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RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 11Proceedings of th e hawaiian entom ological soci ety (2023) 55:11–27
Rediscoveries and Presumed Extinctions of Hawaiian
Leaf-roller Moths (Lepidoptera: Tortricidae)
Kyhl A. Austin and Daniel Rubinoff
Entomology Section, Depar tment of Plant and Environmental Protection Sciences, University
of Hawaii, 310 Gilmore Hall, 3050 Maile Way, Honolulu, Hawaii, 96822-2231, USA.
Corresponding author: kaaustin@hawaii.edu
Abstract. Three species of endemic Hawaiian leaf-roller moths were rediscovered
after a 90-year gap in collection records. Twenty-one other species are presumed
extinct, with no collection records from the past 50 years. Remarks concerning the
type localities of four species (Cydia chlorostola, C. gypsograpta, C. parapteryx,
and Pararrhaptica leucostichas) described by Meyrick are given. A list of all
described endemic Hawaiian Tortricidae is provided with corresponding years for
when each species was last reliably seen or collected as well as the type locality
(when known). For each species, we include a list of known or hypothesized host
plants, which we hope will spur future rediscoveries and conservation efforts for
this group which has, to date, been ignored in conservation planning.
Key words. Conservation, endangered species, endemic, insects, host plant
The Hawaiian Islands are commonly
referred to as the “extinction capital of the
world,” but literature supporting this claim
is largely focused on the most well-studied
and often charismatic taxa (Régnier et al.
2015, Paxton et al. 2018, Rønsted et al.
2022). In contrast, there are exceedingly
few studies on extinction rates in the Ha-
waiian insect fauna. The rst and only
comprehensive attempt was undertaken in
the 1980s (Gagné 1982, Gagné and Chris-
tensen 1985). With funding from the U.S.
Fish and Wildlife Service, Wayne Gagné
and colleagues assessed approximately
800 endemic arthropod species to deter-
mine their conservation status, including
22 species of Omiodes (Crambidae) and
a handful of endemic macrolepidoptera.
Haines et al. (2004) subsequently reported
the rediscovery of ve species of Omiodes
on Hawaiʻi island considered extinct by
Gagné (1982), offering hope that similar
rediscoveries could be made. Understand-
ably, these researchers could not include
all endemic Lepidoptera in their studies
due to the sheer numbers of described spe-
cies and the lack of data for the majority of
them. Since then, taxonomic monographs
have alluded to possible extinctions in a
select few genera: Scotorythra (Heddle
20 03), Thyrocopa ( Med ei ros 2009),
Cydia (Oboyski 2011), and Philodoria
(Kobayashi et al. 2021), but no order-wide
assessment has been made to date.
Though we investigated only a small
fraction of the approximately 6,000
described endemic arthropod species
(Nishida 2002, Hembry et al. 2021), which
includes 930 Lepidoptera (Nishida 2002;
Austin and Rubinoff 2022), our assessment
provides an importa nt glimpse into the dire
situation for many endemic insects.
As part of an ongoing effort to catalogue
possible extinctions in endemic Hawaiian
Lepidoptera, we visited museum collec-
tions, reviewed literature, and conducted
eld work throughout the state in order
to build a comprehensive database of all
12 austin and Rubinoff
species, including the year each species
was last reliably seen or collected. Dozens
of “rediscoveries” of species have been
documented to date (in prep.), but because
of the paucity of existing literature and
identified material in museum collec-
tions, it is often difcult to determine
if species were actually “rediscovered”
or are relatively common and have been
simply overlooked and unidentied by
entomologists until now. Towards that end,
we present a snapshot of the Hawaiian Tor-
tricidae, historically a diverse family with
a disproportionate number of independent
colonizations and subsequent radiations
(Hembry et al. 2021), and which appears
to be particularly hard-hit by extinctions
in the past century.
There are 66 described endemic species
of Hawaiian Tortricidae, along with dozens
of undescribed species. Four authors are
responsible for the named species: A.G.
Butler (1844–1925), Lord Walsingham
(1843–1919), E. Meyrick (1854 –1938),
and O.H. Swezey (1869–1959). Of these,
only Swezey lived in Hawaiʻi. Butler and
Walsingham based their descriptions on
material sent to London by R.C.L. Perkins
(1866–1955). Meyrick used material col-
lected by both Perkins and Swezey. No
native Tortricidae have been described
since 1946, although many introduced spe-
cies have been reported or released since
then (Davis 1959, 1962, Davis and Krauss
1966, Pogue 1988, Miller and Hodges
1995, Austin and Rubinoff 2022, 2023).
The Hawaiian Tortricidae are a diverse
group of endemic insects, having inde-
pendently colonized the Hawaiian Islands
at least ten times (Zimmerman 1978,
Hembry et al. 2021), but possibly more,
making them the most successful colonists
of any family of Hawaiian Lepidoptera,
and exceeding virtually all other native
terrestrial arthropod families (Zimmer-
man 1948). Most genera, including several
undescribed ones, are highly host-specic,
utilizing a single family of native plants,
feeding typically as leaf-tiers or -rollers
on leaves, but some occur in fruits, stems,
seeds, and bark. These specialized feed-
ing habits (often on uncommon or rare
plant species), coupled with their appar-
ent vulnerability to introduced predators
and parasitoids (naked larvae, only rarely
internal feeders) seem to have made them
exceptionally prone to extinction.
Materials and Methods
The following museum collections were
examined to establish the most recent
year each species was collected: Bernice
Pa uahi Bishop Muse um (BPB M), Hawa iʻi
Department of Agriculture (HDOA), and
University of Hawaiʻi Insect Museum
(UHIM). For some species, only the
holotype, housed in the Natural History
Museum, London (NHMUK, formerly
the British Museum of Natural History),
is known. Identications were made using
original descriptions, supplemented by
Zimmerman (1978). Genitalia dissections,
when necessary, were prepared using
standard techniques and slide-mounted,
following Landry (2007). Label data are
copied verbatim except for “USA: Ha-
waii,” collection-event specic codes, and
museum accession/catalogue numbers,
which are omitted. Slashes (/) separate
data labels when multiple labels are pres-
ent on the same specimen.
Several species are currently placed in
incorrect genera or are members of unde-
scribed genera; these species are written
with their current genus in quotation
marks. Manuscripts are currently being
prepared to transfer these species to the
correct genus and describe new genera as
necessary. Host data were gathered from
Zimmerman (1978), Oboyski (2011), and
museum specimens. Hypothesized host
plants are based on the known host plants
of congeners or presumed congeners in
the case of misplaced species. Evenhuis
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 13
(2007) was used to identify type localities
based on the dates provided on Perkins’
data labels; localities identied in such a
way are written in brackets in Table 1.
The arbitrary date of 1972 was selected
as the cut-off date for presuming a species
extinct, representing 50 years since the last
known specimen was collected. Though
many species are reported from multiple
islands (Zimmerman 1978), we only list
the island containing the type locality
as correct, as most species appear to be
single-island endemics, with undescribed
sister species on other islands. A few spe-
cies (e.g., Bactra straminea, Crocidosema
blackburnii, Cryptophlebia illepida) ap-
pear to occur on multiple islands, and it
is unclear whether they represent endemic
species or early introductions from un-
known source regions.
Results
Pararrhaptica leopardellus
(Walsingham)
Rediscovery
This species was originally described
from a single male collected in 1895 at
Kahōluamanu, Kauaʻi by R.C.L. Perkins.
J.A. Kusche collected a male and female
at the same locality in 1920, which KAA
identified in BPBM. No other known
specimens have been collected since. Here
we report it from six specimens collected
on the Nuʻalolo Trail in Kōkeʻe State Park.
Larvae are suspected to feed on Myrsine
lanaiensis (Primulaceae), based on the
abundance of typical Pararrhaptica-like
feeding damage on leaves of M. lanaiensis
in the immediate vicinity of the collecting
locality of the most recent specimens.
Material examined. Ka uaʻ i: 1♂, 2♀♀,
Kōkeʻe State Park, Nuʻalolo Trail; 22.1310,
–159.6607; 1160 m; 22–23 iii 2022; K. A.
Austin; LED bucket trap / DNA extrac-
tions KA0493, KA0494, KA0497 / KAA
diss. #0738(♀), #0739(♀), #0742(♂)
(U HIM). 3♂♂, same as previous except
22.1312, -159.6606; 1165m / DNA extrac-
tions KA0496, KA0498, KA0499 / KAA
diss. #0741, #0743, #0744 (UHIM).
Spheterista oheoheana (Swezey)
Rediscovery
This species was previously known only
from the type series of ve specimens
(four males, one female) reared “from
larvae and pupae found in dead twigs” of
Polyscias kavaiensis (Araliaceae) at Hale-
manu, Kauaʻi in 1932 (Swezey 1933). All
specimens of the type series are in poor
condition, with wings heavily rubbed and
not spread. We report a single female from
the Nuʻalolo Trail. A few Polyscias ka-
vaiensis were observed nearby, although
no dead twigs were noticed.
Spheterista oheoheana is one of three
species of Hawaiian Tortricidae to have
been included in a USFWS Candidate
Notice of Review (USFWS 1984), mean-
ing it has been considered for possible
protection under the Endangered Species
Act. It was listed as a category 2 candidate
species, meaning that “conclusive data on
biological vulnerability and threat are not
currently available” to support inclusion.
The other two species included in the
review, also category 2, S. pterotropiana
and S. reynoldsiana, have been collected
in recent years (although S. reynoldsiana
may be ext i nct on Oʻa hu). All th r e e sp e cie s
were removed from consideration in 1996
when the USFWS discontinued the use of
category 2 (USWFS 1996).
Material examined. Kauaʻi: 1♀,
Kōkeʻe State Park, Nuʻalolo Trail; 22.1310,
-159.6607; 1160m; 22-23 iii 2022; K. A.
Austin; LED bucket trap / DNA extraction
KA0556 / KAA diss. #0787 (UHIM).
Spheterista tetraplasandra (Swezey)
Rediscovery
This species was originally described
from two males reared from fruits of “Tet-
raplasandra” [=Polyscias] (Araliaceae)
14 austin and Rubinoff
in 1918 from Kaumuahona1 and Wailupe,
Oʻahu (Swezey 1920). A few additional
specimens were collected in the following
years, most recently a small series reared
from “Tetraplasandra” [=Polyscias] at
Puʻu Kaua, Oʻahu in 1932. One specimen
is known from Kauaʻi, but this may rep-
resent a closely related undescribed spe-
cies. We report three specimens from two
separate locations in the central Koʻolau
Mountains of Oʻahu.
Material examined: Oʻahu: 1♂, Ho-
nolulu Watershed For[est] Res[erve], nr.
summit of Moanalua Ridge Trail along
K[oʻolau] S[ummit] T[rail]; 21.3945,
-157.8243; 830m; larva coll[ected] 22
viii, pupa[ted] 24 viii, adult ecl[osed] 1 ix
2021; K. A. Austin, K. Faccenda / HOST:
Polyscias (Tetraplasandra) oahuensis
(Araliaceae), leaet-tier / DNA extraction
KA0297 / KAA diss. #0595 (UHIM).
2♂♂, ʻEwa For[est] Res[erve], Manana
Ridge, inside fenced area; 21.4502,
-157.8887; 575m; 24-25 xi 2021; K. A.
Austin, K. Faccenda; LED bucket trap
/ DNA extractions KA0387, KA0388 /
KAA diss. #0650, #0651 (UHIM).
The following species are presumed
extinct owing to the absence of records
over the past 50 years:
Cydia chlorostola (Meyrick)
Presumed extinct
This species is known only from the
female holotype, supposedly collected
in 1909 by R.C.L. Perkins from Waialua,
Oʻa hu (but se e com m ent s below). Mey r ick
(1932) suggested that it was introduced
along with a hypothesized leguminous
host, possibly from Asia. Zimmerman
(1978) disagreed and considered it en-
demic. Oboyski (2011) agreed with Zim-
merman and compared it to several
Canavalia-feeding Hawaiian Cydia. It
appears very similar to C. parapteryx
(Meyrick), also only known from Oʻahu.
Cydia crassicornis (Walsingham)
Presumed extinct
This species is known only from two
males collected by R.C.L. Perkins in
1892 at 4000 ft near Kona, Hawaiʻi island
(Walsingham 1907). Although possibly a
color form of the koa-feeding C. walsing-
hamii (Butler), Oboyski (2011) considered
it distinct based on subtle differences in
the male genitalia. Perkins’ eld notes for
September 1892 indicates that he “col-
lected … by sifting dead leaves at the foot
of a big koa tree” (Evenhuis 2007). The
forewings are very similar to C. conspicua
(Walsingham), but the absence of a “sex
pouch” (Zimmerman 1978) on the ventral
surface of the male hindwing readily
separates it from that species.
Cydia gypsograpta (Meyrick)
Presumed extinct
This species is known only from the
male holotype supposedly collected by
R.C.L. Perkins near Honolulu, Oʻahu in
1908 (Meyrick 1932; but see comments
below). Oboyski (2011) found it difcult
to distinguish from Canavalia-feeding
Hawaiian Cydia.
Cydia obliqua (Walsingham)
Presumed extinct
This species is known from three
female specimens collected at Hualālai
(5000 ft) and Kona (4000 ft) on Hawaiʻi
island by R.C.L. Perkins in 1892. Oboyski
(2011) considered it possibly just an ex-
treme color form of the widespread and
polymorphic C. plicatum (Walsingham),
but refrained from synonymizing it
because of signicant differences in the
female genitalia. The collection date given
for the two female types from Hualālai (5
viii 1892) corresponds to a journal entry
in which Perkins wrote, “there were two
Tortricids on Koa” (Evenhuis 2007).
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 15
Eccoptocera osteomelesana (Swez ey)
Presumed extinct
This species is the most recent na-
tive Hawaiian tortricid to be described
(Swezey 1946). It has been reared from
leaves of Osteomeles anthyllidifolia (Ro-
saceae) and is known from small series
of specimens collected in Mānoa and
Wailupe Valley on Oʻahu and was most
recently collected on the campus of UH
Mānoa in 1969. However, it may have been
introduced, as it more closely resembles
some Pacic and Australian Eccoptocera
than native Hawaiian ones. It is also no-
table that no other entomologist collected
this species prior to F.X. Williams in
1929, despite it having been reared and
collected in such heavily urbanized places.
No specimens have been collected by us
despite extensive searching near large
patches of Osteomeles in the southern
Koʻo lau Mou ntai n s and ext ens ive tra ppi ng
in areas it might be expected.
Nuritamburia phyllanthana (Swezey)
Presumed extinct
This species is known from the type
series of seven specimens reared from
Phyllanthus distichus (Phyllanthaceae)
from Malamalama Ridge, Oʻahu in 1916
and 1925 (Swezey 1940). No other speci-
mens are known, and targeted collecting
near P. distichus on Oʻah u did not re su lt in
any specimens. The hostplant has not been
encountered frequently by us, although
Wagner et al. (1999) considered it “locally
common in mesic forest, often on steep
slopes or ridge tops, sometimes in dry
shrubland” on Kauaʻi, Oʻahu, Molokaʻi,
Lānaʻi, West Maui, and rarely on East
Maui.
Paraphasis perkinsi (Walsingham)
Presumed extinct
This enigmatic genus and species is
known only from the male collected in
1894 from 3000– 4000 ft elevation on
Kauaʻi by R.C.L. Perkins. This area has
been consistently surveyed by us and
other entomologists over the past century.
Walsingham (1907) originally described
Paraphasis in Tineidae, but Zimmer-
man (1978) treated it as a tortricid. The
illustration of the head in Zimmerman
(1978) appears to show a basally scaled
proboscis, a character only very rarely
seen in Tortricidae (Diakonoff 1977). In
addition, the wing venation and genitalia
would be highly unusual for a tortricid. In
the absence of a more compelling family
placement, we consider it a tortricid for
the purposes of this paper.
Pararrhaptica capucina (Walsingham)
Presumed extinct
This species was described from two
females: the holotype collected in 1893
at 4000 ft on Molokaʻi and a question-
ably associated paratype collected in
1892 at 3000 ft near Kona, Hawaiʻi island
(Walsingham 1907). The two specimens
are likely not conspecic, as most Parar-
rhaptica appear to be single island en-
demics. Several other high islands host
similar-looking Pararrhaptica, including
P. chlorippa (Meyrick) on Oʻahu (see be-
low). Howev er, in recent years on Molok aʻi
we have collected a species very similar
to the type of P. capucina in genitalia but
externally more closely resembling P. lon-
giplicatus (Walsingham), described from
Maui, suggesting that perhaps only one
species is involved and that the holotype of
P. capucina is just an unusual color form
of a more widespread Maui Nui species.
Pararrhaptica chlorippa (Meyr ick)
Presumed extinct
This species was described from a
single male specimen reared from Myr-
sine lessertiana (P rimulaceae) by Swezey
in 1911 from Mt. Olympus, Oʻahu. Perkins
collected a small series of what appear
to be this species in 1901 and 1908 from
16 austin and Rubinoff
various locations in the Koʻolau Moun-
tains. The associated females appear very
similar to the holotype of P. capucina
(Walsingham), known from Molokaʻi. It
appears likely that these two species were
closely-related. A similar, undescribed
sp e cies occ urs on Kau aʻi an d is still ext a nt.
Pararrhaptica dermatopa (Mey rick)
Presumed extinct
This species was described from two
males and a female reared from three
different dates from Myrsine lessertiana
(Primulaceae) by Swezey in 1911 on Mt.
Olympus and Tantalus, Oʻahu. Meyrick
(1932) wrote that a second, larger female
from Mt. Olympus “may be the same spe-
cies,” but we have not been able to locate
this specimen.
Pararrhaptica lysimachiae (Swez ey)
Presumed extinct
This species is known only from the
male holotype, supposedly reared from
leaves of Lysimachia hillebrandi va r.
venosa [=L. venosa] (Pr imulaceae) by
Swezey in 1932 on the “Kalalau Trail” at
an elevation of 3800 ft. The collecting site
may refer to the trail leading down from
the Kalalau Lookout at Kōkeʻe State Park.
Zimmerman (1978) listed Lysima chia
glutinosa as an additional host, but we
are not sure of his justication, as we
are only aware of the holotype for this
species. Perhaps Zimmerman (1978)
believed Swezey’s initial plant identica-
tion was incorrect; Lysimachia venosa is
an exceedingly rare species, known only
from three specimens from the summit of
Waiʻaleʻale (Marr and Bohm 1997) until
its rediscovery in 2012 (Wood 2013). On
the other hand, Lysmiachia glutinosa oc-
curs somewhat commonly along the upper
portion of the trail leading down from the
Kalalau Lookout at Kōkeʻe State Park
(KAA pers. obs.).
Pararrhaptica lysimachiana (Swezey)
Presumed extinct
This species is known only from the
female holotype, reared from leaves of
Lysimachia rotundifolia [=L. hillebran dii]
(Primulaceae) by Swezey at Puʻu Hāpapa
on Oʻahu in 1927. Collecting by KAA
at the type locality did not result in any
specimens, despite the host plant’s pres-
ence in the immediate vicinity, and many
island-wide surveys.
Pararrhaptica punctiferanus
(Walsingham)
Presumed extinct
This species was described from two
specimens: the holotype male collected by
Perkins in 1894 at 5000 ft on Haleakalā,
Maui and a questionably associated female
paratype, collected in the same year at
4000 ft elevation on Molokaʻi. The female
paratype likely represents an undescribed
species, as most Pararrhaptica appear to
be single island endemics. Like most other
Pararrhaptica, it likely fed on Myrsine
spp. (Primulaceae).
Pararrhaptica subsenescens
(Walsingham)
Presumed extinct
This species is known from a single
female collected by Perkins in 1893 on
Molokaʻi (Walsingham 1907). No further
locality data are given on the label, but the
date (5 viii 1893) corresponds to a journal
entry by Perkins from “near Waikolu”
(Evenhuis 2007). We have collected in this
general area and along the Hanalilolilo
Trail but have not found this species. Like
other Pararrhaptica, it likely fed on leaves
and shoots of Myrsine spp. (Primulaceae).
“Pararrhaptica” trochilidanus
(Walsingham)
Presumed extinct
This species is known from three
specimens: the holotype female and two
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 17
paratype males collected by Perkins at
elevations of 3000–4000 ft on Molokaʻi.
It appears closely related to “P.” fuscoviri-
dis (Walsingham) from Lānaʻi and “P.”
lichenoides (Walsingham) from Hawaiʻi
island and likely fed on Psychotria spp.
(Rubiaceae) or Nestegis sandwicensis
(Oleaceae) like other members of this
undescribed genus.
Spheterista argentinotata
(Walsingham)
Presumed extinct
This species is known from two fe-
males collected by Perkins in Olaʻa and
Hilo, Hawaiʻi island in 1895 and 1896,
respectively (Walsingham 1907). In the
absence of known males, it is difcult to
ascertain for certain whether it belongs
to Spheterista or another genus. If Sph-
eterista, it likely fed on leaves or fruit of
Cheirodendron or Polyscias (Araliaceae).
“Spheterista” cassia (Swezey)
Presumed extinct
This species may have been introduced.
It does not belong in Spheterista and more
closely resembles some Australian tortri-
cids, such as Merophyas Common, than
any native Hawaiian species. It has been
reared from leaves of Senna gaudachaudii
(Fabaceae), an indigenous species that also
occurs in Australia and other islands in
the Pacic. There are large series of this
species in HDOA and BPBM, suggesting
it was once common on several islands. It
has not been collected since 1920; we have
not encountered its host plant in the wild.
“Spheterista” avocincta
(Walsingham)
Presumed extinct
This species was described from a
single female collected by Perkins in
1892 near Kona, Hawaiʻi island at an
elevation of 4000 ft. Walsingham (1907)
described the male of this species as
Capua trigonifer, later synonymized by
Zimmerman (1978). Records of this spe-
cies on islands other than Hawaiʻi island
are likely erroneous. Capua santalata
Swezey, described from Oʻahu and cur-
rently considered a junior synonym of “S.”
avocincta, is still extant and probably
deserves its species status restored. True
“S.” avocincta appears to be endemic to
Hawaiʻi island and has not been collected
since 1911. It probably fed on Santalum
spp. (Santalaceae) like its close relatives.
Spheterista pernitida (Walsingham)
Presumed extinct
This species is known from a single
fema l e col le c ted at Olaʻa on Hawa iʻi isla n d
in 1896 by Perkins (Walsingham 1907).
However, in the absence of males, it is
difcult to determine whether it belongs
to Spheterista or another genus. If Sph-
eterista, it likely fed on leaves or fruit of
Cheirodendron or Polyscias (Araliaceae).
The male mentioned by Walsingham
(1907) as being “closely allied to perni-
tida” is almost certainly not conspecic.
The male genitalia closely resemble those
of the Urticaceae-feeding “Spheterista”
(“S.” infaustana, “S.” urerana).
“Spheterista” urerana (Swezey)
Presumed extinct
This species was described from a
series of four specimens reared from
twigs of “Urera sandwicensis” [= U.
glabra] (Urticaceae) by Swezey in 1914
on Tantalus, Oʻahu (Swezey 1915). Zim-
merman (1978) designated a lectotype and
paralectotype and removed the latter from
the cork mount. It does not appear he was
able to locate the other two specimens in
the series. We were unable to locate the
paralectotype, though it is supposedly
deposited in the BPBM. Females are un-
known.
This species may be a synonym of “S.”
infaustana (Walsingham), described from
18 austin and Rubinoff
Kauaʻi. Alternatively, the type series could
be an unusual color form of a widespread
extant species on Oʻahu closely related to
“S.” infaustana but provisionally consid-
ered undescribed by KAA. It is still com-
mon on parts of Tantalus where Pipturus
and Urera grow.
“Spheterista” xanthogona
(Walsingham)
Presumed extinct
This species is only known from two
females, collected by Perkins in 1892 near
Kona, Hawaiʻi island at 4000 ft elevation
(Walsingham 1907). It appears to belong
to the same Santalum-feeding group of
“Spheterista” as “S.” avocincta (Wals -
ingham), from Hawaiʻi island, and “S.”
avopicta (Walsingham), from Kauaʻi.
Journal entries by Perkins around the
collection dates of these two moths men-
tion the abundance of Santalum (Sant a-
laceae) in the area as well as Acacia koa
(Fabaceae) and Myoporum sandwicense
(Scrophulariaceae) (Evenhuis 2007), al-
though the exact location is unclear.
Additional comments
For three species described by Mey-
rick in 1932 (i.e., Cydia gypsograpta, C.
parapteryx, and Pararrhaptica leucos-
tichas), “Honolulu” is given as the type
lo ca l ity. Cydia chlorostola, also described
by Meyrick in the same publication, was
from “Waialua.” These four species sup-
posedly collected or reared by Perkins
between 1906 and 1909 were almost
certainly collected elsewhere in the ar-
chipelago.
By 1904, Perkins had started working
for the Hawaiian Sugar Planters Asso-
ciation (HSPA) in Honolulu and stopped
traveling to neighboring islands (Evenhuis
2007). However, his biocontrol partner,
Albert Koebele, went to other islands
and collected and brought back larvae
for rearing by Perkins (Evenhuis, pers.
comm.). Specimens of these four species,
now in the NHMUK, have different labels
than Perkins’ Fauna Hawaiiensis material
collected in the 1890s, and the handwrit-
ing on them does not appear to be his. We
believe Meyrick mistakenly interpreted
these package’s points of origin as the
type locality; post ofces existed in both
Waialua and Honolulu at this time and
were frequented by Perkins and colleagues
(see Evenhuis 2007).
With the exception of C. parapteryx, all
of these species were previously known
only from the type series. Oboyski (2011)
identied additional material from Oʻahu
as C. parapteryx, including additional
HSPA reared material from “Honolulu”
and various eld stations as well as wild-
caught and -reared specimens from the
Waiʻanae and Koʻolau Mountains from
more specic localities. A species identi-
cal to the type series of Pararrhaptica
leucostichas has been collected com-
monly on Hawaiʻi island in recent years,
suggesting that the true type locality for
it is no t Oʻahu. Simila rly, a spe cie s close ly
resembling C. gypsograpta was collected
recently by the authors in Hawaiʻi Vol-
canoes National Park, but we hesitate to
identify it condently as such at present.
Cydia chlorostola closely resembles some
of the Canavalia-feeding Cydia species
known from Hawaiʻi island, Maui, Oʻahu,
and Kauaʻi and may be a synonym of one
of them.
These four species appear to be the
only Hawaiian species of Lepidoptera
described by Meyrick in such a manner.
However, other authors may have made
the same mistake. For example, Agrotis
diplosticta (Hampson) was similarly de-
scribed from “Waialua,” despite all other
collections of this species being from the
mountains of Oʻahu and Kauaʻi. As far as
we are aware, all other Meyrick species
were based on Fauna Hawaiiensis mate-
rial overlooked by Walsingham (1907) or
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 19
sent to London by Swezey and included
data labels. We suggest treating the type
locality of these four species as unknown,
as well as similar occurrences. It would
be useful to attempt to nd HSPA rearing
records to nd out where these specimens
may have been collected before being
shipped to London.
Discussion
The rediscovery of the three species
reported here are cause for hope that fu-
ture rediscoveries can be made, although
there is limited cause for optimism given
the apparent extinction of 21 other species,
representing nearly a third of the native
Hawaiian tortricid fauna. Several of the
species we found in recent years likely
are on the verge of becoming critically
endangered. As native Hawaiian plants
disappear, the available habitat and hosts
for the moths also recede. Given the un-
abated onslaught of invasive species, it is
only a matter of time before more moth
species are lost forever. This will have
cascading effects on native ecosystems
across the state as tortricids represent one
of the most signicant native herbivore
groups in Hawaiʻi and a major source of
food for native forest birds (Banko et al.
2022) and bats (Pinzari et al. 2019). Gen-
era such as Eccoptocera and Spheterista
can still be abundant in pristine wet forest
and indicate the importance of tortricid
moths in native ecosystems. Such species
might serve as useful indicators of overall
forest health.
Without rapid intervention, many rare
species will soon blink out of existence –
without much notice or fanfare – as the
disappearance of the aforementioned 21
species indicates. A critical rst step may
be federal listing for the most vulnerable,
but still extant, species in order to allocate
resources to better understand their cur-
rent ranges, life histories, and most sig-
nicant threats, as these are all virtually
unknown for the majority of native moth
species. For native moths in general, the
most critical component of their conserva-
tion is likely the protection and restoration
of their host plants and habitat.
Acknowledgments
We thank Jim Boone (BPBM), Jeremy
Frank (BPBM), Janis Matsunaga (HDOA),
and Coquelicot Shirey (Hawaiʻi Volcanoes
National Park) for allowing us to examine
material under their care and facilitating
loans. We thank Neal Evenhuis for provid-
ing helpful comments on Perkins’ life and
work. We thank Ryan Peralta (Oʻahu DO-
FAW), Mapuana O’Sullivan (Kauaʻi DO-
FAW), Lance DeSilva (Maui DOFAW),
Cynthia King (Hawaiʻi State DLNR), Dee-
na Gary (University of Hawaiʻi at Mānoa),
Karl Magnacca (Army Natural Resources
Program, Oʻahu), Russell Kallstrom (The
Nature Conservancy, Molokaʻi), Kerri Fay
(The Nature Conservancy, Maui), and
Adam Radford (The Nature Conservancy,
Maui) for providing collection permits,
site access, and logistics. We thank Will
Haines, Matt J. Medeiros, Camiel Dooren-
weerd, Michael San Jose, Brad Reil, Kevin
Faccenda, and Malia Staab for help with
eldwork. We thank John Brown and Matt
J. Medeiros for their careful review of this
manuscript and their useful feedback. This
study was supported, in part, by USDA
Cooperative Agreement PPA 3.0877.03:
“Diagnostic tools to identify exotic Tor-
tricidae that threaten U.S. agriculture.
Additional funding was provided by The
College of Tropical Agriculture and Hu-
man Resources (CTAHR), University of
Hawaiʻi at Mānoa; and USDA Cooperative
State Research, Education and Exten-
sion (CSREES), Grant/Award Number:
HAW00942-H.
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Footnote
1This place name, now out of use, refers
to “the ridge leading up to Kōnāhuanui,
forming the east side of Nuʻuanu Valley,
and in particular the portion on the east
side of Lulumahu Stream” (Magnacca
2013).
22 austin and Rubinoff
Bactra straminea
(Butler 1881)
2022 UHIM Bolboschoenus, Carex,
Machaerina
Oʻahu “salt marshes near Honolulu”
Crocidosema blackburnii
(Butler 1881)
2022 UHIM Abutilon, Sida Maui Makawao
Crocidosema leprarum
(Walsingham 1907)
2022 UHIM Sesuvium Molokaʻi sea level
Crocidosema marcidellum
(Walsingham 1907)
2022 UHIM Abutilon, Hibiscus Oʻahu Waiʻanae Mountains [exact location unclear]
Cryptophlebia illepida
(Butler 1882)
2022 UHIM polyphagous Oʻahu [“in the neighborhood of Honolulu”, see com-
ments in Zimmerman 1978]
Cydia chlorostola †
(Meyrick 1932)
1909
NHMUK
Canavalia? Oʻahu* Waialua*
Cydia conspicua
(Walsingham 1907)
2022 UHIM Acacia koa Maui Haleakalā, 5000 ft
Cydia crassicornis †
(Walsingham 1907)
1892
NHMUK
Acacia koa? Hawaiʻi Kona [vicinity of Pulehua?], above 4000 ft
Cydia falsifalcellum
(Walsingham 1907)
2005 Oboyski
2011
Canavalia, Vicia Hawaiʻi Olaʻa, 2000 ft
Cydia gypsograpta †
(Meyrick 1932)
1908
NHMUK
Canavalia? Oʻahu* Honolulu*
Cydia latifemoris
(Walsingham 1907)
2022 UHIM Sophora chrysophylla Maui Haleakalā crater
Last Type locality
Species collected Source Host(s) (island) Type locality
Table 1. List of Hawaiian Tortricidae with corresponding years for which each species was last seen and known (without question mark) or
hypothesized (with question mark) host plants. Comments in brackets regarding type localities are based on Perkins’ eld notes transcribed by
Evenhuis (2007). †presumed extinct (has not been collected in 50+ years) *see comments in text regarding the type locality for this species.
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 23
Cydia montanum
(Walsingham 1907)
2022 UHIM Acacia koa Hawaiʻi Kīlauea
Cydia obliqua †
(Walsingham 1907)
1892
NHMUK
Sophora chrysophylla?
Acacia koa?
Hawaiʻi Hualālai, 5000 ft
Cydia parapteryx
(Meyrick 1932)
2022 UHIM
Canavalia, Strongylodon
Oʻahu* Honolulu*
Cydia plicatum
(Walsingham 1907)
2022 UHIM Sophora chrysophylla Hawaiʻi Kona [Kanahaha], 4000 ft
Cydia rupennis
(Butler 1881)
2022 UHIM Acacia koa Oʻahu [“mountains of Oʻahu”, see comments in Zimmer-
man 1978]
Cydia storeella
(Walsingham 1907)
1988 UHIM Fabaceae? Maui Haleakalā, 5000 ft
Cydia walsinghamii
(Butler 1882)
2022 UHIM Acacia koa Oʻahu [“in the neighborhood of Honolulu”, see com-
ments in Zimmerman 1978]
Eccoptocera foetorivorans
(Butler 1881)
2022 UHIM Metrosideros Oʻahu “Mountains of Oʻahu”
Eccoptocera osteomelesana †
(Swezey 1946)
1969 UHIM Osteomeles anthyllidi-
folia
Oʻahu Woodlawn, Mānoa Valley
Macraesthetica rubiginis
(Walsingham 1907)
2022 UHIM Unknown Oʻahu Waiʻanae Mountains, [>1300 ft]
Mantua fulvosericea
(Walsingham 1907)
2022 UHIM Xylosma, Flueggea Molokaʻi [gulch SE of Makakupaʻia], 3500 ft
Nuritamburia chlorocalla
(Walsingham 1907)
2022 UHIM Perrottetia Hawaiʻi Olaʻa
Last Type locality
Species collected Source Host(s) (island) Type locality
24 austin and Rubinoff
Nuritamburia metallurgica
(Walsingham 1907)
2022 UHIM Perrottetia, Xylosma,
Myrsine
Molokaʻi 3000 ft
Nuritamburia phyllanthana †
(Swezey 1940)
1925 HDOA Phyllanthus distichus Oʻahu Malamalama Ridge
“Nuritamburia” semicinereana
(Swezey 1913)
2022 UHIM Unknown Hawaiʻi Kīlauea
Nuritamburia thoracina
(Walsingham 1907)
2022 UHIM Perrottetia Kauaʻi Kahōluamanu, 4000 ft
“Panaphelix” asteliana
Swezey 1932
2007 UHIM Astelia menziesiana Oʻahu Kaʻala
Panaphelix marmorata
Walsingham 1907
2022 UHIM Unknown Maui Haleakalā [exact location unclear, but above
Perkins’ usual camp], 6000 ft
Paraphasis perkinsi †
Walsingham 1907
1894
NHMUK
Unknown Kauaʻi Mts, [vicinity of Kahōluamanu?], 3000–4000 ft
Pararrhaptica capucina †
(Walsingham 1907)
1893
NHMUK
Myrsine?
Molokaʻi
[exact location unclear, but above Perkins’ camp
at Makakupaʻia, perhaps near the “boggy plateau”
he references elsewhere in his journal], 4000 ft
Pararrhaptica chlorippa †
(Meyrick 1928)
1935 BPBM Myrsine lessertiana Oʻahu Mt. Olympus
Pararrhaptica dermatopa †
(Meyrick 1928)
1928 BPBM Myrsine lessertiana Oʻahu Mt. Tantalus
Pararrhaptica falerniana
(Walsingham 1907)
2022 UHIM Myrsine? Molokaʻi [exact location unclear, gulch above
Makakupaʻia], 4000 ft
Pararrhaptica fuscocinereus
(Swezey 1913)
2000 UHIM Myrsine? Hawaiʻi Kīlauea
Last Type locality
Species collected Source Host(s) (island) Type locality
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 25
“Pararrhaptica” fuscoviridis
(Walsingham 1907)
1981 BPBM Psychotria? Nestegis? Lānaʻi [near Waipaʻa], 2000 ft
Pararrhaptica leopardellus
(Walsingham 1907)
2022 UHIM Myrsine lanaiensis? Kauaʻi Kahōluamanu, 4000 ft
Pararrhaptica leucostichas
(Meyrick 1932)
2022 UHIM Myrsine lessertiana Oʻahu* Honolulu*
“Pararrhaptica” lichenoides
(Walsingham 1907)
2001 UHIM Psychotria, Nestegis? Hawaiʻi Kīlauea
Pararrhaptica longiplicatus
(Walsingham 1907)
2022 UHIM Myrsine Maui Haleakalā, 5000 ft
Pararrhaptica lysimachiae †
(Swezey 1933)
1932 BPBM Lysimachia Kauaʻi Kalalau Trail, 3800 ft
Pararrhaptica lysimachiana †
(Swezey 1946)
1927 BPBM Lysimachia Oʻahu Puʻu Hāpapa
Pararrhaptica notocosma
(Meyrick 1928)
2012 UHIM Myrsine lessertiana Oʻahu Mt. Olympus
Pararrhaptica perkinsana
Walsingham 1907
2012 UHIM Myrsine? Maui Haleakalā, 5000 ft
Pararrhaptica punctiferanus †
(Walsingham 1907)
1894 UHIM Myrsine? Maui Haleakalā, 5000 ft
Pararrhaptica pycnomias
(Meyrick 1928)
2012 UHIM Myrsine lessertiana Oʻahu Wailupe
Pararrhaptica sublichenoides
(Swezey 1913)
2022 UHIM Myrsine lessertiana Hawaiʻi Kīlauea
Last Type locality
Species collected Source Host(s) (island) Type locality
26 austin and Rubinoff
Pararrhaptica subsenescens †
(Walsingham 1907)
1894
NHMUK
Myrsine? Molokaʻi [“near Waikolu”]
“Pararrhaptica” trochilidanus †
(Walsingham 1907)
1893
NHMUK
Psychotria? Nestegis? Molokaʻi [gulch SE of Makakupaʻia], 3500 ft
Spheterista argentinotata †
(Walsingham 1907)
1896
NHMUK
Cheirodendron? Polys-
cias?
Hawaiʻi Olaʻa
“Spheterista” cassia †
(Swezey 1912)
1920 BPBM Senna gaudichaudii Oʻahu Kaʻena Pt.
“Spheterista” avocincta †
(Walsingham 1907)
1911 HDOA Santalum? Hawaiʻi Kona [vicinity of Pulehua?], 4000 ft
“Spheterista” avopicta
(Walsingham 1907)
2022 UHIM Santalum Kauaʻi Mts, [vicinity of Kahōluamanu?], 3000–4000 ft
Spheterista fulva
(Walsingham 1907)
1979 BPBM Polyscias? Kauaʻi Mts, [vicinity of Kahōluamanu?], 3000–4000 ft
Spheterista glaucoviridana
(Walsingham 1907)
2009 UHIM Cheirodendron? Polys-
cias?
Kauaʻi Kahōluamanu, 4000 ft
“Spheterista” infaustana
(Walsingham 1907)
2022 UHIM Pipturus Kauaʻi Mts, [vicinity of Kahōluamanu?], 3000–4000 ft
Spheterista ochreocuprea
(Walsingham 1907)
2022 UHIM Polyscias? Kauaʻi Mts, [vicinity of Kahōluamanu?], 3000–4000 ft
Spheterista oheoheana
(Swezey 1933)
2022 UHIM Polyscias Kauaʻi Halemanu
Spheterista pernitida †
(Walsingham 1907)
1896
NHMUK
Cheirodendron? Polys-
cias?
Hawaiʻi Olaʻa
Last Type locality
Species collected Source Host(s) (island) Type locality
RediscoveRies an d pResumed e xtinctions of Hawaiian leaf-RolleR mot Hs 27
Spheterista pleonectes
(Walsingham 1907)
2022 UHIM Cheirodendron Hawaiʻi Kīlauea
Spheterista pterotropiana
(Swezey 1933)
2022 UHIM Polyscias kavaiensis Kauaʻi Halemanu
Spheterista reynoldsiana
(Swezey 1920)
2009 UHIM Polyscias sandwicensis Oʻahu Wailupe
Spheterista tetraplasandra
(Swezey 1920)
2022 UHIM Polyscias oahuensis Oʻahu “Kaumuahona” (see Magnacca 2013)
“Spheterista” urerana †
(Swezey 1915)
1914 BPBM Urera glabra Oʻahu Mt. Tantalus
Spheterista variabilis
(Walsingham 1907)
2022 UHIM Cheirodendron? Molokaʻi Kahanui
“Spheterista” xanthogona †
(Walsingham 1907)
1892
NHMUK
Santalum? Hawaiʻi Kona [vicinity of Pulehua?], 4000–5000 ft
Last Type locality
Species collected Source Host(s) (island) Type locality
28 austin and Rubinoff