors and Management Strategies. Island Press,
Washington, D.C., pp. 93–122.
Cowie, R.H., N.L. Evenhuis, and C.C. Christensen. 1995.
Catalog of the native land and freshwater molluscs of
the Hawaiian Islands. Backhuys Publishers, Leiden. vi þ
248 pp.
Curry, P.A., N.W. Yeung, K.A. Hayes, W.M. Meyer, III, A.D.
Taylor, and R.H. Cowie. 2016. Rapid range expansion of
an invasive predatory snail, Oxychilus alliarius (Miller
1822), and its impact on endemic Hawaiian land snails.
Biological Invasions 18: 1769–1780.
Didham, R.K., J.M. Tylianakis, N.J. Gemmell, T.A. Rand, and
R.M. Ewers. 2007. Interactive effects of habitat modifica-
tion and species invasion on native species decline.
Trends in Ecology and Evolution 22(9): 489–496.
Durkan, T.H., N.W. Yeung, W.M. Meyer, III, K.A. Hayes, and
R.H. Cowie. 2013. Evaluating the efficacy of land snail
survey techniques in Hawaii: implications for conservation
throughout the Pacific. Biodiversity and Conservation
22: 3223–3232.
Fehe
´r, Z. 2013. Melanopsis parreyssii. The IUCN Red List of
Threatened Species 2013: e.T155737A4835365. http://dx
.doi.org/10.2305/IUCN.UK.2011-1.RLTS.T155737A4835
365.en. Accessed 23 June 2016.
Gaston, K.J. and R.M. May. 1992. Taxonomy of taxonomists.
Nature 356: 281–282.
Gerlach, J. 2007. Short-term climate change and the extinction
of the snail Rhachistia aldabrae (Gastropoda: Pulmonata).
Biology Letters 3: 581–585.
Gerlach, J. 2009. Rhachistia aldabrae. The IUCN Red List of
Threatened Species 2009: e.T168122A6454921.
Gerlach, J. 2016. Icons of Evolution: Pacific Island Tree-Snails
of the Family Partulidae. Phelsuma Press, Cambridg,
334 pp.
Groombridge, B. (ed.) 1994. 1994 IUCN Red List of Threat-
ened Animals. IUCN, Gland, lvi þ286 pp.
Hadfield, M.G. 1986. Extinction in Hawaiian achatinelline
snails Malacologia 27: 67–81.
Hadfield, M.G., S.E. Miller and A.H. Carwile. 1993. The
decimation of endemic Hawai’ian [sic] tree snails by alien
predators. American Zoologist 33: 610–622.
IISE (International Institute for Species Exploration. 2017.
SOS Reports. http://www.esf.edu/species/SOS.htm. Accessed
15 February 2017.
IUCN. 1990. 1990 IUCN Red List of Threatened Animals.
IUCN, Gland and Cambridge, xxiv þ[12] þ192 pp.
IUCN. 2012. IUCN Red List Categories and Criteria. Version 3.1.
Second edition. IUCN, Gland and Cambridge, iv þ32 pp.
IUCN. 2016. The IUCN Red List of Threatened Species.
Version 2016-3. http://www.iucnredlist.org. Accessed
9 December 2016.
James, A.F., R. Brown, K.A. Weston, and K. Walker. 2013.
Modelling the growth and population dynamics of the
exiled Stockton coal plateau landsnail, Powelliphanta
augusta. New Zealand Journal of Zoology 40: 175–185.
Page 12 THE NAUTILUS, Vol. 131, No. 1
Ko
¨hler, F., Mary [B.] Seddon, A.E. Bogan, D.V. Tu, P. Sri-
Aroon and D. Allen. 2012. The status and distribution of
freshwater molluscs of the Indo-Burma region. In: D.J.
Allen, K.G. Smith and W.R.T. Darwall (compilers)
The Status and Distribution of Freshwater Biodiversity
in Indo-Burma. IUCN, Cambridge, UK, and Gland,
Switzerland, pp. 66–88.
Lee, T.E. 2014. A simple numerical tool to infer whether a
species is extinct. Methods in Ecology and Evolution
5: 791–796.
Leakey, R.E. and R. Lewin. 1996. The Sixth Extinction.
Biodiversity and its Survival. Weidenfeld and Nicolson,
London, 288 pp.
Ligaszewski, M., A. Łysak, and Z. Mach-Paluszkiewicz. 2007.
Reproductive performance of Helix pomatia (Gastropoda:
Pulmonata: Helicidae) and survival of its hatchlings under
farm conditions. American Malacological Bulletin 22: 1–6.
Lomborg, B. 2001. The Skeptical Environmentalist: Measuring
the Real State of the World. Cambridge University Press,
Cambridge, xxiii þ515 pp.
Lydeard, C., R.H. Cowie, W.F. Ponder, A.E. Bogan, P.
Bouchet, S. Clark, K.S. Cummings, T.J. Frest, O.
Gargominy, D.G. Herbert, R. Hershler, K. Perez, B. Roth,
M. Seddon, E.E. Strong, and F.G. Thompson. 2004.
The global decline of nonmarine mollusks. BioScience
54: 321–330.
May, R.M. 2011. Why worry about how many species and their
loss? PLoS Biology 9: e1001130.
Meyer, W.M., III, J.A. Eble, K. Franklin, R.B. McManus,
S.L. Brantley, J. Henkel, P.E. Marek, W.E. Hall, C.A.
Olson, R. McInroy, E.M. Bernal Loaiza, R.C. Brusca,
and W. Moore. 2015. Ground-dwelling arthropod com-
munities of a sky island mountain range in southeastern
Arizona, USA: obtaining a baseline for assessing the
effects of climate change. PLoS ONE 10(9): e0135210.
Mora, C., D.P. Tittensor, S. Adl, A.G.B. Simpson and B. Worm.
2011. How many species are there on earth and in the
ocean? PLoS Biology 9(8): e1001127.
Morris, R. 2010. An unfortunate Experiment. Forest and Bird
337: 14–18.
Mrosovsky, N. 1997. IUCN’s credibility critically endangered.
Nature 389: 436.
Murray, J., E. Murray, M.S. Johnson and B. Clarke. 1988.
The extinction of Partula on Moorea Pacific Science
42: 150–153.
Neto, N.A.L., R.A. Voeks, T.L.P. Dias, and R.R.N. Alves. 2012.
Mollusks of Candomble
´: symbolic and ritualistic impor-
tance. Journal of Ethnobiology and Ethnomedicine 8: 10.
Neubert E., C. Che
´rel-Mora, and P. Bouchet. 2009. Polytypy,
clines, and fragmentation: the bulimes of New Caledonia
revisited (Pulmonata, Orthalicoidea, Placostylidae).
Me
´moires du Muse
´um National d’Histoire Naturelle
198: 37–131.
Neubert, E., M.B. Seddon, and D. Allen. 2017. IUCN review
workshop on terrestrial molluscs of Europe successfully
held in Uppsala. Tentacle 25: in press.
Neves, R. J. 1999. Conservation and commerce: management
of freshwater mussel (Bivalvia: Unionoidea) resources in
the United States. Malacologia 41: 461–474.
Novacek, M.J. 2001. The Biodiversity Crisis: Losing What
Counts. New Press, New York. 224 pp.
Nyoagbe, L.A., V. Appiah, J. Nketsia-Tabiri, D. Larbi and I.
Adjei. 2016. Evaluation of African giant snails (Achatina
and Archachatina) obtained from markets (wild) and breed-
ing farms. African Journal of Food Science 10: 94–104.
Osemeobo G.J. 1991. Effects of common property resource
utilization on wildlife conservation in Nigeria. GeoJournal
23: 241–248.
Pearce, T.A. and M.E. Paustian. 2013. Are temperate land
snails susceptible to climate change through reduced
altitudinal ranges? A Pennsylvania example. American
Malacological Bulletin 31: 213–224.
Peters, H., B.C. O’Leary, J.P. Hawkins, K.E. Carpenter and
C.M. Roberts. 2013. Conus: first comprehensive conser-
vation Red List assessment of a marine gastropod mollusc
genus. PLoS ONE 8(12): e83353. doi:10.1371/journal
.pone.0083353
Peters, H., B. O’Leary, J.P. Hawkins and C.M. Roberts. 2015.
Identifying species at extinction risk using global
models of anthropogenic impact. Global Change Biology
21: 618–628.
Pippard, H. 2012. Data evaluation begins for Pacific Island
land snails. Tentacle 20: 37.
Possingham, H.P., S.J. Andelman, M.A. Burgman, R.A.
Medellı
´n, L.L. Master, and D.A. Keith. 2002. Limits to
the use of threatened species lists. Trends in Ecology and
Evolution 17: 503–507.
Re
´gnier, C., B. Fontaine, and P. Bouchet. 2009. Not knowing,
not recording, not listing: numerous unnoticed mollusk
extinctions. Conservation Biology 23: 1214–1221.
Re
´gnier, C., G. Achaz, A. Lambert, R.H. Cowie, P. Bouchet
and B. Fontaine. 2015a. Mass extinction in poorly known
taxa. Proceedings of the National Academy of Sciences
112(25): 7761–7766.
Re
´gnier, C., P. Bouchet, K.A. Hayes, N. W. Yeung, C.C.
Christensen, D.J.D. Chung, B. Fontaine, and R. H.
Cowie. 2015b. Extinction in a hyperdiverse endemic
Hawaiian land snail family and implications for the under-
estimation of invertebrate extinction. Conservation
Biology 29: 1715–1723.
Ricciardi, A., R.J. Neves, and J. B. Rasmussen. 1998.
Impending extinction of North American freshwater
mussels (Unionoida) following zebra mussel (Dreissena
polymorpha) invasion. Journal of Animal Ecology 67:
613–619.
Richling, I. and P. Bouchet. 2013. Extinct even before scientific
recognition: a remarkable radiation of helicinid snails
(Helicinidae) on the Gambier Islands, French Polynesia.
Biodiversity and Conservation 22: 2433–2468.
Rosenberg, G. 2014. A new critical estimate of named species-
level diversity of the Recent Mollusca. American Malaco-
logical Bulletin 32: 308–322.
Sartori, A.F., O. Gargominy, and B. Fontaine. 2013. Anthropo-
genic extinction of Pacific land snails: a case study of Rurutu,
French Polynesia, with description of eight new species
of endodontids (Pulmonata). Zootaxa 3640: 343–372.
Sartori, A.F., O. Gargominy, and B. Fontaine. 2014. Radiation
and decline of endodontid land snails in Makatea, French
Polynesia. Zootaxa 3772(1): 1–68.
Seddon, M.[B.] 2011. A sampled approach to Red Listing:
freshwater mollusc assessment. Tentacle 19: 50–51.
Seddon, M.[B.] 2014. Annual report of the IUCN SSC Mollusc
Specialist Group 2013. Tentacle 22: 39–41.
Seddon, M.[B.] 2015. Progress on listing. Tentacle 23: 37.
Seddon, M.B., U
¨. Kebapc¸i, M. Lopes-Lima, D. van Damme
and K.G. Smith. 2014. Freshwater molluscs. In: K.G.
Smith, V. Barrios, W.R.T. Darwall and C. Numa (eds.).
The Status and Distribution of Freshwater Molluscs in
the Eastern Mediterranean. IUCN, Cambridge, Malaga
and Gland, pp. 43–56.
R.H. Cowie et al., 2017 Page 13
ˆrbu, I. and A.M. Benedek. 2016. Requiem for Melanopsis
parreyssii or the anatomy of a new extinction in Romania.
Tentacle 24: 26–28.
ˆrbu, I., A. Gagiu and A.M. Benedek. 2013. On the brink of
extinction: fate of the Pet¸ea thermal lake (Romania) and
its endemic species. Tentacle 21: 35–37.
Solem, A. 1990. How many Hawaiian land snail species are
left? and what we can do for them. Bishop Museum Occa-
sional Papers 30: 27–40.
Solem, A. 1976. Endodontoid land snails from Pacific Islands
(Mollusca: Pulmonata: Sigmurethra). Part 1. Family
Endodontidae. Field Museum of Natural History,
Chicago, xii þ508 pp.
Strayer, D.L. and H.M. Malcom. 2007. Effects of zebra
mussels (Dreissena polymorpha) on native bivalves:
the beginning of the end or the end of the beginning?
Journal of the North American Benthological Society
26: 111–112.
Strayer, D.L., J.A. Downing, W.R. Haag, T.L. King, J.B.
Layzer, T.J. Newton and S.J. Nichols. 2004. Changing
perspectives on pearly mussels, North America’s most
imperiled animals. BioScience 54: 429–439.
Thompson, C.J., T.E. Lee, L. Stone, M.A. McCarthy and M.A.
Burgman. 2013. Inferring extinction risks from sighting
records. Journal of Theoretical Biology 338: 16–22.
Trewick, S.A. 2005. Taxonomic status of the land snail
Powelliphanta “Augustus”—evidence from mitochondrial
DNA data. Unpublished report to the New Zealand
Department of Conservation, Hokitika, 4 pp.
Trewick, S.A., K.J. Walker and C.J. Jordan. 2008. Taxonomic
and conservation status of a newly discovered giant
landsnail from Mount Augustus, New Zealand. Conserva-
tion Genetics 9: 1563–1575.
Triantis, K.A., P.A.V. Borges, R.J. Ladle, J. Hortal, P. Cardoso,
C. Gaspar, F. Dinis, E. Mendonc¸a, L.M.A. Silveira,
R. Gabriel, C. Melo, A.M.C. Santos, I.R. Amorim, S.P.
Ribeiro, A.R.M. Serrano, J.A. Quartau, and R.J. Whittaker.
2010. Extinction debt on oceanic islands. Ecography 33:
285–294.
Va
´zquez, A.A., J. Sa
´nchez, E. Martı
´nez, and A. Alba. 2016.
Facilitated invasion of an overseas invader: human
mediated settlement and expansion of the giant African
snail, Lissachatina fulica, in Cuba. Biological Invasions
DOI 10.1007/s10530-016-1266-3, on line early
Walker, K.J., S.A. Trewick, and G.M. Barker. 2008.
Powelliphanta augusta, a new species of land snail, with a
description of its former habitat, Stockton coal plateau,
New Zealand. Journal of the Royal Society of New
Zealand 38: 163–186.
Wells, S.M., R.M. Pyle, and N.M. Collins. 1983. The IUCN
Invertebrate Red Data Book. IUCN, Gland. [ii] þi–l þ
632 pp.
Whitmore, N. 2016. Harnessing local ecological knowledge for
conservation decision making via Wisdom of Crowds: the
case of the Manus green tree snail Papustyla pulcherrima.
Oryx 50(4): 684–692.
WoRMS Editorial Board. 2017. World Register of Marine
Species. Available from http://www.marinespecies.org at
VLIZ. Accessed 15 February 2017. doi:10.14284/170
Yildirim, M.Z., U
¨. Kebapc¸ i, and B.A. Gu¨mu¨ . 2004. Edible
snails (terrestrial) of Turkey. Turkish Journal of Zoology
28: 329–335.
Zimmermann, G., O. Gargominy, and B. Fontaine. 2009.
Quatre espe
`ces nouvelles d’Endodontidae (Mollusca,
Pulmonata) e
´teints de Rurutu (I
ˆles Australes, Polyne
´sie
franc¸aise). Zoosystema 31: 791–805.
Page 14 THE NAUTILUS, Vol. 131, No. 1
Table A1. Species considered extinct, extinct in the wild or critically endangered (possibly extinct) by Re
´gnier et al. (2009) and/or
the Red List (IUCN, 2016) but now known or thought to be extant (or unrankable). DD – data deficient, VU – vulnerable,
EN – endangered, CR – critically endangered, EW – extinct in the wild, EX – Extinct.
Species Red List
Re
´gnier
et al.
References indicating
species is extant
ACHATINELLIDAE
Achatinella livida Swainson, 1828 EX Extant M.G. Hadfield, pers. comm.
Auriculella uniplicata (Pease, 1868) EX Extant N.Y. Yeung, pers. comm.
Perdicella fulgurans Sykes, 1900 EX Extant M.G. Hadfield, pers. comm.
Perdicella maniensis (Pfeiffer, 1856) EX Extant M.G. Hadfield, pers. comm.
Perdicella zebrina (Pfeiffer, 1856) EX Extant M.G. Hadfield, pers. comm.
CERASTIDAE
Rhachistia aldabrae (Martens, 1898) EX EX Battarbee, 2014
CYCLOPHORIDAE
Cyclophorus horridulum (Morelet, 1882) EX EX? Abdou et al., 2004
Cyclosurus mariei Morelet, 1881 EX Abdou et al., 2004
HYDROBIIDAE
Belgrandiella zermanica Radoman, 1973 VU EX Slapnik and Lajtner, 2011
Bracenica spiridoni Radoman, 1973 EN EX Pes
ˇic
´, 2010a; Pes
ˇic
´and Glo
¨er, 2013
Islamia zermanica Radoman, 1973 CR(PE) EX Beran et al., 2016
Marstonia castor (Thompson, 1977) CR EX? Johnson et al., 2013
Mercuria globulina (Letourneux and Bourguignat, 1887) EX Glo
¨er et al., 2015
Tanousia zrmanjae (Brusina, 1866) CR EX Beran, 2011; Falniowski, 2011a
Vinodolia fiumana Radoman, 1973 EN EX Szarowska et al., 2013; Falniowski and Seddon, 2014
Vinodolia fluviatilis (Radoman, 1973) EN EX Beran, 2011; Falniowski, 2011b
Vinodolia gluhodolica (Radoman, 1973) EN EX Pes
ˇic
´, 2010b; Glo
¨er and Pes
ˇic
´, 2014
LITHOGLYPHIDAE
Clappia cahabensis Clench, 1965 EX Johnson et al., 2013
Somatogyrus alcoviensis Krieger, 1915 EX Johnson et al., 2013
Somatogyrus amnicoloides Walker, 1915 EX EX “unrankable”, Johnson et al., 2013
Somatogyrus crassus Walker, 1904 CR(PE) “unrankable”, Johnson et al., 2013
Somatogyrus currierianus Lea, 1863 CR(PE) “unrankable”, Johnson et al., 2013
Somatogyrus hendersoni Walker, 1909 CR(PE) “unrankable”, Johnson et al., 2013
Somatogyrus humerosus Walker, 1906 CR(PE) “unrankable”, Johnson et al., 2013
Somatogyrus nanus Walker, 1904 CR(PE) “unrankable”, Johnson et al., 2013
LITTORINIDAE
Littoraria flammea (Philippi, 1847) EX EX Dong et al., 2015
NEOCYCLOTIDAE
Incerticyclus martinicensis (Shuttleworth, 1857) EX EX Delannoye et al., 2015
PARTULIDAE
Partula leefei Smith, 1897 EX Gerlach, 2016
Partula nodosa Pfeiffer, 1853 EW EW Gerlach, 2016
Samoana annectens (Pease, 1864) DD EX Gerlach, 2016
Samoana diaphana (Crampton and Cooke, 1953) EN EX Gerlach, 2016
Samoana inflata (Reeve, 1842) EX Gerlach, 2016
PHYSIDAE
Physella columbiana Keep, 1887 EX? Johnson et al., 2013
Physella hemphilli Taylor, 2003 EX? Johnson et al., 2013
PLANORBIDAE
Rhodacmea filosa (Conrad, 1834) CR EX O
´Foighil et al., 2011
PLEUROCERIDAE
Elimia lachryma (Reeve, 1861) EX Johnson et al., 2013
Elimia melanoides (Conrad, 1834) EX Minton et al., 2003; Johnson et al., 2013
Elimia mutabilis (Lea, 1862) EX? Johnson et al., 2013
Elimia troostiana (Lea, 1838) CR(PE) Johnson et al., 2013
Elimia varians (Lea, 1861) VU EX Cordeiro and Perez, 2012
Elimia vanuxemiana Lea, 1843 EX Johnson et al., 2013
Leptoxis compacta (Anthony, 1854) EX EX Whelan et al., 2012
(Continued)
APPENDIX
R.H. Cowie et al., 2017 Page 15
Table A1. (cont.)
Species Red List
Re
´gnier
et al.
References indicating
species is extant
Leptoxis foremanii (Lea, 1843) EX EX Johnson et al., 2013
STREPTAXIDAE
Gulella mayottensis (Connolly, 1885) EX Abdou et al., 2004
VIVIPARIDAE
Tchangmargarya yangtsunghaiensis
(Tchang and Tsi, 1949)
CR(PE) Zhang et al., 2015
UNIONIDAE
Pleurobema taitianum (Lea, 1834) EN EX Cummings and Cordeiro, 2012
Table A2. Species listed on the Red List (IUCN, 2016) or by Re
´gnier et al. (2009) as extinct or critically endangered (possibly
extinct) but now considered as subspecies or synonyms of other species, as nomina dubia, or that are undescribed. These are
excluded from the present analysis.
Species Red List Re
´gnier et al.
Source for subspecies/synonym/
undescribed status
HYDROBIIDAE
Bythiospeum dubium
(Geyer, 1904)
CR(PE) Synonym of Bythiospeum acicula;
Richling et al., 2016
Bythiospeum gonostoma
(Geyer, 1905)
CR(PE) Synonym of Bythiospeum acicula;
Richling et al., 2016
Bythiospeum putei
(Geyer, 1904)
CR(PE) Synonym of Bythiospeum acicula;
Richling et al., 2016
Bythiospeum turritum
(Clessin, 1877)
CR(PE) Synonym of Bythiospeum acicula;
Richling et al., 2016
Pseudamnicola desertorum
(Bourguignat, 1862)
EX 2010 Synonym of Pseudamnicola letourneuxiana;
Glo
¨er et al., 2010
ORTHALICIDAE
Bulimulus sp. nov. ‘josevillani CR(PE) undescribed; IUCN (2016)
Bulimulus sp. nov. ‘krameri CR(PE) undescribed; IUCN (2016)
Bulimulus sp. nov. ‘nilsodhneri CR(PE) undescribed; IUCN (2016)
Bulimulus sp. nov. ‘tuideroyi CR(PE) undescribed; IUCN (2016)
Bulimulus sp. nov. ‘vanmoli CR(PE) undescribed; IUCN (2016)
PARTULILDAE
Partula callifera Pffeiffer, 1857 EX Subspecies of Partula dentifera;
Gerlach, 2016
Partula candida Crampton, 1956 EX Subspecies of Partula dentifera;
Gerlach, 2016
Partula cedista Crampton, 1956 EX Subspecies of Partula dentifera;
Gerlach, 2016
Partula citrina Pease, 1866 EX Subspecies of Partula dentifera;
Gerlach, 2016
Partula formosa Garrett, 1884 EX Subspecies of Partula dentifera;
Gerlach, 2016
Partula imperforata Pfeiffer, 1877 EX Subspecies of Partula dentifera;
Gerlach, 2016
Partula raiatensis Garrett, 1884 EX Synonym of Partula imperforata;
Gerlach, 2016
PLEUROCERIDAE
Elimia timida timida Goodrich, 1942 EX Dillon and Robinson, 2011
UNIONIDAE
Nodularia cariei (Germain, 1919) EX 1996 EX Synonym of Coelatura aegyptiaca;
Graf and Cummings, 2009
Unio madagascariensis Sganzin, 1842 EX 2016 Nomen dubium; Graf and Cummings, 2009
Unio malgachensis Germain, 1911 EX 2016 Nomen dubium; Graf and Cummings, 2009
Page 16 THE NAUTILUS, Vol. 131, No. 1
Table A3. Terrestrial species considered extinct (EX), possibly extinct (EX?) or extinct in the wild (EW) in the present study,
compared with their status as evaluated by Re
´gnier et al. (2009), and on the Red List (IUCN, 2016). Red List categories are extinct
(EX), extinct in the wild (EW), critically endangered (possibly extinct) (CR(PE)), critically endangered (CR), and data deficient
(DD); the date of the IUCN evaluation follows the status. EX? is treated as equivalent to CR(PE). A dash indicates the species was
not evaluated. Sources are only provided if the status in this study differs from the later of IUCN (2016) and Re
´gnier et al. (2009).
Species explicitly described as fossil or subfossil are asterisked.
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
ACHATINELLIDAE
Achatinella abbreviata Reeve, 1850 EX 1996 EX EX
Achatinella apexfulva (Dixon, 1789) EX? B.S. Holland, pers. comm., 2016
Achatinella buddii Newcomb, 1854 EX 1990 EX EX
Achatinella caesia Gulick, 1858 EX 1990 EX EX
Achatinella casta Newcomb, 1854 EX 1990 EX EX
Achatinella cestus Newcomb, 1854 EX? USFWS, 1993
Achatinella decora (Fe
´rrusac, 1821) EX 1990 EX EX
Achatinella dimorpha Gulick, 1858 EX 1990 EX EX
Achatinella elegans Newcomb, 1854 EX 1996 EX EX
Achatinella juddii Baldwin, 1895 EX 1996 EX EX
Achatinella juncea Gulick, 1856 EX 1996 EX EX
Achatinella lehuiensis Smith, 1873 EX 1990 EX EX
Achatinella papyracea Gulick, 1856 EX 1990 EX EX
Achatinella phaeozona Gulick, 1856 EX? USFWS 1993
Achatinella rosea Swainson, 1828 EX USFWS 1993
Achatinella spaldingi Pilsbry and Cooke, 1914 EX 1990 EX EX
Achatinella stewartii (Green, 1827) EX? B.S. Holland, pers. comm., 2016
Achatinella thaanumi Pilsbry and Cooke, 1914 EX 1990 EX EX
Achatinella valida Pfeiffer, 1855 EX 1990 EX EX
Achatinella viridans Mighels, 1845 EX? USFWS, 1993
Achatinella vittata Reeve, 1850 EX USFWS, 1993
Achatinella vulpina (Fe
´rrusac, 1821) EX USFWS, 1993
Auriculella expansa Pease, 1868 EX 1994 EX EX
Hotumatua anakenana Kirch et al., 2009 EX Kirch et al., 2009
Lamellidea monodonta (Pilsbry and Hirase, 1904) EX 1994 EX EX
Lamellidea nakadai (Pilsbry and Cooke, 1915)*EX 1994 EX
Newcombia canaliculata (Baldwin, 1893) EX B.S. Holland, pers. comm., 2016
Newcombia gagei Severns, 2009* EX Severns, 2009
Newcombia perkinsi Sykes, 1896 EX B.S. Holland, pers. comm., 2016
Newcombia pfeifferi (Newcomb, 1853) EX B.S. Holland, pers. comm., 2016
Newcombia philippiana (Pfeiffer, 1857) EX 1994 EX
Newcombia sulcata (Pfeiffer, 1857) EX B.S. Holland, pers. comm., 2016
Partulina variabilis (Newcomb, 1854) EW
Partulina confusa (Sykes, 1900) CR 1996 EX EX
Partulina crassa (Newcomb, 1854) EX 1986 EX EX
Partulina dubia (Newcomb, 1853) EX B.S. Holland, pers. comm., 2016
Partulina horneri (Baldwin, 1895) EX EX
Partulina montagui Pilsbry, 1913 EX 1986 EX EX
Partulina semicarinata (Newcomb, 1854) EW
Perdicella carinella (Baldwin, 1906) EX B.S. Holland, pers. comm., 2016
Perdicella ornata (Newcomb, 1854) EX B.S. Holland, pers. comm., 2016
Perdicella thwingi (Pilsbry and Cooke, 1914) EX B.S. Holland, pers. comm., 2016
Perdicella zebra (Newcomb, 1855) EX 1994 EX EX
Tornelasmias capricorni Iredale, 1944 EX 1996 EX EX
AMASTRIDAE
Amastra abavus Hyatt and Pilsbry, 1911) EX? Re
´gnier et al., 2015
Amastra aemulator Hyatt and Pilsbry, 1911) EX? Re
´gnier et al., 2015
Amastra affinis (Newcomb, 1854) EX Re
´gnier et al., 2015
Amastra albocincta Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Amastra albolabris (Newcomb, 1854) EX 1994 EX EX Re
´gnier et al., 2015
Amastra amicta Smith, 1873 EX? Re
´gnier et al., 2015
(Continued)
R.H. Cowie et al., 2017 Page 17
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Amastra anthonii (Newcomb, 1861) EX Re
´gnier et al., 2015
Amastra antiqua (Baldwin, 1895)* EX Re
´gnier et al., 2015
Amastra assimilis (Newcomb, 1854) EX Re
´gnier et al., 2015
Amastra aurostoma Baldwin, 1896) EX? Re
´gnier et al., 2015
Amastra badia (Baldwin, 1895) EX? Re
´gnier et al., 2015
Amastra baldwiniana Hyatt and Pilsbry, 1911 EX Re
´gnier et al., 2015
Amastra biplicata (Newcomb, 1854) EX? Re
´gnier et al., 2015
Amastra borcherdingi Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra breviata Baldwin, 1895) EX? Re
´gnier et al., 2015
Amastra caputadamantis Hyatt and Pilsbry, 1911* EX Re
´gnier et al., 2015
Amastra conica Baldwin, 1906* EX Re
´gnier et al., 2015
Amastra conifera Smith, 1873 EX? Re
´gnier et al., 2015
Amastra cornea (Newcomb, 1854) EX 1994 EX EX Re
´gnier et al., 2015
Amastra crassilabrum (Newcomb, 1854) EX 1994 EX EX Re
´gnier et al., 2015
Amastra cyclostoma (Baldwin, 1895) EX Re
´gnier et al., 2015
Amastra davisiana Cooke, 1908 EX Re
´gnier et al., 2015
Amastra decorticata Gulick, 1873 EX? Re
´gnier et al., 2015
Amastra delicata Cooke, 1933 EX Re
´gnier et al., 2015
Amastra durandi Ancey, 1897 EX? Re
´gnier et al., 2015
Amastra dwightii Cooke, 1933 EX Re
´gnier et al., 2015
Amastra elegantula Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra elephantina Cooke, 1917* EX Re
´gnier et al., 2015
Amastra elliptica Gulick, 1873 EX? Re
´gnier et al., 2015
Amastra elongata (Newcomb, 1853) EX 1996 EX EX Re
´gnier et al., 2015
Amastra eos Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Amastra extincta (Pfeiffer, 1856)* EX Re
´gnier et al., 2015
Amastra farcimen (Pfeiffer, 1857) EX Re
´gnier et al., 2015
Amastra flavescens (Newcomb, 1854) EX Re
´gnier et al., 2015
Amastra flemingi Cooke, 1917* EX Re
´gnier et al., 2015
Amastra forbesi Cooke, 1917*EX 1996 EX EX Re
´gnier et al., 2015
Amastra fossilis Baldwin, 1903* EX Re
´gnier et al., 2015
Amastra fragilis Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Amastra fragosa Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra fraterna Sykes, 1896 EX? Re
´gnier et al., 2015
Amastra globosa Cooke, 1933* EX Re
´gnier et al., 2015
Amastra gouveii Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra grayana (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Amastra gulickiana Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra hawaiiensis Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra hitchcocki Cooke, 1917* EX Re
´gnier et al., 2015
Amastra humilis (Newcomb, 1855) EX? Re
´gnier et al., 2015
Amastra hutchinsonii (Pease, 1862) EX? Re
´gnier et al., 2015
Amastra implicata Cooke, 1933 EX Re
´gnier et al., 2015
Amastra inflata (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Amastra inopinata Cooke, 1933* EX Re
´gnier et al., 2015
Amastra irwiniana Cooke, 1908 EX Re
´gnier et al., 2015
Amastra johnsoni Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra juddii Cooke, 1917 EX Re
´gnier et al., 2015
Amastra kalamaulensis Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Amastra kauaiensis (Newcomb, 1860) EX Re
´gnier et al., 2015
Amastra kaunakakaiensis Pilsbry and Cooke, 1914 EX Re
´gnier et al., 2015
Amastra knudsenii (Baldwin, 1895) EX Re
´gnier et al., 2015
Amastra laeva Baldwin, 1906 EX Re
´gnier et al., 2015
Amastra lahainana Pilsbry and Cooke, 1914 EX Re
´gnier et al., 2015
Amastra lineolata (Newcomb, 1853) EX? Re
´gnier et al., 2015
Amastra luctuosa (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Amastra luteola (Fe
´rrusac, 1825) EX? Re
´gnier et al., 2015
Amastra magna (Adams, 1851) EX? Re
´gnier et al., 2015
(Continued)
Page 18 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Amastra makawaoensis Hyatt and Pilsbry, 1911 EX Re
´gnier et al., 2015
Amastra malleata Smith, 1873 EX? Re
´gnier et al., 2015
Amastra mastersi (Newcomb, 1854) EX? Re
´gnier et al., 2015
Amastra melanosis (Newcomb, 1854) EX Re
´gnier et al., 2015
Amastra metamorpha Pilsbry and Cooke, 1914 EX Re
´gnier et al., 2015
Amastra mirabilis Cooke, 1917 EX Re
´gnier et al., 2015
Amastra modesta (Adams, 1851) EX? Re
´gnier et al., 2015
Amastra modicella Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra moesta (Newcomb, 1854) EX? Re
´gnier et al., 2015
Amastra montagui Pilsbry, 1913 EX? Re
´gnier et al., 2015
Amastra montana Baldwin, 1906 EX Re
´gnier et al., 2015
Amastra montivaga Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra morticina Hyatt and Pilsbry, 1911* EX Re
´gnier et al., 2015
Amastra mucronata (Newcomb, 1853) EX? Re
´gnier et al., 2015
Amastra nana (Baldwin, 1895) EX? Re
´gnier et al., 2015
Amastra nannodes Cooke, 1933 EX Re
´gnier et al., 2015
Amastra neglecta Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Amastra nigra (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Amastra nubifera Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra nubigena Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Amastra nubilosa (Mighels, 1845) EX? Re
´gnier et al., 2015
Amastra nucleola (Gould, 1845) EX Re
´gnier et al., 2015
Amastra nucula Smith, 1873 EX? Re
´gnier et al., 2015
Amastra obesa (Newcomb, 1853) EX? Re
´gnier et al., 2015
Amastra oswaldi Cooke, 1933 EX? Re
´gnier et al., 2015
Amastra ovatula Cooke, 1933* EX Re
´gnier et al., 2015
Amastra pagodula Cooke, 1917* EX Re
´gnier et al., 2015
Amastra paulula Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra peasei Smith, 1873 EX? Re
´gnier et al., 2015
Amastra pellucida (Baldwin, 1895) EX 1994 EX EX Re
´gnier et al., 2015
Amastra perversa Hyatt and Pilsbry, 1911* EX Re
´gnier et al., 2015
Amastra petricola (Newcomb, 1855) EX? Re
´gnier et al., 2015
Amastra pilsbryi Cooke, 1913 EX? Re
´gnier et al., 2015
Amastra porcus Hyatt and Pilsbry, 1911 EX 1994 EX EX Re
´gnier et al., 2015
Amastra porphyrostoma (Pease, 1869) EX? Re
´gnier et al., 2015
Amastra praeopima Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra problematica Cooke, 1933 EX? Re
´gnier et al., 2015
Amastra pullata (Baldwin, 1895) EX? Re
´gnier et al., 2015
Amastra pusilla (Newcomb, 1854) EX? Re
´gnier et al., 2015
Amastra reticulata (Newcomb, 1854) EX 1994 EX EX Re
´gnier et al., 2015
Amastra ricei Cooke, 1917 EX Re
´gnier et al., 2015
Amastra rubida Gulick, 1873 EX? Re
´gnier et al., 2015
Amastra rubristoma Baldwin, 1906 EX? Re
´gnier et al., 2015
Amastra rugulosa Pease, 1870 EX Re
´gnier et al., 2015
Amastra seminigra Hyatt and Pilsbry, 1911 EX Re
´gnier et al., 2015
Amastra seminuda Baldwin, 1906 EX? Re
´gnier et al., 2015
Amastra senilis Baldwin, 1903* EX Re
´gnier et al., 2015
Amastra sericea (Pfeiffer, 1859 EX? Re
´gnier et al., 2015
Amastra similaris Pease, 1870 EX Re
´gnier et al., 2015
Amastra sinistrorsa Baldwin, 1906* EX Re
´gnier et al., 2015
Amastra sola Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra soror (Pfeiffer, 1868) EX Re
´gnier et al., 2015
Amastra spaldingi Cooke, 1908 EX Re
´gnier et al., 2015
Amastra sphaerica Pease, 1870 EX Re
´gnier et al., 2015
Amastra spicula Cooke, 1917 EX? Re
´gnier et al., 2015
Amastra subcornea Hyatt and Pilsbry, 1911* EX Re
´gnier et al., 2015
Amastra subcrassilabris Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra subobscura Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
(Continued)
R.H. Cowie et al., 2017 Page 19
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Amastra subrostrata (Pfeiffer, 1859 EX 1994 EX EX Re
´gnier et al., 2015
Amastra subsoror Hyatt and Pilsbry, 1911 EX 1994 EX EX Re
´gnier et al., 2015
Amastra sykesi Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra tenuilabris Gulick, 1873 EX? Re
´gnier et al., 2015
Amastra tenuispira (Baldwin, 1895) EX 1994 EX EX Re
´gnier et al., 2015
Amastra textilis (Fe
´rrusac, 1825) EX? Re
´gnier et al., 2015
Amastra thaanumi Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra thurstoni Cooke, 1917* EX Re
´gnier et al., 2015
Amastra transversalis (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Amastra tricincta Hyatt and Pilsbry, 1911 EX? Re
´gnier et al., 2015
Amastra tristis (Fe
´rrusac, 1825) EX? Re
´gnier et al., 2015
Amastra turritela (Fe
´rrusac, 1821) EX? Re
´gnier et al., 2015
Amastra ultima Pilsbry and Cooke, 1914 EX Re
´gnier et al., 2015
Amastra umbilicata (Pfeiffer, 1856) EX 1996 EX EX Re
´gnier et al., 2015
Amastra umbrosa (Baldwin, 1895) EX? Re
´gnier et al., 2015
Amastra undata (Baldwin, 1895) EX? Re
´gnier et al., 2015
Amastra uniplicata (Hartman, 1888) EX? Re
´gnier et al., 2015
Amastra variegata (Pfeiffer, 1849) EX? Re
´gnier et al., 2015
Amastra vetusta (Baldwin, 1895)* EX Re
´gnier et al., 2015
Amastra violacea (Newcomb, 1853) EX? Re
´gnier et al., 2015
Amastra viriosa Cooke, 1917 EX Re
´gnier et al., 2015
Amastra whitei Cooke, 1917 EX Re
´gnier et al., 2015
Armsia petasus (Ancey, 1899) EX? Re
´gnier et al., 2015
Carelia anceophila Cooke, 1931 EX 1990 EX EX Re
´gnier et al., 2015
Carelia bicolor (Jay, 1839) EX 1990 EX EX Re
´gnier et al., 2015
Carelia cochlea (Reeve, 1849)*EX 1990 EX EX Re
´gnier et al., 2015
Carelia cumingiana (Pfeiffer, 1855) EX 1990 EX EX Re
´gnier et al., 2015
Carelia dolei Ancey, 1893*EX 1990 EX EX Re
´gnier et al., 2015
Carelia evelynae Cooke and Kondo, 1952*EX 1990 EX EX Re
´gnier et al., 2015
Carelia glossema Cooke, 1931 EX 1990 EX EX Re
´gnier et al., 2015
Carelia hyattiana Pilsbry, 1911 EX 1990 EX EX Re
´gnier et al., 2015
Carelia kalalauensis Cooke, 1931 EX 1990 EX EX Re
´gnier et al., 2015
Carelia knudseni Cooke, 1931 EX 1990 EX EX Re
´gnier et al., 2015
Carelia lirata Cooke, 1931*EX 1990 EX EX Re
´gnier et al., 2015
Carelia lymani Cooke, 1931*EX 1990 EX EX Re
´gnier et al., 2015
Carelia mirabilis Cooke, 1931*EX 1990 EX EX Re
´gnier et al., 2015
Carelia necra Cooke, 1931*EX 1990 EX EX Re
´gnier et al., 2015
Carelia olivacea Pease, 1866 EX 1990 EX EX Re
´gnier et al., 2015
Carelia paradoxa (Pfeiffer, 1854) EX 1990 EX EX Re
´gnier et al., 2015
Carelia periscelis Cooke, 1931 EX 1990 EX EX Re
´gnier et al., 2015
Carelia pilsbryi Sykes, 1909 EX 1990 EX EX Re
´gnier et al., 2015
Carelia sinclairi Ancey, 1892*EX 1990 EX EX Re
´gnier et al., 2015
Carelia tenebrosa Cooke, 1931 EX 1990 EX EX Re
´gnier et al., 2015
Carelia turricula (Mighels, 1845) EX 1990 EX EX Re
´gnier et al., 2015
Laminella alexandri (Newcomb, 1865) EX Re
´gnier et al., 2015
Laminella bulbosa (Gulick, 1858) EX? Re
´gnier et al., 2015
Laminella citrina (Pfeiffer, 1848) EX? Re
´gnier et al., 2015
Laminella concinna (Newcomb, 1854) EX? Re
´gnier et al., 2015
Laminella depicta (Baldwin, 1895) EX? Re
´gnier et al., 2015
Laminella gravida (Fe
´russac, 1825) EX? Re
´gnier et al., 2015
Laminella kuhnsi (Cooke, 1908) EX? Re
´gnier et al., 2015
Laminella picta (Mighels, 1845) EX? Re
´gnier et al., 2015
Laminella remyi (Newcomb, 1855) EX? Re
´gnier et al., 2015
Laminella straminea (Reeve, 1850) EX? Re
´gnier et al., 2015
Laminella tetrao (Newcomb, 1855) EX? Re
´gnier et al., 2015
Laminella venusta (Mighels, 1845) EX? Re
´gnier et al., 2015
Leptachatina accineta (Mighels, 1845) EX? Re
´gnier et al., 2015
Leptachatina acuminata (Gould, 1847) EX Re
´gnier et al., 2015
(Continued)
Page 20 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Leptachatina anceyana Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina antiqua Pease, 1870 EX Re
´gnier et al., 2015
Leptachatina approximans Ancey, 1897 EX? Re
´gnier et al., 2015
Leptachatina arborea Sykes, 1900 EX? Re
´gnier et al., 2015
Leptachatina attenuata Cooke, 1911 EX Re
´gnier et al., 2015
Leptachatina baldwini Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina balteata Pease, 1870 EX Re
´gnier et al., 2015
Leptachatina brevicula (Pease, 1869) EX Re
´gnier et al., 2015
Leptachatina callosa (Pfeiffer, 1857) EX? Re
´gnier et al., 2015
Leptachatina captiosa Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina cingula (Gould, 1847) EX? Re
´gnier et al., 2015
Leptachatina compacta (Pease, 1869) EX Re
´gnier et al., 2015
Leptachatina concolor Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina conicoides Sykes, 1900 EX? Re
´gnier et al., 2015
Leptachatina conspicienda Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina convexiuscula Sykes, 1900 EX? Re
´gnier et al., 2015
Leptachatina cookei Pilsbry, 1914 EX Re
´gnier et al., 2015
Leptachatina corneola (Pfeiffer, 1846) EX? Re
´gnier et al., 2015
Leptachatina coruscans (Hartman, 1888) EX? Re
´gnier et al., 2015
Leptachatina costulata (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina costulosa Pease, 1870 EX? Re
´gnier et al., 2015
Leptachatina deceptor Cockerell, 1927* EX Re
´gnier et al., 2015
Leptachatina defuncta Cooke, 1910* EX Re
´gnier et al., 2015
Leptachatina dimidiata (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Leptachatina dormitor Pilsbry and Cooke, 1914* EX Re
´gnier et al., 2015
Leptachatina dulcis Cooke, 1911 EX Re
´gnier et al., 2015
Leptachatina emerita Sykes, 1900 EX? Re
´gnier et al., 2015
Leptachatina exilis (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina exoptabilis Cooke, 1910* EX Re
´gnier et al., 2015
Leptachatina extensa Pease, 1870 EX? Re
´gnier et al., 2015
Leptachatina fossilis Cooke, 1910* EX Re
´gnier et al., 2015
Leptachatina fraterna Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina fulgida Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina fumida (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina fusca (Newcomb, 1853) EX? Re
´gnier et al., 2015
Leptachatina fuscula (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina gayi Cooke, 1911 EX Re
´gnier et al., 2015
Leptachatina glutinosa (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Leptachatina grana (Newcomb, 1853) EX? Re
´gnier et al., 2015
Leptachatina guttula (Gould, 1847) EX? Re
´gnier et al., 2015
Leptachatina haenensis Cockerell, 1927* EX Re
´gnier et al., 2015
Leptachatina henshawi Sykes, 1903 EX Re
´gnier et al., 2015
Leptachatina hyperodon Pilsbry and Cooke, 1914* EX Re
´gnier et al., 2015
Leptachatina illimis Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina imitatrix Sykes, 1900* EX Re
´gnier et al., 2015
Leptachatina impressa Sykes, 1896 EX? Re
´gnier et al., 2015
Leptachatina irregularis (Pfeiffer, 1856) EX Re
´gnier et al., 2015
Leptachatina isthmica Ancey and Sykes, 1899* EX Re
´gnier et al., 2015
Leptachatina knudseni Cooke, 1910 EX Re
´gnier et al., 2015
Leptachatina konaensis Sykes, 1900 EX Re
´gnier et al., 2015
Leptachatina kuhnsi Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina labiata (Newcomb, 1853) EX? Re
´gnier et al., 2015
Leptachatina laevigata Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina laevis Pease, 1870 EX Re
´gnier et al., 2015
Leptachatina lagena (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina lanaiensis Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina lanceolata Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina leiahiensis Cooke, 1910* EX Re
´gnier et al., 2015
(Continued)
R.H. Cowie et al., 2017 Page 21
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Leptachatina lenta Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina leucochila (Gulick, 1856) EX Re
´gnier et al., 2015
Leptachatina longiuscula Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina lucida Pease, 1870 EX Re
´gnier et al., 2015
Leptachatina maniensis (Pfeiffer, 1855) EX? Re
´gnier et al., 2015
Leptachatina marginata (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina mcgregori Pilsbry and Cooke, 1914 EX Re
´gnier et al., 2015
Leptachatina microdon Pilsbry and Cooke, 1914 EX? Re
´gnier et al., 2015
Leptachatina molokaiensis Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina morbida Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina nematoglypta Pilsbry and Cooke, 1914) EX? Re
´gnier et al., 2015
Leptachatina obsoleta (Pfeiffer, 1857) EX Re
´gnier et al., 2015
Leptachatina obtusa (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Leptachatina octogyrata (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina omphalodes (Ancey, 1899) EX? Re
´gnier et al., 2015
Leptachatina opipara Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina optabilis Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina oryza (Pfeiffer, 1856)* EX Re
´gnier et al., 2015
Leptachatina ovata Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina pachystoma (Pease, 1869) EX Re
´gnier et al., 2015
Leptachatina perforata Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina perkinsi Sykes, 1896 EX? Re
´gnier et al., 2015
Leptachatina petila (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina pilsbryi Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina praestabilis Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina pulchra Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina pumicata (Mighels, 1845) EX? Re
´gnier et al., 2015
Leptachatina pupoidea Cooke, 1911 EX Re
´gnier et al., 2015
Leptachatina pyramis (Pfeiffer, 1846) EX? Re
´gnier et al., 2015
Leptachatina resinula (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina saccula (Hartman, 1888) EX? Re
´gnier et al., 2015
Leptachatina sagittata Pilsbry and Cooke, 1914 EX Re
´gnier et al., 2015
Leptachatina sandwicensis (Pfeiffer, 1846) EX? Re
´gnier et al., 2015
Leptachatina saxatilis (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina sculpta (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Leptachatina scutilus (Mighels, 1845) EX? Re
´gnier et al., 2015
Leptachatina semipicta Sykes, 1896 EX? Re
´gnier et al., 2015
Leptachatina simplex (Pease, 1869) EX Re
´gnier et al., 2015
Leptachatina smithi Sykes, 1896 EX? Re
´gnier et al., 2015
Leptachatina somniator Pilsbry and Cooke, 1914* EX Re
´gnier et al., 2015
Leptachatina stiria (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina striata (Newcomb, 1861) EX? Re
´gnier et al., 2015
Leptachatina striatula (Gould, 1845) EX Re
´gnier et al., 2015
Leptachatina subcylindracea Cooke, 1910* EX Re
´gnier et al., 2015
Leptachatina subovata Cooke, 1910 EX? Re
´gnier et al., 2015
Leptachatina subula (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina succincta (Newcomb, 1855) EX? Re
´gnier et al., 2015
Leptachatina supracostata Sykes, 1900 EX? Re
´gnier et al., 2015
Leptachatina tenebrosa Pease, 1870 EX Re
´gnier et al., 2015
Leptachatina tenuicostata (Pease, 1869)* EX Re
´gnier et al., 2015
Leptachatina terebralis (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina teres (Pfeiffer, 1856) EX? Re
´gnier et al., 2015
Leptachatina thaanumi Cooke, 1911 EX? Re
´gnier et al., 2015
Leptachatina triticea (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina turrita (Gulick, 1856) EX? Re
´gnier et al., 2015
Leptachatina vana Sykes, 1900 EX? Re
´gnier et al., 2015
Leptachatina varia Cooke, 1910 EX Re
´gnier et al., 2015
Leptachatina ventulus (Fe
´russac, 1825) EX? Re
´gnier et al., 2015
(Continued)
Page 22 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Pauahia artata (Cooke, 1911 EX? Re
´gnier et al., 2015
Pauahia chrysallis (Pfeiffer, 1855) EX? Re
´gnier et al., 2015
Pauahia tantilla (Cooke, 1911) EX? Re
´gnier et al., 2015
Planamastra digonomorpha (Ancey, 1889) EX? Re
´gnier et al., 2015
Planamastra peaseana Pilsbry, 1911 EX? Re
´gnier et al., 2015
Planamastra spaldingi Cooke, 1933 EX? Re
´gnier et al., 2015
Tropidoptera alata (Pfeiffer, 1856) EX Re
´gnier et al., 2015
Tropidoptera discus Pilsbry and Vanatta, 1905 EX Re
´gnier et al., 2015
Tropidoptera heliciformis (Ancey, 1890) EX? Re
´gnier et al., 2015
Tropidoptera rex (Sykes, 1904) EX? Re
´gnier et al., 2015
Tropidoptera wesleyi (Sykes, 1896) EX? Re
´gnier et al., 2015
AMPHIBULIMULIDAE
Eudolichotis euryomphala (Jonas, 1844) EX? EX?
Eudolichotis sinuata (Albers, 1854) EX? EX?
Plekocheilus pulicarius (Reeve, 1848) EX? EX?
Plekocheilus succinoides (Petit, 1840) EX? EX?
ANNULARIIDAE
Parachondria basicarinata (Pfeiffer, 1855) EX? Watters, 2014
ARIOPHANTIDAE
Ariophanta thyreus (Benson, 1852) EX? EX?
Hemiplecta neptunus Pfeiffer, 1854 EX? EX?
Vitrinula chaunax (Pilsbry and Hirase, 1904) EX 1994 EX
Vitrinula chichijimana (Pilsbry and Hirase, 1905) EX 1994 EX EX
Vitrinula hahajimana (Pilsbry and Hirase, 1905) EX 1994 EX EX
ASSIMINEIDAE
Conacmella vagans Hirase, 1907 EX EX
Cyclomorpha secessa Bouchet and Abdou, 2003 EX EX
Electrina succinea (Sowerby, 1846) EX EX
Garrettia rotella (Pease, 1868) EX? Brook, 2010; F. Brook, pers.
comm, 2016
Kubaryia pilikia Clench 1948 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Omphalotropis bassinblancensis Griffiths and
Florens, 2004
–EXEX
Omphalotropis dupontiana Nevill, 1878 EX EX
Omphalotropis ingens (Mousson, 1870) CR(PE) 2012 EX?
Omphalotropis margarita (Pfeiffer 1851) EX EX
Omphalotropis maxima Madge, 1939 EX EX
Omphalotropis multilirata (Pfeiffer, 1852) EX EX
Omphalotropis plicosa (Pfeiffer, 1852) EX 1994 EX
Omphalotropis quittorensis Griffiths and
Florens, 2004
–EXEX
Omphalotropis rotumana Smith, 1897 EX EX
Omphalotropis vacoasensis Griffiths and
Florens, 2004
–EXEX
BOTHRIEMBRYONTIDAE
Leucocharis loyaltiensis (Souverbie, 1879) EX 1994 EX EX
Leucocharis porphyrocheila (Dautzenberg and
Bernier, 1901)
EX 1994 EX EX
Placostylus cuniculinsulae Cox, 1872 EX 1996 EX EX
Placostylus koroensis (Garrett, 1872) CR(PE) 2012 EX? EX?
BRADYBAENIDAE
Calocochlia cailliaudi Deshayes, 1839 EX? EX?
Calocochlia chlorochroa Sowerby, 1841 EX? EX?
Calocochlia cumingii Pfeiffer, 1842 EX? EX?
Chloraea fragilis (Sowerby, 1841) EX? EX?
Euhadra murayamai Habe, 1976 DD EX Ministry of the Environment
Government of Japan, 2016
(Continued)
R.H. Cowie et al., 2017 Page 23
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Euhadra nachicola Kuroda, 1929 DD EX Ministry of the Environment
Government of Japan, 2016
Euhadra sadoensis (Pilsbry and Hirase, 1903) DD EX Ministry of the Environment
Government of Japan, 2016
Helicostyla carbonaria (Sowerby, 1842) EX? EX?
Helicostyla collodes (Sowerby 1841) EX EX
Helicostyla cunctator (Reeve, 1849) EX EX
Helicostyla daphnis (Broderip, 1841) EX EX
Helicostyla moreleti (Pfeiffer, 1890) EX EX
Helicostyla pfeifferi Semper, 1877 EX EX
Helicostyla phloiodes (Pfeiffer, 1842) EX EX
Helicostyla pilsbryi (Hidalgo 1890) EX EX
Helicostyla propitia (Fulton, H.C. 1907) EX? EX?
Helicostyla velata (Broderip, 1841) EX? EX?
Helicostyla zebuensis (Broderip, 1841) EX? EX?
Mandarina luhuana (Sowerby, 1839) EX EX
BULIMULIDAE
Bulimulus achatellinus (Forbes, 1850) CR(PE) 2003 EX?
Bulimulus adelphus (Dall, 1917) CR(PE) 2003 EX?
Bulimulus brunoi von Ihering, 1917 EX EX
Bulimulus deridderi (Coppois, 1985) CR(PE) 2003 EX?
Bulimulus duncanus (Dall, 1893) CR(PE) 2003 EX?
Bulimulus eos (Odhner, 1951) CR(PE) 2003 EX?
Bulimulus lycodus (Dall, 1917) CR(PE) 2003 EX?
Bulimulus saeronius (Dall, 1917) CR(PE) 2003 EX?
Bulimulus tanneri (Dall, 1895) CR(PE) 2003 EX?
Naesiotus arnaldoi (Lanzieri and Rezende, 1971) EX? Salvador et al., 2013
CAMAENIDAE
Aegista inexpecta Kuroda and Minato, 1977 EX EX
Amphidromus dohrni (Pfeiffer, 1863) EX? EX?
Amphidromus metabletus Moellendorff, 1900 EX? EX?
Amphidromus sinensis (Benson, 1851) EX? EX?
Satsuma fausta (Pilsbry, 1902) EX Association of Wildlife Research and
EnVision Conservation Office, 2015
CERASTIDAE
Pachnodus curiosus Gerlach, 2003 EX 2009 EX EX
Pachnodus ladiguensis Gerlach, 2003 EX 2009 EX EX
Pachnodus velutinus (Pfeiffer, 1868) EX 2009 EX EX
Rachis comorensis (Morelet, 1881) EX 1994 EX? EX?
Rachis sanguineus (Barclay, 1857) EX 1994 EX EX
Rachistia vesiculatus (Benson, 1859) EX EX
CHAROPIDAE
Charopa perryi Smith, 1897 EX EX
Charopa rotumana Smith 1897 EX EX
Damonita geminoropiformis Climo, 1981 EX Spencer et al., 2009
Discocharopa aperta (Moellendorff, 1888) EX EX
Helenoconcha leptalea (Smith, 1892) EX 1994 EX EX
Helenoconcha minutissima (Smith, 1892) EX 1994 EX EX
Helenoconcha polyodon (Sowerby, 1844) EX 1994 EX EX
Helenoconcha pseustes (Smith, 1892) EX 1994 EX EX
Helenoconcha sexdentata (Smith, 1893) EX 1994 EX EX
Helenodiscus bilamellata (Sowerby, 1844) EX 1994 EX EX
Helenodiscus vernoni (Smith 1892) EX 1994 EX EX
Hirasea biconcava Hirase, 1907 EX EX
Hirasea diplomphalus Pilsbry, 1902 EX EX
Hirasea eutheca Hirase, 1907 EX EX
Hirasea goniobasis Pilsbry, 1902 EX EX
Hirasea hypolia Hirase, 1907 EX EX
(Continued)
Page 24 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Hirasea insignis Pilsbry and Hirase, 1904 EN EX Ministry of the Environment
Government of Japan, 2016
Hirasea major Pilsbry, 1902 EX EX
Hirasea operculina (Gould, 1859) EN EX Ministry of the Environment
Government of Japan, 2016
Hirasea sinuosa Pilsbry, 1902 EX EX
Lauopa mbalavuana Solem, 1983 CR(PE) 2012 EX?
Libera subcavernula (Tryon, 1887) EX 1994 EX EX
Libera tumuloides (Garrett, 1872) EX 1994 EX EX
Mautodontha acuticosta (Garrett, 1884) EX 1994 EX EX
Mautodontha consimilis (Pease, 1868) EX 1994 EX EX
Mautodontha consobrina (Garrett, 1884) EX 1994 EX EX
Mautodontha maupiensis (Garrett, 1872) EX 1994 EX EX
Mautodontha parvidens (Pease, 1861) EX 1994 EX EX
Mautodontha punctiperforata (Garrett, 1884) EX 1994 EX EX
Mautodontha saintjohni Solem, 1976 EX 1994 EX EX
Mautodontha subtilis (Garrett, 1884) EX 1994 EX EX
Mautodontha unilamellata (Garrett, 1874) EX 1994 EX EX
Mautodontha zebrina (Garrett, 1874) EX 1994 EX EX
Mocella elliottae (Climo, 1969) EX Spencer et al., 2009
Mocella spelaeus (Climo, 1971) EX Spencer et al., 2009
Propilula cyclaria (Morelet, 1875) EX EX
Sinployea canalis (Garrett, 1872) EX 1994 EX EX
Sinployea decorticata (Garrett, 1872) EX 1994 EX EX
Sinployea harveyensis (Garrett, 1872) EX 1994 EX EX
Sinployea muri Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm, 2016
Sinployea otareae (Garrett, 1872) EX 1994 EX EX
Sinployea planospira (Garrett, 1881) EX 1994 EX EX
Sinployea proxima (Garrett, 1872) EX 1994 EX EX
Sinployea rudis (Garrett, 1872) EX 1994 EX EX
Sinployea tenuicostata (Garrett, 1872) EX 1994 EX EX
Sinployea titikaveka Brook, 2010 EX? EX Brook, 2010; F. Brook, pers.
comm, 2016
Sinployea tupapa Brook, 2010 EX? EX Brook, 2010; F. Brook, pers.
comm, 2016
Sinployea youngi (Garrett, 1872) EX 1994 EX EX
Taipidon anceyana (Garrett, 1887) EX 1994 EX EX
Taipidon marquesana (Garrett, 1887) EX 1994 EX EX
Taipidon octolamellata (Garrett, 1887) EX 1994 EX EX
Trachycystis rariplicata (Benson, 1887) EX EX
Zelandiscus elevatus (Climo, 1978) EX Climo, 1981
CHRONIDAE
Trochochlamys ogasawarana (Pilsbry, 1902) EX EX
CLAUSILIDAE
Neophaedusa spelaeonis Kuroda and Minato, 1975 DD DD? EX Ministry of the Environment
Government of Japan, 2016
COCHLICELLIDAE
Monilearia pulverulenta (Lowe, 1861) CR(PE) 2011 EX?
CYCLOPHORIDAE
Cyclophorus acutimarginatus (Sowerby, 1842) EX? EX?
Cyclophorus stenomphalus (Pfeiffer, 1846) EX? EX?
Nobuea kurodai Minato and Tada, 1978 DD 1996 EX Ministry of the Environment
Government of Japan, 2016
DIPLOMMATINIDAE
Diplommatina alata (Crosse 1866) CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Diplommatina aurea Beddome 1889 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
(Continued)
R.H. Cowie et al., 2017 Page 25
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Diplommatina gibboni Beddome 1889 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Opisthostoma decrespignyi (H. Adams, 1865) CR(PE) 2004 EX? EX?
Opisthostoma otostoma Boettger, 1893 CR(PE) 2004 EX? EX?
Palaina albata (Beddome 1889) CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Palaina patula (Crosse 1866) CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Palaina platycheilus (Beddome 1889) CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Palaina pupa Crosse 1866 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Plectostoma charasense (Tomlin, 1948) CR(PE) 2014 EX?
Plectostoma dindingense Liew et al., 2014 CR(PE) 2014 EX?
Plectostoma sciaphilum (van Benthem
Jutting, 1952)
EX 2014 EX Schilthuizen and Clements, 2008;
Liew et al., 2014
Plectostoma turriforme (van Benthem Jutting, 1952) CR(PE) 2014 EX?
DISCIDAE
Keraea garachicoensis (Wollaston 1878) EX EX
DYAKIIDAE
Dyakia clippeus (Mousson, 1849) EX? EX?
ENDODONTIDAE
Aaadonta angaurana Solem 1976 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Aaadonta kinlochi Solem 1976 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Aaadonta pelewana Solem 1976 CR(PE) 2012 EX? R.J. Rundell, pers. comm., 2016
Anceyodonta alternata Cooke and Solem, 1976 EX EX
Anceyodonta andersoni Cooke and Solem, 1976 EX EX
Anceyodonta constricta Cooke and Solem, 1976 EX EX
Anceyodonta densicostata Cooke and Solem, 1976 EX EX
Anceyodonta difficilis Solem, 1976 EX EX
Anceyodonta ganhutuensis Cooke and Solem, 1976 EX EX
Anceyodonta gatavakensis Abdou and Bouchet, 2000 EX EX
Anceyodonta hamyana (Ancey, 1889) EX EX
Anceyodonta labiosa Solem, 1976 EX EX
Anceyodonta obesa Solem, 1976 EX EX
Anceyodonta sexlamellata (Pfeiffer, 1845) EX EX
Anceyodonta soror Solem, 1976 EX EX
Anceyodonta subconica Solem and Cooke, 1976 EX EX
Anceyodonta umbilicata Abdou and Bouchet, 2000 EX EX
Australdonta anneae Sartori et al., 2013 EX Sartori et al., 2013
Australdonta collicella Zimmermann et al., 2009 EX
Australdonta ectopia Solem, 1976 EX EX
Australdonta florencei Sartori et al., 2013 EX Sartori et al., 2013
Australdonta magnasulcatissima
Zimmermann et al., 2009
EX Zimmerman et al., 2009
Australdonta microspiralis Zimmermann et al., 2009 EX Zimmermann et al., 2009
Australdonta oheatora
Sartori, Gargominy and Fontaine, 2013 EX Sartori et al., 2013
Australdonta pakalolo
Sartori, Gargominy and Fontaine, 2013 EX Sartori et al., 2013
Australdonta pharcata Solem, 1976 EX EX
Australdonta pseudplanulata Solem, 1976 EX EX
Australdonta rimatarana Solem, 1976 EX EX
Australdonta sibleti Sartori et al., 2013 EX Sartori et al., 2013
Australdonta sulcata Zimmermann et al., 2009 EX Zimmermann et al., 2009
Australdonta teaae Sartori et al., 2013 EX Sartori et al., 2013
Australdonta tubuaiana Solem, 1976 EX EX
Endodonta apiculata (Ancey, 1889) CR(PE) 2000 EX?
Gambiodonta agakauitaiana Solem
and Cooke, 1976
–EXEX
Gambiodonta grandis Cooke and Solem, 1976 EX EX
Gambiodonta mangarevana Solem and Cooke, 1976 EX EX
Gambiodonta mirabilis Cooke and Solem, 1976 EX EX
(Continued)
Page 26 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Gambiodonta pilsbryi Cooke and Solem, 1976 EX EX
Gambiodonta tumida Cooke and Solem, 1976 EX EX
Hirasea planulata Pilsbry and Hirase, 1903 EX 1994 EX EX
Kleokyphus callimus Solem, 1976 EX Sartori et al., 2014
Kleokyphus cowiei Sartori et al., 2014 EX Sartori et al., 2014
Kleokyphus hypsus Solem, 1976 EX Sartori et al., 2014
Mautodontha aurora Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha ceuthma Solem, 1976 EX EX
Mautodontha domaneschii Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha harperae Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha makateaensis Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha occidentalis Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha passosi Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha rarotongensis (Pease, 1870) EX? Brook et al., 2010; F. Brook,
pers. comm., 2016
Mautodontha spelunca Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha temaoensis Sartori et al., 2014 EX Sartori et al., 2014
Mautodontha virginiae Sartori et al., 2014 EX Sartori et al., 2014
Minidonta anatonuana Solem, 1976 EX EX
Minidonta aroa Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta arorangi Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta bieleri Sartori et al., 2013 EX Sartori et al., 2013
Minidonta boucheti Sartori et al., 2013 EX Sartori et al., 2013
Minidonta extraria Cooke and Solem, 1976 EX EX
Minidonta flammulina Abdou and Bouchet, 2000 EX EX
Minidonta gravacosta Solem, 1976 EX EX
Minidonta haplaenopla Solem, 1976 EX EX
Minidonta iota Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta kavera Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta macromphalus Preece, 1998 EX EX
Minidonta manuaensis Solem, 1976 EX EX
Minidonta micra Solem and Cooke, 1976 EX EX
Minidonta micraconica Solem, 1976 EX EX
Minidonta ngatangiia Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta perminima Abdou and Bouchet, 2000 EX EX
Minidonta planulata Solem, 1976 EX EX
Minidonta pue Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta rutaki Brook, 2010 EX Brook, 2010; F. Brook, pers.
comm., 2016
Minidonta simulata Solem and Cooke, 1976 EX EX
Minidonta sulcata Solem, 1976 EX EX
Minidonta taravensis Solem and Cooke, 1976 EX EX
Minidonta taunensis Solem and Cooke, 1976 EX EX
Minidonta vallonia Abdou and Bouchet, 2000 EX EX
Pseudohelenoconcha spurca (Sowerby, 1844) EX 1994 EX EX
Pseudolibera aubertdelaruei Sartori et al., 2014 EX Sartori et al., 2014
Pseudolibera cookei Sartori et al., 2014 EX Sartori et al., 2014
Pseudolibera elieporoii Sartori et al., 2014 EX Sartori et al., 2014
Pseudolibera extincta Sartori et al., 2014 EX Sartori et al., 2014
Pseudolibera lillianae Solem, 1976 EX Sartori et al., 2014
Pseudolibera matthieui Sartori et al., 2014 EX Sartori et al., 2014
Pseudolibera paraminderae Sartori et al., 2014 EX Sartori et al., 2014
(Continued)
R.H. Cowie et al., 2017 Page 27
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Pseudolibera parva Sartori et al., 2014 EX Sartori et al., 2014
Pseudolibera solemi Sartori et al., 2014 EX Sartori et al., 2014
Rikitea insolens Cooke and Solem, 1976 EX EX
Rikitea tapinoptyx Abdou and Bouchet, 2000 EX EX
Thaumatodon multilamellata (Garrett, 1872) EX 1994 EX EX
ENIDAE
Mirus hachijoensis (Kuroda, 1945) DD EX Ministry of the Environment
Government of Japan, 2016
EUCONULIDAE
Advena campbelli (Gray, 1834) EX 1996 EX EX
Aukena endodonta Bouchet and Abdou, 2001 EX EX
Aukena tridentata (Baker, 1940) EX EX
Coneuplecta turrita (Semper, 1873) CE(PE) 2012 EX?
Cookeana anathesis Baker, 1938 EX EX
Cookeana vindex Baker, 1938 EX EX
Diastole matafaoi Baker, 1938 EX 1996 EX
Diastole rurutui Baker, 1938 EX EX
Fanulena perrugosa Iredale, 1945 EX 1996 EX EX
Lamprocystis rurutuana Baker, 1938 EX EX
Microcystis adusta Baker, 1938 EX EX
Microcystis andersoni Baker, 1938 EX EX
Microcystis kondoi Baker, 1938 EX EX
Nancibella quintalia (Cox, 1870) EX 1996 EX EX
Philonesia pyramidalis Preece, 1998 EX EX
Philonesia weisleri Preece, 1998 EX EX
Quintalia flosculus Cox, 1866 EX 1996 EX EX
Quintalia stoddartii Gray, 1834 EX 1996 EX EX
GASTROCOPTIDAE
Campolaemus perexilis (Smith, 1892) EX 1994 EX EX
Gastrocopta chichijimana Pilsbry, 1916 EX 1994 EX
Gastrocopta ogasawarana Pilsbry, 1916 EX 1994 EX
GASTRODONTIDAE
Atlantica engonata (Shuttleworth, 1852) EX EX
Atlantica retexta (Shuttleworth, 1852) EX EX
Atlantica textilis (Shuttleworth, 1852) EX EX
Janulus pompylius (Shuttleworth, 1852) EX EX
Poecilozonites reinianus (Pfeiffer, 1863) EX EX
HELICARIONIDAE
Caldwellia philyrina Morelet, 1873 EX 1996 EX
Ctenoglypta newtoni (Nevill, 1871) EX 1994 EX EX
Ctenophila aigretteianum Griffiths, 2000 EX EX
Dancea bewsheriana (Morelet, 1875) EX EX
Dupontia affouchensis Griffiths, 2000 EX EX
Dupontia proletaria (Morelet, 1860) EX 1996 EX
Epiglypta howinsulae (Cox, 1873) EX EX
Erepta chloritiformis Griffiths and Vincent, 2004 EX EX
Erepta nevilli (H. Adams, 1867) EX 1994 EX EX
Erepta pyramidalis Griffiths and Florens, 2004 EX EX
Erepta thiriouxi (Germain, 1918) EX EX
Erepta wendystrahmi Griffiths, 2000 EX EX
Harmogenanina linophora (Morelet, 1860) EX 1994 EX
Harmogenanina subdetecta Germain, 1921 EX 1994 EX
Hirasiella clara Pilsbry, 1902 EX EX
Pachystyla rufozonata (H. Adams), 1867 EX 1994 EX
Pachystyla waynepagei Griffiths, 2000 EX EX
Plegma duponti (Morelet, 1866) EX EX
Pseudophasis nevilli (H. Adams, 1867) EX EX
HELICIDAE
(Continued)
Page 28 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Hemicycla modesta (Fe
´russac, 1821) CR(PE) 2011 EX?
HELICINIDAE
Alcadia guadeloupensis (Sowerby, 1842) EX EX
Nesiocina abdoui Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Nesiocina gambierensis Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Nesiocina grohi Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Nesiocina mangarevae Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Nesiocina pauciplicata Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Nesiocina pazi (Crosse 1865) EX Richling and Bouchet, 2013
Nesiocina superoperculata Richling
and Bouchet, 2013
EX Richling and Bouchet, 2013
Nesiocina trilamellata Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Nesiocina unilamellata Richling and Bouchet, 2013 EX Richling and Bouchet, 2013
Ogasawarana arata Pilsbry, 1902 DD EX Ministry of the Environment
Government of Japan, 2016
Ogasawarana capsula Pilsbry, 1902 DD EX Ministry of the Environment
Government of Japan, 2016
Ogasawarana chichijimana Minato, 1980 EX EX
Ogasawarana discrepans Pilsbry, 1902 DD EX Ministry of the Environment
Government of Japan, 2016
Ogasawarana habei Minato, 1980 EX EX
Ogasawarana metamorpha Minato, 1980 EX EX
Ogasawarana nitida Minato, 1980 DD EX Ministry of the Environment
Government of Japan, 2016
Ogasawarana rex Minato, 1980 EX EX
Orobophana carinacosta Preece, 1998 EX EX
Pleuropoma hendersoni Preece, 1998 EX EX
Pseudotrochatella undulata (Morelet, 1878) EX EX
HOLOSPIRIDAE
Holospira piloceri (Pfeiffer, 1841) EX EX
Coelostemma richardi Thompson, 1971 EX EX
HYGROMIIDAE
Discula lyelliana (Lowe, 1852) CR(PE) 2011 EX EX?
Discula tetrica (Lowe, 1852) CR(PE) 2011 EX?
Geomitra delphinuloides (Lowe, 1860) CR(PE) 2011 EX?
Geomitra grabhami (Wollaston, 1878) CR(PE) 2011 EX EX?
Helicopsis paulhessei (Lindholm, 1936) EX 2011 EX
Montserratina becasis (Rambur, 1868) CR(PE) 2011 EX?
Pseudocampylaea loweii (Fe
´russac, 1835) EX 1996 EX EX
Trochoidea picardi (Haas, 1955) EX 1996 EX EX
LAURIIDAE
Leiostyla abbreviata Lowe, 1852 CR(PE) 2011 EX?
Leiostyla cassida (Lowe, 1831) CR(PE) 2011 EX?
Leiostyla gibba Lowe, 1852 CR(PE) 2011 EX?
Leiostyla lamellosa Lowe, 1852 EX 2011 EX EX
Leiostyla simulator (Pilsbry, 1923) CR(PE) 2011 EX?
MEGALOMASTOMATIDAE
Madgeaconcha sevathiani Griffiths
and Florens, 2004
CR(PE) 2014 EX EX?
NEOCYCLOTIDAE
Amphicyclotulus guadeloupensis
de la Torre, et al., 1942
EX 1996 EX EX
Incerticyclus cinereus (Droue
¨t, 1859) EX 1996 EX EX
ODONTOSTOMIDAE
Tomigerus gibberulus (Burrow, 1815) EX 1996 EX EX
Tomigerus turbinatus (Pfeiffer, 1845) EX 1996 EX EX
OLEACINIDAE
Oleacina guadeloupensis (Pfeiffer, 1857) EX 1996 EX EX
(Continued)
R.H. Cowie et al., 2017 Page 29
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Oleacina paivana (Pfeiffer, 1866) EX? EX?
OREOHELICIDAE
Oreohelix florida Pilsbry, 1939 EX EX
PARMACELLIDAE
Cryptella tamaranensis Hutterer, 1990 EX? EX?
Parmacella gervaisii Moquin-Tandon, 1850 EX EX
PARTULIDAE
Eua globosa Pilsbry and Cooke, 1934 CR(PE) 2012 EX?
Palaopartula leucothoe (Semper, 1865) CR(PE) 2012 EX?
Partula arguta (Pease, 1866) EX 1996 EX EX
Partula atilis Crampton, 1956 EX 1994 EX EX
Partula aurantia Crampton, 1932 EX 1988 EX EX
Partula auriculata Broderip, 1832 EX 1994 EX EX
Partula bilineata Pease, 1866 EX 1996 EX EX
Partula clarkei Gerlach, 2016 EX Gerlach, 2016
Partula cootei Gerlach, 2016 EX Gerlach, 2016
Partula crassilabris Pease, 1866 EX 1994 EX EX
Partula cuneata Crampton, 1956 EX 1994 EX EX
Partula cytherea Cooke and Crampton, 1930 EX 1996 EX EX? Gerlach, 2016
Partula dentifera Pfeiffer, 1853 EW 1996 EW EX Gerlach, 2016
Partula desolata Bauman and Kerr, 2013* EX Bauman and Kerr, 2013;
Gerlach, 2016
Partula diminuta Adams, 1851 EX Gerlach, 2016
Partula dolichostoma Crampton, 1956 EX 1994 EX EX
Partula dolorosa Crampton and Cooke, 1953 EX 1996 EX EX
Partula eremita Crampton and Cook, 1953 EX 1996 EX EX
Partula faba (Gmelin, 1791) EW 1996 EW EX Gerlach, 2016
Partula garrettii Pease, 1865 EX 2009 EW EW
Partula guamensis (Pfeiffer, 1846) CR(PE) 2012 EX Gerlach, 2016
Partula hebe (Pfeiffer, 1846) EW 1996 EW EW
Partula jackieburchi (Kondo, 1981) EX 1996 EX EX
Partula labrusca Crampton and Cooke, 1953 EX 2009 EW EW Gerlach, 2016
Partula langfordi Kondo, 1970 CR 1996 EX Kerr, 2013; Bauman and
Kerr, 2013; Gerlach, 2016
Partula leptochila Crampton, 1956 EX 1994 EX EX
Partula levistriata Crampton, 1956 EX 1994 EX EX
Partula lugubris Pease, 1865 EX 2009 EX EX
Partula lutea Lesson, 1831 EX 1994 EX EX
Partula magistri Gerlach, 2016 EX Gerlach, 2016
Partula makatea Gerlach, 2016* EX Gerlach, 2016
Partula mirabilis Crampton, 1924 EW 1996 EW
Partula mooreana Hartman, 1880 EW 1996 EW EW
Partula navigatoria (Pfeiffer, 1849) EX 2009 EX EW Gerlach, 2016
Partula pearcekellyi Gerlach, 2016 EX Gerlach, 2016
Partula planilabrum Pease, 1864 EX 1996 EX EX
Partula producta Pease, 1865 EX 1994 EX EX
Partula protracta Crampton, 1956 EX 1994 EX EX
Partula remota Crampton, 1956 EX 1994 EX EX
Partula rosea Broderip, 1832 EW 2009 EW
Partula rufa Lesson, 1831 EX? Gerlach, 2016
Partula sagitta Crampton and Cooke, 1953 EX 1996 EX EX
Partula salifana Crampton, 1925 EX 1994 EX EX
Partula suturalis Pfeiffer, 1855 EW 2009 EW EW
Partula tohiveana Crampton, 1924 EW 1996 EW EW
Partula tristis Crampton and Cooke, 1953 EW 1996 EW EX Gerlach, 2016
Partula turgida (Pease, 1865) EX 1994 EX EX
Partula umbilicata Pease, 1866 EX 1996 EX EX
Partula varia Broderip, 1832 EW 2009 EX EW Gerlach, 2016
(Continued)
Page 30 THE NAUTILUS, Vol. 131, No. 1
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Samoana cramptoni Pilsbry and Cooke, 1934 CR(PE) 2012 EX?
Samoana minuta (Pfeiffer, 1856) EX? Gerlach, 2016
Samoana pilsbryi Gerlach, 2016 EX? Gerlach, 2016
PLEURODONTIDAE
Discolepis desidens (Rang, 1834) EX 1996 EX EX
Polydontes perplexa (Pfeiffer, 1850) EX? EX?
Polydontes undulata (Fe
´russac, 1821) EX? EX?
POLYGYRIDAE
Vespericola ohlone Roth, 2003 EX? Roth, 2003
POMATIIDAE
Tropidophora carinata (Born, 1780) EX EX
Tropidophora desmazuresi (Crosse, 1873) EX 1994 EX
Tropidophora icterica (Sowerby, 1847) EX EX
Tropidophora lienardi Morelet, 1876 EX EX
Tropidophora mauritiana (H. Adams, 1867) EX EX
Tropidophora scabra (H.Adams, 1867) EX EX
Tropidophora semilirata (Morelet, 1881) EX 1994 EX Evaluated as Tropidophora
semilineata’ by IUCN (2016)
Tropidophora vincentflorensi Griffiths, 2000 EX EX
PRISTILOMATIDAE
Gyralina hausdorfi Riedel, 1990 EX EX
PUNCTIDAE
Punctum mokotoense Abdou and Bouchet, 2000 EX EX? Abdou and Bouchet, 2000
PUPILLIDAE
Pupilla obliquicosta Smith, 1892 EX 1994 EX EX
RHYTIDIDAE
Delos gardineri Smith, 1897 CR(PE) 2012 EX EX? Barker, 2012
Schizoglossa major Powell, 1938 EX Spencer et al., 2009
STREPTAXIDAE
Conturbatia crenata Gerlach, 2001 CR(PE) 2009 EX?
Gibbus lyonetianus Pallas, 1780 EX 1994 EX EX
Gonidomus newtoni (Adams, 1867) EX 1994 EX EX
Gonospira adamsiana Nevill, 1871 EX EX
Gonospira cirneensis Madge, 1946 EX EX
Gonospira helodes (Morelet, 1875) EX EX
Gonospira majuscula (Morelet, 1878) EX EX
Gonospira mondraini (H. Adams, 1868) EX EX
Gonospira nevilli Adams, 1867 EX 1994 EX
Gulella mamellensis Griffiths, 2000 EX EX
Microstrophia abnormala Griffiths, 2004 EX EX
Microstrophia baideri Griffiths, 2004 EX EX
Plicadomus newtoni (H. Adams, 1867) EX EX
STROPHOCHEILIDAE
Anthinus multicolor Rang, 1831 EX? EX?
Anthinus turnix (Gould 1846) EX? EX?
Gonyostomus egregius (Pfeiffer, 1845) EX? EX?
Gonyostomus goniostoma (Wood, 1828) EX? EX?
Megalobulimus cardosoi (Morretes, 1952) EX 1996 EX EX
SUBULINIDAE
Chilonopsis blofeldi Forbes, 1852 EX 1994 EX EX
Chilonopsis exulatus (Benson, 1850) EX 1994 EX EX
Chilonopsis helena Quoy and Gaimard, 1833 EX 1994 EX EX
Chilonopsis melanoides (Wollaston, 1892) EX 1996 EX EX
Chilonopsis nonpareil*(Perry, 1811) EX 1994 EX EX
Chilonopsis subplicatus (Sowerby, 1852) EX 1994 EX EX
Chilonopsis subtruncatus (Smith, 1892) EX 1994 EX EX
Chilonopsis turtoni (Smith, 1892) EX 1994 EX EX
Vegrandinia trindadensis (Breure and Coelho, 1976) EX?
(Continued)
R.H. Cowie et al., 2017 Page 31
Table A3. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
SUCCINEIDAE
Succinea atollica Hertlein and Allison, 1968 EX EX
Succinea rotumana Smith, 1897 CR(PE) 2012 EX EX? Barker, 2012
UROCOPTIDAE
Anoma adamsi Pilsbry, 1904 EX? EX?
Anoma alboanfractus (Chitty, 1853) EX? EX?
Anoma dohriana (Pfeiffer, 1871) EX? EX?
Anoma flexuosa (Pfeiffer, 1866) EX? EX?
Anoma fuscolabris (Chitty, 1853) EX? EX?
Anoma gossei (Pfeiffer, 1846) EX? EX?
Anoma gracilis (C. B. Adams, 1851) EX? EX?
Anoma integra (C.B. Adams, 1851) EX? EX?
Anoma jarvisi Pilsbry, 1903 EX? EX?
Anoma nitens (Chitty, 1853) EX? EX?
Anoma prunicolor (Chitty, 1853) EX? EX?
Anoma pulchella (Chitty, 1853) EX? EX?
Anoma pulla (Chitty, 1853) EX? EX?
Anoma radiata (Chitty, 1853) EX? EX?
Anoma solida (C. B. Adams, 1851) EX? EX?
Anoma tricolor (Pfeiffer, 1847) EX? EX?
UROCYCLIDAE
Colparion madgei Laidlaw, 1938 EX 1994 EX EX
Malagarion borbonica (Morelet, 1860) EX? EX?
Zingis radiolata Martens, 1878 CR(PE) 2004 EX?
VERTIGINIDAE
Lyropupa perlonga (Pease, 1871) EX 1994 EX EX
Nesopupa turtoni (Smith, 1892) EX 1994 EX EX
Vertigo bermudensis Pilsbry, 1919 EX? EX?
Vertigo marki Gulick, 1904 EX? EX?
ZONITIDAE
Zonites embolium Fuchs and Ka
¨ufel, 1936 EX EX
Zonites santoriniensis Riedel and Norris 1987 EX EX Kornilios et al., 2009
Zonites siphnicus Fuchs and Ka
¨ufel 1936 EX EX Kornilios et al., 2009
Page 32 THE NAUTILUS, Vol. 131, No. 1
Table A4. Freshwater species considered extinct (EX), possibly extinct (EX?) or extinct in the wild (EW) in the present study,
compared with their status as evaluated by Re
´gnier et al. (2009), and on the Red List (IUCN, 2016). Red List categories are extinct (EX),
extinct in the wild (EW), critically endangered (possibly extinct) (CR(PE)), critically endangered (CR), least concern (LC), and data
deficient (DD); the date of the IUCN evaluation follows the status. EX? is treated as equivalent to CR(PE). A dash indicates the species
was not evaluated. Sources are only provided if the status in this study differs from the later of IUCN (2016) and Re
´gner et al. (2009).
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
GASTROPODA
AMNICOLIDAE
Amnicola rhombostoma Thompson, 1968 EX? EX?
Lyogyrus bakerianus (Pilsbry, 1917) EX? Johnson et al., 2013
ASSIMINEIDAE
Pseudogibbula cara
Pilsbry and Bequaert, 1927
CR(PE) 2010 EX?
Valvatorbis mauritii
Bequaert and Clench, 1936
CR(PE) 2010 EX?
BITHYNIIDAE
Gabbiella barthi (Brown, 1980) CR(PE) 2016 EX?
Gabbiella matadina Mandahl-Barth, 1968 CR(PE) 2010 EX?
Soapitia dageti Binder, 1961 CR(PE) 2010 EX?
BYTHINELLIDAE
Bythinella eutrepha (Paladilhe, 1867) CR(PE) 2010 EX?
Bythinella gibbosa
(Moquin-Tandon, 1856)
EX 2010 EX
Bythinella limnopsis
Letourneux and Bourguignat, 1887
EX 2010 EX
Bythinella mauritanica
Letourneux and Bourguignat, 1887
EX 2010 EX
Bythinella microcochlia
Letourneux and Bourguignat, 1887
EX 2010 EX
Bythinella punica
Letourneux and Bourguignat, 1887
EX 2010 EX
COCHLIOPIDAE
Dyris amazonicus (Haas, 1949) EX? EX?
Heleobia peiranoi (Weyrauch, 1963) EX Rumi et al., 2006
Heleobia spinellii (Gredler, 1859) EX 2010 EX
Heleobia steindachneri (Westerlund, 1902) EX EX
Heleobia sublineata (Pilsbry 1911) EX Rumi et al., 2006
Juturnia brunei (Taylor, 1987) CR(PE) 2012 EX Hershler et al., 2014
Littoridina gaudichaudii Souleyet, 1852 EX 1996 EX? EX?
Sioliella effusa Haas, 1949 EX? EX?
Tryonia hertleini (Drake, 1956) EX EX
Tryonia santarosae Hershler et al., 2014 EX Hershler et al., 2014
Tryonia shikueii Hershler et al., 2014 EX Hershler et al., 2014
GLACIDORBIDAE
Glacidorbis costatus Ponder and Avern, 2000 EX EX
HYDROBIIDAE
Alzoniella galaica (Boeters and Rolan, 1988) CR(PE) 2011 EX?
Antibaria notata (Frauenfeld, 1865) EX EX
Belgrandia moitessieri (Bourguignat, 1866) CR(PE) 2010 EX?
Belgrandia varica (Paget, 1854) CR(PE) 2010 EX EX? Prie
´, 2010
Belgrandiella boetersi
Reischu¨ tz and Falkner, 1998
CR(PE) 2010 EX?
Belgrandiella cavernica Boettger, 1957 CR(PE) 2014 EX?
Belgrandiella intermedia (Boeters, 1970) EX 1996 EX EX
Belgrandiella kreisslorum Reischu¨ tz, 1997 CR(PE) 2010 EX?
Belgrandiella multiformis
Fischer and Reischu¨ tz, 1995
CR(PE) 2010 EX?
Bracenica spiridoni Radoman, 1973 EX EX
Bythiospeum pfeifferi (Clessin, 1890) EX? EX?
Dalmatinella fluviatilis Radoman, 1973 EN 2011 EX EX Evaluated as Vinodolia fluviatilis
by IUCN (2016)
(Continued)
R.H. Cowie et al., 2017 Page 33
Table A4. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Dianella schlickumi Schu¨ tt, 1962 CR(PE) 2011 EX EX?
Falsipyrgula beysehirana (Schu¨ tt, 1965) CR(PE) 2014 EX?
Graecoanatolica brevis Radoman, 1973 CR(PE) 2014 EX? Kebapc¸ i et al., 2012
Graecoanatolica conica Radoman, 1973 CR(PE) 2014 EX? Kebapc¸ i et al., 2012
Graecoanatolica macedonica
Radoman and Stanovic, 1978
EX 2002 EX EX
Hydrobia anatolica Schu¨ tt, 1965 CR(PE) 2014 EX?
Hydrobia gracilis Morelet, 1880 EX 2010 EX
Islamia ateni (Boeters, 1969) EX 2011 EX
Islamia bendidis Reischu¨ tz, 1988 CR(PE) 2011 EX?
Islamia epirana (Schu¨ tt, 1962) EX EX
Islamia graeca Radoman, 1973 CR(PE) 2011 EX EX
Islamia hadei (Gittenberger, 1982) CR(PE) 2011 EX EX
Islamia pseudorientalica Radoman, 1973 CR(PE) 2014 EX?
Kirelia carinata Radoman, 1973 CR(PE) 2014 EX
Marstonia olivacea Pilsbry, 1895 EX 2000 EX EX
Mercuria punica
(Letourneux and Bourguignat, 1887)
CR(PE) 2010 EX?
Nanivitrea alcaldei (Jaume and Abbott, 1947) EX? Va
´zquez Perera and Perera
Valderrama, 2010
Nanivitrea helicoides (Gundlach, 1865) EX? Va
´zquez Perera and Perera
Valderrama, 2010
Neohoratia coronadoi (Bourguignat, 1870) EX? Arconada and Ramos, 2006
Ohridohauffenia drimica (Radoman, 1964) EX 1994 EX EX
Ohridohauffenia minuta (Radoman, 1955) CR(PE) 2010 EX?
Potamopyrgus acus Haase, 2008 CR(PE) 2013 EX?
Pseudamnicola barratei
Letourneux and Bourguignat, 1887
EX 2010 EX
Pseudamnicola doumeti
Letourneux and Bourguignat, 1887
EX 2010 EX
Pseudamnicola latasteana
Letourneux and Bourguignat, 1887
EX 2010 EX
Pseudamnicola letourneuxiana
(Bourguignat, 1862)
EX 2010 EX
Pseudamnicola macrostoma (Ku¨ ster, 1853) EX EX
Pseudamnicola oudrefica
(Letourneux and Bourguignat, 1887)
EX 2010 EX
Pseudamnicola ragia
Letourneux and Bourguignat, 1887
EX 2010 EX
Pseudamnicola singularis
Letourneux and Bourguignat, 1887
EX 2010 EX
Pseudoislamia balcanica Radoman, 1979 CR 2011 EX EX
Pyrgulopsis brandi (Drake, 1953) EX EX
Pyrgulopsis carinata Hershler, 1998 EX EX
Pyrgulopsis coloradensis Hershler, 1998 EX? Center for Biological Diversity
et al., 2009
Pyrgulopsis nevadensis (Stearns, 1833) EX 2000 EX EX
Pyrgulopsis ruinosa Hershler, 1998 EX EX
Pyrgulopsis torrida Hershler et al., 2016 EX? Hershler et al., 2016
Radomaniola curta (Ku¨ ster, 1853) LC 2010 EX EX
Sardohoratia sulcata Manganelli et al., 1998 CR(PE) 2010 EX?
Tanousia zrmanjae (Brusina, 1866) CR(PE) 2011 EX EX?
Trichonia kephalovrissonia Radoman, 1973 DD 2011 EX EX Evaluated as Heleobia
steindachneri by IUCN (2016)
(Continued)
Page 34 THE NAUTILUS, Vol. 131, No. 1
Table A4. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Trichonia trichonica Radoman, 1973 CR 2011 EX EX
Turcorientalia hohenackeri (Ku¨ ster, 1853) VU 2011 EX EX
Vinodolia fiumana Radoman, 1973 EN 2014 EX EX
Vinodolia gluhodolica (Radoman, 1973) EN 2010 EX EX
Vinodolia lacustris (Radoman, 1973) CR 2010 EX Albrecht et al., 2012
Vinodolia matjasici (Bole, 1961) CR 2010 EX EX
Zaumia sanctizaumi(Radoman, 1964) CR(PE) 2010 EX?
IRIDINIDAE
Aspatharia divaricata (Martens, 1897) CR(PE) 2016 EX?
Chambardia letourneuxi EX 2010 EX
LITHOGLYPHIDAE
Clappia umbilicata (Walker, 1904) EX 2000 EX EX
Fluminicola minutissimus Pilsbry, 1907 EX ? Johnson et al., 2013
Fluminicola nuttallianus Lea, 1838 EX EX ? Johnson et al., 2013
Somatogyrus crassilabris Walker, 1915 EX 2000 EX EX? Johnson et al., 2013
Somatogyrus wheeleri Walker, 1915 EX 2000 EX EX? Johnson et al., 2013
LYMNAEIDAE
Erinna aulacospira (Ancey, 1899) DD EX? Johsnon et al., 2013
Galba cyclostoma (Walker, 1908) EX? Johnson et al., 2013
Galba perpolita (Dall, 1905) EX? Johnson et al., 2013
Galba tazewelliana (Wolf, 1870) EX? Johnson et al., 2013
Galba vancouverensis (F.C. Baker, 1939) EX? Johnson et al., 2013
Lantzia carinata (Jousseaume, 1872) CR(PE) 2016 EX?
Lymnaea plicata Hylton Scott 1953 EX Rumi et al., 2006
Stagnicola neopalustris (F.C. Baker, 1911) EX? Johnson et al., 2013
Stagnicola petoskeyensis (Walker, 1908) EX? Johnson et al., 2013
Stagnicola pilsbryi Hemphill, 1890 EX 2012 EX EX
Stagnicola utahensis (Call, 1884) CR(PE) 2012 EX Center for Biological Diversity
et al., 2009
MELANOPSIDAE
Melanopsis germaini Pallary, 1939 CR(PE) 2014 EX?
Melanopsis infracincta Martens, 1874 CR(PE) 2014 EX?
Melanopsis khabourensis Pallary, 1939 CR(PE) 2014 EX?
Melanopsis pachya Pallary, 1939 CR(PE) 2014 EX?
Melanopsis parreyssii (Philippi, 1847) CR 2011 EX
ˆrbu and Benedek, 2016
MOITESSIERIIDAE
Henrigirardia wienini (Girardi, 2001) CR(PE) 2010 EX?
Iglica gratulabunda (Wagner, 1910) CR(PE) 2010 EX?
Paladilhiopsis janinensis Schu¨ tt, 1962 CR(PE) 2011 EX EX?
Spiralix corsica Bernasconi, 1994 CR(PE) 2010 EX?
NERITIDAE
Neritina tiassalensis Binder, 1955 CR(PE) 2010 EX?
PACHYCHILIDAE
Sulcospira martini (Schepmann, 1898) EX? EX?
Sulcospira pisum (Brot, 1868) EX? Marwoto and Isnaningsih, 2012
Sulcospira sulcospira (Mousson, 1849) DD 2011 EX? EX? Marwoto and Isnaningsih, 2012
PHYSIDAE
Physella microstriata (Chamberlain and
Berry, 1930)
EX 2000 EX EX
PLANORBIDAE
Amphigyra alabamensis Pilsbry, 1906 EX 2000 EX EX
Ceratophallus concavus
(Mandahl-Barth, 1954)
CR(PE) 2016 EX ?
Glyptophysa oconnori (Cumber, 1941) DD 2011 EX Spencer et al., 2009
Neoplanorbis carinatus Walker, 1908 EX 2000 EX EX
Neoplanorbis smithi Walker, 1908 EX 2000 EX EX
Neoplanorbis tantillus Pilsbry, 1906 EX 2012 EX EX
Neoplanorbis umbilicatus Walker, 1908 EX 2000 EX EX
(Continued)
R.H. Cowie et al., 2017 Page 35
Table A4. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Planorbella columbiensis (F.C. Baker, 1935) EX? Johnson et al., 2013
Planorbella multivolvis (Case, 1847) EX 2000 EX EX
Planorbella traskii (Lea, 1856) EX Johnson et al., 2013
Rhodacmea hinkleyi (Walker, 1908) EX? Johnson et al., 2013
Vorticifex solida (Dall, 1870) EX? Johnson et al., 2013
PLEUROCERIDAE
Athearnia crassa (Haldeman, 1841) EX 1996 EX EX
Elimia brevis (Reeve, 1860) EX 2000 EX EX
Elimia clausa (Lea, 1861) EX 2000 EX EX
Elimia fusiformis (Lea, 1861) EX 2000 EX EX
Elimia gibbera (Goodrich, 1922) EX 2000 EX EX
Elimia hartmaniana (Lea, 1861) EX 1994 EX EX
Elimia impressa (Lea, 1841) EX 1994 EX EX
Elimia jonesi (Goodrich, 1936) EX 1994 EX EX
Elimia laeta (Jay, 1839) EX 1994 EX EX
Elimia macglameriana (Goodrich, 1936) EX 2000 EX EX
Elimia pilsbryi (Goodrich, 1927) EX 1994 EX EX
Elimia pupaeformis (Lea, 1864) EX 1994 EX EX
Elimia pupoidea (Anthony, 1854) EX Johnson et al., 2013
Elimia pygmaea (Smith, 1936) EX 1994 EX EX
Gyrotoma excisa (Lea, 1843) EX 2000 EX EX
Gyrotoma lewisii (Lea, 1869) EX 2000 EX EX
Gyrotoma pagoda (Lea, 1845) EX 2000 EX EX
Gyrotoma pumila (Lea, 1860) EX 2000 EX EX
Gyrotoma pyramidata
(Shuttleworth, 1845)
EX 2000 EX EX
Gyrotoma walkeri (Smith, 1924) EX 2000 EX EX
Leptoxis clipeata (Smith, 1922) EX 2000 EX
Leptoxis formosa (Lea, 1860) EX 2000 EX EX
Leptoxis ligata (Anthony, 1860) EX 2000 EX EX
Leptoxis lirata (Smith, 1922) EX 2000 EX EX
Leptoxis minor (Hinkley, 1912) EX Johnson et al., 2013
Leptoxis occultata (Smith, 1922) EX 2000 EX EX
Leptoxis showalterii (Lea, 1860) EX 2000 EX EX
Leptoxis torrefacta (Goodrich, 1922) EX 2000 EX EX
Leptoxis trilineata (Say, 1829) EX Johnson et al., 2013
Leptoxis vittata (Lea, 1860) EX 2000 EX EX
Lithasia hubrechti Clench, 1956 EX Johnson et al., 2013
Lithasia jayana (Lea, 1841) EX Johnson et al., 2013
POMATIOPSIDAE
Pomatiopsis hinkleyi Pilsbry, 1896 EX? Johnson et al., 2013
TATEIDAE
Beddomeia tumida Petterd, 1889 CR(PE) 2011 EX EX? Clark, 2011
Fluvidona dulvertonensis
(Tennison-Woods, 1876).
EX 1996 EX EX
Leiorhagium solemi
Haase and Bouchet, 1998
EX 2011 EX? EX?
Posticobia norfolkensis (Sykes, 1900) EX 1996 EX EX
Potamolithus concordianus Parodiz, 1966 EX Rumi et al., 2006
THIARIDAE
Aylacostoma brunneum
Vogler and Peso, 2014
EW M.G. Quintana, pers. comm., 2016
Aylacostoma chloroticum
Hylton-Scott, 1953
EW 2000 EW EW M.G. Quintana pers. comm., 2016
Aylacostoma guaraniticum
Hylton-Scott, 1953
EW 2000 EW EX Rumi et al., 2006; M.G. Quintana
pers. comm., 2016
Aylacostoma stigmaticum
Hylton-Scott, 1953
EW 2000 EW EX Rumi et al., 2006; M.G. Quintana
pers. comm., 2016
(Continued)
Page 36 THE NAUTILUS, Vol. 131, No. 1
Table A4. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Melanoides agglutinans
(Bequaert and Clench, 1941)
CR(PE) 2010 EX?
VALVATIDAE
Valvata klemmi Schu¨ tt, 1962 EN 2011 EX EX
Valvata virens Tryon, 1863 EX? Johnson et al., 2013
VIVIPARIDAE
Bellamya phthinotropis (Martens, 1892) CR(PE) 2016 EX?
Viviparus bermondianus (d’Orbigny, 1842) EX EX
BIVALVIA
CORBICULIDAE
Corbicula linduensis Bollinger, 1914 EX? EX?
Corbicula subplanata Martens, 1897 EX? EX?
DREISSENIDAE
Dreissena caspia Eichwald, 1855 CR(PE) 2011 EX?
MYCETOPODIDAE
Anodontites moricandi (Lea, 1860) EX? EX?
SPHAERIIDAE
Eupera crassa (Mandahl-Barth, 1954) CR(PE) 2016 EX?
Pisidium betafoense Kuiper, 1953 CR(PE) 2016 EX?
UNIONIDAE
Alasmidonta mccordi Athearn, 1964 EX 2000 EX EX
Alasmidonta robusta Clarke, 1981 EX 2000 EX EX
Alasmidonta wrightiana (Walker, 1901) EX 2000 EX EX
Coelatura rothschildi
(Neuville and Anthony, 1906)
CR(PE) 2016 EX?
Cuneopsis demangei Haas, 1929 CR(PE) 2011 EX?
Elliptio nigella (Lea, 1852) CR 2012 EX EX
Epioblasma arcaeformis (Lea, 1831) EX 2000 EX EX
Epioblasma biemarginata (Lea, 1857) EX 2000 EX EX
Epioblasma flexuosa (Rafinesque, 1820) EX 2000 EX EX
Epioblasma haysiana (Lea, 1834) EX 2000 EX EX
Epioblasma lenior (Lea, 1842) EX 2000 EX EX
Epioblasma lewisii (Walker, 1910 EX 2000 EX EX
Epioblasma othcaloogensis (Lea, 1857) CR(PE) 2012 EX?
Epioblasma personata (Say, 1829) EX 2000 EX EX
Epioblasma propinqua (Lea, 1857) EX 2000 EX EX
Epioblasma sampsonii (Lea, 1862) EX 2000 EX EX
Epioblasma stewardsonii (Lea, 1852) EX 2000 EX EX
Epioblasma turgidula (Lea, 1858) EX 2000 EX EX
Germainaia geayi (Germain, 1911) EX 2016 EX
Lamprotula crassa (Wood, 1815) CR(PE) 2011 EX?
Lamprotula liedtkei Rolle, 1904 CR(PE) 2011 EX?
Lamprotula nodulosa (Wood, 1815) CR(PE) 2011 EX?
Lampsilis binominata Simpson, 1900 EX 2000 EX EX
Medionidus mcglameriae
van der Schalie, 1939
EX 2000 EX EX
Obovaria haddletoni (Athearn, 1964) CR(PE) 2012 EX?
Pleurobema altum (Conrad, 1854) EX 2000 EX EX
Pleurobema avellanum Simpson, 1900 EX 2000 EX EX
Pleurobema bournianum (Lea, 1840) EX 2000 EX EX
Pleurobema chattanoogaense (Lea, 1858) EX EX
Pleurobema curtum Lea, 1859 CR(PE) 2012 EX?
Pleurobema flavidulum (Lea, 1861) EX 2000 EX
Pleurobema hagleri Frierson, 1900 EX 2000 EX
Pleurobema hanleyianum (Lea, 1852) CR 2012 EX EX
Pleurobema johannis (Lea, 1859) EX 2000 EX
Pleurobema murrayense (Lea, 1868) EX 2000 EX
Pleurobema nucleopsis (Conrad, 1849) EX 2000 EX
(Continued)
R.H. Cowie et al., 2017 Page 37
Table A4. (cont.)
Species Red List
Re
´gnier et al.,
2009
This
study
Source for revised
status; comments
Pleurobema perovatum (Conrad, 1834) EX 2000 EX
Pleurobema troschelianum (Lea, 1852) EX 2000 EX
Pleurobema verum (Lea, 1861) EX 2000 EX EX
Quadrula tuberosa (Lea, 1840) CR 1996 EX EX Evaluated as Theliderma tuberosa
by IUCN (2016)
Table A5. Marine species considered extinct (EX) or possibly extinct (EX?) in the present study, compared with their status as
evaluated by Re
´gnier et al. (2009), and on the Red List (IUCN, 2016). Dashes indicate that the species was not evaluated. A source is
only provided for species of Conidae, for which the status in this study differs from that of IUCN (2016) and Re
´gner et al. (2009).
Species Red List Re
´gnier et al., 2009 This study Source for revised status
CONIDAE
Conasprella sauros (Garcia, 2006) DD EX? Peters et al., 2013
Conus bellulus Rola
´n, 1990 EX? Peters et al., 2013
Conus colmani Ro
¨ckel and Korn, 1990 DD EX? Singleton, 2007
LOTTIIDAE
Lottia alveus Conrad, 1831 EX 1994 EX EX
NACELLIDAE
Collisella edmitchelli Lipps, 1966 EX 1996 EX EX
POTAMIDIDAE
Cerithideopsis fuscata (Gould, 1857) EX EX
Page 38 THE NAUTILUS, Vol. 131, No. 1
LITERATURE CITED (APPENDIX)
Abdou A. and P. Bouchet. 2000. Nouveaux gaste
´ropodes
Endodontidae et Punctidae (Mollusca, Pulmonata)
re
´cemmente
´teints de l’archipel des Gambier (Polyne
´sie).
Zoosystema 22: 689–707.
Abdou, A., I. Muratov, and P. Bouchet. 2004. Mollusques
terrestres de Mayotte: ele
´ments pour l’inventaire des
ZNIEFF. Muse
´um national d’Histoire naturelle, Paris,
45 pp.
Albrecht, C., T. Hauffe, K. Schreiber, and T. Wilke. 2012.
Mollusc biodiversity in a European ancient lake system:
lakes Prespa and Mikri Prespa in the Balkans.
Hydrobiologia 682: 47–59.
Arconada, B. and M.-A. Ramos. 2006. Revision of the genus
Islamia Radoman, 1973 (Gastropoda, Caenogastropoda,
Hydrobiidae), on the Iberian Peninsula and description
of two new genera and three new species. Malacologia
48: 77–132.
Association of Wildlife Research and EnVision Conservation
Office, 2015. Search System of Japanese Red Data. http://
www.jpnrdb.com/index.html Accessed 22 December 2016.
Barker, G. 2012. Delos gardineri. The IUCN Red List of
Threatened Species 2012: e.T195520A2383269.
Barker, G. 2012. Succinea rotumana. The IUCN Red List of
Threatened Species 2012: e.T177716A1495957.