RECORDS OF THE WESTERN AUSTRALIAN MUSEUM
34
055–114 (2019) DOI: 10.18195/issn.0312-3162.34(2).2019.055-114
Endemic species of Christmas Island,
Indian Ocean
D.J. James1, P.T. Green2, W.F. Humphreys3,4 and J.C.Z. Woinarski5
1
73 Pozieres Ave, Milperra, New South Wales 2214, Australia.
2
Department of Ecology, Environment and Evolution, La Trobe University, Melbourne,
Victoria 3083, Australia.
3
Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia.
4
School of Biological Sciences, The University of Western Australia, 35 Stirling Highway, Crawley,
Western Australia 6009, Australia.
5
NESP Threatened Species Recovery Hub, Charles Darwin University, Casuarina,
Northern Territory 0909, Australia,
Corresponding author: john.woinarski@cdu.edu.au
ABSTRACT – Many oceanic islands have high levels of endemism, but also high rates of extinction,
such that island species constitute a markedly disproportionate share of the world’s extinctions.
One important foundation for the conservation of biodiversity on islands is an inventory of endemic
species. In the absence of a comprehensive inventory, conservation effort often defaults to a focus
on the better-known and more conspicuous species (typically mammals and birds). Although this
component of island biota often needs such conservation attention, such focus may mean that less
conspicuous endemic species (especially invertebrates) are neglected and suffer high rates of loss.
In this paper, we review the available literature and online resources to compile a list of endemic
species that is as comprehensive as possible for the 137 km2 oceanic Christmas Island, an Australian
territory in the north-eastern Indian Ocean. This objective is helped by impressive biodiversity
inventories made within a decade of the island’s first human settlement (1888) that provide a
reasonable baseline from which to measure the changes associated with the island’s colonisation
and development. However, there are some notable challenges in compiling this inventory: the spate
of surveys that preceded and immediately followed the island’s settlement has not been matched
subsequently; many groups have not been sampled, or sampled only superficially; the taxonomic
fate of some of the species initially described from the island is opaque; some endemic taxa are
of contested taxonomic rank; and demonstrating endemicity is difficult given that there has been
relatively little sampling in the nearest lands (Java and nearby islands, about 350 km distant from
Christmas Island).
We conclude that at least 253 species are endemic to Christmas Island (including 17 vascular
plants, 27 molluscs, 15 crustaceans, 150 insects and 21 vertebrates). There has been a high rate of
extinction of the island’s endemic mammal and reptile faunas, with at least six of the 10 endemic
species now extinct or extinct in the wild. In the last decade, an endemic mammal and an endemic
reptile species became extinct, and two endemic reptile species became extinct in the wild. Given
the array of introduced species and other environmental disruptions now present on the island, it
is highly plausible that many endemic species in less conspicuous or charismatic groups are now
imperilled or already extinct; indeed, we conclude that more than 50 endemic species have not been
reported for >100 years. Hence, the recognised number of extinct and of threatened species on this
island is likely to be severely under-estimated. Although most of the endemic vertebrate species are
listed as threatened (or extinct), only one of the c. 200 endemic invertebrate species is formally listed
as threatened. This lack of listing is likely to severely understate the conservation plight of many
species, and most would merit recognition as threatened.
KEYWORDS: conservation, endemism, extinction, fungi, invertebrates, island, lichen, plants, vertebrates
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
56
INTRODUCTION
Oceanic islands support a disproportionately large
share of the world’s biodiversity relative to their total
area (Kier et al. 2009; Tershy et al. 2015). However,
island-endemic species also make up a disproportionate
share of the world’s modern extinctions, including the
majority of recent extinctions in many taxonomic groups
(Blackburn et al. 2004; Brooke et al. 2007; Sax and
Gaines, 2008; Loehle and Eschenbach, 2012; Tershy
et al. 2015; Doherty et al. 2016; McCreless et al. 2016;
Woinarski et al. 2018). Hence, oceanic islands are a
global priority for conservation attention.
Conservation management for any island is likely to
be most effective and appropriately directed if it rests
upon a robust knowledge base of the island’s biodiversity
attributes, such as the number and identity of its
endemic species; and that its most imperilled species are
recognised as such. Where such information is lacking
or limited, there may be a high risk of species declining
or going extinct before managers are aware of the
species or aware of the need for remedial management
action (Régnier et al. 2009; Régnier et al. 2015a; Régnier
et al. 2015b). Such risks may be particularly severe for
endemic species that are relatively inconspicuous and
uncharismatic and those in relatively poorly-known
taxonomic groups, such as most invertebrates (DinizFilho et al. 2010; Diniz-Filho et al. 2013): in contrast,
island birds and mammals are generally well-known and
the focus of most conservation attention.
Here we consider the biodiversity of the 137 km 2
Christmas Island, an Australian external territory in
the tropical eastern Indian Ocean, and its conservation
prioritisation. There is a marked taxonomic bias in the
global and national (Australian, under the Environment
Protection and Biodiversity Conservation Act) listing
of threatened species (Walsh et al. 2012), which is
particularly pronounced in the case of Christmas
Island. Of its 14 endemic terrestrial vertebrate species,
twelve are listed as threatened or extinct (Table 1),
and a further five endemic subspecies of terrestrial
vertebrates are also listed as threatened at the national
level. For plants, one of the island’s 16 endemic species
is listed as threatened, along with one endemic variety
and two local populations of plant species with much
more widespread distributions. In contrast, only one of
Christmas Island’s c. 200 endemic invertebrate species
is listed as threatened globally and none are listed
nationally. In part, this disparity reflects contrasts in the
amount of knowledge available for different taxonomic
groups, but it is also a consequence of a pervasive
tendency to highlight the conservation concerns of
more charismatic taxonomic groups at the expense of
those with less public appeal (Cardoso et al. 2011a;
Cardoso et al. 2011b; Walsh et al. 2012). In the case
of Christmas Island biodiversity, listing as threatened
under Australian legislation provides some notable
conservation benefits: listed species tend to be regularly
monitored, such that information on population trends
is routinely available; the potential impacts upon them
of proposed developments must be considered through
legislated environmental impact assessment processes;
recognised threats to them are the subject of management
investment and control; and they are explicitly considered
in spatial land-use planning exercises (Butz 2004; Hill
2004a; Hill 2004b; Schulz 2004; Schulz and Lumsden
2004; Cogger 2006). In contrast, species that may
be equally imperilled but are not formally listed as
threatened are often neglected, compounding a lack
of knowledge that will constrain their likelihood of
being listed as threatened and perpetuating a lack of
conservation attention.
We aim to provide an inventory of species endemic
to Christmas Island, as a basis for more comprehensive
consideration of its biodiversity conservation values and
needs. Such an inventory is of interest and importance
for several reasons. First, and unusually, a substantial
(albeit not exhaustive) inventory of the plants and
animals of the island was compiled at about the time
of the island’s initial settlement, from the late 1880s
(Boulenger 1887, 1889; Butler 1887, 1889; Dendy 1887;
Pocock 1887, 1900; Sharpe 1887, 1900; Thomas 1887,
1889; Waterhouse 1887; Gahan 1889; Lister 1889; Smith
1889; Ridley 1891, 1906a,b; Waterhouse et al. 1900).
Part of the explicit purpose of this early collection work
was to provide an exemplary baseline against which
changes in the island’s biota (due to settlement and human
influence) could subsequently be assessed (Andrews
1900c; Andrews 1909): ‘It has not hitherto been possible
to watch carefully the immediate effects produced by
the immigration of civilized man – and the animals and
plants which follow in his wake – upon the physical
conditions and upon the indigenous fauna and flora of
an isolated oceanic island. I hope to arrange that this
shall be done in the case of Christmas Island’ (Murray
1900). To some extent, this current paper facilitates some
such assessment of the post-settlement fate of the island’s
endemic biota. Regrettably, calamity befell the next major
collection, by Carl Gibson-Hill collected over a 2-year
period from 1938–40: ‘the greater part of the material
obtained, covering the majority of the insect groups, was
destroyed, unidentified, when the Kuala Lumpur Museum
was bombed in 1945’ (Gibson-Hill 1949b).
Second, from the time of the earliest biological
sampling, the distinctiveness and endemicity of the
island’s biota was recognised. For example, the first
visiting naturalist, J.J. Lister in 1887, reported that ‘the
most striking factor is the peculiarity of the fauna’ (Lister
1889).
Third, the island has a highly unusual ecological
context, largely determined by extremely high densities
of a single invertebrate species, the red crab Gecarcoidea
natalis (O’Dowd and Lake 1991; Green 1997; Green et al.
1997, 1999).
However, since the 1980s, the island’s ecology has
been subverted through ‘invasional meltdown’ (Green
et al. 2001, 2011) due to the extensive establishment
of supercolonies of the invasive yellow crazy ant
Anoplolepis gracilipes, which have encompassed up to
ENDEMIC SPECIES OF CHRISTMAS ISLAND
25–30% of the island at times (Abbott 2006). There have
been subsequent episodes of control with broad-scale
application of insecticide followed by rebounds in colony
extent (Green and O’Dowd 2009). Such disruption by
the yellow crazy ants may be expected to have caused
appreciable detriment to many endemic species. In
response to the threat posed by crazy ants, managers
implemented broad-scale baiting with the insecticide
Fipronil. This has had serious non-target impacts
elsewhere (Peveling et al. 2003; Gibbons et al. 2015; Van
der Sluijs et al. 2015). However, baiting on Christmas
Island has been implemented strategically with stringent
and substantial measures to mitigate non-target impacts
customised for the local conditions (e.g. see EPBC Act
Referral Notices and Decisions 2002/722, 2009/5016 and
2012/6438 covering the three aerial baiting operations
in 2002, 2009 and 2012 respectively; http://epbcnotices.
environment.gov.au/referralslist/). Three targeted studies
(Weeks and McColl 2011; Weeks 2013; Stork et al. 2014)
found no substantial evidence of non-target impacts on
invertebrates, although they did not assess impacts on
most endemic invertebrates specifically, at least in part
because there was no comprehensive listing of these then
available.
Many other plant and animal species – including
invasive species known to have caused declines or
extinctions in a wide range of island-endemic species
elsewhere (e.g. black rat Rattus rattus, house cat Felis
catus, giant centipede Scolopendra subspinipes, wolf
snake Lycodon capucinus, giant African land snail
Achaetina fulica) (Amori and Clout 2003; Doherty et
al. 2016) – have also been introduced, deliberately or
inadvertently, to the island and many of these are likely to
have had detrimental impacts on at least some Christmas
Island endemic species. Furthermore, since its settlement,
the island has had an almost continuous history of
phosphate mining (with about 25% of the island’s original
vegetation cleared for mining), and mining operations
continue. Frequent proposals to extend mining operations
require ongoing assessments of the likely impact upon
biodiversity (particularly listed threatened species) from
potential further forest clearing and mining operations
(Frydenberg 2018). To properly assess the potential
immediate and cumulative impacts of such proposed
habitat loss it is necessary to be as explicit as possible
about all the biodiversity assets at risk, and their status
and their locations. Mining and other developments have
also affected marine environments (such as damage to
coral reefs for shipping infrastructure and from ships,
spills of phosphate into marine environments, urban
storm water run-off and oil spills), but these impacts are
generally little documented (Hobbs et al. 2014a).
As a consequence of these detrimental factors, there
has been a very high rate of loss of endemic vertebrates,
including the extinction of at least three and probably four
of the island’s five endemic mammal species (Wyatt et al.
2008; Woinarski et al. 2017) and the recent (since 2009)
extinction (or extinction in the wild) of three of its four
endemic lizard species (Smith et al. 2012; Andrew et al.
57
2018). This gives reasonable grounds for concern that
endemic species in less well-known groups might have
become extinct, or are now highly imperilled, with such
loss and decline going unnoticed.
The primary objective of this study is to compile a
list of species endemic to Christmas Island. This is not
a straightforward exercise, for several reasons. First,
there is no comprehensive list of species that have been
recorded from the island, from which endemic, native
but non-endemic, and introduced species can readily be
categorised: an exemplary case where this has been done
is for invertebrates of Lord Howe Island (Cassis et al.
2003). The lack of such an inventory for Christmas Island
also renders it difficult to compare proportional levels of
endemicity among different taxonomic groups. Second,
although some species have been recorded only from
Christmas Island, it remains possible that some also exist
in less well-sampled areas in the nearest land masses
(e.g. Java, at c. 350 km distant) or more distantly. It is
therefore possible that an inventory of species currently
known only from Christmas Island may over-estimate
the real extent of endemicity, but this is a general
qualification in many such biogeographical analyses
(Whittaker et al. 2008). Third, taxonomic boundaries are
poorly resolved for many Christmas Island taxa. Highly
conservative (mostly historical) taxonomy has seemingly
treated several taxa as subspecies of more widely-ranging
species rather than endemic species (James and McAllan
2014). In general, where such Christmas Island taxa of
contested or indeterminate rank have been scrutinised
using modern genetic or other systematic approaches,
they have been shown to be valid species (Norman et
al. 1998; Ng and Davie 2012; Orchard 2012; Eldridge et
al. 2014; Rheindt et al. 2017). However, there is at least
one contrary case where presumed endemic species have
been shown to be not specifically distinct (Ohlsen et al.
2015). Fourth, many Christmas Island species have been
recorded only in samples taken around the beginning
of the twentieth century, and for some of these species
there has been little subsequent taxonomic appraisal, so
it is challenging to determine whether these described
species remain valid, or whether taxa then considered
to be referrable to more wide-ranging species may in
fact be endemic. Fifth, although there have been some
notable surveys of Christmas Island for many taxonomic
groups, there are many groups that are unsampled or
little sampled: this is likely to cause a potentially marked
under-estimation in our tally of endemic species. Sixth,
there has been some confusion in the literature relating to
endemic species on this island, with attribution of some
of its endemic species to a once-similarly named island
(now Kiritimati), in the republic of Kiribati in the Pacific
(Alcover et al. 1998). Given these caveats and constraints,
we recognise that the list of endemic species developed
here is an interim compilation, and we would expect and
hope that it will be modified with ongoing research.
Endemicity is also a somewhat nuanced concept.
We include here as endemic: (i) some species that
breed only on Christmas Island, but disperse widely
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
58
elsewhere in the non-breeding season (i.e. breeding
endemic), such as Christmas Island frigatebird Fregata
andrewsi (James and McAllan 2014; Tirtaningtyas and
Hennicke 2015); (ii) one species that has been extirpated
from all parts of its formerly extensive breeding range
other than Christmas Island (i.e. neoendemic), being
Abbott’s booby Papasula abbotti (Morris-Pocock et
al. 2012); and (iii) species originally occurring only
on Christmas Island but from which populations have
been translocated elsewhere (i.e. original endemic),
for example, Christmas Island white-eye Zosterops
natalis (Woinarski et al. 2014). With respect to this last
category, family and business connections, resource
needs and logistical tractability stimulated importation
from Christmas Island to the Cocos (Keeling) islands
of soil and timber, inadvertently with associated
invertebrates in the late nineteenth and early twentieth
centuries (Gibson-Hill 1950; Tweedie 1950), as well as
some deliberate introductions of bird species (GibsonHill 1949a). Although we note some endemic subspecies
in this account, our focus is on endemic species, largely
because there is very uneven treatment of subspecies
across taxonomic groups.
This study is of intrinsic interest for biodiversity
conservation on one island. But it is likely that at
least some of the findings, particularly in relation
to biases in threatened species’ listing, will have
generalisations applicable to many other islands.
Furthermore, the results from this study are likely to
contribute to ongoing analyses of variation in the extent
of endemism and its loss, and the factors associated
with such variation, across sets of islands (Adler 1994),
with such analyses currently constrained for some
taxonomic groups by the relatively few islands with such
information available.
METHODS
STUDY AREA
Christmas Island is a solitary sea-mount island, of
volcanic origin (Hoernle et al. 2011), in the tropical
eastern Indian Ocean (at 10°25'S and 105°43'E) (Figure 1).
It is the sole emergent peak within a large chain of
volcanoes that formed about 80 million years ago, with
subsequent volcanic reactivation 40–35 million years
ago (Trueman 1965), and possibly also between five and
three million years ago (Borissova 1994). It has likely
been continuously emergent for at least three million,
and possibly up to ten million, years (Grimes 2001;
Humphreys and Eberhard 2001; Namiotko et al. 2004)
2004), but with the most recent analysis indicating
about 5.7–4.5 million years (Ali and Aitchison 2020).
Its basaltic core is now mostly capped by a sequence of
Tertiary limestones up to 250 metres thick. It is a rugged
island, with a highest elevation of 361 metres. It has no
outlying islets, and is distant in all directions from any
other land mass. The main vegetation is tall rainforest.
Freshwater environments are limited but diverse, and
there is a complex mix of subterranean environments
(Grimes 2001; Humphreys 2014). About one-quarter of
the island has now been cleared as part of a phosphate
mining venture that has operated almost continuously
since the island’s settlement in 1887. A national park was
established in 1980, and now encompasses 63% of the
island (Director of National Parks 2014).
Biological inventory of the island has been episodic
and incomplete. Following the initial spate of collections
around the first decade of the island’s settlement (see
references in the Introduction), there was little sampling
until some notable collecting in the 1930s (Chasen
1933a,b; Gibson-Hill 1947a,b,c,d,e,f,g), followed by
another hiatus until more sampling for some taxonomic
groups in the decades since the 1970s. The intermittent
and variable nature of sampling for most taxonomic
groups largely precludes the use of species’ accumulation
curves to assess the extent to which sampling is likely to
have detected most or all species present.
ASSESSING ENDEMICITY
By definition, species (and other taxa) endemic to
Christmas Island must occur there and nowhere else,
other than the nuanced exceptions discussed above.
Any species described from elsewhere (i.e. the type
locality is not Christmas Island) is ineligible. Thus to
compile a list of presumed or known endemic species,
involves (mostly) the following steps: (1) documenting all
species originally described from Christmas Island; (2a)
determining if these species are valid (i.e. that they have
not been lumped or dissolved into other species; and (2b)
determining that they (and their synonyms) do not occur
beyond Christmas Island. Of course, it is not necessarily
so simple, because some species collected from Christmas
Island may have been considered conspecific with species
originally described from elsewhere, but subsequent
taxonomic review has shown (or may in future show)
that such assumed conspecificity is wrong. Furthermore,
our process will detect only those species that have been
collected on, and described from, Christmas Island. It is
likely that there are many endemic species that have never
been collected, and many that have been collected but not
yet described.
Step 1: For several taxonomic groups, there have been
recent comprehensive accounts of species recorded
from Christmas Island with these accounts explicitly
noting the number and identity of endemic species: these
inventories include birds (James and McAllan, 2014), ants
(Framenau and Thomas 2008), vascular plants (Claussen
2005), lichens (McCarthy 2018), molluscs (Tan and Low
2014), crabs (Orchard 2012), fish (Hobbs et al. 2014b),
scale insects (Neumann et al. 2013, 2016), centipedes
(Waldock and Lewis 2014) and subterranean fauna
(Humphreys 2014). Unless we found compelling evidence
to the contrary, or there have been subsequent taxonomic
assessments, we used such recent reviews as the basis for
recognising endemic species in these groups.
For other taxonomic groups, we used as the first step
in the listing of endemic species a collation by James
(2005) and James and Milly (2006) of species for which
the type specimen was collected from Christmas Island
and for which they found no subsequent records from
ENDEMIC SPECIES OF CHRISTMAS ISLAND
59
A
B
FIGURE 1
Location maps for Christmas Island. A) general location; B) map showing place names.
elsewhere, though we recognise that these compilations
contained errors and omissions. The 2005–06 collations
indicated that 225 species were nominally endemic to
Christmas Island. We searched available taxonomic
literature for new records of Christmas Island species
published since these 2005–06 compilations as well as
for additional species that may have been overlooked in
2005–06. We also extracted all records from the Atlas of
Living Australia (‘ALA’) spatial database (see below), to
cross-check for additional species that may be endemic to
Christmas Island that were not included in the 2005–06
compilation.
Steps 2a and 2b: Making use of substantial recent
increases in access to biodiversity information on-line,
we searched three ‘primary’ databases for information
on all of these nominally endemic species, to assess
whether they are still recognised as valid species with
spatially-explicit records only from Christmas Island: the
Global Biodiversity Information Facility (https://www.
gbif.org/ ; hereafter cited as ‘GBIF’), the ALA (https://
www.ala.org.au/) and the Australian Fauna Directory
(https://biodiversity.org.au/afd/home ; hereafter ‘AFD’).
As scholarly as these databases may be, they do contain
omissions and errors, and sometimes contradict one
60
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
another, so when they provided inadequate information
we investigated as deeply as possible. We searched other
relevant accessible taxonomic data bases (e.g. Mesibov
2010; Bieler et al. 2018; McCarthy 2018; Orthoptera
Species File, 2018; Sierwald and Spelda 2018; Van Soest
et al. 2018), used online search-engines and investigated
the primary literature. We cross-checked all relevant
animal taxa against the 31 published volumes of the
‘Zoological Catalogue of Australia’ (1983 to 2006: AGPS
Canberra pre-1994, and ABRS and CSIRO Publishing
Canberra from 1994), but they are cited specifically only
when they provide additional information to their online
successor, AFD. GIS-based assessments were impractical,
as most of the records from early collections of Christmas
Island biota (often the only records of these species)
lacked precise locational information, so these records are
typically not mapped in national or global databases.
As a further line of inquiry, we also searched through
references in Google Scholar using combinations of
‘Christmas Island’ and family names.
Based on results from such searches, we allocated
species to three categories: (i) confirmed endemic (Table
2), (ii) cited as endemic or probably so by collectors, but
not yet described (Table 3), or (iii) previously claimed as
endemic but now shown not to be (Table 4). Omission
proved to be the most challenging issue. Under the
taxonomic codes, newly described species are valid (if
they meet naming rules) until a reviewer shows them
to be the same as an earlier described species. Some
species described from Christmas Island appear never
to have been mentioned again, at least in the sources we
found. Whilst such cases have never been invalidated by
taxonomic revision, it is also difficult to state confidently
that they are valid endemic species. Furthermore,
taxonomy is appropriately not fixed, and future revisions
will undoubtedly change taxonomic positions, as they
have in the past.
CONSERVATION STATUS OF ENDEMIC SPECIES
We noted and tabulated the global (i.e. IUCN Red List)
and Australian (i.e. under the Environment Protection and
Biodiversity Conservation Act, 1999) conservation status
for all endemic species, as at February 2019 (Table 1).
As a potential indicator of current conservation status,
we also attempted to identify the most recent record
for every endemic species, and we list that date when
it was more than 50 years ago. For this we have drawn
on the published literature, the online databases and
several invertebrate surveys that are informally published
(Campbell 1968; CSIRO Division of Entomology 1990;
Surman 2004; James 2005, 2007; Framenau and Waldock
2006; James and Milly 2006; Kessner 2006; C. Pink,
pers. comm.). This assessment also has some notable
interpretational caveats, as the year of collection is often
not given for records in the biodiversity databases we
searched, and a lack of recent records for a taxon may
simply be due to a lack of appropriate survey. Where
possible, we also report on any other description of
the abundance or distribution of endemic species on
Christmas Island.
RESULTS
In the sections that follow, we document the inventory
and taxonomic material relevant to endemic species,
by major taxonomic groups. For some groups we could
locate no relevant information.
Kingdom EUBACTERIA
Phylum FIRMICUTES
CLASS BACILLI
A major disease threat emerged recently to the captive
breeding program for two endemic Christmas Island
reptile species. This invariably fatal disease was found
to be due to infection with a previously unknown
Enterococcus species, although the report documenting
its occurrence did not formally describe it (Rose et al.
2017). This unusual bacterium was recorded not only
from individuals in the captive breeding program for
threatened endemic lizards, but also in individuals of
Christmas Island populations of introduced lizards
(Rose et al. 2017). Although the paper documenting
its existence noted that there were no records of it
from elsewhere, the current available information is
insufficient to categorise it as native to Christmas Island,
let alone endemic.
Kingdom PROTISTA
We acknowledge that the ‘Kingdom Protista’ is a
diverse assemblage of eukaryotic organisms that are
not plants, animals or fungi, but that are not all closely
related (Keeling et al. 2009). However, it is beyond our
scope to clarify such issues.
Phylum APICOMPLEXA
CLASS ACONOIDASIDA
A recent study has examined the blood parasites of
some seabirds on Christmas Island and described a
new species, Haemoproteus valkiunasi (Haemosporida:
Haemoproteidae [although with family name given
as Plasmodiidae in ALA]) from samples taken from
Christmas Island frigatebirds Fregata andrewsi
(Merino et al. 2012). All of the limited known records
of this parasite species are from this endemic frigatebird
species, so as currently known this parasite is also an
endemic species.
Kingdom PROTOZOA
Phylum MYCETOZOA
CLASS MYXOMYCETES
These are fungus-like organisms known as plasmodial
or acellular slime moulds. Ten species in four genera
across four orders have been recorded from Christmas
Island. According to GBIF all ten species have global
distributions and so none are regarded as endemic.
C.W. Andrews collected Arcyria obvelata (as A. flava),
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Lycogala epidendrum (as L. miniatum) and Stemonitis
splendens in 1897–98, and H.D. Yorkston collected the
other species in 1986–87.
CLASS PROTOSTELIOMYCETES
These are fungus-like organisms known as slime
moulds, and regarded as primitive within that group.
They occur mostly on bark and rotting wood. Two
species have been recorded from Christmas Island,
neither of which is regarded as endemic. Ceratiomyxa
fruticulosa is globally distributed (GBIF), but
C. sphaerosperma has a markedly disjunct distribution,
with all known records from Africa and the Americas,
except for a single Asian occurrence on Christmas
Island. Both species were collected by H.D. Yorkston in
1986–87.
Kingdom FUNGI
The earliest collections of fungi on Christmas Island
were made by J.J. Lister in 1887 (three species identified
by M.C. Cooke in Hemsley 1890), followed by C.W.
Andrews in 1897–98 (22 species identified by V. H.
Blackman and A.L. Lister in Andrews 1900), and by
H.N. Ridley in 1904 (50 species identified by M.G.
Massee in Ridley 1906). The fate of these collections is
largely unknown. Only five collections (three species) by
Andrews and possibly one of Ridley’s collections (one
species) are discoverable through the Kew Fungarium
online catalogue.
There followed a collecting hiatus until D.A.
Powell, a naturalist working for the British Phosphate
Commission, collected around a dozen species in 1968
(Reid 1969), and Powell and his assistant K.C. H’ng
continued to make collections in the 1980s and early
1990s. Powell sent most of his material to Kew (K), but
there are only 15 Powell/Powell and H’ng collections
discoverable through the Kew Fungarium online
catalogue, and several in Canberra (CANB) and Perth
(PERTH).
Shivas and Hilton (1990) compiled a checklist of the
fungi of Christmas Island, based on a 1-week survey in
1986, further collections by H.D. Yorkston in 1986–87,
and collation of the limited previous collections. The
compilation comprised c. 120 taxa, including 58 not
previously recorded: 97 of the taxa were determined to
species level (of which only three were described from
Christmas Island), and 24 were determined only to
genus level.
Following these collections, H. Lepp of the Australian
National Herbarium made extensive collections in 2000
which were lodged mostly at CANB, followed by N.L.
Bougher with the Western Australian Herbarium in
2016, who lodged most of his specimens at PERTH. The
tally of non-lichenised fungi known from the island now
stands at 132 species.
Three species from the early collecting period
1888–1904 were described as new at the time. The
first was the earthstar Geastrum andrewsii (Blackman
61
1900). The only collection of this species was by
Andrews in 1897–98, from which the holotype was
designated. It has not been collected since, although
a photograph by Peter Goh for the month of March in
the 1991 Christmas Island Natural History Association
calendar may be this species. It is still regarded as
endemic to the island (Shivas and Hilton 1990), but is
not listed in ALA and is listed as ‘doubtful’ in GBIF.
Massee described two new species from Ridley’s
1904 collections (Ridley 1906a). The type of Poria
chlorina is lodged at the New York Botanical Garden
Herbarium, but this taxon has since been synonymised
with the much more widespread Ceriporia mellea
(Berk. and Broome) Ryvarden 1978. Ridley (1906a)
also listed ‘a small white agaric’ Favolus albidus
Massee as a new species, but Shivas and Hilton (1990)
noted that this species was actually described in 1902
by Massee from specimens collected in Thailand, and
the placement of an agaric in the genus Favolus is
problematic anyway.
Two species in the earthstar genus Radiigera were
described by Reid (1986) from material collected on
Christmas Island in 1983 and 1984, and thought then
to be known only from there. However, Domínguez de
Toledo and Castellano (1996) examined the holotype
of Radiigera asperata D.A. Reid and determined
it to be an immature, unopened Geastrum species.
Given the only Geastrum species known from the
island is the endemic G. andrewsii, in all likelihood
it was that species in which case it has been collected
several decades after Andrews’ original 1897–98
collection. Radiigera asperata was never recorded as
occurring elsewhere, and so the name can no longer be
applied. Domínguez de Toledo and Castellano (1996)
also regarded Radiigera termitariicola D.A. Reid as
synonymous with Phialastrum barbatum (Dissing and
Lange) Sunhede 1989, itself originally described as
Geastrum barbatum Dissing and Lange (1962) from the
Congo. Based on this synonymy, R. termitariicola can
no longer be regarded as endemic to Christmas Island.
In addition to Geastrum andrewsii, there appears to be
only one other non-lichen fungus endemic to Christmas
Island. Aecidium alchorneae-rugosae Gjaerum and D.A.
Reid 1983 (Pucciniaceae) is common all over the island
on its host Alchornea rugosa (Euphorbicaeae), especially
in drier areas (PG pers. obs.). Neither of the two
described endemic fungi species is listed as threatened
nationally or globally.
LICHENS
McCarthy (2018) collated checklists of lichens for all
of Australia’s main oceanic islands. He listed 101 taxa
(100 species) for Christmas Island, and stated that four
of these species are endemic: Lithothelium quiescens,
Strigula elixii, Strigula natalis and Trichothelium
oceanicum. These names are recognised in GBIF and all
(of the few) locational records in that database for these
four species are from Christmas Island. Lithothelium
quiescens is corticolous and Trichothelium oceanicum
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
62
is follicolous, and both are widespread and common
(McCarthy 2001a,c). Strigula elixii and S. natalis are
saxicolous and scarce, with the latter known from one
site only. None of the four described endemic species is
listed as threatened nationally or globally.
The list of species from Christmas Island (McCarthy
2018) is mostly based on results from a recent (2000)
survey (McCarthy, 2001a,b,c; McCarthy and Elix, 2002).
There are few historical records and hence no indication
of changes in status. Although the 2000 sampling study
was extensive, McCarthy and Elix (2002) considered
that the known inventory of 100 species was a major
under-estimate of the actual number of species present:
the tally ‘does not include least 10 unidentified species
each of Graphidaceae, Thelotremataceae, Pyrenulaceae
and other saxicolous and corticolous pyrenolichens,
as well as numerous sterile and apothecial crusts on
limestone, bark, soil and leaves. The addition of 50 or
more taxa that were not collected during our visit (a
conservative estimate considering the forest canopy
flora and much of the southern half of the island remain
largely unexplored) would bring the total close to 250’
(McCarthy and Elix 2002).
ALGAE
Gepp and Gepp (1905) listed 21 species and one
indeterminate species of marine algae from a collection
made by H.N. Ridley in October 1904 from flying Fish
Cove and Waterfall Cove. This included one new species
of red algae, Halymenia polyclada, which remains a
valid species, but has since been recorded from the
Seychelles (GBIF; Table 4). Other than being mentioned
again in Ridley (1906a), we have found no further
records of this species from Christmas Island, but it is
not listed as threatened nationally or globally.
Kingdom PLANTAE
Phylum BRYOPHYTA
Gepp and Gepp (1905) listed 15 species of mosses
(Class Bryopsida) from collections made by C.W.
Andrews in 1897–98 and H.N. Ridley in October 1904.
These are mostly widespread species in Southeast Asia.
However, their account contained two new species
described by Fleischer as Ectropothecium micronesiense
and Isopterygium jelink. Both are recognised as valid
species by ALA and GBIF, and neither database list
records from elsewhere. Streimann and Curnow
(1989) also treated both species as valid and endemic
to Christmas Island (Table 2), even though Gepp and
Gepp (1905) stated that Isopterygium jelinki occurs on
Sumatra. Neither endemic species of moss has been
reported since 1904, but neither is listed as threatened,
nationally or globally.
Gepp and Gepp (1905) also reported one widespread
species of liverwort (Class Hepaticopsida) but we
are not aware of any reports of hornworts (Class
Anthocerotopsida) from Christmas Island, or of any
collections since 1904.
Vascular Plants
The vascular f lora of Christmas Island is well
inventoried, with extensive early collections at about the
time of the island’s settlement (Hemsley 1890; Ridley,
1891, 1906a,b), and some notable more recent surveys and
inventories, including studies to assess the conservation
status of some species (Du Puy 1988, 1993; Holmes and
Holmes 2002; Claussen 2005; Green et al. 2010). Claussen
(2005) reported that there are 213 species of native plant
on the island.
Although the early collections led Ridley (1906a)
to list 34 species of vascular plants as endemic to the
island, about half are no longer considered so (Table 4).
The 17 species still recognised as endemic are: Abutilon
listeri, Arenga listeri, Asystasia alba, Brachypeza
archytas, Colubrina pedunculata, Dicliptera maclearii,
Flickingeria nativitatis, Grewia insularis, Hoya aldrichii,
Illigera elegans, Ischaemum nativitatis, Pandanus
christmatensis, Pandanus elatus, Peperomia rossii,
Phreatia listeri, Pittosporum nativitatis and Zeuxine
exilis. With a few exceptions, all are species recognised
by GBIF, and for most species all locational data in
GBIF are from Christmas Island. The exceptions are that
GBIF includes no locational data for Pandanus elatus;
it includes a record from Kiribati (Pacific Ocean) for
Flickingeria nativitatis, which we discount as a case of
mistaken island name; and there are early records for
‘Indonesia’ (without explicit locational information) for
Dicliptera maclearii, Arenga listeri and Hoya aldrichii,
which we assume simply represent historically imprecise
geographical descriptors. Pittosporum nativitatis is
treated in GBIF as a synonym of P. ferrugineum subsp.
ferrugineum, but is maintained as a valid species endemic
to Christmas Island in ALA and CHAH Australian Plant
Census, consistent with its reinstatement as a species in a
recent review of the genus (Cayzer et al. 2000).
Two species for which Christmas Island is the type
locality and which have long been considered endemic
(Claussen 2005) are no longer recognised as such.
Asplenium listeri is now recognised to be much more
widely distributed (Ohlsen et al. 2015) and Zehneria alba
has been subsumed in the more widespread Z. mucronata
(CHAH Australian Plant Census: http://www.chah.gov.au/
apc/about-APC.html, but is still recognised as specifically
distinct in GBIF). Ardisia pulchra is recognised by GBIF
as a valid species endemic to Christmas Island, but is
subsumed within the more wide-ranging A. sanguinolenta
in CHAH’s Australian Plant Census and ALA. Likewise,
Saprosma nativitatis is recognised by GBIF as a valid
species endemic to Christmas Island, but is subsumed
within the more wide-ranging Amaracarpus pubescens in
CHAH’s Australian Plant Census and ALA.
There are records for all but two endemic plant
species within the last 50 years, notably including the
re-discovery of Zeuxine exilis in 2009 more than 100
years after its only previous collection in 1904 (Green et
al. 2010). However, Peperomia rossii is known only from
the 1898 type collection. Only one endemic plant species
is listed as threatened: Arenga listeri is listed globally as
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Endangered. Three native plants occurring on Christmas
Island (Asplenium listeri, Pneumatopteris truncata and
Tectaria devexa var. minor) are listed as threatened
nationally, but all also occur elsewhere (Table 3).
Kingdom ANIMALIA
Eleven animal phyla reported from Christmas Island
are considered below. Phylum Chaetognatha has been
reported in the plankton by Davies and Beckley (2010), but
no species have been identified. Nineteen more phyla
have not been reported at all. The nomenclature and
taxonomic sequence of animals follows AFD unless
stated otherwise.
Phylum PORIFERA
The Porifera of Christmas Island is poorly studied
and has not been reviewed for over 100 years. Dendy
(1887) described one species (Table 4). Kirkpatrick
(1900) listed 32 species from Andrews’ collection and
described seven of them as new. He later described one
more species, Murrayona phanolepis (Kirkpatrick 1910),
which is now recognised as the sole representative of a
monotypic genus and family (Murrayonidae), but it is
not endemic to Christmas Island. Sponges collected by
the Western Australian Museum in 1987 have not been
identified (Marsh and Fromont 2000), and the Raffles
Museum of Biodiversity Research expedition in 2010–12
did not report on Porifera (Tan et al. 2014a). Hooper
and Wiedenmayer (1994) updated the taxonomy and
nomenclature, but provided no new records. Although
sponges do not form a prominent feature of the island’s
coral reefs (Marsh and Fromont 2000), it is probable that
the species complement is more diverse than currently
known, particularly considering that all of the reported
collections come from a single locality, Flying Fish Cove.
Of the nine species whose type locality is Christmas
Island, Tethya affinis has been synonymised with Tethya
deformis (Hooper and Wiedenmayer 1994) and six have
been recorded beyond Christmas Island (ALA, AFD,
GBIF: see Table 4). Haliclona irregularis (Kirkpatrick
1900), described from Christmas Island, has three
homonyms described from elsewhere, H. irregularis
(Kieschnick 1896), H. irregularis (Czerniavsky 1880) and
H. irregularis (Brøndsted 1924) (Van Soest et al. 2018).
Following Burton (1959), we recognise two endemic
species of Porifera from Christmas Island, Oceanapia
sessilis and Haliclona innominata, both of which are
in the Class Demospongiae (Table 2). Although we are
aware of no records for these endemic species since 1897–
98, neither is listed as threatened, nationally or globally.
Phylum CNIDARIA
The surface of Christmas Island is dominated by
limestone karst that was built in part by coral growth
over millions of years, and living corals are obviously
essential elements of existing coral reefs, but ironically,
the Cnidaria of Christmas Island are largely neglected
(Beeton et al. 2010). Hard corals have received some little
63
attention recently (Done and Marsh 2000; Richards and
Hobbs 2014) but other Cnidarians are barely documented.
Richards and Hobbs (2014) listed 169 known species of
Scleractinia coral and five species of non-scleractinian
coral (three Hydrozoa and two Octocoralia). Their species
accumulation curves were close to asymptotic, although
the southern coast was not surveyed. Meanwhile, soft
corals, which dominate the southern coast, have received
no study. Hobbs et al. (2013) listed three widespread
Actinaria anemone species, but they did not attempt to be
comprehensive. Tan et al. (2014b) recorded an additional
Actinaria from submarine caves that was identified to
the genus Edwardsiella, for which no species are known
in the tropics (GBIF). Nektonic forms have not been
reported. Planktonic forms were reported by Davies and
Beckley (2010) only as ‘Cnidaria’. There is no evidence
that Christmas Island has any endemic species of
Cnidaria (Richards and Hobbs 2014), with the possible
exception of the Edwardsiella sp. Nevertheless, the
composition of the coral community is biogeographically
unique, but it is threatened by local pollution and invasive
species (Richards and Hobbs 2014).
Phylum PLATYHELMINTHES
Platyhelminthes have been found on Christmas
Island only recently, and little has been published on
them. Humphreys and Eberhard (2001) recorded a
single unidentified species of ‘free-living’ flatworm
(Turbellaria) from Jane-up Well, which was thought
to be styogobitic. James (2007) recorded two morphospecies of terrestrial ‘turbillarians’. One of these might
be the widespread Caenoplana coerulea Moseley 1877
(Tricladida, Geoplanidae), whilst the other might be a
turbillarian or a misidentified nemertean (DJ pers. obs.).
No parasitic Platyhelminthes have been recorded.
Phylum NEMERTEA
Nemerteans have not been described from Christmas
Island, but specimens of at least one terrestrial
species (Monostilifera) have been collected and
await determination (PG pers. obs.). It may be close
to Geonemertes pelaensis Semper 1863, which has a
distribution encompassing islands in the western Indian
Ocean, to the south-east of India and South-East Asia
(Moore et al. 2001). It is also possible that the Christmas
Island taxon is a new and endemic species. No marine
nemerteans have been reported.
Phylum BRACHIOPODA
A single species of Brachiopod has been recorded
from Christmas Island, and it was described as a new
species, Thecidellina blochmanni by Dall (1920). Marsh
and Fromont (2000) stated that it had not been recorded
elsewhere, but GBIF lists a specimen from Guam and
AFD includes Madagascar in the distribution. Therefore,
it is not considered endemic to Christmas Island. It
was collected at least as recently as 1987 (Marsh and
Fromont 2000).
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
64
Phylum MOLLUSCA
The molluscs of Christmas Island are diverse and have
been well-studied compared to most invertebrate groups.
There have been two significant modern studies, the first
by the Western Australian Museum (Wells and SlackSmith 2000) and the second by the Raffles Museum of
Biodiversity Research (Tan and Low, 2014). There is
also a popular book depicting a good selection of marine
mollusc species (Wells et al. 1990), though it is now
dated.
A remarkable collection was made incidentally by Mr
R. Kirkpatrick of the British Museum while dredging
for the Pharetronid sponge Murrayona phanolepis, off
North East Point, probably in January 1916. A sample
of dredged sand rich in ‘small’ (length < 5 mm) snails
passed to the Australian Museum from which Iredale
(1917) described five new species in four new genera,
and about 40 years later Laseron described 34 new
species and eight new genera (Laseron 1956a,b, 1958).
Most of these remain valid taxa, and while many of
the species are now known to have wider distributions,
some seem to be endemic. However, few of the species
and none of the endemic ones have been recorded from
Christmas Island again.
Tan and Low (2014) reviewed all previous studies
except for an unpublished report of a survey of
terrestrial mollusc species (Kessner 2006). They listed
the known Christmas Island fauna as 760 species from
130 families and four classes (four species in three
families of Polyplacophora; 104 species in 20 families
of Bivalvia; 640 species in 101 families of Gastropoda,
and 12 species in six families of Cephalapoda). Still,
their species accumulation curve suggests that the total
fauna is considerably larger. They listed the species
they considered to be endemic to Christmas Island:
one Bivalvia, 38 Gastropoda, no Polyplacophora and
no Cephalapoda. However, there is low agreement
on taxonomy and distribution among authorities (Tan
and Low 2014; Bieler et al. 2018) and taxonomic
databases (ALA; AFD; GBIF). Schwartziella lata is
synonymised with Zebina triticea and Schwartziella
oceanica is synonymised with Zebina ephamilla by
AFD and ALA. Eleven Gastropoda species from Tan
and Low’s list of endemics have locational records
beyond Christmas Island in either ALA, AFD or GBIF
and therefore are not considered endemic here (Table 4).
The remaining one Bivalvia and 26 Gatropoda species
(seven terrestrial and 19 marine) have type localities at
Christmas Island and are not recorded from elsewhere,
so are taken here to be endemic (Table 2), with the
following considerations. Five species (Semperula
insularis, Succinea solitaria, Lamprocystis mabelae,
Lamprocystis mildredae and Lamprocystis normani)
are not recognised in any of ALA, AFD or GBIF, but
we surmise that they have been overlooked rather than
synonymised and follow Gomes and Thomé (2004)
and Tan and Low (2014) in recognising them as valid.
Tomlin (in Laidlaw 1935) considered there was no
difference between L. mildredae and L. normani, but
Kessner (2006) recently collected and distinguished
both. In addition, we note an additional apparently
endemic species described by Laseron (1956b) that
was not included in Tan and Low (2014): Cyclonidea
carina is recognised as a valid species in ALA and
GBIF, although with no locational records. Also, four
undescribed putative species have been reported that
might be endemic Gastropoda species (Table 3).
Eighteen of the 27 known endemic Mollusca species
have not been certainly recorded for over 50 years
(Table 2). This is concerning given that the group has
been surveyed comparatively well. Nevertheless, none
of the endemic Mollusca species are listed as threatened,
nationally or globally.
Phylum ANNELIDA
There has been very little reporting of annelids
from Christmas Island. Gates (1935) provided the only
available list of Oligochaeta, which was 13 species.
One of these, Polypheretima brevis (Rosa 1898)
was originally described from the Island (Andrew’s
collection) as Pheretima brevis, but it has since been
recorded from Tonga (Easton 1984) (Table 4).
Humph reys (2014) repor ted th ree species of
Polychaeta from anchialine environments, a Syllidae sp.
a Nerilla sp. (Nerillidae) and a Prionospio sp. The last is
considered to be a new species, based on an unpublished
molecular phylogeny by K. Worsaae (pers. comm.) at
the University of Copenhagen (Table 3). No members of
the Class Hirudinida (leeches) have been recorded. No
annelid species from the Island are listed as threatened,
nationally or globally.
Phylum ARTHROPODA
The arthropods are the largest and most diverse
phylum of the animal Kingdom. Representatives of all
four subphyla have been recorded on Christmas Island.
Phylum ARTHROPODA
Subphylum Crustacea
Two of the five crustacean classes (Branchiopoda and
Remipia) have not been recorded at Christmas Island.
The remaining three classes are treated below.
CLASS OSTRACODA
Ostracoda have recently been discovered on Christmas
Island in anchialine systems and various freshwater
environments including subterranean streams, springs
and tufa flows, but not in the marine environment
( Hu mph reys a nd Ebe rha rd 20 01; Hu mph reys
2014). Their diversity is surprising, their endemism
extraordinary and their biogeography astounding
(Humphreys 2014). About six species in four families
have been discerned, but while the exact number is
clouded by taxonomic uncertainty, it is evident that
greater diversity will emerge with further sampling and
sorting. In just two decades three new species and two
ENDEMIC SPECIES OF CHRISTMAS ISLAND
new genera have been discovered and described (Table 2),
namely: Humphreysella baltanasi, Microceratina
martensi and Isabenula humphreysi (Namiotko et al.
2004; Kornicker et al. 2006; Humphreys et al. 2009;
Rossetti et al. 2011). In addition Vestalenula sp. E was
recognised as new by Rossetti et al. (2011), but left in
open nomenclature because only one individual was
known, and Humphreys (2014) recognised a new species
of Penthesilenula (Table 3). No ostracod species from
the Island are listed as threatened, nationally or globally.
CLASS MAXILLOPODA
Murray and Andrews made a small collection of
marine plankton in 1908 that was rich in Copepoda,
from which 12 new species and a new genus were
described (Farran 1911, 1913). Not surprisingly, none
are endemic (Table 4). Three of these have been
synonymised into wider ranging species and nine are
accepted species but have wider distributions (Table 4).
A subterranean copepod considered to be a new
species and genus in the family Arietellidae is very
likely endemic (Bruce and Davie 2006; Humphreys
2014).
CLASS MALACOSTRACA
Five orders of the large and diverse crustacean Class
Malacostraca recorded at Christmas Island are treated
below. Davies and Beckley (2010) collected planktonic
forms of three other orders (Cumacea, Mysidacea, and
Euphausiacea), but these were identified only to Order
level. These and seven more orders never reported from
Christmas Island are not considered further here.
Order Stomatopoda
Ahyong (2014) listed eight species of mantis
shrimps known from Christmas Island. One of these,
Chorisquilla quinquelobata, was described from
Christmas Island as Gonodactylus quinquelobatus from
specimens collected by Harms in 1933. Ahyong (2014)
considered it to be endemic to Christmas Island, and
this treatment is supported by AFD, ALA and GBIF. A
single female Chorisquilla quinquelobata was collected
by hand from Thundercliff Cave in 2008, which was the
first record since 1933 (Ahyong 2014). However, it is not
listed as threatened, nationally or globally.
Order Amphipoda
No endemic species are known, although the fauna
is not thoroughly documented. Specimens of the genus
Leucothoe (Leucothoidae) have been recorded in
anchialine systems, and of the genus Floresorchestia
(Talitridae) in damp caves and adjacent to anchialine
pools (Humphreys 2014)
Order Isopoda
There is very little literature on isopods from
Christmas Island. One species, Tylos nudulus, was
described by Budde-Lund (1906) from Andrews’
1897–98 collections, although Andrews (1900c) made
65
no reference to terrestrial (littoral) isopods. This
species is still considered to be valid and endemic to
Christmas Island (GBIF, AFD, Hurtado et al. 2014). We
have found no records subsequent to the type series.
However, James (2007) reported three morpho-species
of isopods, one littoral and two terrestrial species.
Humphreys (2014) listed 16 terrestrial oniscidean isopod
species from six families that have been recorded in
subterranean environments on the island, including
seven putative undescribed species (four Myrmecodillo,
two Papuaphiloscia and one Elumoides) (Table 3). No
endemic isopod species is listed as threatened.
Order Thermosbaenacea
The Thermosbaenacea is a small order restricted
to subterranean waters. It was previously known in
Australia from a single species (AFD), Halosbaena
tulki, which is from a cave in tropical Western
Australia (Poore and Humphreys 1992). Specimens of a
Halosbaena have recently been discovered in caves on
Christmas Island that represent an undescribed species
(Humphreys 2014) (Table 3).
Order Decapoda
The crab fauna of Christmas Island is widely and
justifiably celebrated for its diversity, abundance and
ecological importance. There have been notable recent
reviews and popular accounts (Hicks et al. 1990;
Orchard, 2012). Many species are considered endemic,
with this number increasing because of surveys in
specialised subterranean habitats (Naruse and Ng 2014;
Tan et al. 2014b) and because recent taxonomic scrutiny
has raised to specific status some Christmas Island
populations formerly considered to be part of more
wide-ranging species (Ng and Davie 2012).
Pocock (1889) described the first crab species from
Christmas Island, the red crab, as Hylaeocarcinus
natalis (now Gecarcoidea natalis), which has
subsequently been shown to be a pivot of the island’s
ecology (Green et al. 1996). Andrews (1900a) reported
on five species from his 1897–98 collection, all of
which he attributed to species already described from
elsewhere. However, the blue crab he recognised as
Cardisoma carnifex has subsequently been recognised
as specifically distinct (Discoplax celeste) and endemic
to the island (Ng and Davie 2012), with subsequent
taxonomic change to Tuerkayana celeste (Guinot et
al. 2018). Andrews’ subsequent collection in 1908,
mostly of intertidal species, was reported on by Calman
(1909). This comprised about 67 species, of which four
species were described as new: Lioxanthodes alcocki,
Sesarma murrayi, Hyastenus andrewsi and H. uncifer,
with Lioxanthodes described as a new genus. Two of
these species have since been synonymised with species
earlier described from elsewhere, and the other two
subsequently also recorded from elsewhere (Table 4).
Collections in 1932–33 by J.W. Harms, in 1932 by M.
Tweedie and in 1938–40 by Carl Gibson-Hill resulted
in further substantial advances in the inventory of
66
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
the Christmas Island crab fauna, with 13 new species
described by Ward (1934), including the new genera
Tweedieia and Proechinoecus, two new species
described by Balss (1934), one by Gordon (1935), and
one by Shen (1936). Of these newly-described species,
only two, Karstarma jacksoni and Echinoecus sculptus,
are still considered endemic to Christmas Island (Tables
2 and 4). Oddly, both GBIF and ALA list a 1932 record
of Echinoecus sculptus from Kiribati, Pacific Ocean, but
this is likely to be a geographic error, as the collection
date coincides with the time at which Tweedie collected
the type material on Christmas Island. Tweedie (1947)
provided a then-comprehensive inventory of the c. 50
species of Brachyura recorded from Christmas Island,
including some substantial taxonomic re-assessments of
species previously named from there.
Morgan (2000) listed a total of 204 species of
decapods, many of which were not determined to
species level. However, he did not attempt to identify
endemic species. Since then there has been a series
of surveys, particularly of the island’s subterranean
environments, and further taxonomic reviews, that have
added many more species, including the discovery or
taxonomic recognition of five species of crabs (Davie
and Ng 2012, 2013; Ng and Davie 2012; Naruse and Ng
2014), a stygobitic prawn, Macrobrachium xmas (Fujita
et al. 2015), and an anchialine cave shrimp, Procaris
noelensis (Bruce and Davie 2006), all considered
endemic to Christmas Island (Table 2). One of the
newly discovered crabs, Christmaplax mirabilis, also
represents an endemic genus (Naruse and Ng 2014).
In addition, since the inventory published by Morgan
(2000), there have been several recent reviews of
components of the Christmas Island decapod fauna.
Mendoza et al. (2014) compiled records of 83 species
of Xanthidae from Christmas Island, of which 30 were
additions to the known local fauna, but none were
considered endemic. Ng and Naruse (2014) compiled
records of seven species of reef lobster (Palinuridae,
Scyllaridae and Enoplometopidae), of which one was
locally new but none were endemic. Osawa (2014) listed
nine species of Porcelanidae, including five (or four)
new records but no endemics. Tan et al. (2014b) reported
on the Decapod (and other) fauna of submarine and
associated anchialine caves of Christmas Island, noting
particularly that there were at least three undescribed
and presumed endemic species, including a new genus
of Paguridae (Table 3). This brings the total species list
of decapods to c. 249.
We conclude that ten described Decapod species
are endemic to Christmas Island (Table 2). Most of
these represent recent discoveries or the consequences
of recent taxonomic revision. Echinoecus sculptus
(Pilumnidae) is recognised as valid and known only
from the type locality by Davie (2002); it is symbiotic
with the intertidal echinoid Colobocentrotus atratus. All
of the endemic species have been recorded recently, but
some are known from very few specimens and a small
number of locations on the island: for example, Procaris
noelensis is known from only a single specimen.
The endemicity of Christmas Island’s iconic red
crab Gecarcoidea natalis is not clearly resolved: we
treat it as endemic species, consistent with most recent
assessments (e.g. Orchard 2012; Morris et al. 2018).
However, it also occurs on the Cocos (Keeling) islands
(Tweedie 1950), where it is patchily abundant on Pulu
Keeling (North Keeling Island) (JW pers. obs.) and
scarce in the northern parts of the southern atoll (DJ
pers. obs.; IAW McAllan pers. comm.). It has been
known from the Cocos (Keeling) islands group for more
than a century (Wood Jones 1909). This occurrence
beyond Christmas Island may be a result of deliberate
or accidental introduction, or sporadic colonisation.
Tweedie (1950) asserted that ‘there is no doubt that
they have been introduced (to Cocos (Keeling)
islands) with soil imported from there (Christmas
Island)’. Alternatively, Orchard (2012) noted that
‘low numbers … occur on North Keeling Island’ and
considered that they derived from ‘larvae that drifted
from Christmas Island. North Keeling red crabs have
never been observed to breed’. GBIF also includes two
1970s records of this species from islands off western
Thailand, but these records are unlikely to represent a
breeding population. Populations of Gecarcoidea natalis
on Christmas Island have been conspicuously and
severely depleted by supercolonies of yellow crazy ants
since the first supercolony was reported in 1989 (Green
and O’Dowd 2009).
The Christmas Island crab fauna is unusually famous,
charismatic and ecologically significant, and has
attracted international scientific interest. Other decapods
are significant examples of ancient lineages that, along
with other crustaceans, are shedding new light on
evolution and biogeography in the marine environments
of the early Mesozoic. However, no species is listed as
threatened, globally or nationally.
Phylum ARTHROPODA
Subphylum Chelicerata
The chelicerate Class Arachnida is treated below; the
Class Picnogonida has not been recorded.
CLASS ARACHNIDA
The arachnid fauna is poorly known, despite
collections in 1887, 1897–98, 1939, 1964, 1989 and 2005
(Pocock 1889, 1900; Savory 1943, 1947; Campbell 1968;
CSIRO Division of Entomology 1990; Framenau and
Waldock 2006). Seven subterranean fauna collections
were made between 1987 and 2013, but the fauna is still
considered poorly known (Humphreys 2014). A recent
review of spiders is unfortunately incomplete and not
formally published (Framenau and Waldock 2006),
and the taxonomy of the Araneae is poorly resolved in
the wider region. Framenau and Waldock (2006) listed
37 Araneae species, but also 40–50 additional taxa
not identified beyond genus. Humphreys (2014) listed
eight troglomorpic spiders, of which three species are
additional to the above. Also recorded are two species of
ENDEMIC SPECIES OF CHRISTMAS ISLAND
67
Scorpiones, at least eight species of Pseudoscorpiones,
one species of Amblypygida and one species of
Schizomida (Savory 1943, 1947; CSIRO Division of
Entomology 1990; Humphreys 2014). The Acari have
been little studied and never reviewed. Neumann (1904)
described a single species of tick, Ixodes nitens, from
a specimen of the endemic Rattus macleari. Campbell
(1968) noted the presence of dog ticks Rhipicephalus
sanguineus on domestic dogs and/or cats. More
recently, Humphreys and Eberhard (2001) reported two
indeterminate Acarina from subterranean environments.
An arthropod voucher collection made from 2004–06
contained 11 morpho-species of mites and two of ticks
(James 2007). Thus there are 54 identified species of
Arachnida from the island and possibly 50–60 further
species that have been collected but not identified fully.
type collection in 1939, but a filistatid spider (not
determined to species level) reported by (CSIRO
Division of Entomology 1990) may represent this
species. Heteropoda listeri was collected in 1897–98 and
1939 but not since. Idioctis xmas is rarely encountered
and might be known only from the type collection taken
in 1983. Hormurus polisorum is rare but was collected
as recently as 2006 (ALA). No species of Arachnida
from Christmas Island is listed as threatened, nationally
or globally.
Nine arachnid species have been described from
Christmas Island: one Amblypygida, one Acari, one
Scorpiones, two Pseudoscorpiones and four Araneae
(Tables 2 and 4). Of these, one species is not endemic.
Metawithius murrayi, described by Pocock (1900), has
subsequently been recorded widely in Indonesia and
the Andaman and Nicobar Islands (Harvey 2015). In
their original description of the whip scorpion Charon
gervais, Harvey and West (1998) speculated that it may
have been introduced to Christmas Island from Java, but
we found no records beyond Christmas Island. GBIF
lists a record of Ariadna natalis from Natal, South
Africa, but this is almost certainly an error resulting
from the specific epithet. ALA and GBIF list records
of Idioctis xmas from Queensland, Australia, but this
is also almost certainly an error resulting from the
location where the specimens are housed. Accordingly,
we recognise Charon gervaisi and these last two
Areneae species, along with Filistata gibsonhilli and
Heteropoda listeri, the tick Ixodes nitens (recognised
also as a valid endemic species in a recent global review:
Guglielmone et al. (2010)), the scorpion Hormurus
polisorum, and the pseudoscorpion Paratemnoides
pococki as endemic species, being valid species never
recorded beyond Christmas Island (Table 2). In addition,
three undescribed taxa have been reported that might
be endemic Arachnida species (Table 3). Given the
poor taxonomic resolution of the group, there might be
more undiscovered endemic species. For example, an
eyeless troglomorphic spider, tentatively identified as
Olin platnicki Deeleman-Reinhold 2001, differs from
the type population (in Sulawesi) that has ‘reduced eyes’
(Humphreys 2014). An Acarina parasitic on O. platnicki
might also be endemic (Humphreys 2014).
CLASS CHILOPODA
Ixodes nitens is known only from a few individuals
collected from the last specimens of Rattus macleari,
before it became extinct between 1901 and 1904
(Durham 1908; Green, 2014), so the tick presumably
became extinct at the same time (Mihalca et al.
2011; Colwell et al. 2012). Ariadna natalis has not
been recorded since the type collection in 1897–98.
Filistata gibsonhilli has not been recorded since the
Subphylum Myriapoda
Two classes of myriapods (Chilopoda and Diplopoda)
have been recorded from Christmas Island and two have
not (Pauropoda and Symphyla).
The centipede fauna of Christmas Island was reviewed
by Waldock and Lewis (2014). The collections have been
limited, and several key specimens are incomplete and
hence difficult to determine. Waldock and Lewis (2014)
listed ten species from historical sources and recent
records, but four of these were identified only to genus.
Cryptops inermipes, described from Christmas Island
by Pocock (1889), is still recognised as a valid species
(Lewis 2011). It is known definitively only from the type
specimen, but two other incomplete specimens from
Christmas Island, one collected in 1887 and identified as
C. hortensis (Pocock 1889) and the other in 2006, may
be referrable to it (Waldock and Lewis 2014).
CLASS DIPLOPODA
The Christmas Island millipede fauna was reviewed
by Jeekel (2006) and Mesibov (2010). From specimens
collected by Lister in 1887, Pocock (1889) described
two species, Cylindrodesmus hirsutus and Spirostreptus
exocoeti, the former constituting a new genus, albeit
with other congeneric species subsequently reported
from elsewhere (Pocock 1900). Both of these species,
and another (the non-endemic Orthomorpha coarctata),
were also reported by Pocock (1900) from Andrews’
1897–98 collections. Cylindrodesmus hirsutus has since
been found to have a wide distribution beyond Christmas
Island (Jeekel 2006).
A collection in 1933 by J.W. Harms was documented
by Jeekel (2006), who noted the previously reported three
species and added a further five species (Leptogoniulus
sorornus, Solaenaulus butteli, Prosopodesmus jacobsoni,
Monographis sp. and an undetermined Siphonotidae
species). Of these, at least the first three are widely
distributed beyond Christmas Island and may have
been introduced there. Another wide-ranging species,
Trigoniulus corallianus, was earlier recorded from the
island by Jeekel (2001). Jeekel (2006) also transferred
S. exocoeti to Hypocambala, a treatment followed by AFD.
There has been little subsequent collection. For
example CSIRO Division of Entomology (1990) did not
document any diplopod records in their invertebrate
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
68
survey of the island, but that survey resulted in the
lodgement of four specimens of Asiomorpha coarctata
(=Orthomorpha coarctata) in the ANIC. However,
collections in 1998 resulted in the description of two
additional cave-dwelling species: Lophoturus speophilus
and L. humphreysi (Nguyen Duy-Jacquemin 2014).
The online database Millibase (Sierwald and Spelda
2018) recognises S. exocoeti (as Iulomorpha exocoeti,
with synonym Hypocambala exocoeti), Lophoturus
speophilus and L. humphreysi as valid species endemic
to Christmas Island. Mesibov (2010) characterised
Hypocambala exocoeti as a ‘tramp’ species, and
Mesibov (2018) noted it as introduced to Australia;
likewise, the AFD includes the enigmatic comment
‘introduced from ?’. Given that it was collected
on Christmas Island prior to the island’s human
colonisation, and that it has not been found elsewhere,
we consider instead that it is native and endemic to
Christmas Island.
Whereas Lophoturus speophilus and L. humphreysi
were collected in 1998 (from < 10 specimens and the
holotype only, respectively), we are aware of no records
of Hypocambala exocoeti since Harms’ collections
of 1933 (Jeekel 2006). Notwithstanding the lack of
documented records for Hypocambala exocoeti for
more than 80 years, and the few specimens and highly
localised distribution of the two endemic Lophoturus
species, none of the three endemic species is listed as
threatened, nationally or globally.
Subphylum Hexapoda
All three classes of hexapods (Collembola, Entognatha
and Insecta) are recorded from Christmas Island.
CLASS COLLEMBOLA
There are no endemic species known. About 850
specimens belonging to 15 morpho-species of springtails
were identified from the CSIRO collection in 1989
(CSIRO Division of Entomology, 1990). Although only
one was determined to species level, it was considered
that no genera and probably no species are endemic to
Christmas Island.
CLASS ENTOGNATHA
Of the two orders within Entognatha (Diplura and
the Protura), only the former has been recorded from
Christmas Island, with a single species collected
from subterranean habitats in 1989 (Humphreys and
Eberhard 2001). It was considered to be a new species
of Cocytocampa (Family Campodeidae) and likely to be
endemic (Table 3).
CLASS INSECTA
Within the Class Insecta 21 Orders reported from
Christmas Island are treated below. There are no
reports from Christmas Island of four orders, namely
Archaeognatha (bristletails), Plecoptera (stoneflies),
Megaloptera (alderflies), or Mecoptera (scorpion-flies).
Order Zygentoma
The silverfish (formerly Order Thysanura) are poorly
represented on Christmas Island. CSIRO Division
of Entomology (1990) collected two (specimens? of)
Lepismatidae under bark near North West Point that
were not determined to species. James (2007) reported
a single representative of Lepismatidae, not determined
to species. Humphreys and Eberhard (2001) collected
a troglobitic species of Metrinura (Nicoletiidae)
considered likely to be a new (and endemic) species
(Table 3); this collection represented a range extension
of the genus from New Caledonia and eastern Australia.
Order Odonata
There are no endemic species. One widespread,
migratory species of Odonata, Pantala flavescens is
common on Christmas Island (Kirby 1900e; CSIRO
Division of Entomology 1990). Two other widespread
species were reported by Kirby (1900e), but not since.
Order Phasmida
Only one stick insect species has been recorded
on Christmas Island. Kirby (1889) described the
phasmid species, Clitumnus stilpnoides, from three
specimens in Lister’s 1887 collections, with a further
four specimens collected by Andrews in 1899–1900
(Kirby 1900f). This species, now Ramulus stilpnoides,
is recognised as a distinct species in AFD, ALA and
GBIF and the few distributional records in those sources
are from Christmas Island only: it is widely accepted
as a Christmas Island endemic (CSIRO Division of
Entomology 1990; Rudolf and Brock 2017).
No other identified species of phasmids have been
reported from Christmas Island. However from
Andrews’ 1898–99 collection, Kirby (1900f) noted
that in addition to R. stilpnoides, ‘there are two other
Phasmidae in Mr. Andrews’ collection, apparently
belonging to the allied genus Entoria …, but hardly in
sufficiently good condition to describe, though probably
new.’ We can find no further account of this putative
species and assume that the specimens may have
represented poorly preserved examples of R. stilpnoides.
Rudolf and Brock (2017) considered that R. stilpnoides
‘has very rarely been reported, except for old records
pre-1900, and the only current record of the species
is a photo of a live female, which was received in
September 2015’. Consequently, it was recognised in
2017 as Vulnerable by the IUCN (Rudolf and Brock,
2017), the only Christmas Island invertebrate species
recognised nationally or globally as threatened (Table
1). Its presumed ongoing decline was inferred to be due
to predation by black rats and yellow crazy ants (Rudolf
and Brock 2017). However, there are many additional
records of this species. ‘About 30 specimens’ were
collected in the CSIRO survey of 1989 (CSIRO Division
of Entomology 1990). In 2004, it was considered to be
widespread but uncommon in forest habitat (Surman
2004), and four voucher specimens were collected
(James 2005).
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Order Trichoptera
The only account of Trichoptera from Christmas
Island is from a recent freshwater invertebrate survey by
Weeks & McColl (2011), who reported two undetermined
species of caddisflies (Trichoptera; Leptoceridae and
Hydroptilidae). They did not assess endemicity.
Order Mantodea
A single mantid species has been recorded from
Christmas Island. From material collected by Andrews
in 1897–98, Kirby (1900f) described Hierodula dispar
as ‘a very distinct species’. However, it was subsequently
synonymised with the more widely distributed H.
patellifera, so it is not endemic to Christmas Island
(AFD; ALA; GBIF). About 50 specimens were collected
in 1989 (CSIRO Division of Entomology 1990).
Order Ephemeroptera
The only account of Ephemeroptera from Christmas
Island is from a recent freshwater invertebrate survey
by Weeks and McColl (2011), who reported at least
one undetermined species of mayfly (Ephemeroptera;
Caenidae, Tasmanocoenis sp.). They did not assess
endemicity, but noted that this species was abundant.
Order Blattodea
Few species of this order have been collected from
Christmas Island. The cockroach fauna was reviewed by
Roth (2000), who reported 11 species, of which ten were
widespread (described from elsewhere) and one was a
cave-dwelling species in a monotypic genus known only
from Christmas Island, Metanocticola christmasensis
(Roth 1999). Subsequently Humphreys and Eberhard
(2001) reported that ‘a second troglobitic cockroach of
the family Blattellidae also represents an undescribed
genus L. Roth, pers. comm. 1998’. However this taxon
apparently has not yet been described.
Of the ten non-endemic cockroach species reported
from Christmas Island, it is challenging to determine
which are native and which are introduced. Two were
collected on the island in 1887 before any substantial
settlement (Kirby 1889) and another four by Andrews in
1897–98, so these may all be native (Kirby 1900f).
The termite (Isoptera) fauna of Christmas Island is
poorly documented. Andrews’ collection of 1897–98
included one taxon not determined to species level
(Kirby 1900f). From his 1932 collections, Tweedie
(1933) added another species, again not determined to
species level. Gibson-Hill (1947d) noted three species
were present in 1939–40, of which two were identified
to genus only and the third not to that level. The CSIRO
survey of 1989 collected six termite species identified to
genus only (CSIRO Division of Entomology 1990).
Order Embioptera
There are no endemic species recognised. CSIRO
Division of Entomology (1990) collected a single
specimen of an adult web-spinner from the Family
Oligotomidae. James (2007) recorded two morphospecies identified only to the level of order.
69
Order Orthoptera
The Orthoptera fauna is small but poorly surveyed
with no recent reviews. CSIRO listed 12 taxa but
identified only seven to species level (CSIRO Division
of Entomology 1990), and James (2007) listed 18
morpho-species but identified only six to species level.
Seven species have been described from the island, all
by Kirby (Kirby 1889, 1900f). Of these, Epacromia
rufostriata has been synonymised with the more
widespread Aiolopus thalassinus (Orthoptera Species
File 2018); and Cyrtacanthacris disparilis has been
synonymised at species level but remains an endemic
subspecies, namely Valanga nigricornis disparilis
(GBIF; Table 4), although it is still recognised as a valid
species in ALA. The other five species, Ectadoderus
f lavipalpis (now Ornebius f lavipalpis), Gryllacris
rufovaria, Phisis listeri (now Paraphisis listeri), Psyra
pomona (now Psyrana pomona) and Primnia orientalis
(now Nisiocatantops orientalis) are still recognised in
at least one of GBIF, ALA or AFD or the Orthoptera
Species File (2018) and have not been recorded from
elsewhere, so are accepted here as endemic species
(Table 2). In addition, two undescribed putative
species have been reported that might be endemic
Orthoptera species (CSIRO Division of Entomology
1990; Humphreys and Eberhard 2001) (Table 3). Dirsh
and Uvarov (1953) created the new monotypic genus
Nisiocatantops for Primnia orientalis, and therefore an
endemic genus as well.
Ornebius flavipalpis has not been recorded since
Andrews collected the types in 1897. Nisiocatantops
orientalis was last recorded in 1964 (Campbell 1968),
and the other three endemics have been recorded
recently (James 2007). None are listed as threatened,
nationally or globally.
Order Dermaptera
Lister’s 1887 collection included one species,
described by Kirby (1889) as Labidura nigricornis,
but this species was later subsumed in the more wideranging Chelisoches morio. From material collected
by Andrews in 1897–98, Kirby (1900f ) identified
eight species, including his previously described L.
nigricornis, two species described from elsewhere,
one taxon identified to genus only and four new
species: Labia murrayi, L. incerta, L. indistincta
and L. subarmata (now Anisolabis subarmata).
Two of these, L. incerta and L. intermedius, were
subsequently synonymised with a third, Labia murrayi,
now Paralabella murrayi (Steinmann 1989). The
1989 CSIRO survey collected about 130 specimens,
comprising six species: of these the wide-ranging
Euborellia stali was recorded previously by Kirby
(1900f), one other was identified as a wide-ranging
species, three to genus only and one only to family level.
Neither of the previously described endemic species was
recorded. Collections in 2004 reported three morphospecies (James 2005, 2007).
70
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Paralabella murrayi and Anisolabis subarmata are
still recognised as valid species with type localities of
Christmas Island and no occurrences elsewhere. Neither
species has been recorded since 1897–98, but neither is
listed as threatened, nationally nor globally.
Order Zoraptera
This small order (c. 30 species globally) was not
known from Australia until CSIRO Division of
Entomology (1990) collected two specimens from
leaf litter on Christmas Island that were subsequently
described as Zorotypus lawrencei (New 1995). Although
it is closely related to several species in Indonesia, it
is currently recognised as a valid species (AFD, ALA,
GBIF) endemic to Christmas Island (AFD). It has not
been recorded since 1989, but is not listed as threatened.
Order Psocodea
AFD treats the Phthiraptera (‘true’ lice) and the
Psocoptera (barklice and booklice) as higher taxa
in the Order Psocodea. There are few reports of the
Phthiraptera from Christmas Island. Most bird species
carry avian lice of two sub-orders, Amblycera and
Ischnocera (Murray et al. 1990). Kirby (1900d) listed
a single specimen collected by C.W. Andrews that was
large and thought to be from a seabird, but which was
not identified beyond ‘Mallophaga’ (now the Sub-orders
Amblycera and Ischnocera). Durham (1908) observed
that two Christmas Island f lying-foxes Pteropus
(melanotus) natalis were infested with a ‘louse-like
parasite’. The endemic birds of Christmas Island may
carry endemic lice, but checklists of lice on Australian
birds (e.g. Murray et al. 1990) have provided no specific
details for Christmas Island birds.
Surprisingly, there were no documented collections
of Psocoptera until Hill and Robertson made small
collections in 1983 and the 1989 CSIRO survey collected
177 specimens (CSIRO Division of Entomology, 1990).
From these collections Smithers (1995) identified 33
species, from which he described ten new species and
two new genera. Although there has been no relevant
taxonomic review since, all these species and genera
are recognised as valid in AFD, ALA and GBIF, with
no records of any beyond Christmas Island. Therefore
we treat them as ten endemic species and two endemic
genera (Table 2). James (2007) reported 36 morphospecies of Psocoptera collected from 2004 to 2006.
They were not examined by a specialist, but it suggests
a possibility that the fauna may be larger than currently
documented. No species are listed as threatened.
Order Thysanoptera
There has been little collecting and documentation for
this group and no endemic species are recognised. The
1989 CSIRO survey collected 25–30 species of thrips
in 21 genera, but none were identified to species level
(CSIRO Division of Entomology 1990). James (2007)
recorded eight morpho-species identified only to the
level of order.
Order Hemiptera
There has been no comprehensive review of the
Hemipteran fauna of Christmas Island. However,
Neumann et al. (2016) and Neumann et al. (2018) listed
endemic species in some Hemipteran families, as part
of a risk assessment for the introduction of parasitoid
wasps as a biological control agent for infestations of the
introduced yellow crazy ant Anoplolepis gracilipes.
The first collection was of five species by Lister
in 1887, all of which were described by Kirby (1889)
as new: Lygaeus subrufescens, Oxypleura calypso,
Ricania flavicostalis, R. affinis and R. hyalina. Of
these, Neumann et al. (2018) and CSIRO Division of
Entomology (1990) recognised O. calypso, Ricania
flavicostalis (now Varcia flavicostalis) and R. hyalina
(now Salona oceanica) as valid endemic species.
Ricania affinis (now V. affinis) is still recognised
as a valid species (GBIF, ALA): although CSIRO
Division of Entomology (1990) commented that it is
‘shared with Australia’, there are no locational records
beyond Christmas Island in GBIF or ALA. Lygaeus
subrufescens (now Leptocoris subrufescens) is no longer
considered endemic, with Göllner-Scheiding (1980)
recognising L. s. flava from the Caroline Islands in the
Pacific.
Andrews’ 1897–98 collections increased the number
of known species to 17, with 10 new species described
and only two species in the collection having previously
been recorded from elsewhere (Kirby 1900a,b). Of the
ten species described from this collection, nine are
still recognised as valid endemic species: Pentatoma
grossepunctatum (now Plautia grossepunctata)
(endemicity noted by Cassis and Gross (2002)),
Brachyrhynchus lignicolus (now Neuroctenus lignicolus)
(endemicity noted by Cassis and Gross (2002)), Ricania
flavifrontalis (endemicity noted in ALA), Paurostauria
delicata (endemicity noted in ALA, and a monotypic
genus), Nogodina subviridis (now Sassula subviridis)
(endemicity noted in ALA), Bidis aristella (now
Ugyops aristella) (endemicity noted by AFD and
Fennah (1964)), Clovia eximia (endemicity noted in
ALA and Neumann et al. (2018)), Issus andrewsi (now
Distiana andrewsi) (accepted as an endemic species
in review by Fennah (1954)) and Idiocerus punctatus
(now Batracomorphus punctatus) (endemicity noted in
AFD). Andrews’ collections were further considered
by Distant (1901), who described five more species, of
which two are still recognised as valid and endemic
species: Nysius spectabilis (with endemicity noted
in Malipatil (2010) and Cassis and Gross (2002)) and
Lethaeus maculatus (now Elasmolomus maculatus)
(with endemicity noted by AFD and Cassis and Gross
(2002)). Of the three other species described by Distant
(1901) from this collection, Geocoris vestitus was
accepted as a valid endemic species by Cassis and
Gross (2002) but was synonymised with the more
widespread G. jucundus by Kondorosy (2016); Pamera
insignis (now Paraeucosmetus insignis) was considered
as a valid endemic species by Cassis and Gross (2002)
ENDEMIC SPECIES OF CHRISTMAS ISLAND
but Chandra and Kushwaha (2014) considered that it
occurred more widely (to India); and Pamera andrewsi
was subsequently synonymised with the wide-ranging
Remaudiereana nigriceps (ALA).
The next collection was made by F. Harms in 1933,
with this documented by Izzard (1933), who also
reviewed the previous published studies of the island’s
Hemipteran fauna. Izzard (1933) described a further
ten species, and Harms’ collection brought the tally
of Hemipterans recorded from Christmas Island to 56
species, of which 28 were then considered endemic. Of
the species described by Izzard (1933), seven are still
recognised as valid endemic species: Lygis aldrichi
(now Tayloriligus aldrichi) (ALA, GBIF), Lygis murrayi
(now Tayloriligus murrayi) (ALA, GBIF), Cylapofulvius
listeri (now Peritropis listeri) (ALA, GBIF, and
recognised as a valid endemic species by Cassis and
Gross (1995)), Laccocoris montandoni (ALA, GBIF),
Andrewsiella oceanica (ALA, GBIF, constituting a
monotypic genus), Delphacodes muirianus (ALA,
GBIF) and Erythroneura harmsi (ALA, GBIF). Nearly
40 years after being collected by Harms, Hishimonus
festivus was described from that material by Knight
(1970), but there have been subsequent records from
west Java (Fletcher and Dai 2013).
In 1964, a CSIRO survey of invertebrates of potential
health and agricultural concern resulted in collection
of several previously-unrecorded, but widespread,
Hemipteran species presumed to have been introduced
to Christmas Island (Campbell 1968). The more
comprehensive CSIRO survey of 1989 collected about
5600 Hemipteran specimens categorised into about
140 morpho-species, with most not named (CSIRO
Division of Entomology 1990). However, about 20 wideranging and probably introduced species were recorded
for the first time, and the collection documented the
continued presence of nine previously-described species
recognised as endemic to Christmas Island. In a targeted
and comprehensive survey of scale insects (Coccoidea)
on the island, Neumann et al. (2016) recorded 28 species,
none of which were endemic, and most of which were
probably recent introductions.
Two further species have been described from
Christmas Island: Matigocoris insularis by Kormilev
(1983), which was still accepted as a valid endemic
species by Cassis and Gross (2002); and Oliarus
trispiralis by Löcker et al. (2006).
Twenty-four Hemipteran species are recognised as
endemic to Christmas Island (Table 2), of which three
have not been recorded since 1897–98, four not since
1933 and four not since 1964. No species are listed as
threatened on global or national lists.
Order Neuroptera
The Order Neuroptera (including lacewings and antlions) is poorly studied on Christmas Island, with six
species identified to date (New 1991). C.W. Andrews
collected two species that were described as new by
Kirby (1900e). Formicaleo morpheus has since been
71
dissolved under Distoleon somnolentus (New 1991)
(Table 4), whereas Myrmeleon iridescens is still
recognised as a valid species (New 1991), endemic to
Christmas Island (AFD). The CSIRO 1989 collection
contained six taxa, five wide-ranging species and one
probably undescribed species (Malada sp. nov.; Table
3), but not Myrmeleon iridescens (CSIRO Division of
Entomology 1990; New 1991). James (2007) recorded 13
morpho-species of winged adults (not identified beyond
order), mostly collected in light-traps.
Myrmeleon iridescens was described from a single
specimen collected in 1897–98, and has not been
recorded again, but it is not listed as threatened.
Order Coleoptera
Many beetles have been described from Christmas
Island, and most are still recognised as endemic, but
there has been no detailed review. As with many other
taxonomic groups, the most important collections were
by Maclear in 1887 (Waterhouse 1887), Lister in 1887
(Gahan 1889), Andrews in 1898–99 (Waterhouse et al.
1900), Tweedie in 1932 (Tweedie 1933) and CSIRO in
1989 (CSIRO Division of Entomology 1990), with very
limited other occasional collecting.
From ‘several specimens’ of beetles collected by
Maclear, Waterhouse (1887) described two species,
Chrysodema simplex and Piezonotus discoidalis
(subsequently transferred to the endemic genus
Rhyncholobus), both of which are still recognised as
valid species. Rhyncholobus discoidalis has no records
other than from Christmas Island (ALA, GBIF).
However, C. simplex was subsequently reported from
Cocos (Keeling) islands by Gibson-Hill (1950), but
it is likely that this represented an importation from
Christmas Island, with ‘teak’ Berrya trees.
Lister’s collection comprised about 70 specimens
representing about 20 species, with 10 not identified
to species and four described as new by Gahan (1889):
Paraegus listeri (now Aegus listeri), Ceresium nigrum,
Monohammus nativitatis (now Acalolepta nativitatis)
and Praonetha perplexa (now Pterolophia perplexa).
All are still recognised as valid species, and only one of
these has since been recorded beyond Christmas Island
(AFD, ALA, GBIF): the exception is A. nativitatis,
which was also reported from Cocos (Keeling) islands
by Gibson-Hill (1950), probably with importation from
Christmas Island of soil and timber, as for C. simplex.
Slipinski and Escalona (2016) also recognised Ceresium
nigrum as a valid endemic species.
Andrews’ 1897–98 collection was appreciably larger,
comprising 93 species, of which 12 were determined
only to genus, six were the species previously described
from Christmas Island, 25 were attributed to species
described from elsewhere and an impressive 50 were
newly described (Waterhouse et al. 1900). Of those 50
species, at least 35 are still recognised as valid species
in both ALA and GBIF (unless stated otherwise below)
and have never been recorded other than for Christmas
Island: Apatenia apicalis, Litocerus jordani (recognised
72
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
in GBIF), Xenocerus nativitatis (GBIF), Paranobium
posticum (now Clada posticum) (ALA), Neoptinus
parvus (with endemicity noted by Belles and Lawrence
(1990)), Bothrideres strigatus (now Ascetoderes
strigatus) (ALA), Orychodes andrewsi, Chrysobothris
andrewsi (GBIF), Ceresium quadrimaculatus (with
endemicity noted by Slipinski and Escalona (2016)),
Examnes affinis (now Ceresium affnis, with endemicity
noted by Slipinski and Escalona (2016)), Aegocidnus
exiguus (now Sciades exigua), Prinobius coxalis
(now Anomophysis coxalis), Olenecamptus basalis,
Rhyparida modesta, Rhyparida rossi, Cossonus
variipennis, Pachyops incertus, Phloeophagosoma
dubium (now Rhyncholosoma dubium), Camptorhinus
c r inipe s, Acic ne mi s a n dre wsi, R hy n cholob u s
andrewsi, Rhyncholobus rossi, Rhyncholobus vittatus,
Dryophthorus assimilis, Tetrigus murrayi, Anchastus
discoidalis, Megapenthese andrewsi, Hololepta
malleata, Plat ysoma lignarium (now Plat ylister
lignarium), Figulus rossi (with endemicity noted by
Monte et al. (2016)), with a possible record from Cocos
(Keeling) islands (Gibson-Hill 1950; Cassis et al. 1992)
likely to be a result of introductions from Christmas
Island), Laius tibialis (with endemicity noted by Liu et
al. (2015)), Shoguna striata (ALA), Protaetia andrewsi
(noted as ‘known only from type locality’ in AFD),
Phileurus convexus (now Eophileurus convexus) and
Oniscomorpha marmorata (now Leperina marmorata,
per ALA and noted as a valid species endemic to
Christmas Island by Kolibáč (2013)). Although there are
putative unpublished records for Apomecyna nigritarsis
from Java (http://www.cerambycoidea.com/forum/topic.
asp?TOPIC_ID=14880), ALA and GBIF recognise the
species as valid and include no records from beyond
Christmas Island.
We could find no documentation in GBIF or ALA
for several other species described from Christmas
Island by Waterhouse et al. (1900), but consider them
as valid and endemic (Table 2). Epliachna nativitatis
(now Henosepilachna nativitatis) is recognised as a valid
species restricted to Christmas Island by Jadwiszczak
and Wegrzynowicz (2003); Paederus listeri is retained
as a valid species by Frank (1988); Bradymerus
seminitidus is retained as a valid species endemic to
Christmas Island by Schawaller (2006); Amarygmus
funebris is retained as a valid species endemic to
Christmas Island by Bremer (2007); Opatrum dubium
(now Gonocephalum dubium) is retained as a valid
species endemic to Christmas Island by Iwan et al.
(2010). Demotina lateralis, Toxicum antilope, Sessinia
andrewsi and Nyctobates carbonaria (as Promethis
carbonaria) were recognised as valid species and
collected by CSIRO Division of Entomology (1990),
although a global review of Toxicum did not list antilope
(Doyen et al. 1989). CSIRO Division of Entomology
(1990) also recognised as valid species, but did not
collect, Psylliodes tenuepunctata and Sessinia listeri.
We consider the remaining species described by
Waterhouse et al. (1900) as no longer valid or not
endemic to Christmas Island. Rhabdocnemis fausti is
subsumed in the more wide-ranging R. obscurus in
ALA; Stelidota orientalis is subsumed in the more wideranging Omosita nigrovaria (GBIF, ALA); and Shoguna
polita is lumped in the more wide-ranging Shoguna
termitiformis (ALA, GBIF).
After Andrews’ 1897–98 collections, the next
published report on beetles was a brief account of a
small collection made by Tweedie in 1932 (Tweedie
1933). This comprised 28 species, of which 18 had
been described previously from Christmas Island,
seven had been described from elsewhere, two were not
determined to species, and one, Phelipara subvittata
(Cerambycidae), was newly described. It is still
recognised as a valid species with no known records
beyond Christmas Island.
The entomological survey by CSIRO in 1989 dwarfed
all previous survey efforts for beetles, collecting ‘about
10,000’ specimens representing 350 species’ (CSIRO
Division of Entomology 1990). This sampling included
representation of 77 of the c. 90 species recorded in the
1897–98 sampling (Waterhouse et al. 1900), and with
other previously-unpublished minor samples brought
the known tally of beetles on the island to 390 species.
CSIRO Division of Entomology (1990) also reported
that 14 species collected in the 1897–98 sampling
(Waterhouse et al. 1900) were not collected in 1989
and that ‘most of these are moderate-sized insects …
that would not be overlooked in general collecting and
may no longer occur on the island: however, given the
small amount of time devoted to surveying the fauna,
sampling error is more likely ’. About 37 of the species
collected in 1989 had been described earlier from
Christmas Island, and about 59 had been described from
elsewhere. Most (253) of the beetle species collected
during the 1989 survey were diagnosed only to genus,
and to our knowledge most of these collections have not
subsequently been described or identified to species.
However, there are some exceptions, with material
collected in the CSIRO survey of Christmas Island
used as the basis for the description of the endemic
Psammorpha lawrencei by Stebnicka (1994) (a blind
and flightless species, constituting a monotypic genus,
considered endemic to the island: Stebnicka and
Howden (1996)), Paratrichapus christmasensis (SouzaGonçalves et al. 2019) and Cephennomicrus lawrencei
(Jałoszyński 2017).
The only subsequent notable collection from
Christmas Island is of a single ‘remarkable’ beetle
collected from a cave in 2006, and described as
Lymnastis brooksi (Carabiidae) by Baehr (2008).
Table 2 lists 58 beetle species that we consider are
valid species reported only from Christmas Island.
Given that the CSIRO collection of 1989 also reported
253 undetermined species, it is highly likely that the
actual number of endemics is appreciably higher than
the 58 listed in Table 2. Rhyncholobus is an endemic
genus of large and brightly coloured weevils containing
four species, and it represents the most likely example of
autochthonous radiation to have occurred on Christmas
Island. Ten of the endemic species have not been
ENDEMIC SPECIES OF CHRISTMAS ISLAND
reported since 1897–98 (Table 5). Notwithstanding these
long periods without records, none of the endemic beetle
species is listed as threatened.
Order Strepsiptera
There are no endemic species recognised, and only a
single individual has been collected, in 1989. This was
a single specimen of stylops parasitising a delphacid
hemipteran, deduced to be in either Halictophagidae or
Elenchidae (CSIRO Division of Entomology 1990).
Order Diptera
The Diptera fauna of Christmas Island has been far
from comprehensively documented. Andrews collected
between 30 and 40 species in 1898–99, but noted that
these ‘unfortunately have not yet been determined’.
We can find no record of any subsequent substantial
collecting until 90 years later, when CSIRO collected
about 5000 specimens, ‘but only a few complete
identifications could be made’ (CSIRO Division of
Entomology 1990). These were sorted to about 185
morpho-species, but only 22 of these were determined
to species level and 63 to genus. The fauna now includes
many introduced human-commensal pests such as house
flies, fruit flies and mosquitoes (Campbell 1968; CSIRO
Division of Entomology 1990).
Nine species of Diptera have been described from
Christmas Island between 1889 and 1994 and eight of
these are considered endemic species (Table 2). Laphria
nigrocaerulea (Kirby, 1889) is a junior homonym of
Laphria nigrocaerulea (van der Wulp 1872) from New
Guinea, so was renamed Orthogonis christmasensis
by Daniels (2012); it is considered valid and endemic
(ALA, AFD, GBIF) and is still abundant (DJ pers. obs.).
The marine tipulid flies Cymatopus calcaratus and
Cymatopus longipilus were described by Parent (1935)
from a collection by Harms in 1933: the most recent
review of the genus considered both to be still valid
and endemic to Christmas Island (Evenhuis 2005). The
flower fly Syritta maritima was described by Hull (1944),
also from Harms’ 1933 collection, and the most recent
review of the genus concluded that it is still regarded as
a valid species endemic to Christmas Island (Lyneborg
and Barkemeyer 2005). Thompson (2014) reviewed the
family Syrphidae on Christmas Island, and concluded
that Syritta maritima was the only endemic species
among seven reported species, but considered it may be
extinct, with no records since the original collection in
1933. The ‘remarkable’ Poecilosomella pectiniterga was
described by Deeming (1964) with its holotype from
Andrews’ 1897 collection and other material from Biak
Island, with subsequent records from other locations.
However, the most recent review split the species and, as
redefined, considered that Poecilosomella pectiniterga
is restricted to Christmas Island (Hayashi 1997).
Micrepimera punctipennis described from a specimen
collected on Christmas Island in 1983 by Matile (1990) is
still recognised as a valid species with no records beyond
Christmas Island (GBIF). When described it comprised
a monotypic genus, but other species in the genus have
been described subsequently (Ševčík and Papp 2011).
73
Stilbomyia juncunda (now Hemipyrellia jucunda)
(Kirby 1889) is recognised as a valid species in GBIF
(but not ALA or AFD) with no records from elsewhere.
It was last collected by CSIRO in 1964 (Campbell
1968), and was explicitly noted as not collected
(notwithstanding a large collection of dipterans) by
CSIRO in 1989 (CSIRO Division of Entomology 1990).
Lissocephala powelli described from Christmas Island
by Carson and Wheeler (1973) is recognised as a valid
species in GBIF (but not ALA or AFD) with no records
from elsewhere. There are many recent records of
Lissocephala powelli on its host crabs (Orchard 2012).
Amblypsilopus natalis, described by Bickel (1994) from
material collected by CSIRO in 1989, is recognised as
a valid species endemic to Christmas Island in ALA,
AFD and GBIF. The mosquito Aedes andrewsi (Edwards
1927) has been recorded from Indonesia and Australia
(GBIF), and is not recognised by ALA or AFD, so is not
accepted here (Table 4).
None of the endemic dipteran species is listed as
threatened, nationally or globally.
Order Siphonaptera
The last specimens of the endemic Maclear’s rat Rattus
macleari were collected by Durham in 1902 (Durham
1908; Pickering and Norris 1996; Green 2014). Fleas
collected from these specimens were described as a new
species, Xenopsylla nesiotes (Jordan and Rothschild
1908). The species is accepted by AFD, ALA and GBIF
with no records beyond Christmas Island. Maclear’s rat
was the only known host for the flea, so the flea almost
certainly became extinct, unceremoniously, when the rat
did (Kwak 2018), between 1902 and 1907, shortly before
it was described by science. Maclear’s rat is listed as
extinct nationally and globally, but its host-specific flea is
not formally listed as extinct nationally or globally.
Order Lepidoptera
The first collection of Christmas Island butterflies,
by Maclear in 1887, was largely destroyed in transit,
but two specimens were intact and both were named
as new species by Butler (1887): Vadebra macleari and
Terias amplexa. In 1887, Lister collected five species
including the two previously described Christmas
Island species, a new species (Hypolimnas listeri) and
two previously described from elsewhere (Butler 1889).
Andrews collected another four species in 1897–98,
with one of these described as a new species by Butler
(1900): Charaxes andrewsi. No butterflies have been
described from the island subsequently, although other
more wide-ranging species (including some visitors)
have been added to the species complement (GibsonHill 1947e; Moulds and Lachlan 1987; Johnson and
Wilson 2018), which now numbers 29 species (Johnson
and Wilson 2018).
Of the four butterfly species for which Christmas
Island is the type locality, only one, Charaxes andrewsi,
is still regarded as a valid species endemic to Christmas
Island (e.g. ALA) (Table 2), although it has at times been
considered a subspecies of Eriboea pyrrhus (Pendlebury
74
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
1933; Gibson-Hill 1947e). The three other butterfly taxa
have had a buffeted taxonomic history. Terias amplexa
was relegated to a subspecies of Eurema hecabe by
Corbet and Pendlebury (1932), with this treatment
followed by Pendlebury (1933) and Gibson-Hill (1947e),
but treated as a distinct species by Moulds and Lachlan
(1987), then reduced to a subspecies of Eurema alitha
by Yata (1995), a treatment followed by ALA. Vadebra
macleari is still accepted by GBIF but synonymised with
the more wide-ranging Euploea climena in ALA; and
Hypolimnas listeri is synonymised with the more wideranging H. bolina (e.g. in GBIF) (Table 4).
In contrast to the relatively small and reasonably well
resolved butterfly fauna, the moth fauna of Christmas
Island is large and very inadequately catalogued, and
there has been no recent comprehensive review. Lister’s
collection in 1887 included five species (Butler, 1889),
of which two were described as new (Eudragana
limbata – with a new genus established for it, although
this was soon subsumed in Bocula (Hampson 1900)
– and Pyralis listeri), and one taxon (Hydrillodes sp.
nov.) considered ‘probably a new species’ but with
insufficient material to describe it (Butler 1889).
Andrews’ collection of 1897–98 was much more
extensive, comprising 74 species comprehensively
documented by Hampson (1900) and Walsingham
(1900). In characterising this material, Hampson (1900)
noted that ‘the distribution of Christmas Island moths
seems to show very clearly that it has no connection
with the Malayan sub-region’. The collection included
all five species previously recorded by Butler (1889)
and 54 species previously described from elsewhere.
It also included 15 new species: Mimeusemia econia,
Hydrillodes vexillefera (with the description noting
also records from beyond Christmas Island), Erastria
griseomixta, Porthesia pulverea, Boarmia scotezonea,
Ephestia scotella, Zinckenia nigerrimalis, Glyphodes
(Phacellura) holophaealis (all described by Hampson
(1900)) and Cosmoclostis quadriquadra, Brenthia
elachista, Simaethis ornaticornis, Tortricomorpha
chlorolepis, Epagoge halysideta, Caenognosis incisa (in
a newly established genus), and Dendroneura punctata
(all described by Walsingham (1900)). Two of Andrews’
specimens were attributed by Walsingham (1900) to
the African species Phycodes adjectella, but these have
subsequently been described by Kallies (2013) as the
Christmas Island endemic Nigilgia browni, with further
specimens collected recently (C. Pink pers. comm.).
A further eight originally unsorted specimens from
Andrews’ collection were described, with Christmas
Island as the type locality, over the next two decades in
more miscellaneous taxonomic reviews: Thalassodes
subviridis by Warren (1905), Maliattha phaeozona
and Amyna crocosticta by Hampson (1910), Armactica
andrewsi and Earias latimargo by Hampson (1912),
Comostolopsis regina by Thierry-Mieg (1915), Scopula
tumiditibia by Prout (1920), and Anomis esocampta by
Hampson (1926). A further description of a Christmas
Island endemic moth from Andrews’ original collection
– of Acontia sollemnis – was made more than 100 years
after the initial collection, by Hacker and Holloway (in
Hacker et al. 2008).
The next substantial collection was of 42 specimens
from nine species of geometrid moths collected by
Tweedie in August-September 1932 and documented by
Prout (1933), who described five new species: Hemithea
(Chlorissa) hyperymna (‘a charming little species’),
Anisodes hypomion, Ecliptopera phaula, Sauris pelagitis
[previously collected and identified by Hampson as the
more wide-ranging S. hirudinata] and Syrrhodia vindex
[previously collected and identified by Hampson as the
more wide-ranging Hyperythra lutea]. Prout also felt
‘reluctantly compelled to leave ... unnamed a distinct
Scopula species that Hampson had earlier listed as an
undetermined Craspedia species’.
A more substantial collection (of about 1740 moth
specimens) was gathered by Gibson-Hill in 1939–40 that
nominally included representation of 174 species, but the
description of this collection reported only on 74 of these
species, noting that ‘it is regrettable that the study of the
collection cannot be completed now, but the limitations
imposed as a result of the war render this impossible’
(Pendlebury 1947): probably as a consequence of that
interruption, and his death in 1943, no new species were
described from this collection.
CSIRO’s collection of Lepidoptera in 1990 included
about 900 specimens representing about 185 species
(of which about 16 were butterf lies), with species
attributions not made for most moths (CSIRO Division
of Entomology, 1990). This collection included nine
species previously described as endemic (Zinckenia
nigerrimalis (as Spoladea nigerrimalis), Glyphodes
(Phacellura) holophaealis (as Diaphania holophaealis),
Thalassodes subviridis, Comostolopsis regina, Anisodes
hypomion, Maliattha phaeozona, Amyna crocosticta,
Earias latimargo and Armactica andrewsi. However,
most of the collection was sorted to family or genus level
only. CSIRO Division of Entomology (1990) also noted
that 22% of the 185 species were represented in their
collection by only single individuals, and interpreted this
to mean that much of the fauna remained uncollected,
most likely including more species of endemics. From
2004 to 2006, James (2007) collected 141 morphospecies of moths, but only five of these were identified to
named species.
Of the 32 moth species described from Christmas
Island, the status (whether or not still considered a
valid species, and whether or not reported subsequently
away from Christmas Island) of many is unclear.
Here, we consider 24 as endemic (Table 2). Nineteen
of these are regarded as valid species in GBIF with no
records beyond Christmas Island: Ephestia scotella,
Ruttelerona scotozonea (= Boarmia scotezonea),
Hymenia nigerrimalis (= Zinckenia nigerrimalis),
Cosmoclostis quadriquadra, Brenthia elachista, Moca
chlorolepis (= Tortricomorpha chlorolepis), Opogona
punctata (= Dendroneura punctata), Thalassodes
subviridis (= Pelagodes subviridis), Armactica andrewsi,
Bocula limbata, Earias latimargo, Comostolopsis
regina, Scopula tumiditibia (with endemicity noted
ENDEMIC SPECIES OF CHRISTMAS ISLAND
by Sihvonen (2005), Loboschiza halysideta, Nigilgia
browni, Hemithea hyperymna, Anisodes hypomion,
Ecliptopera phaula and Sauris pelagitis. One is listed
as a valid (though ‘doubtful’) species in GBIF, with no
records beyond Christmas Island: Lithacodia griseomixta
(= Erastria griseomixta). Three are not listed in GBIF
but recognised elsewhere as valid and without records
beyond Christmas Island: Diaphania holophaealis
(http://globiz.pyraloidea.org), Choreutis ornaticornis
(= Simaethis ornaticornis) (http://choreutidae.myspecies.
info/taxonomy/term/66) and Amyna crocosticta (http://
www.nic.funet.fi/pub/sci/bio/life/insecta/lepidoptera/
ditrysia/noctuoidea/noctuidae/acontiinae/amyna/index.
html). The recently described Acontia sollemnis has not
been reported from elsewhere (Hacker et al. 2008).
Five of the 25 accepted endemic lepidopteran species
have not been reported since 1897–98 and a further
six not since 1939 (Table 2). Notwithstanding these
long periods without records, none of the endemic
lepidopteran species is listed as threatened, either
nationally or globally.
Order Hymenoptera
A comprehensive recent review, based on intensive
and extensive field surveys and a re-appraisal of
historic records listed 52 species of ants from the
island (Framenau and Thomas 2008). None of these is
considered endemic. Most of the ant species reported
from the island are introduced (Framenau and Thomas
2008). Christmas Island is the type locality for three ant
species – Camponotus melichlorus, Leptogyne harmsi
and Pachycondyla christmasi (Kirby 1889; Donisthorpe
1935), but all three species are now known to also occur
naturally in other areas (Framenau and Thomas 2008).
The number of native species is indeterminate, but only
two (presumed native, but not endemic) ant species were
recorded immediately before and after the island’s first
settlement (Kirby 1889; Andrews 1900c).
There has never been a comprehensive treatment of
the bees and wasps of Christmas Island. Collecting
commenced pre-settlement with J.J. Lister in 1887 with
two species (Euodynerus polyphemus and Polistes
(Gyrostoma) balder) from this collection described
by Kirby (1889). Five further species (Homalictus
andrewsi, Patellapis (Pachyhalictus) binghami,
Ophion flavocephalus (now Enicospilus flavocephalus),
Megachile nivescens, Megachile rotundipennis) were
described from Andrews’ 1897–98 collections (Kirby
1900c), and another (Lithurgus andrewsi) from the same
collection was described by Cockerell (1909). Other than
Enicospilus flavocephalus, all of these species are still
regarded as valid and endemic to Christmas Island (e.g.
Walker 1997).
Carl Gibson-Hill collected further specimens in
1939–40, but most were not subsequently worked over;
however two species were described from this material
about 50 years later. Bouček (1988) described Sirovena
stigma, originally considered to comprise a monotypic
genus until Li et al. (2013) added another species, from
China. Zettel (1990) described Phanerotoma pacifica
75
from Gibson-Hill’s collection, although its specific name
indicates confusion with the other Christmas Island in
the Pacific.
Campbell (1968) recorded 11 non-ant hymenoptera
in a survey focussed on insects of medical, veterinary,
agricultural and forestry significance, but notably did
not list the honey bee Apis mellifera, dating its likely
introduction to the island to sometime during the next 20
years to 1989, when ANIC-CSIRO collected about 2700
specimens representing 299 species (CSIRO Division of
Entomology 1990). Eight taxa were named to species,
105 to genus, and 190 as unnamed genera (CSIRO
Division of Entomology 1990). These collections have
scarcely been worked upon, having yielded just five
additional species records for the island, and five species
new to science, including three endemics: Dendrosotinus
insularis (Belokobylskij et al. 2004), Neoheterospilus
insularis (known from a single specimen: Belokobylskij
(2006)) and Oxyscelio caesitas (Burks et al. 2013).
James (2007) reported 68 morpho-species of wasps
collected by Parks Australia in 2004 to 2006, but
identified only Polistes balder to species level. More
recently, a large collection of parasitoid wasps was made
by the CESAR Consulting Group (Weeks 2013) as part
of their assessment of the potential non-target impacts of
fipronil insecticide that has been used to control yellow
crazy ants across the island. Using yellow sticky traps
in 2012, they collected 3,390 specimens of parasitoids
in 19 families. None were identified to genus or species.
Three parasitoid species were identified from the island
as part of the research program supporting the indirect
biological control of the yellow crazy ant (Green et al.
2014), but these are not regarded as native to the island.
The total list of species of non-ant hymenoptera
stands at 34, 12 of which we regard as endemics (Table
2). However it is clear that given the great diversity of
non-ant Hymenoptera on Christmas Island especially
amongst the wasps, and the relative paucity of named
species in the ANIC-CSIRO, Parks Australia and
CESAR collections, there are likely to be more endemics
on the island.
Our list of endemics includes two large predatory wasp
species (Vespidae) that also occur in the Cocos (Keeling)
Islands, 900 km further west. Polistes balder Kirby was
listed by Richards (1978) as occurring there, and he refers
to a specimen in the BMNH but does not give details. We
cannot find this record, so do not know the collection
date. These wasps are large and aggressive and build
conspicuous paper nests, but the species was not recorded
from the islands by Darwin (1845), Wood Jones (1909) or
Gibson-Hill (1950). Gibson-Hill especially was unlikely
to have overlooked this species during his extensive
sampling, so we conclude this species must have been
introduced there from Christmas Island more recently.
Gibson-Hill (1950) gives a good account of how insects
might have been transported from Christmas Island to
the Cocos (Keeling) Islands, in this case perhaps as a
nest attached to some structure and transported on a ship.
Eudynerus polyphemus Kirby was mentioned by CSIRO
Division of Entomology (1990) as being known from
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
76
Home Island in the Cocos (Keeling) islands group, but
gave no further detail. Similar to P. balder, this is a large
and conspicuous species that often builds its mud nests on
buildings, but was not recorded by earlier naturalists. The
same reasoning holds – we conclude it was introduced to
the Cocos (Keeling) Islands from Christmas Island.
Of the 12 recognised endemic species, eight have
been collected relatively recently as part of the 1989
ANIC/CSIRO collections, with Lithurgus andrewsi
being a possible but unconfirmed ninth species (CSIRO
Division of Entomology 1990). Both species of Megachile
were collected by Campbell in 1968, but only single
specimens were collected in 1989 and 2004 and these
were not determined to species level. Finally, the endemic
halictid Patellapis (Pachyhalictus) binghami has not
been collected since Campbell in 1968. Walker (1997)
noted that Josephine Cardale, the CSIRO hymenoptera
specialist on the 1989 expedition, was not able to collect
this halictid despite considerable dedicated effort, and he
speculated that it might be extinct.
Phylum ECHINODERMATA
Marsh (2000) reviewed the Echinodermata (including
four collections by the Western Australian Museum)
and provided a list of 67 species across the five Classes.
She considered that this list was probably not complete
but contained the majority of species present. Tan et
al. (2014b) added a further six species from a limited
survey of submarine caves. The echinoderm fauna is
relatively depauperate, probably owing to a combination
of the island’s small size, relative isolation and limited
diversity of marine habitats (Marsh 2000). No species
are considered endemic to Christmas Island. One species
of crinoid, Comissia pectinifer (now Alloeocomatella
pectinifera), was described from the island by Clark
(1911), but it has since been found to be more widespread
(GBIF; Table 4).
Phylum HEMICHORDATA
Tan et al. (2014b) recorded one species from submarine
caves, but identified it only to the Class Enteropneusta.
Phylum CHORDATA
The classes of the Subphylum Vertebrata are treated
separately below. The only other chordates reported are
salps and doliolids (Tunicata: Thaliacea), which are a
common element of the inshore plankton (Davies and
Beckley 2010).
CLASS PISCES
The fish fauna in waters of and around Christmas
Island has now been well sampled. Early collections
were limited: for example, Andrews (1900c) collected
only 21 species. The inventory increased substantially
with targeted sampling from the 1960s. By 2007, 592
species were recorded (Allen et al. 2007), with a recent
comprehensive review extending this to 681 species
(Hobbs et al. 2014b). Only five of these species occur
in the island’s freshwater systems, three of which
are introduced and two of which are native but occur
widely beyond the island (Hobbs et al. 2014b). Allen
et al. (2007) considered four marine fish species to be
endemic to Christmas Island: Pseudochromis viridis,
Praealticus natalis, Eviota natalis and Aseraggodes
crypticus. The text of Hobbs et al. (2014b) corroborates
that these are the four endemic fish species, but their
accompanying checklist (Hobbs et al. 2014b, Table 1)
also reports two additional recently-described species
as endemic, and known only from their holotypes:
Microbrotula andersoni, collected in 1978 (Schwarzhans
and Nielsen 2011) and Paradiancistrus christmasensis,
collected in 1986 (Schwarzhans and Møller 2011).
Another species, Steeneichthys nativitatis, described
in 1987 from Christmas Island, is now known to be
more widespread (Allen et al. 2007). GBIF and ALA
recognise as valid species all of the six endemics
reported by Hobbs et al. (2014b), but both databases also
have records of Praealticus natalis from well beyond
Christmas Island, extending at least to Guam: so we do
not consider it as a Christmas Island endemic here.
All of the five endemic fish species have been recorded
recently, and none is listed globally or nationally as
threatened.
CLASS AMPHIBIA
There are no native or established amphibians on
Christmas Island. Two species have been intercepted
in cargo shipments on rare occasions (Parks Australia,
unpublished data): Duttaphr ynus melanostictus
(Bufonidae) and Litoria adelaidensis (Hylidae) (DJ, pers.
obs.).
CLASS REPTILIA
The Christmas Island native terrestrial reptile fauna
comprises five lizard species and one blind-snake. The
taxonomic treatment of some of the lizard species has
been unstable (Boulenger 1887, 1889; Smith 1929), but
authorities consistently now recognise Cyrtodactylus
sadleiri, Lepidodactylus listeri, Cryptoblepharus
egeriae and Emoia nativitatis as endemic (Cogger and
Sadlier 1981; Cogger 2014), as well as the blind-snake
Ramphotyphlops exocoeti. All are species recognised by
GBIF, and all locational data in GBIF for these species
are from Christmas Island. The only other native species
is generally considered to be the wide-ranging Emoia
atrocostata (Cogger and Sadlier 1981; Cogger et al. 1983;
Brown 1991), but may merit closer taxonomic scrutiny.
Recent analyses demonstrate that Lepidodactylus listeri,
Cryptoblepharus egeriae and Emoia nativitatis are of
remarkable antiquity, separated from their closest sampled
relatives for 5–20 million years (Oliver et al. 2018).
All five endemic reptile species are considered
threatened at global and national levels (Table 1), having
exhibited severe and rapid decline since the 1980s (Smith
et al. 2012; Andrew et al. 2018). Emoia nativitatis is now
extinct, and Lepidodactylus listeri and Cryptoblepharus
egeriae are extinct in the wild. The Christmas Island
population of Emoia atrocostata was also extirpated over
this period (Andrew et al. 2018).
ENDEMIC SPECIES OF CHRISTMAS ISLAND
CLASS AVES
The birds of Christmas Island have been well-studied
with several reviews, the most recent being James and
McAllan (2014). At the time of colonisation there were
eight seabird and eight landbird species breeding, but
that had increased to nine and 14 species respectively by
1982, due to both colonisation and introduction (Stokes
1988). Four of the seabirds and seven of the landbirds
are endemic taxa, but not all are full species. A further
complication is that the seabird taxa disperse widely
as a routine part of their life histories. Six species are
considered endemic (Table 2).
Of the landbirds, the Christmas Island imperial pigeon
Ducula whartoni, Christmas Island swiftlet Collacalia
natalis, Christmas Island hawk-owl Ninox natalis and
Christmas Island white-eye Zosterops natalis were each
described as full species (Sharpe 1887; Lister 1889).
The Zosterops is the only one to be universally treated
as a full species ever since. However, because it has a
small, introduced subpopulation in the Cocos (Keeling)
Islands (Woinarski et al. 2014), we characterise it here
as an ‘original endemic’. The three other species were
subsumed as subspecies of Wallacean congeners by
Chasen (1933a) and Chasen (1935), and this was followed
internationally, for example by Peters (1940) and locally
by Gibson-Hill (1947g). Ducula whartoni was briefly
considered to be a subspecies of Ducula rosacea, but
otherwise has been regarded as full species by recent
authorities (e.g. Christidis and Boles 2008) and is treated
as such by ALA and GBIF. Ninox natalis was listed
as a subspecies of N. forbesi and then N. squamipila,
before being elevated to a full species again by Norman
et al. (1998). Since Chasen (1933a), Collacalia natalis
has usually been treated as a subspecies of Collacalia
esculenta (e.g. ALA, AFD, GBIF), but also Collacalia
linchi (e.g. Christidis and Boles 2008). A recent study
integrating molecular and morphological data justified its
full species status (Rheindt et al. 2017), which has been
adopted by GBIF.
As with Zosterops natalis, attempts were made by
the Clunies Ross family to translocate populations of
Ducula whartoni, and also the endemic subspecies of
island thrush Turdus poliocephalus erythropleurus
from Christmas Island to islands in the Cocos (Keeling)
group (Gibson-Hill 1949a), but the pigeon and thrush
introductions ultimately failed (Stokes et al. 1984).
The type specimen of Abbott’s booby Papasula abbotti
was collected on Assumption Island in 1893 where it
probably bred. Evidence (mostly from subfossils) indicates
that prehistorically it bred widely in the tropical IndoPacific (James and McAllan 2014). It now breeds only
on Christmas Island. The Christmas Island frigatebird
Fregata andrewsi was described as a full species, and
that has never been challenged. Both of these seabirds
are breeding endemic species that disperse widely when
feeding or not breeding (James and McAllan 2014).
There are also five endemic subspecies of birds. Three
taxa conventionally regarded as subspecies are: ‘golden’
white-tailed tropicbird Phaethon lepturus fulvus (though
77
it has no type locality); Christmas Island emerald dove
Chalcophaps indica natalis; and Christmas Island thrush
Turdus poliocephalus erythropleurus. The Christmas
Island goshawk Accipiter hiogaster natalis is sometimes
treated as a full species (e.g. James and McAllan 2014). A
fifth endemic subspecies, Fregata minor listeri, is widely
overlooked due to a nomenclatural error (see James and
McAllan 2014), so further taxonomic review is warranted.
Above species level, there is one endemic genus of
birds, Papasula (Olson and Warheit 1988), which is
monotypic.
Conservation-wise, the birds have fared well compared
with the other vertebrate groups, with no extinctions
despite some declines (Garnett et al. 2011; James and
McAllan 2014). All endemic taxa are recorded regularly
(James and McAllan 2014); nonetheless, 11 endemic bird
taxa are listed as threatened, that being three species
globally, and seven species and four subspecies nationally
(Table 1).
CLASS MAMMALIA
Collections around the time of the island’s settlement
reported five terrestrial native mammal species (two
rodents, two bats and a shrew) from Christmas Island.
Rattus nativitatis and R. macleari have been consistently
accepted as valid species and endemic (Thomas 1887,
1889; Andrews 1900b; Jackson and Groves 2015). The
shrew has had a mercurial taxonomic history, but the
most recent review confirmed its status as an endemic
species Crocidura trichura (Eldridge et al. 2014), and
this is conventionally accepted (Jackson and Groves
2015). The island’s only insectivorous bat was described
as an endemic species, Pipistrellus murrayi by (Andrews
1900b), but was subsequently subsumed within the more
wide-ranging P. tenuis (Koopman 1973). This treatment
is no longer accepted, and P. murrayi is now universally
accepted as a valid species endemic to Christmas Island
(Jackson and Groves 2015). The island’s flying-fox was
described as an endemic species, Pteropus natalis by
Thomas (1887), but reduced to an endemic subspecies
of the more widespread P. melanotus by Chasen
(1940). This conclusion has been accepted by some
but not all subsequent accounts, with a recent analysis
remaining equivocal (Phalen et al. 2017). The most recent
taxonomic review of Australian mammals accepted it as
a valid species endemic to Christmas Island (Jackson and
Groves, 2015), but noted a need for further taxonomic
assessment. Hence, the terrestrial native mammal species
of Christmas Island comprise either five endemic species,
or four endemic species and one endemic subspecies.
The Christmas Island mammal assemblage has not
fared well. The two Rattus species became extinct within
two decades of the island’s settlement (Andrews 1909;
Wyatt et al. 2008; Green 2014). The pipistrelle became
extinct in 2009 (Lunney et al. 2011; Martin et al. 2012;
Woinarski et al. 2017). The two remaining species
(the shrew and flying-fox) are recognised globally and
nationally as threatened (Table 1), and, with no records
since 1985, the shrew is probably extinct.
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
78
TABLE 1A
Christmas Island endemic plant and animal species that are listed as threatened at national or global level.
EPBCA-listed status refers to the Australian listing (as at December 2019) under the Environment Protection
and Biodiversity Conservation Act.
Species
National status
Global (IUCN) status
Plants
Endangered
Lister’s palm Arenga listeri
Invertebrates
Vulnerable
Christmas Island stick-insect Ramulus stilpnoides
Reptiles
Lister’s gecko Lepidodactylus listeri
Critically endangered
Extinct (in the wild)
Christmas Island giant gecko Cyrtodactylus sadleiri
Endangered
Endangered
Blue-tailed skink Cryptoblepharus egeriae
Critically endangered
Extinct (in the wild)
Forest skink Emoia nativitatis
Critically endangered
Extinct
Christmas Island blind snake Ramphotyphlops exocoeti
Vulnerable
Endangered
Christmas Island frigatebird Fregata andrewsi
Endangered
Critically endangered
Abbott’s booby Papasula abbotti
Endangered
Endangered
Christmas Island hawk-owl Ninox natalis
Vulnerable
Vulnerable
Maclear’s rat Rattus macleari
Extinct
Extinct
Bulldog rat Rattus navitatis
Extinct
Extinct
Christmas Island pipistrelle Pipistrellus murrayi
Critically endangered
Extinct
Christmas Island shrew Crocidura trichura
Critically endangered
Critically endangered
Birds
Mammals
TABLE 1B
Christmas Island endemic plant and animal subspecies that are listed as threatened at national level (note
that subspecies are not listed at global level).
* Pteropus natalis is treated here as an endemic species, although under both national and global
threatened species listings, it is treated as a subspecies.
Subspecies
National status
Plants
Tectaria devexa var. minor
Endangered
Birds
White-tailed tropicbird Phaethon lepturus fulvus
Endangered
Christmas Island emerald dove Chalcophaps indica natalis
Endangered
Christmas Island goshawk Accipiter (hiogaster) natalis
Endangered
Christmas Island thrush Turdus poliocephalus erythropleurus
Endangered
Mammals
Christmas Island flying-fox Pteropus (melanotus) natalis
Critically endangered*
List of species considered endemic to Christmas Island. Year of last collection is given for species not recorded in the last 50 years, otherwise is given simply as ‘Recent’.
Note that synonyms are not given exhaustively for all species.
Class
Order
Family
Species
Authority
Synonyms
Last known
collection date
Haemosporida
Haemoproteidae
(Plasmodiidae
in ALA)
Haemoproteus valkiunasi
Merino et al., 2012
Recent
Gasteromycetes
Lycoperdales
Geastraceae
Geastrum andrewsii
Blackman, 1900
1897–98
(although see text
for possible more
recent records)
Pucciniomycetes
Pucciniales
Pucciniaceae
Aecidium alchorneae-rugosae
Gjaerum and D.A.Reid, 1983
Recent
Eurotiomycetes
Pyrenulales
Pyrenulaceae
Lithothelium quiescens
P.M.McCarthy, 2001
Recent
Eurotiomycetes
(Dothideomycetes)
Strigulales
Strigulaceae
Strigula elixii
P.M.McCarthy, 2001
Recent
Eurotiomycetes
(Dothideomycetes)
Strigulales
Strigulaceae
Strigula natalis
P.M.McCarthy, 2001
Recent
Lecanoromycetes
Ostropales
Porinaceae
Trichothelium oceanicum
P.M.McCarthy, 2001
Recent
Bryopsida
Hypnales
Hypnaceae
Ectropothecium micronesiense
Fleisch ex Gepp and Gepp, 1905
1904
Bryopsida
Hypnales
Pylaisiadelphaceae
Isopterygium jelinkii
Fleisch ex Gepp and Gepp, 1905
1904
Equisetopsida
Arecales
Arecaceae
Arenga listeri
Beccari, 1891
Recent
Equisetopsida
Asparageles
Orchidaceae
Brachypeza archytas
(Ridley) Garay, 1972
Recent
Equisetopsida
Asparageles
Orchidaceae
Flickingeria nativitatis
(Ridley) J.J. Wood, 1982
Recent
Equisetopsida
Asparageles
Orchidaceae
Phreatia listeri
Rolfe, 1890
Recent
Equisetopsida
Asparageles
Orchidaceae
Zeuxine exilis
Ridley 1906
Equisetopsida
Pandanales
Pandanaceae
Pandanus christmatensis
Martelli 1905
Protista
Aconoidasida
Fungi
ENDEMIC SPECIES OF CHRISTMAS ISLAND
TABLE 2
Lichen
Bryophyta
Vascular Plants
Recent
Recent
79
[also described as
P. nativitatis Ridl.]
Order
Family
Species
Authority
Synonyms
Last known
collection date
Equisetopsida
Pandanales
Pandanaceae
Pandanus elatus
Ridley, 1906
Recent
Equisetopsida
Poales
Poaceae
Ischaemum nativitatis
Jansen ex Renvoize, 1985
Recent
Equisetopsida
Apiales
Pittosporaceae
Pittosporum nativitatis
Baker f., 1900
Recent
Equisetopsida
Gentianales
Apocynaceae
Hoya aldrichii
Hemsley, 1890
Recent
Equisetopsida
Lamiales
Acanthaceae
Asystasia alba
Ridley, 1906
Recent
Equisetopsida
Lamiales
Acanthaceae
Dicliptera maclearii
Hemsley, 1890
1960s
Equisetopsida
Laurales
Hernandaceae
Illigera elegans
Duyfjes, 1994
Recent
Equisetopsida
Malvales
Malvaceae
Abutilon listeri
Baker f., 1893
Recent
Equisetopsida
Malvales
Malvaceae
Grewia insularis
Ridley, 1906
Recent
Equisetopsida
Piperales
Piperaceae
Peperomia rossii
Rendle ex Baker f., 1900
1898
Equisetopsida
Rosales
Rhamnaceae
Colubrina pedunculata
Baker f., 1900
Recent
Demospongiae
Haplosclerida
Coelosphaeridae
Oceanapia sessilis
(Kirkpatrick, 1900)
Rhizochalina sessilis
1897–98
Demospongiae
Haplosclerida
Haliclonidae
Haliclona innominata
(Kirkpatrick, 1900)
Reniera innominata
1897–98
Bivalvia
Mytiloida
Mytilidae
Septifer rufolineatus
(Smith, 1911)
Brachydontes
(Hormomya) rufolineatus
Recent
Gastropoda
Orthogastropoda
Seguenziidae
Anxietas perplexa
Iredale, 1917
Gastropoda
Cerithimorpha
Plesiotrochidae
Plesiotrochus fischeri
Smith, 1909
Gastropoda
Cerithimorpha
Scaliolidae
Finella rugosa
(Laseron, 1956)
Gastropoda
Hypsogastropoda
Aclididae
Cyclonidea carina
Laseron, 1956
Gastropoda
Hypsogastropoda
Pickworthiidae
Chrystella islandica
Laseron, 1956
Gastropoda
Hypsogastropoda
Rissoidae
Rissoina isolata
(Laseron, 1956)
Costalynia isolata
1916
Gastropoda
Hypsogastropoda
Rissoidae
Schwartziella delicatula
(Laseron, 1956)
Pandalosia delicatula
1916
Gastropoda
Hypsogastropoda
Rissoidae
Zebina acicula
Laseron, 1956
1916
Gastropoda
Hypsogastropoda
Rissoidae
Zebina constricta
Laseron, 1956
1916
Gastropoda
Hypsogastropoda
Eulimidae
Pyramidelloides viticula
Laseron, 1956
1916
Gastropoda
Littorinimorpha
Assimineidae
Paludinella andrewsiana
(Smith, 1900)
80
Class
Porifera
1932
1932
Obtortio rugosa
1916
1916
1916
Assiminea andrewsiana,
Angustassiminea
andrewsiana
Recent
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Mollusca
Order
Family
Species
Authority
Synonyms
Last known
collection date
Gastropoda
Caenogastropoda
Triphoridae
Mastonia anomala
Laseron, 1958
Gastropoda
Caenogastropoda
Triphoridae
Mastoniaeforis decorata
(Laseron, 1958)
Epiforis decorata
1916
Gastropoda
Caenogastropoda
Triphoridae
Mastoniaeforis radix
(Laseron, 1958)
Epiforis radix
1916
Gastropoda
Caenogastropoda
Triphoridae
Nanaphora minuta
Laseron, 1958
1916
Gastropoda
Caenogastropoda
Triphoridae
Subulophora marginata
Laseron, 1958
1916
Gastropoda
Caenogastropoda
Triphoridae
Subulophora virgina
Laseron, 1958
1916
Gastropoda
Caenogastropoda
Cerithiopsidae
Potenatomus secundus
Laseron, 1956
1916
Gastropoda
Caenogastropoda
Fasciolariidae
Peristernia venusta
Smith, 1911
Recent
Gastropoda
Systellommatophora
Veronicellidae
Semperula insularis
Thomé, 1983
Recent
Gastropoda
Stylommatophora
Succineidae
Succinea solitaria
Smith, 1887
Recent
Gastropoda
Stylommatophora
Vertiginidae
Nesopupa proscripta
(Smith, 1905)
Gastropoda
Stylommatophora
Euconulidae
Kaliella cruda
Smith, 1909
Gastropoda
Stylommatophora
Euconulidae
Lamprocystis mabelae
(Smith, 1889)
Ariophanta mabelae,
Microcystis mabelae,
Sitala mabelae
1932
Gastropoda
Stylommatophora
Euconulidae
Lamprocystis mildredae
(Smith, 1889)
Ariophanta mildredae,
Microcystis mildredae,
Sitala mildredae
Recent
Gastropoda
Stylommatophora
Euconulidae
Lamprocystis normani
(Smith, 1889)
Ariophanta normani,
Microcystis normani,
Sitala normani
Recent
Ostracoda
Halocyprida
Thaumatocyprididae
Humphreysella baltanasi
Kornicker ex Humphreys
et al., 2009
Danielopolina baltanasi
Recent
Ostracoda
Podocopida
Cytheruridae
Microceratina martensi
Namiotko et al., 2004
Recent
Ostracoda
Podocopida
Darwinulidae
Isabenula humphreysi
Rossetti et al., 2011
Recent
Stomatopoda
Protosquillidae
Chorisquilla quinquelobata
(Gordon, 1935)
Gonodactylus
quinquelobatus
Recent
Malacostraca
Decapoda
Gecarcinidae
Tuerkayana celeste
(Ng and Davie, 2012)
Discoplax celeste
Recent
Malacostraca
Decapoda
Gecarcinidae
Gecarcoidea natalis
(Pocock 1889)
Hylaeocarcinus natalis
Recent
1916
Jaminia proscripta
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
Recent
Recent
Ostracoda
Stomatopoda
Malacostraca
Decapoda
81
Order
Family
Species
Authority
Synonyms
Last known
collection date
Malacostraca
Decapoda
Grapsidae
Karstarma jacksoni
(Balss, 1934)
Sesarma jacksoni
Recent
Malacostraca
Decapoda
Sesarmidae
Chiromantes garfunkel
Davie and Ng, 2013
Recent
Malacostraca
Decapoda
Varunidae
Orcovita orchardorum
Davie and Ng, 2012
Recent
Malacostraca
Decapoda
Varunidae
Orcovita hicksi
Davie and Ng, 2012
Recent
Malacostraca
Decapoda
Christmaplacidae
Christmaplax mirabilis
Naruse and Ng, 2014
Malacostrata
Decapoda
Pilumnidae
Echinoecus sculptus
(Ward, 1934)
Proechinoecus sculptus
Recent
Malacostrata
Decapoda
Palaemonidae
Macrobrachium xmas
Fujita, Davie and Ng, 2015
M. microps (in error)
Recent
Malacostraca
Decapoda
Procarididae
Procaris noelensis
Bruce and Davie, 2006
Recent
Malacostraca
Isopoda
Oniscidae
Tylos nudulus
Budde-Lund, 1906
1897–98
Arachnida
Amblypygi
Charontidae
Charon gervaisi
Harvey and West, 1998
Recent
Arachnida
Acari
Ixodidae
Ixodes nitens
Neumann, 1904
Extinct c. 1904
Arachnida
Pseudoscorpiones
Atemnidae
Paratemnoides pococki
(With, 1907)
Chelifer pococki,
Paratemnus pococki
Recent
Arachnida
Scorpiones
Hormuridae
Hormurus polisorum
(Volschenk, Locket
and Harvey, 2001)
Liocheles polisorum
Recent
Arachnida
Araneae
Barychelidae
Idioctis xmas
Raven, 1988
Recent
Arachnida
Araneae
Segestriidae
Ariadna natalis
Pocock, 1900
1897–98
Arachnida
Araneae
Filistatidae
Filistata gibsonhilli
Savory, 1943
1939
Arachnida
Araneae
Sparassidae
Heteropoda listeri
Pocock, 1900
1939
Scolopendromorpha
Cryptopidae
Cryptops inermipes
Pocock, 1889
1887
Diplopoda
Spirostreptida
Iulomorphidae
Hypocambala exocoeti
(Pocock, 1889)
Diplopoda
Penicillata
Lophoproctidae
Lophoturus speophilus
Nguyen Duy-Jacquemin, 2014
Recent
Diplopoda
Penicillata
Lophoproctidae
Lophoturus humphreysi
Nguyen Duy-Jacquemin, 2014
Recent
Phasmida
Phasmatidae
Ramulus stilpnoides
(Kirby, 1889)
82
Class
Recent
Arachnida
Chilopoda
Diplopoda
Iulomorpha exocoeti,
Spirostreptus exocoeti
1933
Phasmida
Insecta
Clitumnus stilpnoides,
Baculum stilpnoides
Recent
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Chilopoda
Order
Family
Species
Authority
Synonyms
Last known
collection date
Blattodea
Nocticolidae
Metanocticola christmasensis
Roth, 1999
Insecta
Orthoptera
Acrididae
Nisiocatantops orientalis
(Kirby, 1889)
Insecta
Orthoptera
Gryllacrididae
Gryllacris rufovaria
Kirby, 1889
Insecta
Orthoptera
Mogoplisticidae
Ornebius flavipalpis
(Kirby, 1900)
Ecatoderus flavipalpis
1897–98
Insecta
Orthoptera
Tettigoniidae
Paraphisis listeri
(Kirby, 1889)
Phisis listeri
Recent
Insecta
Orthoptera
Tettigoniidae
Psyrana pomona
(Kirby, 1900)
Psyra pomona
Recent
Insecta
Dermaptera
Spongiphoridae
Paralabella murrayi
(Kirby, 1900)
Labia murrayi
1897–98
Insecta
Dermaptera
Anisolabididae
Anisolabis subarmata
(Kirby, 1900)
Labia subarmata
1897–98
Zoraptera
Zorapteridae
Zorotypus lawrencei
New, 1995
Recent
Insecta
Psocodea
Amphientomidae
Stimulopalpus distinctus
Smithers, 1995
Recent
Insecta
Psocodea
Caeciliusidae
Paracaecilius cardaleae
Smithers, 1995
Recent
Insecta
Psocodea
Caeciliusidae
Caecilius singularis
Smithers, 1995
Recent
Insecta
Psocodea
Calopsocidae
Calopsocus yatesi
Smithers, 1995
Recent
Insecta
Psocodea
Lepidopsocidae
Echmepteryx lawrencei
Smithers, 1995
Recent
Insecta
Psocodea
Myopsocidae
Myopsocus brunneipes
Smithers, 1995
Recent
Insecta
Psocodea
Pachytroctidae
Pachytroctes sinuosus
Smithers, 1995
Recent
Insecta
Psocodea
Pachytroctidae
Tapinella curvatoides
Smithers, 1995
Recent
Insecta
Psocodea
Pseudocaeciliidae
Levucaecilius hilli
Smithers, 1995
Recent
Insecta
Psocodea
Psocidae
Sundapsocus robertsoni
Smithers, 1995
Recent
Insecta
Hemiptera
Cercopidae
Clovia eximia
Kirby, 1900
Recent
Insecta
Hemiptera
Cicadidae
Oxypleura calypso
Kirby, 1889
Blattodea
Insecta
Recent
Orthoptera
Primnia orientalis,
Catantops orientalis, Oxya
orientalis
1964
Recent
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
Dermaptera
Zoraptera
Insecta
Psocodea
Hemiptera
Recent
83
Platypleura calypso,
Poecilosaltria calypso
Order
Family
Species
Authority
Synonyms
Last known
collection date
Insecta
Hemiptera
Cicadellidae
Batracomorphus punctatus
(Kirby, 1900)
Idiocerus punctatus,
Bythoscopus punctatus
Recent
Insecta
Hemiptera
Cicadellidae
Erythroneura harmsi
Izzard, 1936
Recent
Insecta
Hemiptera
Cixiidae
Oliarus trispiralis
Löcker, 2006
Recent
Insecta
Hemiptera
Naucoridae
Laccocoris montandoni
Izzard, 1936
1933
Insecta
Hemiptera
Nogodinidae
Andrewsiella oceanica
Izzard, 1936
Recent
Insecta
Hemiptera
Nogodinidae
Distiana andrewsi
(Kirby, 1900)
Issus andrewsi,
Moniana andrewsi
Recent
Insecta
Hemiptera
Nogodinidae
Sassula subviridis
(Kirby, 1900)
Nogodina subviridis
1964
Insecta
Hemiptera
Nogodinidae
Varcia affinis
(Kirby, 1889)
Ricania affinis,
Nogodina affinis
1964
Insecta
Hemiptera
Nogodinidae
Varcia flavicostalis
(Kirby, 1889)
Ricania flavicostalis
1964
Insecta
Hemiptera
Delphacidae
Delphacodes muirianus
Izzard, 1936
Insecta
Hemiptera
Delphacidae
Ugyops aristella
(Kirby, 1900)
Insecta
Hemiptera
Ricaniidae
Paurostauria delicata
Kirby, 1900
1933
Insecta
Hemiptera
Ricaniidae
Ricania flavifrontalis
Kirby, 1900
Recent
Insecta
Hemiptera
Tropiduchidae
Salona oceanica
(Kirkaldy, 1909)
Ricania hyalina,
Nogodina hyalina
Recent
Insecta
Hemiptera
Miridae
Peritropis listeri
(Izzard, 1936)
Cylapofulvius listeri
1897–98
Insecta
Hemiptera
Miridae
Taylorilygus aldrichi
(Izzard, 1936)
Lygus aldrichi
1933
Insecta
Hemiptera
Miridae
Taylorilygus murrayi
(Izzard, 1936)
Lygus murrayi
1933
Insecta
Hemiptera
Aradidae
Mastigocoris insularis
Kormilev, 1983
Insecta
Hemiptera
Aradidae
Neuroctenus lignicolus
(Kirby, 1900)
Brachyrhynchus lignicolus,
Mezira lignicola
Recent
Insecta
Hemiptera
Lygaediae
Nysius spectabilis
Distant, 1901
Insecta
Hemiptera
Rhyparochromidae
Elasmolomus maculatus
(Distant, 1901)
Lethaeus maculatus
1897–98
Insecta
Hemiptera
Pentatomidae
Plautia grossepunctata
(Kirby, 1900)
Pentatoma grossepunctata
1964
Neuroptera
Myrmeleontidae
Myrmeleon iridescens
Kirby, 1900
Recent
Bidis aristella
Recent
1897–98
Recent
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Recent
Neuroptera
Insecta
84
Class
Order
Family
Species
Authority
Synonyms
Last known
collection date
Insecta
Coleoptera
Ptinidae
Clada posticum
(Gahan, 1900)
Paranobium posticum
Recent
Insecta
Coleoptera
Ptinidae
Neoptinus parvus
Gahan, 1900
Recent
Insecta
Coleoptera
Anthribidae
Apatenia apicalis
Gahan, 1900
Recent
Insecta
Coleoptera
Anthribidae
Litocerus jordani
Gahan, 1900
Recent
Insecta
Coleoptera
Anthribidae
Xenocerus nativitatis
Gahan, 1900
Recent
Insecta
Coleoptera
Bothrideridae
Ascetoderes strigatus
(Arrow, 1900)
Insecta
Coleoptera
Brentidae
Orychodes andrewsi
Gahan, 1900
1964
Insecta
Coleoptera
Buprestidae
Chrysobothris andrewsi
Waterhouse, 1900
Recent
Insecta
Coleoptera
Buprestidae
Chrysodema simplex
Waterhouse, 1887
Recent
Insecta
Coleoptera
Carabiidae
Lymnastis brooksi
Baehr, 2008
Recent
Insecta
Coleoptera
Cerambycidae
Ceresium nigrum
Gahan, 1889
Recent
Insecta
Coleoptera
Cerambycidae
Ceresium quadrimaculatum
Gahan ,1900
Insecta
Coleoptera
Cerambycidae
Ceresium affinis
(Gahan, 1900)
Examnes affinis
Recent
Insecta
Coleoptera
Cerambycidae
Sciades exigua
(Gahan, 1900)
Aegocidnus exiguus
Recent
Insecta
Coleoptera
Cerambycidae
Anomophysis coxalis
(Gahan, 1900)
Prinobius coxalis
Recent
Insecta
Coleoptera
Cerambycidae
Apomecyna nigritarsis
Gahan, 1900
Insecta
Coleoptera
Cerambycidae
Acalolepta nativitatis
(Gahan, 1889)
Insecta
Coleoptera
Cerambycidae
Olenecamptus basalis
Gahan, 1900
Insecta
Coleoptera
Cerambycidae
Pterolophia perplexa
(Gahan, 1889)
Insecta
Coleoptera
Cerambycidae
Phelipara subvittata
Blair, 1933
Recent
Insecta
Coleoptera
Chrysomelidae
Demotina lateralis
Gahan, 1900
Recent
Insecta
Coleoptera
Chrysomelidae
Rhyparida modesta
Gahan, 1900
Recent
Insecta
Coleoptera
Chrysomelidae
Rhyparida rossi
Gahan, 1900
1964
Insecta
Coleoptera
Chrysomelidae
Psylliodes tenuepunctata
Gahan, 1900
1897–98
Insecta
Coleoptera
Ciidae
Paratrichapus christmasensis
Souza-Gonçalves, 2018
Recent
Insecta
Coleoptera
Coccinellidae
Henosepilachna nativitatis
(Arrow, 1900)
Insecta
Coleoptera
Curculionidae
Cossonus variipennis
Gahan, 1900
1897–98
Insecta
Coleoptera
Curculionidae
Pachyops incertus
Gahan, 1900
Recent
Coleoptera
Bothrideres strigatus,
Aeschyntelus strigatus
1897–98
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
Recent
1897–98
Monohamnus nativitatis,
Dihamnus nativitatis
Recent
Recent
Praonetha perplexa
Epilachna nativitatis
Recent
1897–98
85
Order
Family
Species
Authority
Synonyms
Last known
collection date
Insecta
Coleoptera
Curculionidae
Rhyncolosoma dubium
(Gahan, 1900)
Phloeophagosoma dubium
Recent
Insecta
Coleoptera
Curculionidae
Camptorhinus crinipes
Gahan, 1900
Recent
Insecta
Coleoptera
Curculionidae
Acicnemis andrewsi
Gahan, 1900
Recent
Insecta
Coleoptera
Curculionidae
Rhyncholobus andrewsi
Gahan, 1900
Recent
Insecta
Coleoptera
Curculionidae
Rhyncholobus discoidalis
(Waterhouse, 1887)
Insecta
Coleoptera
Curculionidae
Rhyncholobus rossi
Gahan, 1900
Recent
Insecta
Coleoptera
Curculionidae
Rhyncholobus vittatus
Gahan, 1900
1897–98
Insecta
Coleoptera
Curculionidae
Dryophthorus assimilis
Gahan, 1900
Recent
Insecta
Coleoptera
Elateridae
Tetrigus murrayi
Waterhouse, 1900
Recent
Insecta
Coleoptera
Elateridae
Anchastus discoidalis
Waterhouse, 1900
Recent
Insecta
Coleoptera
Elateridae
Megapenthes andrewsi
Waterhouse, 1900
Recent
Insecta
Coleoptera
Histeridae
Hololepta malleata
Lewis, 1900
Insecta
Coleoptera
Histeridae
Platylister lignarium
(Lewis, 1900)
Platysoma lignarium
Recent
Insecta
Coleoptera
Lucanidae
Aegus listeri
(Gahan, 1889)
Paraegus listeri
Recent
Insecta
Coleoptera
Lucanidae
Figulus rossi
Gahan, 1900
Recent
Insecta
Coleoptera
Melyridae
Laius tibialis
Gahan, 1900
Recent
Insecta
Coleoptera
Oedemeridae
Sessinia andrewsi
Arrow, 1900
Recent
Insecta
Coleoptera
Oedemeridae
Sessinia listeri
Arrow, 1900
1932
Insecta
Coleoptera
Rhizophagidae
Shoguna striata
Arrow, 1900
Recent
Insecta
Coleoptera
Scarabaeidae
Eophileurus convexus
(Arrow, 1900)
Insecta
Coleoptera
Scarabaeidae
Protaetia andrewsi
Gahan, 1900
Recent
Insecta
Coleoptera
Scarabaeidae
Psammorpha lawrencei
Stebnicka, 1994
Recent
Insecta
Coleoptera
Staphylinidae
Cephennomicrus lawrencei
Jaloszyńģski, 2017
Recent
Insecta
Coleoptera
Staphylinidae
Paederus listeri
Gahan, 1900
1897–98
Insecta
Coleoptera
Tenebrionidae
Bradymerus seminitidus
Arrow, 1900
Recent
Insecta
Coleoptera
Tenebrionidae
Promethis carbonaria
(Arrow, 1900)
Insecta
Coleoptera
Tenebrionidae
Amarygmus funebris
Arrow, 1900
Insecta
Coleoptera
Tenebrionidae
Gonocephalum dubium
(Arrow, 1900)
Insecta
Coleoptera
Tenebrionidae
Toxicum antilope
Arrow, 1900
Insecta
Coleoptera
Trogossitidae
Leperina marmorata
(Arrow, 1900)
Piezonotus discoidalis
86
Class
Recent
Recent
Nyctobates carbonaria,
Setenis carbonaria
1897–98
Recent
1897–98
Opatrum dubium
Recent
Recent
Oniscomorpha marmorata,
Lepidopteryx marmorata
1897–98
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Phileurus convexus
Order
Family
Species
Authority
Synonyms
Last known
collection date
Insecta
Diptera
Asilidae
Orthogonis christmasensis
(Daniels, 2012)
Laphria nigrocaerulea,
Laphria christmasensis
Recent
Insecta
Diptera
Calliphoridae
Hemipyrellia jucunda
(Kirby, 1889)
Stilbomyia jucunda
1964
Insecta
Diptera
Dolichopodidae
Amblypsilopus natalis
Bickel, 1994
Recent
Insecta
Diptera
Drosophilidae
Lissocephala powelli
Carson and Wheeler, 1973
Recent
Insecta
Diptera
Keroplatidae
Micrepimera punctipennis
Matile, 1990
Recent
Insecta
Diptera
Sphaeroceridae
Poecilosomella pectiniterga
(Deeming, 1964)
1897–98
Insecta
Diptera
Syrphidae
Syritta maritima
Hull, 1944
1933
Insecta
Diptera
Tipulidae
Cymatopus calcaratus
Parent, 1935
1933
Insecta
Diptera
Tipulidae
Cymatopus longipilus
Parent, 1935
1933
Siphonaptera
Pulicidae
Xenopsylla nesiotes
(Jordan and Rothschild, 1908)
Loemopsylla nesiotes
Extinct c. 1904
Insecta
Lepidoptera
Tineidae
Opogona punctata
(Walsingham, 1900)
Dendroneura punctata
1897–98
Insecta
Lepidoptera
Pterophoridae
Cosmoclostis quadriquadra
Walsingham, 1900
Insecta
Lepidoptera
Immidae
Moca chlorolepis
Walsingham, 1900
Imma chlorolepis,
Tortricomorpha chlorolepis
1897–98
Insecta
Lepidoptera
Tortricidae
Loboschiza halysideta
Walsingham, 1900
Polemograptis halysideta,
Epagoge halysideta
Recent
Insecta
Lepidoptera
Brachodidae
Nigilgia browni
Kallies, 2013
Insecta
Lepidoptera
Choreutidae
Brenthia elachista
Walsingham, 1900
Insecta
Lepidoptera
Choreutidae
Choreutis ornaticornis
(Walsingham, 1900)
Simaethis ornaticornis
1897–98
Insecta
Lepidoptera
Nymphalidae
Charaxes andrewsi
Butler, 1900
Polyura andrewsi,
Eriboea pyrrhus andrewsi,
Hypolimnas andrewsi
Recent
Insecta
Lepidoptera
Crambidae
Hymenia nigerrimalis
Hampson, 1900
Zinckenia nigerrimalis,
Spoladea nigerrimalis
Recent
Insecta
Lepidoptera
Crambidae
Diaphania holophaealis
(Hampson, 1900)
Glyphodes holophaealis,
Phacellura holophaealis
Recent
Diptera
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
Siphonaptera
Insecta
Lepidoptera
1897–98
Recent
1897–98
87
Order
Family
Species
Authority
Synonyms
Last known
collection date
Insecta
Lepidoptera
Pyralidae
Ephestia scotella
Hampson, 1900
1939
Insecta
Lepidoptera
Geometridae
Anisodes hypomion
Prout, 1933
Recent
Insecta
Lepidoptera
Geometridae
Comostoloapsis regina
Thierry-Mieg, 1915
Recent
Insecta
Lepidoptera
Geometridae
Ecliptopera phaula
Prout, 1933
Recent
Insecta
Lepidoptera
Geometridae
Hemithea hyperymna
Prout, 1933
Chlorissa hyperymna
1939
Insecta
Lepidoptera
Geometridae
Ruttelerona scotozonea
(Hampson, 1900)
Ruttelerona scotozonea,
Ectropis scotozonea
1939
Insecta
Lepidoptera
Geometridae
Sauris pelagitis
Prout, 1933
Insecta
Lepidoptera
Geometridae
Scopula tumiditibia
Prout, 1920
Insecta
Lepidoptera
Geometridae
Thalassodes subviridis
Warren, 1905
Insecta
Lepidoptera
Erebidae
Acontia sollemnis
Hacker and Holloway, 2008
Insecta
Lepidoptera
Nolidae
Earias latimargo
Hampson, 1912
Insecta
Lepidoptera
Noctuidae
Amyna crocosticta
Hampson, 1910
Recent
Insecta
Lepidoptera
Noctuidae
Armactica andrewsi
Hampson, 1912
Recent
Insecta
Lepidoptera
Noctuidae
Bocula limbata
(Butler, 1889)
Eudragana limbata
Recent
Insecta
Lepidoptera
Noctuidae
Lithacodia griseomixta
(Hampson, 1900)
Erastria griseomixta,
Deltote griseomixta
1939
Insecta
Hymenoptera
Bracondiae
Dendrosotinus insularis
Belokobylskij, Iqbal
and Austin, 2004
Recent
Insecta
Hymenoptera
Braconidae
Neoheterospilus insularis
Belokobylskij, 2006
Recent
Insecta
Hymenoptera
Braconidae
Phanerotoma pacifica
Zettel, 1990
Recent
Insecta
Hymenoptera
Halictidae
Homalictus andrewsi
Kirby, 1900
Halictus andrewsi,
Lasioglossum andrewsi,
Patellapis andrewsi
Recent
Insecta
Hymenoptera
Halictidae
Patellapis binghami
(Kirby, 1900)
Halictus binghami,
Homalictus binghami
1964
Insecta
Hymenoptera
Megachilidae
Lithurgus andrewsi
Cockerell, 1909
Lithurge andrewsi
1897–98
Insecta
Hymenoptera
Megachilidae
Megachile nivescens
Kirby, 1900
1964
Insecta
Hymenoptera
Megachilidae
Megachile rotundipennis
Kirby, 1900
1964
Insecta
Hymenoptera
Pteromalidae
Sirovena stigma
Bouček, 1988
Recent
88
Class
1939
1939
Pelagodes subviridis
Recent
Recent
Earias chromataria
Recent
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Hymenoptera
Order
Family
Species
Authority
Synonyms
Last known
collection date
Insecta
Hymenoptera
Scelionidae
Oxyscelio caesitas
Burks, 2013
Insecta
Hymenoptera
Vespidae
Euodynerus polyphemus
(Kirby, 1889)
Insecta
Hymenoptera
Vespidae
Polistes balder
Kirby, 1889
Recent
Actinopterygii
Perciformes
Pseudochromidae
Pseudochromis viridis
Gill and Allen, 1996
Recent
Actinopterygii
Perciformes
Gobiidae
Eviota natalis
Allen, 2007
Recent
Actinopterygii
Ophidiiformes
Bythitidae
Microbrotula andersoni
Schwarzhans and Nielsen, 2011
Recent
Actinopterygii
Ophidiiformes
Bythitidae
Paradiancistrus christmasensis
Schwarzhans and Møller, 2011
Recent
Actinopterygii
Pleuronectiformes
Soleidae
Aseraggodes crypticus
Randall and Allen, 2007
Recent
Reptilia
Squamata
Typhlopidae
Ramphotyphlops exocoeti
(Boulenger, 1887)
Reptilia
Squamata
Gekkonidae
Cyrtodactylus sadleiri
Wells and Wellington, 1985
Reptilia
Squamata
Gekkonidae
Lepidodactylus listeri
(Boulenger, 1889)
Gecko listeri
Extinct
(in the wild)
Reptilia
Squamata
Scincidae
Cryptoblepharus egeriae
(Boulenger, 1889)
Ablepharus egeriae
Extinct
(in the wild)
Reptilia
Squamata
Scincidae
Emoia nativitatis
(Boulenger, 1887)
Lygosoma nativitatis
Extinct, 2014
Aves
Columbiformes
Columbidae
Ducula whartoni
(Sharpe, 1887)
Recent
Aves
Apodiformes
Apodidae
Collocalia natalis
Lister, 1889
Recent
Aves
Pelecaniformes
Sulidae
Papasula abbotti
(Ridgway, 1893)
Recent
Aves
Pelecaniformes
Fregatidae
Fregata andrewsi
Mathews, 1914
Recent
Aves
Strigiformes
Strigidae
Ninox natalis
Lister, 1889
Recent
Aves
Passeriformes
Timaliidae
Zosterops natalis
Lister, 1889
Recent
Mammalia
Insectivora
Soricidae
Crocidura trichura
Dobson, 1889
1985
Mammalia
Chiroptera
Pteropodidae
Pteropus natalis
Thomas, 1887
Recent
Mammalia
Chiroptera
Vespertilionidae
Pipistrellus murrayi
Andrews, 1900
Extinct, 2009
Mammalia
Rodentia
Muridae
Rattus nativitatis
(Thomas, 1889)
Extinct c. 1904
Mammalia
Rodentia
Muridae
Rattus macleari
(Thomas, 1887)
Extinct c. 1904
Recent
Odynerus polyphemus
Recent
Pisces
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
Reptilia
Typhlops exocoeti
Recent
Recent
Aves
Mammalia
89
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
90
TABLE 3
List of putative undescribed taxa that are potentially new endemic species to Christmas Island.
Class
Order
Family
Affinities
Source
Gastropoda
Hypsogastropoda
Cypraeidae
Lyncina sp. nov.
Meyer (2004) in Tan and Low (2014)
Gastropoda
Architaenioglossa
Cyclophoridae
Leptopoma sp. nov.
Tan and Low (2014)
Gastropoda
Hypsogastropoda
Assimineidae
gen nov. sp. nov.
Kessner (2006)
Gastropoda
Littorinimorpha
Charopidae
Charopa sp.nov.
Laidlaw (1935); Kessner (2006)
Polychaeta
Spionida
Spionidae
Prionospio sp. nov.
Molecular phylogeny by
K. Worsaae (pers. comm.).
Ostracoda
Podocopida
Darwinulidae
Vestalenula sp. E
Rossetti et al. (2011)
Ostracoda
Podocopida
Darwinulidae
Penthesilenula sp. nov.
Humphreys (2014)
Maxillopoda
Calanoida
Arietellidae
gen. nov., sp. nov.
Bruce and Davie (2006)
Malacostraca
Isopoda
Armadillidae
Myrmecodillo sp. nov. 1
Humphreys and Eberhard (2001)
Malacostraca
Isopoda
Armadillidae
Myrmecodillo sp. nov. 2
Humphreys and Eberhard (2001)
Malacostraca
Isopoda
Armadillidae
Myrmecodillo sp. nov. 3
Humphreys (2014)
Malacostraca
Isopoda
Armadillidae
Myrmecodillo sp. nov. 4
Humphreys (2014)
Malacostraca
Isopoda
Eubelidae
Elumoides sp.nov.
Humphreys (2014)
Malacostraca
Isopoda
Philosciidae
Papuaphiloscia sp. nov. 1
Humphreys and Eberhard (2001)
Malacostraca
Isopoda
Philosciidae
Papuaphiloscia sp. nov. 2
Humphreys (2014)
Malacostraca
Thermosbaenacea
Halosbaenidae
Halosbaena sp. nov.
Humphreys (2014)
Malacostraca
Decapoda
Paguridae
gen. nov., sp. nov. 1
Tan et al. (2014b)
Malacostraca
Decapoda
Paguridae
gen. nov., sp. nov. 2
Tan et al. (2014b)
Malacostraca
Decapoda
Diogenidae
Clibanarius sp. nov.
Tan et al. (2014b)
Arachnida
Pseudoscorpiones
Chthoniidae
Tyrannochthonius sp. nov. 1
Humphreys (2014)
Arachnida
Pseudoscorpiones
Chthoniidae
Tyrannochthonius sp. nov. 2
Humphreys (2014)
Arachnida
Schizomida
Hubbardiidae
Apozomus sp. nov.
Humphreys (2014)
Entognatha
Diplura
Campodeidae
Cocytocampa sp. nov. 2
Humphreys and Eberhard (2001)
Insecta
Zygentoma
Nicoletiidae
Metrinura sp. nov.
Humphreys and Eberhard (2001)
Insecta
Blattodea
Blattellidae
gen. nov., sp. nov.
Humphreys and Eberhard (2001)
Insecta
Orthoptera
Mogoplistidae
Mogoplistinae gen? sp. nov.
CSIRO Division of Entomology (1990)
Insecta
Orthoptera
Tettigoniidae
gen?, sp. nov.
Humphreys and Eberhard (2001)
Insecta
Hemiptera
Omaniidae
gen. nov., sp. nov.
CSIRO Division of Entomology (1990)
Insecta
Neuroptera
Chrysopidae
Mallada sp. nov.?
New (1991)
Insecta
Hymenoptera
Crabronidae
Liris sp. nov.
CSIRO Division of Entomology (1990)
Insecta
Hymenoptera
Crabronidae
Tachysphex sp. nov.
CSIRO Division of Entomology (1990)
Insecta
Hymenoptera
Crabronidae
Pison sp. nov. 1
CSIRO Division of Entomology (1990)
Insecta
Hymenoptera
Crabronidae
Pison sp. nov. 2
CSIRO Division of Entomology (1990)
Insecta
Hymenoptera
Vespidae
Subancistrocerus sp. nov.
CSIRO Division of Entomology (1990)
List of species originally described from Christmas Island but no longer considered endemic.
Class
Order
Family
Species
Authority
Reason
Agaricomycetes
Geastrales
Geastraceae
Radiigera asperata
Reid, 1986
= an immature Geastrum sp.
(Domínguez de Toledo and Castellano, 1996)
Agaricomycetes
Geastrales
Geastraceae
Radiigera termitariicola
Reid, 1986
= Phialastrum barbatum (Dissing and Lange)
Sunhede 1962 (per Domínguez de Toledo and
Castellano, 1996)
Agaricomycetes
Polyporales
Polyporaceae
Poria chlorina
Massee, 1906
= Ceriporia mellea (Berk. and Broome) Ryvarden
(ALA, but cf. GBIF that regards it as a valid species)
Agaricomycetes
Polyporales
Polyporaceae
Favolus albidus
Massee, 1902
Previously described from Thailand
(Shivas and Hilton, 1990)
Halymeniales
Halymeniaceae
Halymenia polyclada
Gepp and Gepp, 1905
Recorded from Seychelles (GBIF)
Equisetopsida
Selaginellales
Selaginellaceae
Selaginella rupicola
Ridl.
= S. alutacea Spring (ALA)
Equisetopsida
Polypodiales
Polypodiaceae
Gymnopteris listeri
(Baker) Ridl.
= Leptochilus decurrens Blume (ALA)
Equisetopsida
Polypodiales
Aspleniaceae
Asplenium listeri
Christensen, 1905
Widely distributed (Ohlsen et al., 2015)
Equisetopsida
Asparageles
Orchidaceae
Dendrobium pectinatum
Finet, 1903
New Caledonia (GBIF) [Ridley (1906) may have
erred in his identification of this taxon from
Christmas Island]
Equisetopsida
Asparageles
Orchidaceae
Phreatia congesta
Rolfe, 1890
= Bryobium retusum (Blume) Y.P. Ng and P.J. Cribb
(GBIF)
Equisetopsida
Asparageles
Orchidaceae
Corymbis angusta
Ridl.
= Corymborkis veratrifolia (Reinw.) Blume (GBIF)
Equisetopsida
Poales
Poacea
Panicum andrewsi
Rendle
ex Baker f., 1900
= Panicum trichoides Swartz, 1788 (GBIF)
Fungi
ENDEMIC SPECIES OF CHRISTMAS ISLAND
TABLE 4
Rhodophyta
Florideophyceae
Vascular Plants
Equisetopsida
Poales
Poacea
Panicum clivale
Ridl.
= Setaria clivalis (Ridl.) Veldkamp (GBIF)
Equisetopsida
Apiales
Araliaceae
Heptapleurum natale
Ridl.
= Schefflera elliptica Harms. (GBIF, ALA)
Equisetopsida
Cucurbitales
Cucurbitaceae
Zehneria alba
Ridley, 1906
= Z. mucronata (Blume) Miq. (ALA)
Equisetopsida
Ericales
Primulaceae
Ardisia pulchra
Ridl.
= Ardisia sanguinolenta Blume (ALA)
Equisetopsida
Gentianales
Rubiaceae
Saprosma nativitatis
Baker f., 1900
= Amaracarpus pubescens Blume, 1826 (ALA)
91
Order
Family
Species
Authority
Reason
Equisetopsida
Laurales
Lauraceae
Cryptocarya nativitatis
Rendle
ex Baker f., 1900
= Cryptocarya nitens (Blume)
Koorders and Valeton, 1904 (ALA)
Equisetopsida
Malphighiales
Euphorbiaceae
Claoxylon caerulescens
Ridl.
= Claoxylon indicum Hassk (ALA)
Equisetopsida
Malvales
Malvaceae
Grewia osmoxylon
Ridley, 1906
= Grewia laevigata Vahl (ALA)
Equisetopsida
Myrtales
Myrtaceae
Eugenia gigantea
Ridl.
= Syzygium nervosum DC. (ALA)
Equisetopsida
Ranunculales
Menispermaceaea
Limacia nativitatis
Ridley, 1906
= Pachygone ovata
Miers ex Hook f., and Thomson, 1855 (GBIF)
Equisetopsida
Rosales
Urticaceae
Laportea murrayana
Rendle
ex Baker f., 1900
= Dendrocnide peltata var. murrayana
(Rendle) Chew, 1969 (ALA, GBIF)
Equisetopsida
Sapindales
Rutaceae
Acronychia andrewsi
Baker f., 1900
= Acronychia trifoliolata
Zollinger and Moritzi, 1845 (GBIF, ALA)
Equisetopsida
Sapindales
Sapindaceaea
Tristiropsis nativitatis
Hemsl. ex Ridl.
= Tristiropsis acutangula Radlk. (GBIF)
Demospongiae
Haplosclerida
Callyspongiidae
Callyspongia spinosissima
(Dendy, 1887)
Widely distributed (GBIF)
Demospongiae
Haplosclerida
Microcionidae
Clathria dubia
(Kirkpatrick, 1900)
Recorded from Indonesia (ALA, GBIF)
Demospongiae
Haplosclerida
Haliclonidae
Neopetrosia exigua
(Kirkpatrick, 1900)
Indo-Pacific (GBIF)
Demospongiae
Suberitida
Suberitidae
Pseudosuberites andrewsi
Kirkpatrick, 1900
Indo-Pacific (GBIF)
Demospongiae
Suberitida
Halichondriidae
Haliclona irregularis
(Kirkpatrick, 1900)
Tropical Indian Ocean (Burton, 1959)
Demospongiae
Tethyida
Tethyidae
Tethya affinis
Kirkpatrick, 1900
= Tethya deformis Thiele, 1898 (GBIF)
Calcarea
Murrayonida
Murrayonidae
Murrayona phanolepis
Kirkpatrick, 1911
Pacific Ocean (GBIF)
Brachiopoda
Thecideida
Thecidellinidae
Thecidellina blochmanni
(Dall, 1920)
Guam (GBIF)
Gastropoda
Hypsogastropoda
Eulimidae
Palisadia subulata
Laseron, 1956
Indo-Pacific (ALA, GBIF)
Gastropoda
Hypsogastropoda
Littorinidae
Littorina granicostata
Smith, 1887
= Granulittorina) reticulata (Anton, 1838) (GBIF)
Gastropoda
Hypsogastropoda
Pickworthiidae
Discrevinia balba
Laseron, 1956
Indo-Pacific (ALA)
Gastropoda
Hypsogastropoda
Pickworthiidae
Merelina hians
Laseron, 1956
= Sansoniella minuta
(Hornung and Mermod, 1927) (GBIF)
Gastropoda
Hypsogastropoda
Pickworthiidae
Pickworthia andrewsi
Iredale, 1917
= Sansonia andamanica (Preston, 1908) (ALA)
Gastropoda
Hypsogastropoda
Pickworthiidae
Reynellona natalis
Iredale, 1917
Indo-Pacific (ALA, GBIF)
Gastropoda
Hypsogastropoda
Pickworthiidae
Sansonia kirkpatricki
(Iredale, 1917)
Indo-Pacific (ALA)
Gastropoda
Hypsogastropoda
Pickworthiidae
Sherbornia mirabilis
Iredale, 1917
Indo-Pacific (ALA, GBIF)
92
Class
Sponges
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Invertebrates
Order
Family
Species
Authority
Reason
Gastropoda
Hypsogastropoda
Rissoidae
Alvania isolata
(Laseron, 1956)
Taiwan (ALA, GBIF)
Gastropoda
Hypsogastropoda
Rissoidae
Austrosina quinita
Laseron, 1956
= Rissoina honoluluensis Watson, 1886 (ALA)
Gastropoda
Hypsogastropoda
Rissoidae
Rissonia evanida
(Laseron, 1956)
Indo-Pacific (ALA, GBIF)
Gastropoda
Hypsogastropoda
Rissoidae
Schwartziella lata
Laseron, 1956
= Zebina triticea (Pease, 1861) (ALA)
Gastropoda
Hypsogastropoda
Rissoidae
Pandalosia oceanica
Laseron, 1956
= Zebina ephamilla (Watson, 1886) (ALA)
Gastropoda
Hypsogastropoda
Rissoidae
Stosicia incisa
(Laseron, 1956)
Indo-Pacific (ALA, GBIF)
Gastropoda
Hypsogastropoda
Rissoidae
Zebina isolata
Laseron, 1956
Japan (GBIF)
Gastropoda
Hypsogastropoda
Rissoidae
Zebina linearis
Laseron, 1956
the Philippines (GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Iniforis ordinata
Laseron, 1958
Hawaii (AFD, GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Iniforis progressa
(Laseron, 1958)
Japan (GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Iniforis tuberia
Laseron, 1958
French Polynesia (GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Liniphora restis
Laseron, 1958
Indo-Pacific (GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Mastoniaeforis insulana
(Laseron, 1958)
Indo-Pacific (GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Opimaphora coralina
Laseron, 1958
Indo-Pacific and Caribbean (AFD, ALA, GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Subalophora indianica
Laseron, 1958
Taiwan (AFD) and Reunion Island (GBIF)
Gastropoda
Hypsogastropoda
Triphoridae
Viriola oceanica
Laseron, 1958
Indo-Pacific (ALA, GBIF)
Gastropoda
Hypsogastropoda
Nassariidae
Nassarius exulatus
Smith, 1911
French Polynesia
Gastropoda
Neogastropoda
Muricida
Orania taeniata
Houart, 1995
Indo-Pacific (GBIF)
Gastropoda
Stylommatophora
Succineidae
Succinea listeri
Smith, 1889
Java (Tan and Low, 2014)
Clitellata
Opisthopora
Megascolecidae
Polypheretima brevis
(Rossa, 1898)
Cocos (Keeling) Is. and Tonga (GBIF)
Maxillopoda
Cyclopoida
Corycaeidae
Ditrichocorycaeus andrewsi
(Farran, 1911)
Widely distributed (ALA, GBIF)
Maxillopoda
Cyclopoida
Corycaeidae
Ditrichocorycaeus dubius
(Farran, 1911)
Widely distributed (GBIF)
Maxillopoda
Cyclopoida
Corycaeidae
Corycaeus murrayi
Farran, 1911
= Ditrichocorycaeus asiaticus (Dahl, 1894) (GBIF)
Maxillopoda
Cyclopoida
Corycaeidae
Corycella brevis
Farran, 1911
= Farranula gibbula (Giesbrecht, 1891) (GBIF)
Maxillopoda
Cyclopoida
Corycaeidae
Corycella curta
Farran, 1911
Widely distributed (ALA, GBIF)
Maxillopoda
Cyclopoida
Oithonidae
Oithona attenuata
Farran, 1913
= Oithona nana Giesbrecht, 1893 (ALA),
widespread (GBIF)
Maxillopoda
Cyclopoida
Oithonidae
Oithona decipiens
Farran, 1913
Australia (ALA, GBIF)
Maxillopoda
Cyclopoida
Oithonidae
Oithona fallax
Farran, 1913
Widely distributed (GBIF)
Cyclopoida
Oithonidae
Oithona oculata
Farran, 1913
Widely distributed (GBIF)
Cyclopoida
Oithonidae
Oithona simplex
Farran, 1913
Widely distributed (GBIF)
93
Maxillopoda
Maxillopoda
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
Family
Species
Authority
Reason
Maxillopoda
Cyclopoida
Oithonidae
Oithona vivida
Farran, 1913
Northern Australia (GBIF)
Maxillopoda
Cyclopoida
Oithonidae
Paraoithona pulla
Farran, 1913
N.W. Pacific and Red Sea (Ferrari and Böttger, 1986)
Malacostraca
Decapoda
Paguridae
Paguritta harmsi
(Gordon, 1935)
Widely distributed (GBIF, ALA)
Malacostrata
Decapoda
Cryptochiridae
Opecarcinus granulatus
(Shen, 1936)
Indo-Pacific (AFD, GBIF)
Malacostraca
Decapoda
Grapsidae
Sesarma murrayi
Calman, 1909
= Pachygrapsus minutus
A. Milne-Edwards, 1873 (GBIF)
Malacostrata
Decapoda
Grapsidae
Pachygrapsus natalensis
Ward, 1934
= Pachygrapsus plicatus
(H. Milne-Edwards, 1837) (per Tweedie, 1947) (GBIF)
Malacostrata
Decapoda
Grapsidae
Percnon demani
Ward, 1934
= Percnon planissimum (Herbst, 1804) (GBIF)
Malacostrata
Decapoda
Epialtidae
Hyastenus andrewsi
Calman, 1909
= Micippoides angustifrons
A. Milne-Edwards, 1873 (GBIF)
Malacostrata
Decapoda
Epialtidae
Hyastenus uncifer
Calman, 1909
Indo-Pacific (GBIF)
Malacostrata
Decapoda
Epialtidae
Hyastenus macrospinosus
Ward, 1934
= Tylocarcinus dumerilii
(H. Milne-Edwards, 1834) (GBIF)
Malacostrata
Decapoda
Xanthidae
Pseudoliomera natalensis
Ward, 1934
= Pseudoliomera granosimana
(A. Milne-Edwards, 1865) (GBIF)
Malacostrata
Decapoda
Xanthidae
Liocarpilodes harmsi
(Balss, 1934)
Indo-Pacific (AFD, GBIF)
Malacostrata
Decapoda
Xanthidae
Chlorodopsis natalensis
Ward, 1934
= Liocarpilodes harmsi (Balss, 1934) (GBIF)
Malacostrata
Decapoda
Xanthidae
Tweedieia noelensis
Ward, 1934
= Tweedieia odhneri (Gordon, 1934) (GBIF)
Malacostrata
Decapoda
Xanthidae
Etisus albus
(Ward, 1934)
Kiribati (GBIF)
Malacostrata
Decapoda
Xanthidae
Paramedaeus noelensis
Ward, 1934
Indo-Pacific (ALA, GBIF)
Malacostrata
Decapoda
Xanthidae
Kraussia proporcellana
(Ward, 1934)
= Kraussia rugulosa (Krauss, 1843) (GBIF)
Malacostrata
Decapoda
Xanthidae
Leptodius planus
Ward, 1934
= Leptodius gracilis (Dana, 1852) (GBIF);
Mendoza et al. (2014) considered these distinct but
listed L. planus from Cocos (Keeling) islands
Malacostrata
Decapoda
Xanthidae
Lioxanthodes alcocki
Calman, 1909
French Polynesia (GBIF)
Malacostrata
Decapoda
Xanthidae
Paraxanthias haematostictus
Ward, 1934
= Lachnopodus ponapensis (Rathbun, 1907) (GBIF)
Malacostrata
Decapoda
Xanthidae
Atergatopsis tweediei
(Ward, 1934)
Indonesia (AFD) and Western Australia (ALA)
Malacostrata
Decapoda
Xanthidae
Neoliomera cerasinus
Ng, 2002
Type specimen from Christmas Island, but Ng (2002)
also reported a record from Ryukyus, Japan
Arachnida
Pseudoscorpiones
Withiidae
Metawithius murrayi
(Pocock, 1900)
Indonesia (GBIF)
Diplopoda
Polydesmida
Haplodesmidae
Cylindrodesmus hirsutus
Pocock, 1889
Widely distributed (GBIF)
Insecta
Mantodea
Mantidae
Hierodula dispar
(Kirby, 1900)
= Hierodula patellifera (Serville, 1838) (GBIF)
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Order
94
Class
Order
Family
Species
Authority
Reason
Insecta
Orthoptera
Acrididae
Epacromia rufostriata
Kirby, 1889
= Aiolopus thalassinus tamulus (Fabricius, 1798)
(GBIF)
Insecta
Orthoptera
Acrididae
Cyrtacanthacris disparilis
Kirby, 1889
= Valanga nigricornis disparilis (GBIF)
Insecta
Dermaptera
Chelisochidae
Labidura nigricornis
Kirby, 1889
= Chelisoches morio (Fabricius, 1775) (GBIF)
Insecta
Dermaptera
Forficulidae
Labia indistincta
Kirby, 1900
= Paralabella murrayi (Kirby, 1900) (GBIF)
Insecta
Dermaptera
Forficulidae
Labia incerta
Kirby, 1900
= Paralabella murrayi (Kirby 1900) (GBIF)
Insecta
Hemiptera
Cydnidae
Aethus nitens
Kirby, 1900
= Fromundus pygmaeus (Dallas, 1851) (ALA)
Insecta
Hemiptera
Cicadellidae
Hishimonus festivus
Knight, 1970
Subsequent records from Java
(Fletcher and Dai, 2013)
Insecta
Hemiptera
Cicadellidae
Xestocephalus izzardi
Metcalfe, 1955
Indo-Pacific (Linnavuori, 1960)
Insecta
Hemiptera
Cicadellidae
Xestocephalus minutus
Izzard, 1936
Replacement name = Xestocephalus izzardi
Metcalfe, 1955 (ALA)
Insecta
Hemiptera
Nogodinidae
Ricania hyalina
Kirby, 1889
Replacement name = Salona oceanica
(Kirkaldy, 1909) (GBIF)
Insecta
Hemiptera
Geocoridae
Geocoris vestitus
Distant, 1901
= Geocoris jucundus (per Kondorosy, 2016) (GBIF)
Insecta
Hemiptera
Lygaediae
Nysius andrewsi
Izzard, 1936
= Nysius vinitor Bergroth 1891
(per Malipatil, 2010) (GBIF)
Insecta
Hemiptera
Lygaediae
Nysius dissimilis
Izzard, 1936
= Nysius caledoniae Distant 1920
(per Malipatil, 2010) (GBIF)
Insecta
Hemiptera
Rhyparochromidae
Paraeucosmetus insignis
(Distant, 1901)
India (Chandra and Kushwaha, 2014)
Insecta
Hemiptera
Rhyparochromidae
Pamera andrewsi
(Distant, 1901)
= Remaudiereana nigriceps (ALA)
Insecta
Hemiptera
Rhopalidae
Leptocoris subrufescens
(Kirby, 1889)
subsp. L. s. flavus occurs on Yap, W. Caroline Is.
(Göllner-Scheiding, 1980)
Insecta
Neuroptera
Myrmeleontidae
Formicaleo morpheus
Kirby, 1900
= Distoleon somnolentus (New, 1991) (AFD)
Insecta
Coleoptera
Dryophthoridae
Rhabdocnemis fausti
Gahan, 1900
= Rhabdoscelus obscurus (Boisduval, 1835) (ALA)
Insecta
Coleoptera
Nitidulidae
Stelidota orientalis
Arrow, 1900
= Omosita nigrovaria (Fairmaire, 1849) (ALA)
Insecta
Coleoptera
Rhizophagidae
Shoguna polita
Arrow, 1900
= Shoguna termitiformis (Fairmaire, 1883) (ALA)
Insecta
Diptera
Asilidae
Laphria nigrocaerulea
Kirby, 1889
Replacement name = Orthogonis christmasensis
(Daniels, 2012) (ALA)
Insecta
Diptera
Culicidae
Aedes andrewsi
Edwards, 1927
Indonesia (GBIF)
Insecta
Lepidoptera
Nympahlidae
Vadebra macleari
(Butler, 1887)
= Euploea climena (ALA)
Insecta
Lepidoptera
Nympahlidae
Hypolimnas listeri
Butler, 1889
= H. bolina (GBIF)
ENDEMIC SPECIES OF CHRISTMAS ISLAND
Class
95
Family
Species
Authority
Reason
Insecta
Lepidoptera
Pieridae
Terias amplexa
(Butler, 1887)
= Eurema alitha (ALA)
Insecta
Lepidoptera
Pyralidae
Pyralis listeri
Butler, 1889
= Endotricha puncticostalis (Walker, 1866) (ALA)
Insecta
Lepidoptera
Geometridae
Syrrhodia vindex
Prout, 1933
= Hyperythra rubricata Warren, 1898
mainland Australia (ALA)
Insecta
Lepidoptera
Erebidae
Euproctis pulverea
Hampson, 1900
Widely distributed (GBIF)
Insecta
Lepidoptera
Erebidae
Hydrillodes vexillefera
Hampson, 1900
Records beyond Christmas Island noted in
original description
Insecta
Lepidoptera
Noctuidae
Mimeusemia econia
Hampson, 1900
= Mimeusemia hedya (AFD, ALA)
Insecta
Lepidoptera
Noctuidae
Anomis esocampta
Hampson, 1926
Fiji (Evenhuis 2017)
Insecta
Lepidoptera
Noctuidae
Maliattha phaeozona
Hampson, 1910
Mainland south-eastern Asia (Kocak and Kemal, 2010)
Insecta
Lepidoptera
Tortricidae
Caenognosis incisa
Walsingham, 1900
Mainland Australia and the Philippines (ALA)
Insecta
Hymenoptera
Ichneumonidae
Opion flavocephalus
Kirby, 1900
Widespread (Gauld, 1977); Sri Lanka (GBIF)
Insecta
Hymenoptera
Mymaridae
Anagrus antipodus
S. Triapitsyn, 2001
Occurs on mainland Australia (ALA) and was not
regarded as endemic in the original description
Insecta
Hymenoptera
Scelionidae
Baeus tropaeumusbrevis
Stevens, 2007
Occurs on mainland Australia (ALA) and was not
regarded as endemic in the original description
Insecta
Hymenoptera
Scelionidae
Mantibaria anomala
Kirby, 1900
= M. seefelderiana de Stephani-Perez, 1891
(per Mineo and Szabó, 1978)
Insecta
Hymenoptera
Formicidae
Camponotus melichloros
Kirby, 1889
Indo-Pacific (Framenau and Thomas, 2008)
Insecta
Hymenoptera
Formicidae
Leptogenys harmsi
Donisthorpe, 1935
Indo-Pacific (Framenau and Thomas, 2008)
Insecta
Hymenoptera
Formicidae
Pachycondyla christmasi
(Donisthorpe, 1935)
Indo-Pacific (Framenau and Thomas, 2008)
Crinoidea
Comatulida
Comatulidae
Alloeocomatella pectinifer
(Clark, 1911)
Indo-West Pacific (AFD, ALA)
Actinopterygii
Perciformes
Plesiopidae
Steeneichthys nativitatIs
Allen, 1987
Indo-Pacific (AFD, ALA)
Actinopterygii
Perciformes
Blenniidae
Praealticus natalis
(Regan, 1909)
Indo-Pacific (AFD, ALA)
Aves
Falconiformes
Accipitridae
Urospizias natalis
Lister, 1889
= Accipiter hiogaster (Müller 1841)
(ALA, but cf. GBIF which retains it as a distinct species)
Aves
Columbiformes
Columbidae
Chalcophaps natalis
Lister, 1889
= Chalcophaps indica (Linnaeus 1758)
(ALA, but cf. GBIF which retains it as a distinct species)
Aves
Passeriformes
Turdidae
Turdus erythropleurus
Sharpe, 1887
= Turdus poliocephalus Latham 1801
(ALA, but cf. GBIF which retains it as a distinct species)
Vertebrates
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
Order
96
Class
Proportion of endemic species (our tallies) relative to all species recorded for Christmas Island, for some of the better-studied taxonomic groups. Numbers include
species reported to have become extinct since the island’s settlement in 1887.
* tally likely to include some introduced species.
** note that the tally of the no. of recorded spp. is from a 1990 assessment.
No. of reported
endemic species
No. of reported
native species
% of native species
that are endemic
Molluscs
27
760
3.6
Tan and Low (2014)
Decapod crustaceans
10
249
4.0
Morgan (2000) and subsequent reviews
5
681
0.7
Hobbs et al. (2014)
Fungi
2
132
1.4
Shivas and Hilton (1990) and subsequent collections
Lichen
4
100
4.0
McCarthy (2018)
Vascular plants
17
213
8.0
Claussen (2005)
Beetles**
58
390
15
CSIRO (1990)
Psocoptera
10
33
30
Smithers (1995)
0
?2–4
0
12
34*
35
P.T. Green (unpubl.)
Butterflies
1
29*
3.4
Johnson and Wilson (2018)
Reptiles
5
6
80
Cogger et al. (1983)
Birds (breeding species only)
6
20
30
James and McAllan (2014)
Mammals
5
5
100
Taxonomic group
Source
Marine groups
Fish
ENDEMIC SPECIES OF CHRISTMAS ISLAND
TABLE 5
Terrestrial groups
Ants
Bees and wasps
Framenau and Thomas (2008)
Eldridge et al. (2014)
97
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
98
TABLE 6
Date of last known record for Christmas Island species unreported for 50 years or more.
Decade
No. of spp.
Species
1880s
1
Cryptops inermipes
1890s
29
Geastrum andrewsii, Peperomia rossii, Oceanapia sessilis, Haliclona innominata,
Tylos nudulus, Ariadna natalis, Ecatoderus flavipalpis, Paralabella murrayi, Anisolabis
subarmata, Peritropis listeri, Nysius spectabilis, Elasmolomus maculatus, Ascetoderes
strigatus, Apomecyna nigritarsis, Psylliodes tenuepunctata, Henosepilachna nativitatis,
Cossonus variipennis, Rhyncholobus vittatus, Eophileurus convexus, Paederus listeri,
Amarygmus funebris, Leperina marmorata, Poecilosomella pectiniterga, Opogona
punctata, Moca chlorolepis, Cosmoclostis quadriquadra, Brenthia elachista, Choreutis
ornaticornis, Lithurgus andrewsi
1900s
6
Ectropothecium micronesiense, Isopterygium jelinkii, Ixodes nitens, Xenopsylla nesiotes,
Rattus nativitatis, Rattus macleari
1910s
15
Finella rugosa, Cyclonidea carina, Chrystella islandica, Rissoina isolata, Schwartziella
delicatula, Zebina acicula, Zebina constricta, Pyramidelloides viticula, Mastonia
anomala, Mastoniaeforis decorata, Mastoniaeforis radix, Nanaphora minuta, Subulophora
marginata, Subulophora virgina, Potenatomus secundus
1920s
0
1930s
20
1940s
0
1950s
0
1960s
12
TABLE 7
Anxietas perplexa, Plesiotrochus fischeri, Lamprocystis mabelae, Filistata gibsonhilli,
Heteropoda listeri, Hypocambala exocoeti, Laccocoris montandoni, Paurostauria delicata,
Taylorilygus aldrichi, Taylorilygus murrayi, Sessinia listeri, Syritta maritima, Cymatopus
calcaratus, Cymatopus longipilus, Ephestia scotella, Hemithea hyperymna, Ruttelerona
scotozonea, Sauris pelagitis, Scopula tumiditibia, Lithacodia griseomixta
Dicliptera maclearii, Nisiocatantops orientalis, Sassula subviridis, Varcia affinis, Varcia
flavicostalis, Plautia grossepunctata, Orychodes andrewsi, Rhyparida rossi, Hemipyrellia
jucunda, Patellapis binghami, Megachile nivescens, Megachile rotundipennis
Comparison of numbers of endemic terrestrial species, and their fate, among some of Australia’s oceanic
islands. Extinctions here include species that are formally and validly listed as extinct or extinct in the
wild by the IUCN, Australian or relevant state/territory jurisdictions, with these tallies undoubtedly underestimating the actual number of extinctions. See text for main information sources for Lord Howe and
Norfolk Island groups.
* One additional now extinct species (Aplonis fusca) occurred only on both Lord Howe and Norfolk Islands.
Christmas
Lord Howe
Norfolk
Island area (km )
137
16
36
Island area including satellite islands (km 2)
137
17
38
17
c. 100
c. 38
No. of endemic plant genera
0
5
2
No. of extinct endemic plant spp.
0
1
1
No. of endemic invertebrate spp.
c. 200
c. 600
no known estimate
No. of endemic invertebrate genera
9
c. 40
no known estimate
No. of extinct endemic invertebrate spp.
0
1
1
No. of endemic terrestrial vertebrate spp.
16
5
4
6
4*
2*
2
No. of endemic plant spp.
No. of extinct endemic vertebrate spp.
ENDEMIC SPECIES OF CHRISTMAS ISLAND
DISCUSSION
There are distinctive features of the biodiversity
sampling history of Christmas Island that render
this exercise both easy and difficult. The unusually
extensive taxonomic breadth and sampling intensity
of the early (1887–1908) collections at the onset of
the island’s settlement provide, for many taxonomic
groups, an almost unparalleled baseline inventory.
Those collections help to now determine which
species in the current biota are native and which have
subsequently been introduced, and which species may
have subsequently been rendered extinct. However, for
many taxonomic groups, those early collections have
not been repeated, and without recent sampling it is
now difficult to assess whether some of the long-ago
described species (often represented in collections by the
holotype only or a very few specimens) are still valid,
and the extent to which those species have persisted,
declined or become extinct.
The ascription of endemism proved challenging for
many taxa, because: (i) for many species there were few
records (rendering it difficult to describe the bounds of
their distribution); (ii) for some species described long
ago it was difficult to trace the subsequent taxonomic
treatment; (iii) there is inconsistency in current specific
recognition; and (iv) some taxa considered as endemic in
previous studies are yet to be described.
99
psocid Sundapsocus, the grasshopper Nisiocatantops,
the cockroach Metanocticola, a new genus of copepod
(Arietellidae) (Bruce and Davie 2006) and Papasula
(comprising Abbott’s booby only). The crab family
Christmaplacidae, originally based on one Christmas
Island species, was considered endemic (Naruse and
Ng 2014) until the recent discovery of another species
assigned to the family, from Guam (Mendoza and Ng
2017).
For those taxonomic groups for which reasonably
comprehensive inventories are available, the extent
of endemism is summarised in Table 5. For at least
reptiles and mammals, a majority of native species
present (or formerly present) are endemic. Endemic
species comprise a smaller proportion of marine species,
presumably due to greater dispersal opportunities
in the marine environment. Birds are unusual in
that the number of native breeding species is small,
but the number of colonising, visiting and vagrant
species is proportionally large and growing steadily
(James and McAllan 2014). However, we note that this
comparison across taxonomic groups in the proportion
of endemic species is compromised; for better-studied
groups (the vertebrates and vascular plants), the total
number of native species and of endemic species is
well-documented, whereas neither tally is likely to be
complete for poorly-known groups.
We conclude that at least 253 described species are
endemic to Christmas Island (Table 2). Most are insects
(150 species), with the highest numbers in the orders
Coleoptera (58 species), Lepidoptera (25 species),
Hemiptera (24 species), Hymenoptera (12 species),
and Psocodea (10 species); there are also 27 endemic
species of Mollusca, 17 vascular plants, 15 crustaceans,
eight arachnids, six birds, five fish, five reptiles and
five mammals. A further 34 species were considered
to be endemic, or probably so, by those documenting
their occurrence, but have not yet been formally
described (Table 3). Our tallies for, and the complement
of, endemic species represent a notable advance on
that presented in a recent listing of endemic species
from several Australian islands, including Christmas
Island (Morris et al. 2018). That listing considered that
there were 36 species and five subspecies endemic to
Christmas Island (18 plants, four invertebrates, four
reptiles, ten birds and five mammals). Four of these are
shown here not to be endemic species (Dendrocnide
peltata, Zehneria alba, Asplenium listeri and Papilio
memnon). The disparity between the Morris et al. (2018)
list and our assessment is especially pronounced for
invertebrates.
Some of the endemic species are of particular
evolutionary and biogeographical significance. For
example, the ostracods Humphreysella baltanasi and
Microceratina martensi, the cave shrimp Procaris
noelensis and the cave microshrimp Halosbaena sp. nov.
represent lineages of great antiquity and remarkably odd
biogeography (Page et al. 2018), and may be relicts of
groups originating in the shallows of the Tethys Sea at
least as early as the Mesozoic (225 to 65 million years
ago) (Namiotko et al. 2004; Humphreys et al. 2009). The
relictual endemic Abbott’s booby is the most ancestral
of all the Sulidae, having diverged from all other living
relatives more than 20 million years ago, and to have
existed as a species for at least fifteen million years
(Olson and Warheit 1988; Patterson et al. 2011). This
species, and the Christmas Island frigatebird, both
feature in the top 50 bird species globally for their
combination of evolutionary distinctness and rarity (Jetz
et al. 2014). At least some of the Christmas Island reptile
species are also of great antiquity, reflecting a very long
period of isolation. For example, recent genetic analyses
have demonstrated that the endemic blue-tailed skink,
Lister’s gecko and forest skink diverged from their
nearest living relatives between 5 and 25 million years
ago (Oliver et al. 2018).
Endemism is not restricted to the species level;
there are about ten endemic genera – the crab genus
Christmaplax, the beetle genera Rhyncholobus and
Psammorpha, the hemipteran genera Andrewsiella
and Paurostauria (although the validity of the latter
genus may merit further scrutiny: Fletcher (2008)), the
The number of endemic species listed here is likely
to be a substantial under-estimate of the actual number
of endemic species given that some taxonomic groups
have been subject to no or only cursory inventory. For
example, the total list of wasps and bees known from
Christmas Island is 34 identified species, of which
100
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
we recognise 12 as endemic. However, the CSIRO
expedition of 1989 collected an estimated 299 morphospecies (CSIRO Division of Entomology 1990); Parks
Australia collected 71 morpho-species (James 2007)
and the CESAR Consulting Group collected a very
large number of parasitoid wasp species (Weeks 2013),
almost all un-named. Likewise, the rich Psocoptera
fauna of Christmas Island was largely undocumented
until the 1990s. Most of the recognised species came
from a single collection (in 1989) of 246 specimens,
representing 33 identifiable species that included ten
newly described ones (Smithers 1995). As another
example, recent sampling of freshwater invertebrates
concluded that ‘numerous macroinvertebrate species are
thought to be novel uncharacterised taxa that may be
endemic to Christmas Island’ (V. Pettigrove pers. comm.
in Weeks and McColl 2011).
The history of collections on Christmas Island is
unusual – it is marked by the extremely substantial
sampling undertaken in the 1–2 decades soon after
the island’s settlement in the 1880s, most notably by
Charles Andrews, with very intermittent sampling
subsequent to that (Figure 2). Other than the spurt of
new species described in the period 1887–1919, the rate
of description of endemic species has been reasonably
continuous and shows no sign of reaching an asymptote
(Figure 2). As documented in the accounts above, for
many taxonomic groups there has been little collecting,
for others substantial collections have not yet been
worked through, and for yet others new taxa have been
recognised but not yet described (e.g. Table 3). Hence, it
is likely that many endemic species remain undiscovered
and undescribed. Our review also concluded that
at least 141 species for which Christmas Island is
the type locality are no longer recognised as valid
endemic species (Table 4), mostly because the species
has subsequently been found elsewhere, or because of
taxonomic re-sorting.
On Christmas Island, losses of some endemic species
occurred very rapidly. The endemic Maclear’s rat Rattus
macleari and bulldog rat R. nativitatis were extremely
abundant at the onset of the island’s settlement, but were
rendered extinct within a decade (Andrews 1909; Green
2014). The Christmas Island pipistrelle Pipistrellus
murrayi remained abundant and apparently secure up
to at least the 1980s but declined rapidly thereafter,
culminating in extinction in 2009 (Martin et al. 2012;
Woinarski 2018). Likewise, three of the four endemic
lizards, and another native but not endemic lizard, were
also abundant in the 1980s, but became extinct or extinct
in the wild by 2012 (Smith et al. 2012; Andrew et al.
2018). For some of these vertebrates, the rapidity of loss
outstripped the pace of formal recognition as threatened.
For example, the Christmas Island forest skink Emoia
nativitatis was first listed as a threatened species in
2010 (at global scale) and January 2014 (nationally). This
was at about the time or after its extinction in the wild
in 2010 and shortly before the 2014 death of the last of
three individuals collected in the hope of establishing
a captive breeding program (Andrew et al. 2018). The
belated listing was notwithstanding substantial evidence
over several decades of severe decline (Smith et al.
2012). Likewise, the blue-tailed skink Cryptoblepharus
egeriae was not listed as threatened globally until it
was recognised as extinct in the wild in 2017, again
notwithstanding compelling evidence over several
decades of rapid and severe decline (Smith et al. 2012).
Fourteen endemic species and six endemic subspecies
are listed as threatened at national or global levels (Table
1), including six that are listed as extinct or extinct in
the wild. All of the latter are vertebrates. No endemic
plant or invertebrate species are listed as threatened at
the national level. This listing may now have belatedly
caught up with the imperilled (or extinct) status for most
terrestrial vertebrates, but it has notably failed to provide
adequate (or indeed, any) consideration of the status
of the majority of Christmas Island’s endemic species,
particularly the c. 200 endemic invertebrate species.
Only one of these endemic invertebrates (Ramulus
stilpnoides) has been accorded any conservation status
notwithstanding the absence of records of many species
for more than 100 years, and the likelihood that at least
some of the threats that have caused decline in endemic
plant and terrestrial vertebrate species (e.g. habitat loss,
introduction and proliferation of yellow crazy ants and
black rats) are also likely to have affected endemic
invertebrate species, perhaps to an even greater extent.
Even within some relatively well-known groups, there
has been anomalous treatment of conservation status
assignation. For example, the endemic plant Peperomia
rossii has not been recorded since 1897–98 but has no
assigned conservation status. In contrast, the relatively
abundant and still widespread palm Arenga listeri is
listed as Endangered at the global level.
Although noting the cautionary tale of the rapid recent
decline (from abundant in the late 1980s to extinction
c. 20 years later) of the Christmas Island pipistrelle
and Christmas Island forest skink, the date of the
most recent record may provide some indication of the
current status or at least flag a cause for concern for
some poorly-known endemic species or species-groups.
However, this date is also difficult to determine for many
taxa given incomplete documentation and determination
of some collections. Our assessment indicates that at
least 51 of the 253 endemic species (20%) have not been
reported for at least 100 years, and another 33 endemic
species (13%) have not been reported for 50–100 years
(Table 6). For the two vertebrates (Rattus nativitatis and
R. macleari) in this list of long-lost species, extinction
is validly recognised, but such status has not been
formally recognised for the similarly long-missing
invertebrate and plant species, although relevant reviews
have concluded so for some species, such as the flea
Xenopsylla nesiotes and tick Ixodes nitens – both hosted
by the extinct endemic Rattus macleari (Mihalca et al.
2011; Colwell et al. 2012; Kwak 2018).
CATALOGUE OF TYPE SPECIMENS OF DRAGON LIZARDS
Many of the endemic species that have not been
reported for at least 50 years may now be extinct,
and many others highly imperilled and may be in
need of urgent conservation management response,
lest they too become extinct. However, long periods
without reporting may also represent lack of sampling,
identification challenges, or simply inconspicuousness,
and may be a weak fou ndation for assu ming
imperilment. There are several notable examples of
101
re-discovery of Christmas Island endemic species after
long absences. For example, Ridley’s ground orchid
Zeuxine exilis was rediscovered in 2009 after not being
reported for more than 100 years (Green et al. 2010),
and has now been observed at numerous sites and often
in very large numbers (PG pers. obs.). The Christmas
Island shrew Crocidura trichura was extremely
abundant at the time of the island’s settlement (Andrews
1900c), but virtually disappeared between 1900 and
A
B
FIGURE 2
Date of description of species still recognised as endemic to Christmas Island. A) Number of species (still
recognised as endemic) described per decade; B) Cumulative number of such species.
102
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
1908 and was considered extinct in assessments in 1908
(Andrews 1909) and in the 1930s (Gibson-Hill 1947f).
However, two individuals were recorded in 1958 and
two more were collected in the mid-1980s (Eldridge et
al. 2014), although it has not been reported since. The
Christmas Island blind snake Ramphotyphlops exocoeti
has been reliably recorded only six times since 1901,
and only once since 1986 (Maple et al. 2012). In 2004
one of the Christmas Island jewel weevils, Rhyncolobus
rossi, was ‘rediscovered’ when a single specimen was
collected for the first time since Andrew’s original
collection in 1897–98 (Surman 2004). The conservation
status (or even whether extant or extinct) of such
nebulous species may be particularly difficult to assess.
Nonetheless, some precautionary consideration
of conservation status for poorly-known endemic
species is justifiable, given the range and intensity of
threats continuing to affect biodiversity on the island,
the demonstrated decline and extinction of many
well-known endemic species, and the retrospectivelyrecognised rate of extinctions of poorly-known species
on many other islands (Régnier et al. 2009, 2015a,b).
In the absence of evidence to the contrary, most or
all endemic terrestrial species on Christmas Island
should qualify (and hence be listed, at national and
global scales) as Endangered under IUCN categories
B1ab(ii,iii,v)+2ab(ii,iii,v) – that is, with extent of
occurrence between 100 and 5,000 km 2 (an actual
value of at most 137 km2) [=B1], area of occupancy
between 10 and 500 km2 (an actual value of at most 137
km2, but probably appreciably less given the extent of
habitat lost to mining and other development) [=B2],
occurrence at between one and five locations (an
actual value of one) [=a], and an inferred or projected
continuing decline [=b] in area of occupancy (reflecting
distributional contraction due to ongoing habitat loss
and likely exclusion of the species from areas with
crazy ant super-colonies, or coral bleaching, or other
threats) [=ii], area, extent and/or quality of habitat
(reflecting ongoing loss of habitat and decline in habitat
quality due to those same threats) [=iii], and number
of mature individuals (due to those same threats)
[=v]. This categorisation may not fit for subterranean
species (for which habitat loss or degradation may not
have occurred, or may not be continuing), although
these are likely to have been affected by other threats
arising from the island’s settlement and resource use,
especially groundwater extraction. The assessment
that the majority of Christmas Island’s endemic
species most likely qualify as Endangered under
IUCN categories B1ab(ii,iii,v)+2ab(ii,iii,v) matches
a comparable assessment of endemic invertebrates
on the Azores (Cardoso et al. 2011a). An alternative
approach may be to list many of these poorly-known
endemic species as Data Deficient. However such a
categorisation is not available at the national level under
Australia’s environmental legislation, and provides little
conservation security or obligation for management
prioritisation, and for most criteria (area of occupancy,
extent of occurrence, number of locations, and – less
so – likelihood of continuing decline) there is sufficient
information to exclude the need to use the Data Deficient
category.
Another option for providing some bolstered
conservation security for some Christmas Island
endemic species is by considering these species as
short-range endemics (Harvey et al. 2011) and using
the island’s existing governance matrix (which includes
applicability of some Western Australian law, including
in some environmental matters) to allow for some
explicit protection of such species under Western
Australian legislation (Environmental Protection
Authority 2009). Such listing is not currently available
under national or global conservation status processes,
strategies or law. This approach may be particularly
applicable for Christmas Island’s endemic subterranean
fauna, with comparable species in Western Australia
a particular focus for listing and protection as shortrange endemics. Most Christmas Island endemic species
would readily qualify for such listing, as they meet the
legislated criteria of being terrestrial and freshwater
invertebrates with distributions of less than 10,000 km 2.
In addition to consideration of such listing, a range
of other measures should be taken to help maintain the
endemic species, and hence the biodiversity significance
of this island. Given that introduced species are the
major cause of extinctions in island species globally
(Sax and Gaines 2008; Medina et al. 2011; Harper and
Bunbury 2015; Doherty et al. 2016; McCreless et al.
2016), a priority for conservation of Christmas Island’s
endemic biodiversity is to bolster the current weak
biosecurity efforts and standards (Beeton et al. 2010).
A reference collection of all endemic species (or
at least images of all endemic species) should be
established and maintained, to help raise awareness
of the existence of poorly-known endemic species,
and as a catalyst for reporting records of them: such
a catalogue of images is currently being compiled for
endemic lepidopteran species (C. Pink pers. comm.).
Targeted surveys should be undertaken for endemic
species, particularly those that have not been recorded
for many decades. Targeted surveys are also warranted
more broadly for those taxonomic groups that have
hitherto been subjected to little sampling. Research
should be conducted into poorly-known endemic
species that are likely to be most imperilled, with
such studies seeking especially to assess population
size, distribution, threats and management needs; and
such assessment should then be repeated regularly via
monitoring to evaluate population trends and responses
to management. Many of the species we considered
here to be confirmed or probable endemics may be
of uncertain taxonomic status, with little taxonomic
scrutiny since original collections more than 100 years
ago. For such species, taxonomic re-assessment may be
now long overdue.
CATALOGUE OF TYPE SPECIMENS OF DRAGON LIZARDS
Many of the poorly-known endemic species that
have been unrecorded for more than 50 years have no
precise locational data (often other than ‘Christmas
Island’). This renders such species difficult to consider
in assessments of possible impacts of proposed
developments and in other spatial conservation planning
exercises for the island. To be able to plan for their
conservation, more knowledge about these species is
required. Establishing how to identify all these endemic
species is a first essential step. Only then will it be
possible to start documenting systematic information
on geographical distribution, habitat preferences, foods
sources, life-cycles, etc. Most of the endemic species
are invertebrates and many of these are likely to have
been severely affected by supercolonies of crazy ants,
with some perhaps also affected by the extensive
application of insecticides used as temporary control
measures for these supercolonies. It is most likely that
populations of many endemic invertebrate species
will have been lost from these supercolony areas, but
may have persisted in those areas that have never been
affected by supercolonies. Because of high-resolution
mapping, these areas (that have not yet been exposed
to supercolonies of crazy ants) have been well-defined
(Boland et al. 2011), and such areas should now be
prioritised for survey for ‘lost’ endemic invertebrates
and excluded from ongoing development.
T here have been some not able examples of
consideration of poorly-known endemic species
in conservation management on the island, and
some examples of inadequate consideration. During
consideration of the introduction of a parasitoid wasp
as a biocontrol agent for the scale insects that are the
resource base for development of super-colonies of
crazy ants, managers inventoried hemipterans already
present on the island, assessed which were likely to
be endemic and evaluated the likely impact of the
proposed introduction of parasitoids on those species
(Neumann et al. 2016, 2018), although even in this case,
our review indicates that that previous scrutiny did not
recognise all endemic hemipterans. In contrast, other
than for two species of high profile charismatic crabs,
poorly-known and endemic species were not considered
at all in an assessment of the likely impacts of broadscale application of insecticides to control yellow crazy
ants (Stork et al. 2014), or a recently failed application
for further forest clearing to facilitate new mining
operations (Frydenberg 2018).
Some other conservation measures taken for the
threatened vertebrates have probably also provided
some benefit for poorly-known endemic species, as is
reportedly the case for biodiversity on other islands
(Aslan et al. 2015). These measures include the control
of yellow crazy ants, some environmental constraints on
mining and other developments, and the establishment
of a national park encompassing the majority of
the island. However, other substantial management
investments targeting threats to threatened endemic
103
vertebrates (notably a current program to control
feral cats) are unlikely to provide protection also to
imperilled endemic invertebrate or plant species; and
the establishment of a national park, while providing
a bulwark against mining and vegetation clearance,
is unlikely of itself to have prevented the impacts of
some other threats (notably pests, disease and weeds) to
endemic species.
Because of very variable sampling comprehensiveness
among island groups, it is challenging to compare the
number and fate of endemic species we report here for
Christmas Island with those of other islands. Albeit with
many caveats (notably including variation among islands
in the extent of sampling), we list some comparisons
with the two other biodiversity-rich oceanic islands of
Australia, Lord Howe Island (and its satellite islands)
and Norfolk Island (and its satellite islands), with
information for these two other island groups sourced
mainly from Cassis et al. (2003) and Department of
Environment and Climate Change (NSW) (2007)
for the Lord Howe Island group and Neuweger et al.
(2001), Cogger et al. (2006), Mills (2009) and Director
of National Parks (2010) for the Norfolk Island group.
There are several notable features of this comparison.
Although the number of known endemic invertebrate
and plant species on Christmas Island is substantial,
comparable tallies on Lord Howe and Norfolk are much
higher, notwithstanding their smaller size: and the
number of endemic genera is appreciably higher on Lord
Howe and Norfolk. In contrast, Christmas Island has
many more endemic vertebrate species. The extinction
of endemic vertebrates, but relative security of endemic
plants, is a notable common feature of the three islands.
However, the extent of extinction or persistence of
endemic invertebrates is difficult to assess on all of
these islands.
ACKNOWLEDGEMENTS
We thank Jeff Clausen, Claire Davies, Volker
Framenau, Alistair Graham, John Hueston, Vince
Kessner, Ian McAllan, Ruth Marr, Nicholas Milly, Lisa
Nichols, Max Orchard, Kent Retallick, Chris Surman
and Azmi Yon for their invaluable field and laboratory
contributions in the early stages of this endeavour. Andy
Austin (University of Adelaide), Tim New (La Trobe
University), Robert Mesibov (Ulverstone, Tas), Massimo
Olmi (Tuscia University, Italy), Max Orchard (Coles
Bay, Tas), Gustav Paulay (Florida Museum of Natural
History, USA), Caitlyn Pink and Alasdair Grigg (Parks
Australia), Nick Stevens (Stantec Australia), Melissa
Titelius (Western Australian Museum), Mark Heath,
Ken Walker (Museums Victoria) and Herbert Zettel
(Naturhistorisches Museum, Germany) kindly shared
their expertise with particular species and groups. We
thank also Mark Harvey for his editorial assistance.
JW’s contribution to this research was supported by
the Australian Government’s National Environmental
Science Programme (Threatened Species Recovery Hub).
D.J. JAMES, P.T. GREEN, W.F. HUMPHREYS AND J.C.Z. WOINARSKI
104
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