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The objectives of the New Zealand Threat Classification System (NZTCS) are summarised together with the methods used to classify taxa as Threatened, At Risk, Data Deficient and Extinct. An analysis is provided of the threat rankings of 3838 New Zealand terrestrial invertebrate taxa that were assessed between 2009 and 2011. Overall, 193 taxa were Threatened with 106 Nationally Critical, 30 Nationally Endangered and 57 Nationally Vulnerable. A further 1055 invertebrates were At Risk comprising 35 Declining, six Recovering, 102 Relict and 912 that were Naturally Uncommon. Another 1208 were Data Deficient, seven were Extinct and 1175 were Not Threatened. The remaining taxa examined were classified as either Introduced and Naturalised (190) or Vagrant (11). Factors and attributes that affect the threat status of New Zealand terrestrial invertebrates are discussed.
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New Zealand Entomologist
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Assessing the conservation status
of New Zealand's native terrestrial
invertebrates
I. A.N. Stringer
a
& R. A. Hitchmough
a
a
Department of Conservation , PO Box 10420, Wellington , 6143 ,
New Zealand
Published online: 21 Jun 2012.
To cite this article: I. A.N. Stringer & R. A. Hitchmough (2012) Assessing the conservation status
of New Zealand's native terrestrial invertebrates, New Zealand Entomologist, 35:2, 77-84, DOI:
10.1080/00779962.2012.686309
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New Zealand Entomologist
Vol. 35, No. 2, July 2012, 77–84
Assessing the conservation status of New Zealand’s native terrestrial
invertebrates
I. A. N. Stringer and R. A. Hitchmough
Department of Conservation, PO Box 10420, Wellington 6143, New Zealand
E-mail: rhitchmough@doc.govt.nz
The objectives of the New Zealand Threat Classification System (NZTCS) are summarised together
with the methods used to classify taxa as Threatened, At Risk, Data Deficient and Extinct.An
analysis is provided of the threat rankings of 3838 New Zealand terrestrial invertebrate taxa that were
assessed between 2009 and 2011. Overall, 193 taxa were Threatened with 106 Nationally Critical,
30 Nationally Endangered and 57 Nationally Vulnerable. A further 1055 invertebrates were At
Risk comprising 35 Declining, six Recovering, 102 Relict and 912 that were Naturally Uncom-
mon. Another 1208 were Data Deficient, seven were Extinct and 1175 were Not Threatened. The
remaining taxa examined were classified as either Introduced and Naturalised (190) or Vagrant (11).
Factors and attributes that affect the threat status of New Zealand terrestrial invertebrates are discussed.
Keywords: classification system, endangered, risk, threat status, threatened species
Introduction
The New Zealand Threat Classification System
(NZTCS) was developed to allow objective list-
ing of the conservation status (risk of extinction)
of any taxa with a wild population established
in New Zealand. The system was developed by
a working group (Molloy et al. 2002) led by
the New Zealand Department of Conservation
(DOC) with representatives from a broad range
of interest groups. Implementation of the system
to develop and periodically update published lists
is a national process led by DOC but involv-
ing experts and interest groups from many other
organisations.
Formal listing of threatened species for protec-
tion or prioritisation is a critical action in respon-
sible conservation (New & Sands 2003). Although
the NZTCS listing has no direct or automatic influ-
ence on the legal status or resourcing of work on
threatened species, it does provide vital informa-
tion for processes such as applying legal protection
to species via amendments to the Schedules to
the New Zealand Wildlife Act 1955. Although
threat status is only one of several criteria used
in prioritisation, classifying taxa according to the
risk of extinction allows prioritisation of work and
resources to those species that need them most.
The published lists also form a basis for national
outcome monitoring to measure the effects that
DOC and other agencies have by their manage-
ment of natural heritage. These are indicators that
are reported nationally and internationally (e.g.,
Convention on Biodiversity).
Four previous lists of the status of New Zealand
species including invertebrates have been pub-
lished (Table 1). The first two lists (Molloy & Davis
1992; Molloy et al. 1994) used an assessment sys-
tem intended to directly generate a list ranked by
management priority. Subsequently, Molloy et al.
(2002) developed the NZTCS, a system intended
to categorise taxa by the risk of extinction they
faced, to be a resource for prioritisation but not an
attempt at prioritisation directly. This system was
used for the last two previous lists (Hitchmough
2002; Hitchmough et al. 2007), then the criteria
used to classify threatened species were reviewed
and a revised manual was produced (Townsend
et al. 2008).
ISSN 0077-9962 print/ISSN 1179-3430 online
© 2012 The Entomological Society of New Zealand
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78 I. A. N. Stringer and R. A. Hitchmough
Table 1. Numbers of Threatened terrestrial invertebrates listed by Molloy and Davis (1992), Molloy et al. (1994),
Hitchmough (2002) and Hitchmough et al. (2007).
Category 1992 1994 Category 2002 2007
Highest priority spp. 20 26
Nationally Critical 89 126
Second priority spp. 40 52
Nationally Endangered 87 103
Third priority spp. 21 20
Nationally Vulnerable 811
Serious Decline 16 14
Gradual Decline 33 40
Sparse 62 117
Range Restricted 361 542
Threatened, but few data 15 166
Data Deficient 368 1319
Not seen for some years 4 12
Extinct 410
Total 100 276 1028 2282
Changes were made to the criteria, new At
Risk categories were introduced and some of the
old categories became qualifiers but, overall, the
changes were not fundamental. The new crite-
ria were developed largely to account for rar-
ity and for the very restricted distributions that
many taxa have amongst the islands of the New
Zealand archipelago (de Lange & Norton 2008).
The NZTCS is intended to provide finer detail and
complement the Red Data listings of the Inter-
national Union for the Conservation of Nature,
which are designed to detect rarity and decline
at global and continental scales (IUCN 2010).
The main differences between the system used
here and Molloy et al. (2002) are: replacement of
the Chronically Threatened, Serious Decline
and Gradual Decline categories by the Declin-
ing category; the re-sorting of the uncommon but
non-declining categories formerly listed as Range
Restricted and Sparse into Relict and Natu-
rally Uncommon categories that separate taxa
according to whether the rarity is natural or human-
induced rather than by geographic pattern (Range
Restricted and Sparse are now available as qual-
ifiers in all categories); the introduction of the
new category Recovering; and the development
of a recovery pathway within Nationally Vul-
nerable. The criteria defining all threat categories
have also changed somewhat to ensure complete
coverage of all possible combinations of population
size and trend.
The three Threatened categories clearly form a
series of decreasing risk of extinction from Nation-
ally Critical to Nationally Vulnerable. The
At Risk categories carry a lower or longer-term
risk of extinction; however, no ranking of the cat-
egories within the At Risk umbrella is possible
because they represent different types rather than
different degrees of risk and, within each category,
a quite broad range of degrees of risk is included
(Townsend et al. 2008).
The revised manual (Townsend et al. 2008)
has been used to generate listings for a wide
range of taxonomic groups including vertebrates,
marine invertebrates, plants, bryophytes, and is
now applied to terrestrial invertebrates (Andrew
et al. 2012; Buckley et al. 2012; Leschen et al.
2012; Mahlfeld et al. 2012; Sirvid et al. 2012;
Stringer et al. 2012a; Stringer et al. 2012b; Trewick
et al. 2012; Ward et al. 2012; Yeates et al.
2012). Here we summarise the overall numbers
of terrestrial invertebrates that are now ranked
in each of the new categories and discuss the
changes that have occurred since the last published
list.
Methods
The current lists include both endemic and
non-endemic terrestrial invertebrate taxa found
in New Zealand and its offshore and outly-
ing islands including the Kermadec, Chatham,
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Conservation of New Zealand invertebrates 79
Snares, Antipodes, Bounty, Campbell and Auck-
land Island groups. Where a non-endemic taxon
is listed, only the New Zealand population(s) was
assessed. Usually, only taxa known to be of concern
were assessed but all known Nematoda, Araneae,
Orthoptera, Phthiraptera, spiders and acanthodrilid
and megascolicid earthworm taxa were assessed.
Both taxonomically determinate and indeter-
minate taxa were assessed. Taxonomically deter-
minate taxa are those that are legitimately and
effectively published according to the criteria of
the International Code of Zoological Nomenclature
and generally accepted by relevant experts as dis-
tinct. Taxonomically indeterminate taxa are either
legitimately and effectively published but not gen-
erally accepted as distinct, or are entities yet to
be furnished with a formal name (Townsend et al.
2008). For brevity, we use the term taxa loosely to
include both species and indeterminate entities; the
latter were included because they were considered
so distinctive that they are likely to be described
as species. We take a precautionary approach by
including taxa that have not yet been described but
are considered by experts on current evidence as
likely to be new species. This ensures that they are
not ignored because they lack formal names so that
appropriate conservation management and research
can be directed towards them when necessary.
We incorporated information from the public
and a broader pool of experts not directly involved
in the listing process. Public submissions on the
threat status re-evaluation were invited via the DOC
website (http://www.doc.govt.nz/getting-involved
/consultations/closed/new-listing-of-threatened-
status-of-new-zealand-terrestrial-and-freshwater-
invertebrates/) on 26 May 2009. Submissions
closed on 30 September 2009 so this website is
no longer active. Such submissions were intended
to notify the expert panel members of the existence
of new information that they might otherwise not
have access to which would inform decisions; they
were not intended to be an avenue for lobbying or
voting on proposed changes.
Panels of experts (the authors of articles in this
issue) were created to undertake the re-evaluation.
The role of the expert panel members was to pro-
vide knowledge on their particular field of exper-
tise, to answer queries on listing decisions reached,
to consult with peers, to bring as much infor-
mation as possible to the meetings and to help
prepare the reports. Panels met between 17 Decem-
ber 2009 and 25 August 2011 and placed taxa
into threat categories based on the criteria provided
by Townsend et al. (2008) (see Supplementary
Appendix 1 (Introduction)). Panel meetings were
followed by subsequent telephone and email com-
munication. This process was guided by panel
knowledge, information from invertebrate collec-
tions (e.g., New Zealand Arthropod Collection),
submissions received, and recent publications relat-
ing to taxonomic and population status. Where
there was doubt in placing a given taxon into a
threat category, the provisional assessments of a
panel were referred to other relevant experts sub-
sequent to the panel meetings. The categories that
have not yet included invertebrates are Coloniser
and Migrant. Coloniser invertebrates certainly
exist, but none that fits these categories has yet been
proposed for NZTCS assessment.
Taxa were classified using both status and trend
criteria (see Townsend et al. 2008). Status criteria
(usually area of occupancy of the total population
or total number of populations; rarely estimates of
the total number of mature individuals or number of
mature individuals in the largest population) were
generally considered first followed by an evalu-
ation of the trend criteria (ongoing or predicted
population trend, due to existing threats, measured
usually by area of occupancy and rarely by esti-
mated population size). Change in status for a taxon
can result from one or more of three, non-mutually-
exclusive drivers: i) it has genuinely become more
or less threatened with extinction; ii) new knowl-
edge has allowed a more accurate assessment of its
true status; and iii) the revised categories and cri-
teria mean that the same information puts it into a
different category.
The main abbreviations used are: Threatened:
NC = Nationally Critical;NE= Nationally
Endangered;NV= Nationally Vulnerable.
At Risk:Dc= Declining;Rc= Recovering;
Rl = Relict;NU= Naturally Uncommon.
Other categories: DD = Data Deficient;NT=
Not Threatened; I&N = Introduced and
Naturalised;Vg= Vagrant;Ex= Extinct.
Townsend et al. (2008; see Supplementary
Appendix 1 (Introduction)) also provided a series
of qualifiers to enable additional information
on each taxon to be captured and consid-
ered. Most that applied to invertebrates are self
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80 I. A. N. Stringer and R. A. Hitchmough
evident: Conservation Dependent; Data Poor;
Extreme Fluctuations (of population size);
Island Endemic; One Location; and Sparse.
The Range Restricted qualifier applies to taxa
that are confined to specific substrates, habitats
or geographic areas of less than 1000 km
2
. The
assessment was made by summing the areas of
habitat occupied by all sub-populations, whether
natural or human induced. It is mutually exclu-
sive with the One Location qualifier. Other
terms used: New = taxa added in 2010; SD =
Serious Decline;GD= Gradual Decline;
Syn = synonym; NmD = nomen dubium; NP =
No Longer Present (includes incorrectly recorded
as present in New Zealand, not a valid species,
etc.). For full definitions of abbreviations and
terminology see Townsend et al. (2008).
Parasites were assessed using a slightly differ-
ent process. In most cases little information was
available on the incidences of infection but the
hosts of parasites are often listed, especially for
vertebrate hosts. The expert panels relied strongly
on their own knowledge and experience using
the precautionary principle detailed by Townsend
et al. (2008). In addition, host-specific parasites
were considered to have the same threat ranking
as their hosts unless there was information that
suggested they were even more threatened. Para-
sites with multiple hosts were given the ranking of
the least threatened host. The paucity of published
geographic location information for most parasite
species meant that the One Location qualifier was
largely not applied unless there was strong evidence
that this was likely to be true.
Results
The threat status of 3838 taxa was assessed
(Table 2). Taxa that were Data Deficient (31.5%)
comprised the largest proportion followed by Not
Threatened (30.6%) and Naturally Uncom-
mon taxa (23.8%). Relatively few were in the
other At Risk categories (3.7%) and few were
Threatened (5.0%). Only seven invertebrate
species (0.2% of those classified) were consid-
ered Extinct. Of the taxa classified as Threat-
ened, 54.9% were Nationally Critical followed
by 29.5% that were Nationally Vulnerable
and 15.5% were Nationally Endangered. Nat-
urally Uncommon taxa predominated (86.5%)
amongst the At Risk categories followed by
Relict (9.7%), Declining (3.3%) and Recovering
(0.6%) (Table 2).
Discussion
Threat assessments have now been made for
approximately 26% of the > 13,000 species and
subspecies of terrestrial invertebrates known to
occur in New Zealand (Gordon 2010). Indetermi-
nate taxa that have not yet been formally described
or are of uncertain identity comprise 12% (462
taxa) of all invertebrates so far assessed and, if
these are also included with species known to
occur in New Zealand, then threat rankings are
now available for about 29% of all terrestrial inver-
tebrates. Overall these comprise about 1.5% that
are Threatened, 8% that are At Risk (these com-
prise mostly Naturally Uncommon taxa, 7.2%)
and 9.2% that are Data Deficient. However, we
suggest that the proportion of invertebrates in all
of these categories is likely to increase as more
species continue to be discovered because these
are likely to be rarer than species that have already
been found.
We also note that this proportion includes a few
marine and/or freshwater species in the Araneae,
Coleoptera, Hemiptera and Lepidoptera. We have
also included all Nematoda known to occur in
New Zealand because they are not considered else-
where and we have included the Nematomorpha
amongst the list of known species for complete-
ness even though their threat rankings have yet to
be assessed. We have excluded most marine inver-
tebrates and those that are traditionally considered
to be aquatic such as Tardigrada, Ephemeroptera,
Odonata and Trichoptera even though many have
a terrestrial phase in their life cycles or, as in some
tardigrades, live in wet terrestrial situations. We
have also excluded all Powelliphanta land snails
because the genus is presently being revised and
their threat rankings will be published later.
Changes in threat status since 2007
Over a third (38.3%) of the invertebrate taxa con-
sidered for the present reassessment had not been
ranked previously by Hitchmough et al. (2007)
and a further 96 taxa ranked by Hitchmough et al.
(2007) have since been either synonymised with
other previously ranked taxa, are now considered to
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Conservation of New Zealand invertebrates 81
Table 2. Changes in the numbers of terrestrial invertebrate taxa in each threat category between 2007 (down)
(Hitchmough et al. 2007) and 2010 (across).
2010 Threatened At Risk Other
2007 NC NE NV Dc Rc Rl NU DD I&N Vg NT Ex Total Syn NmD NP
NC 56 3 7 1 1 3 21 3 95 18
NE 5 15 19 3 2 7 7 3 4 65 6
NV 1 3 1 5
SD 1 3 1 1 1 7 1
GD 1 9 9 2 6 1 1 29 1
Sp 4 2 7 7 56 4 28 108 2
RR 9 2 2 49 358 53 28 1 502 13 2
DD 3 2 2 5 129 856 3 282 1282 16 24 6
I&N 3 36 2 41 3
Vg 0
NT 3 2 10 5 1 204 225 1
Ex 3 1 4 8 2
New 32 2 12 9 2 24 345 259 150 10 624 2 1471 6
Total 106 30 57 35 6 102 911 1208 190 11 1175 7 3838 65 24 12
Note: See Methods for abbreviations.
be of dubious taxonomic status, or are now known
to be not present in New Zealand (Table 2). Of
the taxa previously listed by Hitchmough et al.
(2007), 47.8% remained unchanged, 8.2% were
ranked more threatened and 19.8% less threatened.
This accounts for 75.7% of the taxa; 3.9% of the
remaining taxa were synonyms, nomina dubia or
otherwise considered no longer present. The 20.4%
classified in the At Risk categories of Hitchmough
et al. (2007) were reassessed using different crite-
ria from the At Risk categories of Townsend et al.
(2008) and so they cannot be directly compared.
However, 189 of the 518 taxa previously classi-
fied as Range Restricted by Hitchmough et al.
(2007) that occur only on islands or island groups
are now all classified as Naturally Uncommon.
The latter represent 7.7% of all the taxa classified
by Hitchmough et al. (2007). When these are taken
into account the threat classifications of only 12.7%
of the taxa classified by Hitchmough et al. (2007)
changed their threat status because of changes
to the classification system. Most other changes
in status from Hitchmough et al. (2007) resulted
from careful reassessment that usually included
increased information about the distributions of the
taxa.
Only one reduction in threat classification level
was due to intensive conservation management; the
Mercury Islands tusked weta, Motuweta isolata.
This was previously classified as Nationally
Critical by Hitchmough et al. (2007) but is now
classified as Recovering (Trewick et al. 2012). The
successful management involved captive-rearing
and translocation to islands near the source island
from which introduced predatory mammals had
been eradicated (Stringer & Chappell 2008). Other
translocations of threatened invertebrates have also
been recently made for conservation purposes but
these have yet to affect any threat rankings (Sher-
ley et al. 2010; Watts et al. 2008; Table 3).
We note for completeness that conservation man-
agement has occurred involving Powelliphanta
augusta (Pulmonata: Rhytididae) which was clas-
sified as Nationally Critical by Hitchmough et
al. (2007) but it has not yet had its threat status
reassessed.
Of the 10 taxa classified as Extinct by
Hitchmough et al. (2007), six are still con-
sidered Extinct: Mecodema costellum “spelles”
(NZACMcostel01); Mecodema punctellum; Thot-
mus halli; Waitomophylax worthyi (Coleoptera);
Philopteroides xenicus and Rallicola pilgrimi
(Phthiraptera). Two taxa (Placostylus amba-
giosus “Herangi Hill” and Placostylus amba-
giosus “Kohuronaki” (Pulmonata)) have been
synonymised, one species (Leioproctus otautahi
(Hymenoptera)) is now known to be a Vagrant
(Donovan & Maynard 2010) and three taxa are
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82 I. A. N. Stringer and R. A. Hitchmough
Table 3. Translocations made of Threatened inver-
tebrates for conservation management purposes since
2005 (Sherley et al. 2010; Watts et al. 2008; T. Huggins,
unpublished data).
Years of Status Status
Taxon transfer 2007 2010
Coleoptera
Anagotus turbotti 2006 RR NU
Orthoptera
Deinacrida
carinata
2010 NE Rl
Deinacrida rugosa 2007, 2008,
2010
RR Rl
Hemideina ricta 2005 RR NU
Motuweta isolata 2007, 2008,
2011
NC Rc
Pulmonata
Amborhytida
tarangaensis
2006 NV NU
Theridiidae
Latrodectus katipo 2008 SD Dc
Note: See Methods for abbreviations.
now considered Data Deficient: Megacolabus
sculpturatus (Coleoptera); Charopidae sp. 104
(NMNZ M.127901); and Zelandiscus elevata (Pul-
monata).
Changes in threat ranking for many inverte-
brates are inevitable because assessing the threat
ranking of most of them involves much subjectivity
and precise data are usually lacking. Reliable esti-
mates of total population numbers exist for only a
few taxa which have been subjected to focused dis-
tributional surveys such as those made for katipo
spiders, Latrodectus katipo (Patrick 2002), the
Banks Peninsula tree weta, Hemideina ricta (B.
Brown & I. Townsend, unpublished data), or the
rhytitid snail Rhytidarex buddlei (Brook 2002).
Even fewer invertebrates have been regularly mon-
itored as has been done for the Cromwell chafer,
Prodontria lewisii (Ferreira & McKinlay 1999).
There have, however, been more generalised sur-
veys of invertebrates from specific areas or plant
species (e.g., Patrick 2000).
Some vertebrate parasites in groups such as
Phthiraptera and parasitic nematodes are excep-
tions because their threat ranking follows that of
their well-studied vertebrate hosts. However, most
vertebrate species also have only broadly estimated
population sizes. Population estimates, especially
when the species is very cryptic, are often very
subjective. A rare example, where we know the
minimum number of individuals present is afforded
by the land snail, Powelliphanta augusta, which
was originally thought to occur sparsely over about
8.5 ha at the top of Mt Augusta, Stockton Plateau,
Westport, but 6139 individuals were later found by
exhaustive searching after some of the habitat had
been destroyed by opencast coal mining (Walker
et al. 2008). We acknowledge that subjectivity
is always involved where an estimate of popula-
tion size is required in order to determine a threat
category for terrestrial invertebrates. However, in
most instances, area of occupancy was used as an
index of population size rather than any attempt
being made to estimate population size directly.
The criteria triggered for each listing record these
decisions.
Factors affecting the threat status of New
Zealand terrestrial invertebrates
Habitat loss, or modification, and introduced preda-
tors are widely recognised as the main threats
to the native fauna of New Zealand, and these
have largely influenced the threat status of inver-
tebrates. Less obvious threats were the attitudes
of the human colonists and specimen collectors,
but these have reduced today (Watts et al. 2012).
Large tracts of indigenous vegetation have been
replaced by urban areas and exotic vegetation,
particularly pasture and exotic production for-
est, and introduced weed species have invaded
most habitats (Watts et al. 2012). Introduced
vertebrates have modified the remaining native
vegetation to varying extents through trampling,
herbivory and dispersal of weed seeds (Gibb &
Flux 1973), while introduced predators, including
invertebrate predators and parasites, have directly
affected the populations of many native inverte-
brates. This is particularly so for rodents; the few
recorded instances of invertebrate extinction on
the New Zealand mainland occurred after rodent
invasion and involved flightless and large bod-
ied species with a ground active lifestyle (Gibbs
2010; Ramsay 1978). Many direct and indirect
deleterious effects have been recorded on native
New Zealand invertebrates by exotic invertebrates,
even though such changes usually go unnoticed
(Brockerhoff et al. 2010).
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Conservation of New Zealand invertebrates 83
The attributes of New Zealand’s terrestrial
invertebrates that are generally associated with
increased threat risk are large size, flightlessness,
ground dwelling and having reduced protective
defences such as slow escape responses, especially
freezing behaviour, or a lack of defensive secretions
(McGuinness 2001). These and other factors are
discussed in relation to some New Zealand inver-
tebrate groups by four of the papers in this issue
(Leschen et al. 2012; Mahlfeld et al. 2012; Stringer
et al. 2012b; Trewick et al. 2012). There is some
quantitative evidence that large size and flightless-
ness are associated with increased threat risk for
New Zealand Coleoptera, but there is little or no
such association with flightlessness amongst New
Zealand’s Threatened and At Risk Lepidoptera
(Leschen et al. 2012; Stringer et al. 2012b).
Conclusions
The publications in this issue have two primary
aims. First, to assist in the recovery of threatened
species and, second, to prevent their extinction
by identifying Threatened and At Risk taxa.
The main purpose is to warn conservation man-
agers so they can research the causes of decline
if necessary and organise appropriate conserva-
tion management. Taxonomy, robust identification
tools and reliable distributional data are fundamen-
tal to the assessment of threat status for terrestrial
invertebrates yet neither of the authors of this paper
are invertebrate taxonomists. Our roles in these
reassessments were to gather data from experts (the
authors of the following papers) and to ensure con-
sistency in applying the criteria between different
groups. We are grateful to the professional and
non-professional taxonomists who provided this
information, and especially that of the retired or
unpaid professional taxonomists and the knowl-
edgeable amateurs who comprised 56% of the
authors of the papers in this issue. The papers in this
issue clearly show that an enormous amount of sys-
tematic research is still required in order to provide
reliable information for future threat assessments.
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... The type specimen was collected from secondary forest adjacent to extensive primary forest by sifting a mix of leaf litter and rotten branches in a location near a cave system; we do not believe that this species is a troglophile because it was not sampled proximal to the cave entrance. We recommend that O. marskeae be placed on the Department of Conservation Threatened Species List (Leschen et al. 2012) and treated as "range-restricted" and "sparse" using the criteria of Stringer and Hitchmough (2012). This formality will facilitate further collecting activity to discover more about the biology and distribution of O. marskeae. ...
... We recommend that T. lindsayae be placed on the Department of Conservation Threatened Species List (Leschen et al. 2012) and treated as "rangerestricted" and "sparse" using the criteria of Stringer and Hitchmough (2012). This formality will facilitate further collecting activity to discover more about the biology and distribution of T. lindsayae. ...
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Three new genera of Synchitini are described from New Zealand, including two monotypic genera, Oparara Leschen and Lord, new genus (type species Oparara marskeae Leschen and Lord, new species) and Tarphionivea Leschen and Lord, new genus (type species T. lindsayae Leschen and Lord, new species). The New Zealand Bitoma Herbst comprise members of multiple genera, and six are here transferred to Zebitoma Leschen and Lord, new genus (type species Bitoma nana Sharp, 1876), resulting in the following: Zebitoma brouni (Hetschko, 1928), new combination; Zebitoma discoidea (Broun, 1880), new combination; Zebitoma guttata (Broun, 1886), new combination; Zebitoma lobata (Broun, 1886), new combination; Zebitoma nana (Sharp, 1876), new combination; Zebitoma novella (Hetschko, 1929), new combination; Zebitoma picicornis (Broun, 1909), new combination; Zebitoma rugosa (Sharp, 1876), new combination; Zebitoma scita (Broun, 1886), new combination.
... Except in South Africa, little research has been conducted on the status of these invasive populations and how they have affected local ecosystems. Many of these invaded areas contain native invertebrate communities that already face threats from other invasive species, habitat alteration, and climate change [25][26][27]. The establishment of P. dominula throughout more of these regions may add to these threats. ...
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... Of these, 849 are listed as Threatened or At Risk. The robust grasshopper, Brachaspis robustus, is a flightless New Zealand endemic restricted to the Mackenzie Basin, and is classified as Threatened: Nationally Endangered [21,22]. It prefers open, stony habitat, and primarily inhabits the wide dynamic riverbeds and rocky terraces of braided rivers. ...
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... Some countries have well-developed threat listings for insects, and some taxa are well known, with long-term and ongoing surveys of geographic distribution and population levels (Curtis et al. 2015;Schmucki et al. 2016;Ellis et al. 2019). However, well-developed datasets are particularly 1 3 lacking in Southern Hemisphere countries, with major consequences (Cranston 2009;Stringer and Hitchmough 2012;Holwell and Andrew 2015). Firstly, the insects listed in conservation plans often comprise an over-representation of conspicuous taxa which are easier to survey (New and Samways 2013;Juslen et al. 2015;Potockỳ et al. 2018). ...
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Insect conservation is often impeded by a lack of species-specific data leading to difficulties in assessment of conservation and threat status. Utilising specimens and records held within natural history museums may help to overcome such limitations. We used specimens and their associated data from the New Zealand Arthropod Collection to provide a comparative analysis of threatened species and their related non-threatened congeners. A range of insect groups were covered, including families from the five mega-diverse insect orders. Data on the locality, date, habitat, and morphological measurements, were recorded for nearly 25,000 specimens from 460 species, of which 31 are Threatened and 104 are At Risk. Results found the Protected Area Network provided significantly less coverage for insect species classified as Threatened or At Risk. Body size, geographical distribution, and environmental variables were not consistent predictors of the current threat classification system. There were a disproportionately large number of At Risk species found on offshore islands. The current New Zealand threat classification system, based on trends in geographical occurrence and population sizes, does not accurately reflect the conservation needs of most insect species. This has significant implications for the utility of threat assessments for insect conservation.
... We predicted that predator control would lead to increased abundance and distribution of native prey species. For example, many of New Zealand's lizard and invertebrate taxa have declined due to the impacts of mammalian predators (Hitchmough et al. 2010;Stringer & Hitchmough 2012). Here we describe trends in predator populations and native biodiversity following this landscape-scale intervention. ...
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... For example, rare plant and fungal species appear to be particularly sensitive to changes in environment (Avis et al., 2008;Dickie et al., 2009;Dickie and Reich, 2005;McIntyre and Lavorel, 1994), and pollinating insect losses are concentrated among rare species (Powney et al., 2019). Furthermore, most of the terrestrial invertebrate species currently considered to be at risk or threat of extinction in New Zealand are naturally uncommon (Stringer and Hitchmough, 2012). ...
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The effects of land use on soil invertebrates – an important ecosystem component – are poorly understood. We investigated land-use impacts on a comprehensive range of soil invertebrates across New Zealand, measured using DNA metabarcoding and six biodiversity metrics. Rarity and phylogenetic rarity – direct measures of the number of species or the portion of a phylogeny unique to a site – showed stronger, more consistent responses across taxa to land use than widely used metrics of species richness, effective species numbers, and phylogenetic diversity. Overall, phylogenetic rarity explained the highest proportion of land use-related variance. Rarity declined from natural forest to planted forest, grassland, and perennial cropland for most soil invertebrate taxa, demonstrating pervasive impacts of agricultural land use on soil invertebrate communities. Commonly used diversity metrics may underestimate the impacts of land use on soil invertebrates, whereas rarity provides clearer and more consistent evidence of these impacts.
... For example, rare plant and fungal species appear to be particularly sensitive to changes in environment (Avis et al., 2008;Dickie et al., 2009;Dickie and Reich, 2005;McIntyre and Lavorel, 1994), and pollinating insect losses are concentrated among rare species (Powney et al., 2019). Furthermore, most of the terrestrial invertebrate species currently considered to be at risk or threat of extinction in New Zealand are naturally uncommon (Stringer and Hitchmough, 2012). ...
Article
Full-text available
The effects of land use on soil invertebrates – an important ecosystem component – are poorly understood. We investigated land-use impacts on a comprehensive range of soil invertebrates across New Zealand, measured using DNA metabarcoding and six biodiversity metrics. Rarity and phylogenetic rarity – direct measures of the number of species or the portion of a phylogeny unique to a site – showed stronger, more consistent responses across taxa to land use than widely used metrics of species richness, effective species numbers, and phylogenetic diversity. Overall, phylogenetic rarity explained the highest proportion of land use-related variance. Rarity declined from natural forest to planted forest, grassland, and perennial cropland for most soil invertebrate taxa, demonstrating pervasive impacts of agricultural land use on soil invertebrate communities. Commonly used diversity metrics may underestimate the impacts of land use on soil invertebrates, whereas rarity provides clearer and more consistent evidence of these impacts.
... For example, rare plant and fungal species appear to be particularly sensitive to changes in environment (Avis et al., 2008;Dickie et al., 2009;Dickie and Reich, 2005;McIntyre and Lavorel, 1994), and pollinating insect losses are concentrated among rare species (Powney et al., 2019). Furthermore, most of the terrestrial invertebrate species currently considered to be at risk or threat of extinction in New Zealand are naturally uncommon (Stringer and Hitchmough, 2012). ...
Article
Full-text available
The effects of land use on soil invertebrates – an important ecosystem component – are poorly understood. We investigated land-use impacts on a comprehensive range of soil invertebrates across New Zealand, measured using DNA metabarcoding and six biodiversity metrics. Rarity and phylogenetic rarity – direct measures of the number of species or the portion of a phylogeny unique to a site – showed stronger, more consistent responses across taxa to land use than widely used metrics of species richness, effective species numbers, and phylogenetic diversity. Overall, phylogenetic rarity explained the highest proportion of land use-related variance. Rarity declined from natural forest to planted forest, grassland, and perennial cropland for most soil invertebrate taxa, demonstrating pervasive impacts of agricultural land use on soil invertebrate communities. Commonly used diversity metrics may underestimate the impacts of land use on soil invertebrates, whereas rarity provides clearer and more consistent evidence of these impacts.
... New Zealand's invertebrates are one of the least studied groups of biodiversity, with only approximately a third of the known ~13,000 terrestrial taxa assessed for their conservation status and of those, a third are categorised as 'Data Deficient' (Stringer and Hitchmough 2012). This lack of knowledge and research activity in New Zealand invertebrates makes this group one of the largest challenges in the attempt to understand what their risks and vulnerabilities are to climate change (Holwell and Andrew 2014). ...
Technical Report
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Anthropogenic (human-caused) climate change has the potential to radically alter ecosystems throughout the world, including the Auckland region. In particular, climate change poses significant challenges for ecosystem conservation and is predicted to become a major threat to biodiversity in the 21st century. Climate change risk assessments looking at species at risk have become very common overseas, however in New Zealand and particularly in Auckland, assessments of species and ecosystems have been limited. This lack of spatially explicit models of risk to ecosystems and species makes it difficult for science to inform conservation planning and land management. This report represents a first attempt at a risk and vulnerabilities assessment for Auckland’s terrestrial ecosystems and species. In it we assess some of the possible effects of climate change on Auckland’s terrestrial ecosystems, indigenous species, and invasive pest species. As part of this assessment we attempt to identify which species and ecosystems are most at risk of negative impacts from climate change. Uncertainty is the key message with respect to climate change effects on Auckland’s terrestrial species and ecosystems. Overseas research shows that climate impacts are highly variable, depending on the taxonomic group, spatial scales and time periods considered. However, the majority of these studies do indicate alarming consequences for biodiversity. The interaction between native and exotic biodiversity in their response(s) to climate change is particularly important and these inter-species relationships are very complex and varied in time and space. Almost all of Auckland’s indigenous terrestrial ecosystems are already under threat from introduced animal pests and/ or plant pests, and lowland ecosystems are also at risk from ongoing clearance and fragmentation of habitat. Therefore, potentially the most serious and pressing impacts of climate change arise through its interaction with these pre-existing threats. Despite the critical research need, there is almost no specifically-targeted research on how climate change will affect the relative ecological fitness of pest plants versus native species in any of Auckland’s indigenous ecosystems. We used spatial data on the distribution of Auckland’s 48 terrestrial ecosystem types, and published information on previously identified threats, to assess their vulnerability to six different climatic ‘risk factors’. These risk factors were: more than 10 per cent of the regional extent of an ecosystem type being vulnerable to inundation in a 100-year flood; more than 10 per cent of regional extent lying within 50m of the coast; more than 30 percent of regional extent lying within 500m of the coast; ecosystem types with a highly restricted distribution (<200ha total regional extent); ecosystems with their climatic limits in Auckland region; and ecosystems for which climate change had been identified as a specific threat. Twenty (42%) of Auckland’s 48 different indigenous ecosystem types were identified as having at least one risk factor that has the potential to make them more susceptible to the negative impact(s) of from climate change. And this rises to 28 (58%) if climate change effects increase the virulence and/ or spread of myrtle rust and kauri dieback. Some ecosystems – in particular coastal turf, oioi restiad-rushland and reedland, kauri-towai-rata montane podocarp forest, and pohutukawa-puriri broadleaved forest – are exposed to multiple risk factors. It seems reasonable to expect that ecosystems which are exposed to a higher number of risk factors are more likely to be negatively impacted by climate change. However, the relative vulnerability of ecosystems to the different threats is very difficult to assess without better data on exactly how they will respond to specific climatic drivers and stresses. Thirteen per cent of the total area of Auckland’s indigenous ecosystems lie within the inundation zone of a 100-year storm surge event; including 78 per cent of the different ecosystem types. Thirteen (27%) of the ecosystem types have more than half of their regional extent within the 100-year storm surge inundation zone. We also assessed the risk of individual species for a range of different taxa. Overseas studies have shown that a number of bat species are sensitive to climate change, and Auckland’s bats will need enhanced active management to have the resilience to survive these challenges. Auckland’s Herpetofauna are already stressed due to a range of existing threats and are also at risk of climate change impacts. New Zealand’s invertebrates are one of the least studied groups of biodiversity. This general lack of knowledge and research activity in New Zealand invertebrates means the largest challenge is attempting to understand what their risks and vulnerabilities to climate change are. Climate-related changes in Auckland may add a variety of additional stresses to birds. An international assessment of birds found as many as nine per cent of all birds are vulnerable to climate change, and this number may be higher in Auckland given the low conservation status of many of our indigenous bird species. Auckland’s seabirds are already in a delicate state with the majority of the 24 species ‘at risk’ or ‘threatened’ with extinction and the effects of climate change are challenging to assess for seabirds with limited species information available. Our best estimate of the major climate change effects most relevant to seabirds generally shows a low risk for most species in regards to sea level rise and increased pest pressure, but potentially moderate risk from increased severe weather events. The Auckland region is home to at least 63 different threatened plant species and climate change has not been identified as a specific threat for any of these species. However, it is possible that climate change could interact and exacerbate identified threats such as physical disturbance and erosion, drought, fire and/ or weed invasion. The impact of climate change on the relative competitiveness of threatened verses weedy plant species is of crucial importance, as past experience with weed impacts in New Zealand suggests that native plants are often at a competitive disadvantage when compared to introduced species; particularly in disturbed, high-light environments that may become more common due to the impact of more frequent and severe weather events such as droughts and storms. There is a complete absence of quantitative research on key climatic tipping points for Auckland’s ecosystems. This type of data is of critical importance in order to understand the risks of different climate change effects and to inform adaptive management approaches to dealing with the negative impacts of climate change in the future. Therefore, commitment to the collection of long-term ecological datasets is an absolute imperative in order to improve our understanding of how Auckland’s ecosystems will respond to the impact of climate change. However, it will probably be decades before this data is able to provide definitive answers to the many important questions about the impact(s) of climate change on Auckland’s terrestrial ecosystems and species, and how to best manage them. The potential serious impact of climate change on Auckland’s indigenous terrestrial ecosystems, the lack of information about the best response to these impacts, and the time-lag associated with collecting ecosystem data means that action(s) to manage climate change threats will need to be taken before complete information is available. A common approach to this problem is to embed research, and its evaluation, as an interactive part of ecosystem management from its initiation in a process called adaptive management or ‘learning by doing’. Taking a ‘wait and see’ approach in the absence of robust data is not an acceptable outcome.
Article
Full-text available
We evaluate evidence that reducing mammalian predators benefits threatened and declining grasshoppers in the Mackenzie Basin, New Zealand. Long-term population trends of Sigaus minutus are investigated under three control regimes: high intensity, indirect control through prey reduction and no control. We then test whether predator control benefits conservation management of Brachaspis robustus by translocating wild-caught individuals to areas of moderate versus no predator control. A significant positive trend in S. minutus counts occurred under high intensity and indirect control, suggesting that mammalian predator control is beneficial. Differences in the decline of translocated B. robustus were observed between moderate and no predator control release sites but could not be unequivocally attributed to predator densities. We recommend replicated predator control studies be undertaken to develop a predator management strategy which will enable grasshopper recovery, and investigate the potential for meso-predator release and prey-switching under regimes that target specific mammals.
Article
Full-text available
A detailed study of the moths (Lepidoptera) feeding on most species of shrubs and small trees in the genus Olearia section Divaricaster (Family Asteraceae) revealed a particularly rich fauna including eight new species. Furthermore, the plants were found to be complex ecosystems supporting lichens, mosses and algae, which in turn all support their own moth species. Forty-one species of moth, all endemic to New Zealand, were found to feed naturally on this group of plants. Of these, 20 species were found to be specialists, with 17 species confined to this host group, and the other three species oligophagous within Asteraceae. This Olearia moth guild is divided into 30 herbivores and eleven detritivores. New biological, biogeographical, seasonality and conservation information is presented in an annotated list of the 20 specialist moth species. The number of moth species on each Olearia species is noted and discussed. The richness of this fauna in different regions is presented. Several of the host plants are threatened with extinction and the conservation of these hosts is supported on entomological grounds.
Article
Full-text available
Six New Zealand Orthoptera are considered Threatened: Sigaus homerensis is Nationally Critical; Brachaspis robustus and Sigaus “yellow” are Nationally Endangered; while Sigaus “green”, Sigaus “blue” and Hemideina thoracica with a chromosome arrangement of 2n=23/24 are Nationally Vulnerable. Forty taxa are At Risk comprising one Declining (Sigaus minutus), two Recovering (Deinacrida mahoenui, Motuweta isolata), six anostostomatids that are Relict and 31 Naturally Uncommon taxa. The remaining taxa known to occur in New Zealand comprise eight taxa that are Introducedand Naturalised, 19 that are Data Deficient and 94 taxa, including 80 species, that are Not Threatened. Factors influencing the vulnerability of threatened Orthoptera in New Zealand are noted.
Article
Full-text available
A recently discovered and threatened large land snail, Powelliphanta augusta n. sp. is described from Mt Augustus on the western scarp of the Stockton Plateau, North Westland. On shell characters it is readily distinguishable from all other Powelliphanta, with narrow red spiral lines underlying irregular and variable dark reddish‐brown axial bands, a small adult size and sculptured dorsal surface. Differences in the shell morphology of P. augusta, its closest phylogenetic relative P. lignaria, and its closest geographic neighbour P. patrickensis, were investigated using principal components analysis. P. lignaria is much larger than either P. augusta or P. patrickensis, and there are significant differences in shell shape between the three taxa. Earlier mitochondrial DNA sequencing data, which supported specific status of P. augusta, is discussed. The snail's only habitat on the Mt Augustus ridgeline is described, but most has now been removed by coal mining. Snails salvaged before the destruction of their habitat have been taken into captivity or released in the wild outside their natural range.
Article
The katipo (Latrodectus katipo Powell, 1871) is one of two endemic widow spiders in New Zealand. It is a coastal sand-dune specialist and national invertebrate icon that has suffered a dramatic decline in numbers, contraction in geographical range and population fragmentation. This nationwide survey of the katipo located 26 populations in 90 dune systems sampled of a total of 127 sites visited where it could potentially be found. Details of the 90 dune systems are presented, including the number of adult female spiders found within a standard search time, degree of naturalness of the dunes and location. A survey of sites where katipo had been collected previously, based on specimens in collections, showed that they were present at only 46% of these sites. On the basis of this study, 19 key sites for katipo conservation are proposed. The katipo is threatened with extinction for a myriad of reasons: the main factors appear to be loss of habitat and declining quality of the remaining habitat. The dramatic changes to its sand-dune habitat following European settlement were due to stock grazing, disturbance and burning, and continue to this day with commercial forestry, recreational use of dune systems and other activities. The subsequent destabilisation of the dunes led to the introduction of marram grass (Ammophila arenaria), which continues to displace native sand-binding species. Marram grass can now become so dense that katipo web construction may no longer be possible. Based on this research, recommendations for the conservation of the katipo are presented. Using the accepted New Zealand invertebrate classification criteria, the katipo is assessed as a Category B threatened species, i.e. a second-priority threatened species.
Article
The rare Mercury Islands tusked weta, Motuweta isolata (Orthoptera: Anostostomatidae), a large flightless insect originally confined to 13 ha Middle Island in the Mercury Islands, New Zealand, was last seen there in January 2001. Half-grown or larger insects from a captive-breeding programme were released onto nearby Red Mercury Island (34 ♀, 16 ♂) and Double Island (65 ♀, 19 ♂) in 2000 and 2001 to reduce the potential for accidental extinction. Most (108) were released under individual artificial cover objects (ACOs)-clear Perspex discs under plastic plant-pot saucers- and 26 were placed in artificial holes in soil. Usually <10% were found again under ACOs for up to 18 months including 7.5 months as adults. Adults, found in 2005 and 2006, were 1st to 3rd generation island-bred weta (lifespan 1.7-3.2 years). Ongoing monitoring is planned to confirm long-term success. Inbreeding depression is likely so supplementation from Middle Island is required but they may be extinct there. Scraping the soil to expose weta in underground galleries was the best monitoring method. Few were found by searching with lights at night but adults could be located by following other adults equipped with harmonic radar transponders or micro-transmitters.
Article
Pre-human New Zealand had some unusual feeding guilds of birds (e.g. the herbivorous moa fauna), thought to have developed as a result of the absence of a 'normal' mammal fauna. Insectivorous birds, on the other hand, are an integral part of all the world's ecosystems, regardless of the presence or absence of mammals. While it is acknowledged the overall predation impact from birds in New Zealand is unlikely to have differed greatly from elsewhere, the low impact of mammalian insectivores (apart from microbats), coupled with the presence of a specialised avian feeding guild that concentrated on ground-active prey, might have exerted certain unique selection pressures. Do New Zealand invertebrates reflect this? It would be necessary to compare the New Zealand invertebrate fauna with that of mammal-dominated lands in greater detail than is available today before we could assert whether any unique anti-predator characteristics have evolved. Knowledge of the insects that succumbed to extinction when mammals invaded New Zealand should provide clues to avian-adapted features that might have rendered them particularly vulnerable to introduced rodents. Predation by kiwi (Apteryx spp.), an extraordinarily mammal-like nocturnal bird, may to some extent have prepared the invertebrate fauna for the arrival of small mammals.
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The exciting prospect of discovering New Zealand's untapped insect fauna was incentive enough for the early collectors of natural history specimens. Investigation of the fauna began with Joseph Banks on the Endeavour in 1769 and was continued by a devoted group of eighteenth century naturalists who occasionally voiced their concern at the rate of destruction and modification of the indigenous ecosystems as European Imperialism fuelled the development of agriculture for the economic base of the new colony. Although records are sparse, at least two early entomologists expressed their opinions in the early twentieth century. G.V. Hudson aired his concern about the dwindling numbers of native insects in a Presidential Address to the Wellington Philosophical Society (Hudson 1900) and later recollected that: 'every summer throughout the eighties great fires raged on the hills around Wellington and the air was thick with smoke for days together'. The concept of preserving native fauna was then in the minds of very few and those who protested at the reckless destruction of the forest were regarded as cranks. (Hudson 1950, pp. 161-162). Back 'home' in Britain, another entomologist, Commander James J. Walker (1921) in his presidential address to the Entomological Society of London, also drew attention to the potential plight of New Zealand's insect fauna when, after discussing the rich and endemic attributes of the flora that were being lost both to agriculture and weed invasion, he said that 'the disastrous effect on all forms of life of this destruction and replacement (necessitate) immediate and thorough research in what still remains of the endemic animal life of New Zealand, before many of its most interesting forms are lost forever.' However, nothing was done and no consideration was given to the need to conserve invertebrates. Instead, it was the ornithologists who paved the way for the first conservation ethic as they focussed on New Zealand's unusual and dominant bird life. This began with the influential politician, Thomas Potts, who raised awareness for conservation in the 1850s and was eventually in part responsible for the declaration of island reserves for Resolution, Secretary, Little Barrier, and Kapiti islands, gazetted between 1891 and 1897 (Young 2004). Conservation action on behalf of invertebrates did not occur despite increasing knowledge of their uniqueness and significance in understanding southern biogeography issues (Gibbs 2006). © 2012 Springer Science+Business Media Dordrecht. All rights are reserved.
Article
Records of translocations are incomplete or non-existent for many taxa in New Zealand, yet such records are essential for understanding biogeography and providing context for ecological restoration. Here we summarise all known translocations of native bats, reptiles, amphibians and terrestrial invertebrates, based on written records and first-hand verbal accounts. This report lists details of 183 translocations: 2 with bats, 86 with reptiles, 10 with amphibians and 85 with invertebrates (including 44 molluscs, 39 insects, 1 centipede and 1 spider). We acknowledge the likelihood that there are additional translocations we are unaware of and recommend improvements for recording future translocation events and their outcomes in New Zealand by following the Standard Operating Procedure for translocations that is being developed by DOC wherever possible. We also recommend that consideration be given to the minimum number of individuals for release, to limit loss of genetic variation.
Article
The conservation status of Gastropoda, excluding taxa in the genus Powelliphanta and those in freshwater, are listed. Forty-seven taxa are Threatened and comprise 28 that are Nationally Critical, 11 that are Nationally Endangered and eight that are Nationally Vulnerable. A further 259 are At Risk, comprising five Declining, 50 Relict and 204 Naturally Uncommon taxa. In addition, 138 taxa are Data Deficient; no taxa are listed as Extinct. Factors that contribute towards New Zealand landsnails becoming threatened are discussed.
Article
Rhytidarex buddlei is a giant terrestrial landsnail endemic to South West Island in the Three Kings group. The distribution of old, empty shells indicates that this species was widespread over at least the northern half of the island in the 19th and possibly also early 20th centuries. However, at the end of the 20th century its geographic distribution was restricted to c. 0.03 ha on steep western slopes, and the estimated total population contained no more than 100 and probably fewer than 50 individuals. This decline correlated with, and probably resulted from, ecological changes associated with the widespread establishment of pukanui (Meryta sinclairii) forest on South West Island within the last 200 years. The modern snail population was confined to flax (Phormium aff. tenax) thickets and patches of mixed flax‐broadleaved shrubland within an overall mosaic of shrubland and low pukanui forest. The very small size and narrow geographic‐ecological distribution of the remaining population of R. buddlei place it at high risk of extinction from stochastic events, adverse ecological changes, and anthropic disturbances. Historical evidence of decline suggests that the species is unlikely to recover on its own without some form of external assistance. Potential conservation measures are discussed, and a preferred option is identified.